OXFORD

Southern Africa

2ND EDITION

1seases
EDITED ВУ J А W COETZER & R С TUSTIN

,,
 EDITED ВУ J А W COETZER & R С TUSTIN

Coetzer I А w. BVSctHonsJ. MMedVet(Path) тustin R с. BVSc, ммeovet1Path)

Professor and Head ProfesSOf Emeritus
Department ofVeter,nary Trop,cal Diseases Department of Vete11nary Tropical Diseases
Faculty of Veterinary Sc,ence
Unive�1\y of Pretot1a
Privatc Bag ХО4
..
Faculty of Vete11nary Sc1ence
University of Pretoria
P,ivate Bag ХО4
Onderst0POO({, 0110 Onderstepoort, О 110
South
• Afr1ca South Africa
And
Professor ,n Тiopicat Veteronary Mediclne
Department of lnfect,ous Oiseo1ses and lmmunology
Faculty of Veter,nary Me<t1c,ne
Utrecht Un,versily
Р О Вох
• 80163
Ulrecht NL-3508 ТО
The Netherlands

OXFORD
VNl\ 1 t::RSITY l'Rl!SS

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 OXFORD
VN [Vl!'RSl'l'Y PRESS
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Published in South Africa
by Oxfotd University Press Southern Africa, Cape Town
lnfectlous Diseases o[Uvestock
ISBN O 19 578202 X (Set or 3 Volumes)
ISBN O 19 576169 3 (Volume 1)
ISBN O19 576170 7 {Voltune 2)
ISBN O 19 576171 5 (Voltune 3)
© Oxford University Press Southern Africa 2004
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First published 1994
Second edition 200•~
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I ', I, I', .,

While every effort has been made to check drug

dosages in this book, it is still poss.ible that errors have
occurred. Furthermore, dosage schedules are being
continually revised and new side-effects recognized.
For these reasons, l),e reader is srrongly urged 10
consult the drug c.o rnpanies· printed instructions
before administering any of che drugs recommended in
the book. The editors caonm be he.Id responsible for
the rnisuse or misapplicaLion of material in this work.

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vi

Contents
List of authors xiv
Section two: Protozoa I diseases
Introduction xxiii
12 African animal trypanosomoses 251
Preface XXV
13 Dourine 297
Acknowledgements xxvii
14 Tricbomonosis 305
Abridged list of abbreviations
15 Amoebic infections 316

16 Coccidiosis 319

17 Cryptesporidiosis 332
Section one: AspccLs influe ncing the 18 Toxoplasmosis 337
occurrence of infectious diseases

1 Vectors: Tkks
19 Besnoitiosis 351
3

2 Vectors: Tsetse flies

20 Sarcocystosis 360
43
21 Balantidiosis 376
3 Vectors: Muscidae 77
22 Leishmaniosis 378
4 Vectors: Tabanidae 87
23 Neosporosis 382
5 Vectors: Culicoides spp. 93
24 Equine protozoa! myeloencepbalitis 394
6 Vectors: Mosquitoes 137

7 Classification, epidemiology and 153 BABESJOSES

control of arthropod-borne viruses
lntroduction 405
8 Special factors affecting the control 171
of livestock diseases in sub-Saharan 25 Bovine babesiosis 406
Africa
26 Equine piroplasmosis 425
9 The control of infectious diseases 178
of livestock: Making appropriate 27 Porcine babesiosis 435
decisions in different epidemiological
and socioeconomic conditions 28 Ovine babesiosis 438
10 Infectious diseases of animals in 225
sub-Saharan Africa: The wildlife/ THElLERIOSES
livestock interface
Introduction 447
11 Vaccination: An approach to the 239
control of infectious diseases 29 East Coast fever 448

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30 Corridor disease 468

31 Zimbabwe theileriosis 472

I
32 Turning sickness 475 Section four: Vi ral d 1!>~a'il'~

33 Theileria raurotragi infection 478 PARA,vtYXOVIRlDAE

34 Theileria nuttans infection 480 Introduction 627
35 Theileria velifera infection 483 49 Rinderpest 629
36 Theileria separata infection 485 50 Peste des petits ruminants 660
37 Theileria annulata lheileriosis 48.6 51 Parainfluenza type 3 infection 673
38 Theileriosis of sheep and goats 498 52 Bovine respiratory syncytial 677
virus infection
39 Theileria buffe lilorientalis infection 500
53 Hendra virus infection 681

54 Paramyxovi ru s-induced reproductive 687

Section three: R it'ko•lhi,d a!l<I failure and congenital defects in pigs
chlamydia! dis,•a,<,
55 Nipah virus disease 692
Introduction 505
40 Heartwater 507
CALlClVIR1.DAE
41 Lesser-known rickettsias 536
infecting livestock Introduction 699

42 Chlamydiosis 550 56 Vesicular exanthen1a of swine 700

43 Q fever 565 57 Enteric caliciviruses of pigs and cattle 703

44 Eperythrozoonosis 573
RETROVIR.IDAE
45 Bovine haemobartonellosis 581
Introduction 707
46 Potomac hoi;se fever 583
58 Enzootic bovine leukosis 708
ANAPLASMOSES 59 Jaagsiekte 717
Introduction 593 60 Maedi-visna 733
47 Bovine anaplasmosis 594 61 Caprine arthritis-encephalitis 741
48 Ovine and caprine anaplasmosis 617 62 Equine infectious anae1nia 747

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PAPOVAVIRJDAE HERPESVIRIDAE

Introduction 753 Introduction 827

63 Papillomavirus infections 754 76 Equid herpesvirus l and equid 829

herpesvirus 4 infections

OR1'HOMYXOVJR1DAE 77 Equid herpesvirus 2 and equid 860

herpesvirus 5 infections
Introduction 765
78 Equine coital exanthema 868
64 Equine influenza 766
79 Infectious bovine rhinotracheitis/ 875
65 Swine influenza 775 infectious pustular vulvovaginitis and
infectious pustular balanoposthitis

CORONAVIRTDAE 80 Bovine herpesVirus 2 infections 887

Introduction 779 81 Malignant catarrhal fever 895

66 Transmissible gastroenteritis 780 82 Pseudorabies 909
67 Porcine respiratory corona virus 784 83 Suid herpesvirus 2 infection 919
infection

68 Porcine epidemic diarrhoea 787 ARTERMJUDAE

69 Haemagglutinating encephalomyelitis 791 lntroduction 923

virus infection
84 Equine viral arteritis 924
70 Bovine coronavirus infection 795
85 Porcine reproductive and 933
71 Ovine coronavirus infection 803 respiratory syndrome

72 Equine coronavirus infection 804

FLAVTVTRIDAE
PARVOVIRIDAE Introduction 945
Introduction 805 86 Bovine viral diarrhoea and 946
mucosa! disease
73 Porcine parvovirus infection 806
87 Border disease 970
74 Bovine parvovirus infection 815
88 Hog cholera 975
ADENOVIRJDAE 89 Wesselsbron disease 987
Introduction 817 90 Loupingill 995
75 Adenovirus infections 819 91 West Nile virus infection 1004

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TOGAVJRI DAE 104 Epizootic haemorrhagic disease 1227

of deer
Introduction 1013
105 African horse sickness 1231
92 Equine encephalitides caused 1014
by alphaviruses 106 Equine encephalosis 1247

93 Getah virus infection 1023 107 Palyam scrogroup orbivirus 1252

infections

BUNYAVJRfDAB 108 Rotavirus infections 1256

Introduction 1027
POXVIRIDAE
94 Diseases caused by Akabane and 1029
related Simbu-group viruses lnlroduction 1265
95 Rift Valley fever 1037 109 Lumpy skin disease 1268
96 Nairobi sheep disease 107) 110 Sheeppox and goatpox 1277
97 Crimean-Congo haetnorrhagic fever 1077 111 Orf 1282

112 Ulcerative dermatosis 1287

ASFARVIRTDAE
113 Bovine papular stomatitis 12.89
Introduction 1087
114 Pseudocowpox 1291
98 African swine fever 1088
115 Swinepox 1293

RHABDOVIR1DAE 116 Cowpox 1296

Introduction 1121 117 Horsepox 1298

99 Rabies 1123 118 Camelpox 1300

100 Bovine ephemeral fever 1183 119 BuJfalopox 1302

101 Vesicular stomatitis and other 1194

vesiculovirus infections PICORNAVTRIDAE

Introduction 1305
REOVJRIDAE
120 Teschen, Talfan and reproductive 1307
Introduction 1199 diseases caused by porcine
enteroviruses
102 Bluetongue 1201
121 Encepbalo1nyocarditis 1310
103 Ibaraki disease in cattle 1221 virus infection

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122 Swine vesicular disease 1313

123 Equine rhi.novirus infection 1319

124 Bovine rhinovirus infection 1322 Section five: Bacterial diseases

125 Foot-and-mouth disease 1324 SPIROCHAETES

Introduction 1427
BORNAV!RIDAE
132 SWine dysentery 1428
Introduction 1367
133 Borrelia theileri infection 1435
126 Borna disease 1368
134 Borrelia suilla infection 1437

CIRCOV!RIDAE 135 Lyme disease i.n livestock 1440

Introduction 1373 136 Leptospirosis 1445

127 Post-weaning multi-systemic 1374

wasting syndrome in swine AEROBIC/MICRO-AEROPHILIC,
MOTILE, HELICAL/VlBROID
GRAM-NEGATIVE BACTERIA
VIRUS-LIKE AGENTS
Introduction 1457
Introduction 1387
137 Genital campylobacteriosis in cattle 1459
128 Unclassified virus-like agents, 1388
transmissible spongiform 138 Proliferative enteropathies of pigs 1469
encephalopathies and prion diseases
139 Campylobacrer jejuni infection 1479
129 Scrapie 1391

130 Bovine spongiform encephalopathy 1408

GRAM.-NEGATIVE AEROBIC OR
131 Transmissible spongiform 1422 CAPNOPHILIC RODS AND COCCI
encephalopathies related to bovine
spongiform encephalopathy in other Introduction 1485
domestic and captive wild species
140 Moraxellaspp. infections 1487

141 Borderella bronchiseptica infections 1492

Pseudomonas spp. infections

Introduction 1498

14.2 Glanders HiOO

143 Melioidosis 1505

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Brucella spp. infections 158 Actinol1acillus lig11ieresii ·i nfections 1648

Introduction 1507 159 Actinobacillus equuli infections 1652

144 Bovine brucellosis 1510 160 Gram-negative pleomorphic 1655

infections: Actinobacillus se11iinis,
145 Brucella 011is infection 1528 Histophilu.s ouis and Haemophilus
somn11s
146 Brucella melitensis infection 1535
161 Porcine pleuropneumonia 1661
147 Brucella suis infection 1542
162 Actinobacillus suis infections 1670
148 Brucellosis in •.vi.ldlife 1546 Pasteurella and Ma11nhei1nia spp.
infections
FACULTATNELY ANAEROBIC Introduction 1672
GRAM-NEGATIVE RODS
163 Pneun1onic pasteurellosis of cattle 1677
Introduction 1553
164 Haemorrhagic septicaemia 1689
149 Kl.ebsiella spp. infections 1555
165 Pasteurellosis in sheep and goats 1695
150 Escherichia coli infections 1560
166 Porcine pasteurellosis 1704
Salnionella spp. infection s
167 Progressive atrophic rhinitis 1707
Introduction 1578
ANAEROBIC GRAM -NEGATfVE,
151 Bovine salmonellosis 1582 IRREGULAR RODS
152 Ovine and caprine salmonellosis 1594
Introduction 1715
153 Porcine salmoneUosis 1601
168 Fusobticterium 11ecrophorum, 1720
154 Equine salmonellosis 1608
Dichelobacter (Bacteroides) ,wdosus
and Bacteroides spp. infections
155 Yersinia spp. infections 1617
GRAM-POSITIVE COCCI
Haemophilus spp. infections
Introduction 1749
Introduction 1628
Staphylococcus spp. infections
156 Haemophilus parasuis infection 1629
Introduction 1750
157 Haemophilus somnus disease 1634
complex in cattle Staphylococcus aureus infections
169 1754
Actinobacillus spp. infections 170 Exudative epidermitis 1759
Introduction 1646 17) Other Staphylococcus spp. infections 1762

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Streptococcus spp. infections Neurotoxin-producing group of

Clostridium spp.
Introduction 1763
Introduction 1877
172 Strangles 1768
185 Tetanus 1878
173 Streptococcus suis infections 1776
186 Botulism 1885
174 Streptococcus porcinus infections 1781

175 Other Streptococcus spp. infections 1784 REGULAR, NON-SPORING,

GRAM-POSITIVE RODS

Introduction 1903
ENDOSPORE-FORMING GRAM-
POSITIVE RODS AND COCCI 187 Listeriosis 19Q4
Introduction 1787 188 Erys,ipelothrix rhusiopathiae 1908
infections
176 Anthrax 1788

IRREGULAR, NON-SPORJNG,
Clostridium perfringens group infections GRAM-POSITIVE RODS
Introduction 1819 Introduction 1915
177 Clostridium perfringens type A 1829 189 Corynebacterium pseudo- 1917
infections tuberculosis infections
178 Clostridium perfringens type B 1836 190 Corynebacteriurn renale group 1931
infections infections
179 Clostridiuni perfringens type C 1840 191 Bolo disease 1938
infections
192 Actinomyces bovis infections 1942
180 Clostridium perfringens type D 1846
infections 193 Ari:anobacterium pyogenes infections 1946

194 Actinobaculum (Eubacteriurn) 1958

Malignant oedema/gas gangrene suis infections
group of Clostridium spp.
195 Actinomyces hyovaginalis infections 1962
Introduction 1854

181 Clostridium chauvoei infections 1856 MYCOBACTERIA

182 Clostridium novyi infections 1863 Introduction 1965

183 Clostridium septicurn infections 1869 196 Tuberculosis 1973

184 Other clostridial infections 1874 197 Paratuberculosis 1994

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xiii

ACTINOMYCETES
Section six: Mycotic and .iigal discai-es
Introduction 2009
209 Mycoses 2095
198 Nocardiosis 2011
210 Pneumocystosis 2137
199 Rhodococc11s equi infections 2015
211 Protothecosis and other 2140
200 Dermatophilosis 2026 algal diseases

MYCOPlASMAS Section seven: DiseaM' compl1•xci.

and diseases of unknown a<:tiolob'Y
Introduction 2043
212 Epivag 2149
201 Contagious bovine pleuro- 2045
pneumonia 213 Ulcerative balanoposthitis a nd 2152
vulvovaginitis of sheep
202 Contagious caprine pleuro- 2060
pneumonia 214 Ill-thrift 2157
203 Mycoplasmal pneumonia of pigs 2066

204 Mycoplasma1 polyserositis and 2071 Index

arthritis of pigs

205 Mycoplasmal arthritis of pigs 2074

206 Bovine genital mycoplasmosis 2076

UNCLASSIFIED BACTERIA

Introduction 2083

207 Contagious equine merritis 2084

208 Tyzzer's disease 2089

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Xiii

List of authors
Alexander T J L AzlzJ, DVM, MPVM, PhO Bath G F, BVSc
8 The La,\1lls Dep,uty Diree1or Associate Professqr
Clerk Maxwell Road Depar1men1 ofVeterlnary Services Ocpanmem of Production Animal
Cambridge. Cll3 ORU Ministry of Agriculture Malays!a Studies
England Wi$nl:1 Chase Perdana Faculty of Veterinary Sctcnce
Damansarn Heights University of Pretoria
Allan GM, BSc, PhO Kuala l..umpur, 50630 Prl,'lltc Oag X04
Department of Agtlcuhure and "Rural Malaysia Onders1cpoort, 0110
Development (OARD) , South Africa
Vcierinary Sciences Division Babluk LA, PhD. 0Sc
Queens Univctsiry Director Bengls R G. BYS¢, MSc, PhD
DelfaSt, DT4 3SD Vetcrinai:y Infectious Diseases Chief Staie Veterinarian
Nonhern Ireland Organization South African National Department of
University of Saskatchewan Agriculture
Allen GP, PhD 120 Veterinary Road or
Na1lonal Directorate Animal Health
Professor Saskatoon, S7N SE3 PO Box 12
Department ofVe1erlnary Science Canada Kruger Natlonal Park
University of Kenmcl')I Skukuza, 1350
108 M H Gluck l!quine Research Center Babluk S L, PhD South Africa
Le>dngton, KY 40546·0099 Pyxis Gcnomics Inc.
USA 120Vetcrinary Road Be7.uldcnhout J D, BVSc. OVSc
Saskatoon, S7N SE3 PO Box33
Allsopp BA, Ph 0. DIC, MCS Canada. Swnnwater. 0622
Professor of Molecular Parasitology South Africa
Depamnenr ofVe1erinary'll'opieal Barrowman PR, BVSc, DVSc
Diseases Labora1oire Vcprol Blgalke R D, BVSc, DVSc, Olp Appl Par &
Faculty of Veterinary Science 8/ 10 Rue des Aulnaics Ent, Olp Pub Admln
University or Pretoria Magny,en·vexln 23.1 Charles Street
Private Bag X04 9S420 Brooklyn. 0181
Onders1epoon. 01 lO Fm.nee South Africa
South Africa
Barthold S W, DVM, PhD Bishop G C, Deceased
Amln JD Professor of Pathology
Depanment of Veterinary Surgery and Schools of Medicine and Vclerlnary Blasco J M, DVM, PhD
Reproduc1lon Medicine Animnl Health Unit
or
University Maiduguri tenter for Comparative Medicine DGA
PMB 1069 One Shields Avenue PO Box 727
Maiduguri Davis. CA 95616 Zarago7.a, 50080
Nigeria USA Spain
Andersen A A, DVM, PhD BasllaneUo S S, BVSc(I Ions), Bosman P P, Deceased
Avian and Swine Respiratory Diseases MMedVct(Path)
RCS!?arch Unit Cribbles l.11bora1orics Bowen RA. DVM, PhD
National Animal Disease Cemer PO Box 195 Professor
United States Deparm1en1 Qf Agriculture Hamilton. 2001 Departmenl of Biomedical Sciences
Agriculmtal Rescalth Servi.cc New Zealand Colorado Stale University
2300 Dayton Avenue Fon Collins
Ames. IA50010 Bastos A OS. BSc(lionsJ, MSc; PhD C080523
USA Se.nior Lcc:1urcr USA
Oeparuncnt of Zoology and Entomology
Angus RD Mammal Research Institute Bradley R. COE, MSc, OVetMed, FRCVS,
Angus Associa1es University of Pretoria FRCPnth, CbloJ, MTBiol
PO Bcix 156 Pretoria, 0002 Private BSE Consultant
Bancroft South Afric.a Guildford
Idaho UK
USA
Bruckner G I<, BVSt, BAdrnin(Hons)
Oirec.tor of Veterinary Services
Agriculture Wes1em Cape
Private Bag XI
Elsenburg, 7607
SouthAfrlea

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Buergelt C D. OVM. PhO, OACVP Coctzer J AW. BVSc(Honsl. Daniels P \11, BVSc. MSc, PhD. MACVSc
Proressor orvc,crina.ry ~rhology MMcdVet(P:lthl Program Loader
Depamncnr or Pmhobiology Professor and He-dd Dia.gnostic Sciences
CoUegc orve1crinary Medicine Oepattmem orVe1.crina,yTroplcru Diseast-s CSlllO Australian Anhnol Health
University of Florida. Facuhy of Veterinary Science Laboratoiy
PO Box 110880 University of Pretoria Prlvaic Bag 24
Gainesville, FL 3261 t ,0880 Private Bag XO<! Geelong. n:m
OSA Ondemepoort, Ol 10 Austrnllo
South Africa
Bunning M 1., DVM. MPM Davies PC, MA. PhO(Camabl, MB Vet,
/\nd
National Cemer or 1nrcc11ous Oiscascs MRCVS
Cemcrs for Oiseasc.Co,mol and Proressor in Tropical Veterinary Medicine The l\~1i11cry
Prevention Department of 1nrec1ious Diseases and Chlrbury
P0Box20lt7 Immunology Momgomery
Fort ('..otlins, CO 80522-2089 l'ncuhy orVe1erlnury Medicine Powys, SYIS 6 DA
USA Urrech1 Un.ivershy UK
PO Box801 63
Bum>ughs CW, BVSc. MPA. Clr;A Utrcch1 NL-3503TD De Vos A l, BVSc. MMedVe1(Parasi1)
PO Box83 l "hc Netherlands Cen11c Coordlna1or
Dolphin Coast. 4404 Tick fever Hl'search Cemrc
Soulh Africa Collett MC, tlVSc, MMedVct(P.,nh). OT!:, Animal and Plant Meallh Service
M EdfC,\ (). MRCVS Queensland - Departmcn1 or Prima.ry
Bun P J, llSc(Hons). MSc, PhD Pathobiology Group lndus1rles
National lnsl.ltu1c for Communicable Institute orVetcrinary. Animal and 28() (;rindle Road. Waco!
Diseases Biomcdi<"al Sciences Queensland. 4076
Prlvaie BagX4 Massey University AttStralia
Sandrtngham, 213 1 Private &g l I -222
Soulh Afrlca Palmers1on North De V~ V. BVSc, BSc(Honsl
New Zealand ScientilicAdvisor. Wildlire Diseases
Campo MS, PhD PO llox 14724
Professor of Viral Oncology Colly P A. 8\/Sc. BS<· Nelspruh, 1200
Departmem of Veterinary Patholot'l' S1erkspruitVc1erinary Clinic South Africa
lnsthutc of Compara1lve Medicine PO 8ox95t
Glasgow University Ve'terln~ry School Lydenburg, 1120 De Waal OT. BVSc. Dip Oat Met, PhD,
Carscube Es1atc SomhAfrica DipEVPC
Glasgow. G6·l l QII Ocparunem ofVeterinaJ'}' Microbiology
Scodand Connor R J. l)Ve,Med. MVSc and Parasi1ology
Director Faculty ofVe,crinary Medicine
Chanler N, l\Sc, PhD AK'SflRtS l,1d Universt1y College Dublin
Jmervc1 UK Ud 98 High Road Betncld
Walt.o n Manor ~lalton Dublin ,1
Milton Keynes Lancas1er. LA2 fit>U Ireland
Bue.ks, MK7 7AJ England
UK Donachie W. BSc. PhD
Cook R F, tlSc(HOI\S), PhD Deputy Director
Clark I! G, DVM, MVScl. fJACVP Research As~is1am Proressor Moredun RC$ea.rch Jnsti1ute
PraJrle Oiagnosuc ScC\'lccs Departmem orve1erinary Science Pcm lands Stfonce Park
Wes1em College of Veterinary Medicine Uni\'er~i1y of Kemucky Bush Loan
Unlvcrslty of Saskatchewan Gluok ITqulne Research Cemer Penicuik
52 Campus Drive Lcxtngwn. KY 40546 Midlcnhian. EH26 OPZ
Saskatoon USA Scott.ind
Saska1chewan. S7N SB1
Canada Cou.slns DV, llAppSc. PhD, ~t<\SM
Principal Microbiologisl
Clarke f N, BSc !HonsJ. PhD Animal Health Labonuorics
Proressor Wcs\crn Aus1ralian Ocpar1men1 of
Molecular Microbiology and !Meetion ,\grlcuhure
University of Southampton Medical Locked Bag No ,t
School Bcmley Delivery Centre
Mailpoint 814 Bentley, WA 69R3
Southampton, SO 16 6YD Ausrralia
England

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xvi

DrewTW, PhD. MSc, CBiol, MIBiol Ga.rin-Bastu.li B, OVM, PhD Gilsdorf M J, OVM, BS. MSc
He.ad of Mammalian Virology UnltHead .Director of the Na1!onal Cenrer for
Virology Deparlmenl Bactcriol Zoon.oses Unit /\nlmal Flealt.b Programs/Eradication
Ve1erinary l..aboratorics Agency OIE/fAO Brucellos.is Reference and Surveillance Pr.o wwns
(Weybridge) La.bora1ory United S1a1es Oepanmelll of
Woodham Lane French Agency for Food Safety (AFSSA) Agriculture/Animal and Plant Health
New Haw BP67 Jnspecllon Services
Addles tone Maisons-1\lfon, F-94703 An1mal and Planl I lcallb Inspection
Surrey, K'flS 3N8 France Service
IJI( US Depanment of Agriculture
Geraghty R J, OSc, flllMS Veterinary Services
Dubey J P, BVSc & AH, MVSc, PhD Head of Diagnostic Virology 1Jni143
Animal Parasillc DJscuscs Laboratory Animal Health 'n'ust 4700 River Road
USO/\, ARS, LPSI I.an.wades Park Riverdale, MD 20737
BARC· l,!as1 Building 100 I Kcn1ford USA
Beltsville, MO 20705· 2.'iSO Newmarket
USA Suffolk, Cl~ 7UU Godfrold J, OVM, MSc, PhD
England Oepartmc.nr oFBacteriology and
Oumbe,11 I(, MD, DSc(Med), FllCJ'a1h Immunology
PO Box 1933 Gerdes G H, BVSc Veterinary and Agrochemical Research
Somerset Wes1, 7129 Acting Head Centre (VAR)
South Africa l)epanmen1 of Virology Groeselenberg 99
ARC-Onderstepoort Veterinary ln.~tltu1e Uc.de, 0 -1180
Easterday B C, DVM. PhD. Oipl ACVM Privme Bag XOS Belgium
Dean Emeritus and Proft'SSOr Emerhus Onderstepoort, 0110
School ofVc1erinary Medlcinc South Africa Griffin I FT, llA (Mod), PhD
University of Wisconsin Mad,son Professor in Immunology
2015 Linden Drive Geyer A, MSc(Med) UnivcrslryofOmgo
Madison. WI 53706-1102 Virology Dcpanmeot P0Bo<56
USII MEDUNSA Dunedin
P0Boxl73 New Zealand
Edington N, nvsc, PhD, PRCVS, f:RCpalh Ml!DUNSA, 0204
O~panmcllc of PalholQgy and Infectious South Africa Guthrie A I , BVSc. PhD
Oise.asc Director
Royal Veterinary College Gl.b bs E P J, BVSc, PhD, FRCVS. Equine Research Cen1te
Royal College Sl rect Professor of Virology f'aculcy ofVc1erlnary Science
Camden Town Depanmen1 of PatJiobiology University or Prewria
London, M21 O'l'U College QfVc1er!nary Medicine Privaic Bag X04
England Univershy of Florida Onderstepoort, 0110
PO Box 110880 South Africa
llllls JA, OVM, PllO, DACVP. DACVM, Gaincs,1lle, Fl. 32611
OACVI USA Gyles CL, OVM, PhD
Department ofVcterinary Microbiology l'roressor
Western College of Veterinary Medicine GlgiiereS, DVM, PhD, Dlpl ACVIM Department of Pa1hoblology
University or Saskatchewan Assistant Prof~or Ontario Vetennary Colltige
52 Campus Drive Department of Large Animal CUni.cal Untversfty or Guelph
Saskatoon Sciences Guelph
Saskatchewan, S7N 584 College ofVcterinary Medicine Ontario, NIC 2WI
Canada University of Florida C'.snada
PO Box 100136
Erasm.us B J, BVSc Gainesville, FL 326l0 HadaniA. DVM, PhD
PO nox 14958 IJSA Professor Emeritus
Lytlleton, OHO Tapazol Chemical Works
South Africa PO Box 3l65
Rlshon Lezlon, 75102
Fenwick S G, 8VMS, MSc (llquatlc Ver Israel
Med). Stirling. MSc mop Vet Med),
PhD Hallwlrth CV, BSc(l\gric)
Division of Health Sciences Medical Scien1is1
School ofVe1erlna.ry and Biomedical Oepanmcnt ofVlroiogy
Science University or1CwaZu.lu-Na1al
Murdoch University Private Bag 7
Murdoch Congella
l'ertb, 6150 Durban, ~(ll3
Western Australia South Africa

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xvii

Henton M M , BVSc. MMcdWt Irons Pc. BVSc. MMedVet(Gyn), Dip Kita.no Y. PhD
Specialist Veterlnarian ACf Llve&tock Division
Golden Vellab Oepanmem or Production Animal Kagoshima Governmem
P O Box 1537 Studies 10·1 Kamoike-shinmachi
Alberton. 1450 Pacuhy ofVeterinary Science l-agoshima, 890-8577
South Afri.ca Universll)• or Prc1oria Japan
Private Bag X04
Hen S, Deceased Onders1epoori. o 11O Kitching IP, B\/Sc
Somht\frlca Oep.uty Director
lleu.ogS, PhO Provincial Veterinary Laboratory
Research Associate Irvin Ao, MA, )iho, VetM8, FRCVS, Sci), Wcs!ern Cape Oepamnen1 or Agriculture
lnsllture for Virology FRCPath, F1Blol l'rivate !lag X5020
faculty of Veterinary Medicine Formerly Director of Biosciences Stellenbosch, 7599
)ustus-Ueblg-Uni,•crsiiy lnterna1ional u,,cstock Research South Africa
Franlauncrstr 107 ln.stitllle
Ciesscn, 0 -35392 PO Bo,30709 Kitching RP. BVetMed, PhD. MRCVS
Germany Nairobi Director
Kenya Nailonal Centre for Foreign Animal
Hodgins DC, OVM, Phi) Discuses
Research Associate lssel, C ). OVM, PhD Canadian Food Ins pection Agency
Department of Pathobiology Wright-Marke)' Chair of Equine 1015 Arlington Street
Ontario Vetcdnary College lnfeeuous Diseases Suhe 'l'2300
University orGuelph University or Kenrucky Winnipeg
Guelph Gluck Equine llesearch Center Maniloba, R3E 3M4
Ontario, NlG 2WI Kentucky Can3da
Canada Lexington. KY 40546-0099
USA Knowlc, N ) . MPhil
Hodzlc E, PhD, OVM Senior Rcscach Scientist
Associate Research Sele.mist Jackson LA. PhD Department or Exo1ic Disease Control
Center rorComparativc Medicine Immunologist lnsti1ute fqr Animal Health
University of California Tick Fever Rl'Sean:h Centre Pirbright Labora1ory
County Road 98 ~nd Hu1chi~11n Animal and Plant Health Se"ice Ash ltoad
Davis, CA 95616 Queensland - De1>urtmem of l' rlmary Pirbright
USJ\ Industries Woking
280 Grindle Road Surrey, GU24 ONF
Horak I G, BVSc, DVSc, PhD wacol UK
Professor Emeritus Queensland, 4076
Department ofVe1erinnryTropicnJ Australia Kock RA, MA, Vcl MB, MRCVS
Diseases Technical 1\dviser Pnn African
faculty of Veterinary Science Jupp PG, MSc. PhD, 0Sc Programme ror 1hc Control of
University of Pretoria 74 Byron Road Epizootic Dis~-ascs
Privai:e BagX04 L<1mbardy,East EpidemlQlogy Unit
Onderstepoort. 0110 )ohanne~buf1:. 2090 African Union Inter African Bureau
South Africa So111h Africa or Animal Resources
PO Box3D786
Howell P G, BVS<:. OVSc KeUy P I, BVSc, l'hO Nairobi
Professor Emeritus Veturlnary Teaching Hospilal Kenya
Department orvmcrinary Tropital Massey UnMtrsity
Oise= Private lluc 11 222 Krakow.ka S, OVM, PhD, DACVP
Faculty or Veterinary Science Palmerston Norih Departmtm orVeterinary llioSJ;ienccs
University of Pretoria New Zealand Collt-ge ofVetcrlnary Medicine
Private Bag X0'1 Ohio S1atc Univcrsi1y
Onderstepoorc, 0110 Kirkland P D, BVSc, PhD Columbus, OH 432 10
South Africa Virology l~~l>oratory USA
Elizabeth Macarthur Agricultural
Huchttrmeyer.11 F KA, OipVc1Scl, lnstiLuu, Kriek NP J, llVSc, MMedVet (Path)
DrMedVct NSW Agriculture Dean
PO Box 12499 Private Mail llag 8 Facuhy ofVetcrinary Science
Onderstepoorc, 0110 Camden University or Pretoria
South Africa New South Wales. 2570 Private Bag X04
Au.strnlia Ondcrstepoori. O.IJ O
Hunter P, BVSc(l lons) South Africa
Afrlvet
PO Box 12684
Ondemepoon, OllO
South Africa

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xviii

Kydd J H, BSc, M,Sc, PnO Lmtemo,e D II, BSc, llSc(Uons), MSc McDermou J 1. ov~. MPVM, PhD
Centre fo.r Prcvcll!lvc Mcdiciuc P08oxCHJS9 Deputy Dit.cctor General
Animal Heallh Ttust Chisipite lnternattonal Uvestock Research
Lanwades Park Harare Institute
Kentford Zimbabwe PO Box 30709
Newmarket Nairobi, 00100
Sµffol k, CB8 7U\J lucldns AG, BSc, PhD Kenya
England Slr Alexander Robenson Cenire for
Tropical Veterinary Medicine Mead D C. MPH, PhD
Lange AL, BVSc, MMedVet(Path). DVSc Royal (Dick) Scnool o~Veterinary Studies Research Scientist
P0Box6222 University of Edinburgh Southeastern Cooperative Wildlife
Wetgemoed, 7538 t!asterbush Veterinary Centre Disease Study
South Africa Roslin College of Veterinary Medicine
Midlothian. El:125 9RG University of Georgia
LangbansW. DrMedVec Scotland Wildlife Health lluitding
Professor of Animal Sciences ,\thens, GE 30602
Swiss Federal lnslitucc of'l'cchnology Mackay DK), BVetMed, MSc, PhD, USA
lnsrJtute of Animal Sc)cnces MRCVS
Physiology and Animal Husbandry Head of Pirbright Laboratory Melswlnl<el R, MSc.
Schorenscrasse 16 (Sl.o\ C.1) lnstirutc for Animal tlealth Specialist Consultant
Schwer7.enbach, CH 8603 Pirbright Laboratory Department of Untomology
Switterland Ash Road lsthuto Zooprofilauico Sperimentale
Pirbright dell'Abru,.1,0 e def Moll$e 'C. Caporale'
Lau nmson MK. MAtHonsl. VelMB, PhD Wokll'\g Via Campo Boario
Division of Paraclinical Studies Surrey, GU24 ONF Vta del Mullnl 28
University of Edinburgh (Cl'VM) UK Teramo, 64 I00
Easrer Bush Italy
Roslin MacLaclllan NJ, BVSc, PhD, Dip ACVP
Midlothian. EH25 9RG Professor Michel A L. OrMedVet
Scotland Depanmcnt of Pathology, Microbiology Senior Research Veterinnrlan
and Immunology TB Laboratory
Lawrence) A, Dphil, IISC. MRCVS,DTVM School ofVeterinary Medicme Onderstepoorr Veterinary Institute
P0Box8W984 Unlveristy of California Private Bag X05
Borrawdale Davis. Ci\95616 Onderstepoort. 0110
Harare USA Southi\frka
Zimbabwe
M&N C I, BVSc, PhD MoggT D, BVSc(Hons), Phi:), l'ACVSc,
Lawson G H IC, 8\11\ft-S, BSc, PhD. Profossor Emeritus Dip ACVIM. Dip ACVCP
MRCVS Depanment ofVeterlnary Sclence and Institute ofVcterlnary. Animal and
Division ofVcrcrinary Pathology Mkrobiology Biomc<llcal Science
School ofVeterinary Medicine University of Arizona Massey University
Easterbush Ve1erinary Ccnire Tucson. AZ8S721 Private Bag U-222
Roslln \)Si\ l'alme~s ton North
Midlothian, l!H25 9RG New Zealand
Seolland Markus MB, l!A(Hons), 0Sc, MSc,
MSc(M.ed Parasltol). DLSHTM, DIC, Mliller-Dobllcs U U. DrMedVet, MRCVS
LelhTe P-C, OVM, PhD PhD Head ofBi~ecuriiy and Animal Services
lnspecteur G6n~ral V6t6rinairc Director and Honorary Professor Institute for Animal Hea,lth
Conscil ~ncral V~r~rlnairc Biomedica.l Analysis International Plrbright Laboratory
Mlnist~re de !'agriculture, de l'all 27 Old Gloucester Street Ash Road
mentation, de la ~rhc et des l.ondon, WCI N 3XX Pirbright
affaires rurales UK Woking
3 rue Barbel deJouy, 75349 Surrey, Gl,124 ONF
Paris, 07 SP McCollum W H, MS, PhD. Ul(
France Professor
Depanmem ofVet.erinary Science Mumford J A, BSc, PhD. HonAssocRCVS
Love R J, MVSc, PhD University ofKen1uc.ky Director of Science
Faculty of Veterinary :idencc 108 Gluck Equine Research C'.enter Animal Health 'J\'ust
University of Sy,11,·a) Lexington, KY40546·0099 Lanwadcs Parle
Camden US/\ Kent ford
New South Wales. 2570 Newm:1rke1
Australia !>'uffol,k. C08 7UU
England

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Muni, £. OrMedVct Ong 8 L. OVM. MSc Penzhorn OL BVSc. 8Sc(Hon), MAgric,

lnsthu1 rurvergelc.ichcnde CQnrdinator: Ve1crinnry L:,botatory ()Sc
Tropenmedlzin und Pllra~itologic Services Oepartme111 ofVcterinaryTropical
Oniversilat MOnchen Oepartmem of Veterinary Services Diseases
Leopoldsrrasse 5 Minis1ry of Agrocuhurc Malaysia Fawlry ofVeterinary Science
M!lnchtn 40, 0·800 Wismo Chase Pcrdana University of Premrin
G¢rmany .Damansarn Height$ Prlvn1e Sag X04
Kuala Lumpur, 5063Q Onders1epoor1. O11 o
MurrayM J, DVM, MS, l)ipl AC..VlM Malaysia South Africa
Oepo.rtmenl of F.qulnc {\lcdkhtc
Marion duPont Scou liquinc ~lcdkal Osburn BI, OVM, l'hO, Olfl ACVP PeplnM, l'hD
Center Dean Director or ncsenrclt
Virginia-Maryland l\l-gionnl College of School ofVc1cru1ary Medicine French l'ood Safety Agency
Vet.erlna.ry Medicine Univcrsily of California AFSSA
Vi·rginiA Pnlyu~r:.h1'lk lni.1h 111(' Da,,.s, CA95fil~ So1>hla An1l1,nlis
Leesburg. VI 2207S USA Cc<l«x, BPI 11 F-06902
USA France
Palmer JE. VMI>, Oip ACVIM
Nesbit JW. RVSc, MMedVct(l ~uh) As'.~ociate l'rofe.sor of Medkinu Perry 8 0 , OBE, llVM&S. 01'\IM, MSc,
~Main RO-ad Dcpor1mc111 of Clinical Studies DVM&S. l'IICVS
Greyton, 7233 School ofVetccinory Medicine Coordlna1or
South Africa University of Pennsylvania Llves1ock Resourtt~s Group
New Bohon Comer lmernationa1 Livestock Research
N - r J A. BVSc MMedVe1(1'11til) 382 West Street Road 111.lotilUIC
Department of Pnraclinicnl Sclc1>C<''- Kcnneu Square. PA 19352 PO Bo~J0709
Faculry of Veterin11ry Sdrncc l}Si\ Nalrobi, 00100
University of Pretoria Kenya
Private Ba-g X04 l'alon O ), MA, Ve1MB. l'hl). MBCVS
Onders,cpoon. oI Io Head Phelps R J, !1Sc(J\ gric), MSc(Agric). PhD
South Africa l)epartrnem of F.xoric Dlse.,s., Comrol S'1 Hivlcra Couri
lnstilute for 1\ nlnml Health i 64 l' ivc Avenue
Nettleton Pf. MSc, Phi), IIVMS. Mf\CVS Pirhright L.abur:11<1ry Hamre
Moredun Research l11s1irn1e Ash Road Zimbabwe
Pentlands Science Pn.rk Pirbright
Bush Loan Woking Picard JA. BVSe(I Ions)
Penicuik Surrc)•. GU2,t ONF De11.ir1men1 ofVeccrinaryTropical
Edinburgh, EH26 OPZ UK Diseases
S<:otland f'a~ulty of Veterinary Science
Paton MW, BVSc. MACVSc Uni1•crsity of Pretoria
Nevill EM. 8 Sc(/\gric). MScv\gricJ. DS,• Senior Vetcri11a,y Officer Private llag X0'1
PO Box 39498 Exotic Disease Preparcdne.s.~. Animal Qndel'StC)lOOrt, 0110
Howick Welf;;1re S0111h Africa
Auckland, 1703 Wcs1crn Australian Department of
New Zealand Agriculture Plpano E, OVM, Phi)
Locked !lag 4 Professor ofVcwrlnary Parasitology
Newton I R, BVSc. MSc. l'hO, OISIITM, B<?ntlcy Delivery Centre. W1\ 6983 Korct SchQOI ofVc1crin ary Medicine
FRCVS AUStn\lla I le\lrew lfalversity of Jerusalem
Veterinary Epidemiologi.~t POJlox L2
Centre for Preven1lve Medicinu l'enrlth M-l. PhD. 0Sc, SVSc(llonsJ Rchovol. 7100
Animal Healch 1'tusc Laboratory Management Advisor Israel
Lanwades P;uk lnstiluto Nacio11al de Jnvestigacao
Kentford Vctcrinario Potgieu,r PT, PhD
Newmarket CP 1922 Acting Oirectnr
Suffolk, C88 7U U Av de Mocambique ARC·Ondcrs 1epoon Vcrcrlnary Institute
England Km 1.5 Pr'h'tl LC Rag X05
Mn1>uro Ondcrs1cpoor1, OJ 10
Norval RA I, bccea$<>d Mozambique Souch Africa
Odendul MW, BVSc. \\lMedVe1(11ncl), Pensacrt MB, l)VM, MSc. Phi) Potgieter L ND, BVS'c, Ph.D
PhD Professor l;merilus Associate Dean for Hosj,ltal Operations
Regional Veterinarian l.abora1ory ofVe1erinary Virology Colh:ge·ofVetcrinnry Medicine
·Raintiow Farms (Pty) I.Id Faculty ofVetcrhrnry Medicine lTn ivcrsi1y <)(1cnncssee
PO Box 12680 University ofGen1 2407 Ril'cr Dt,ve
Onders1epoort, 0110 Salisburyla.an 133 Rm 0206
South Africa Merell>eke, IHU120 Knoxville. TN 37996-4546
llclgium USJ\

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Pre!ocott J F, Ve!MII, PhD Ron R . Deceased Smith KC. BVSc, PhD
Professor Pa1hologis1
Department of Patbobiology SaJlLJ, MSc. PhD. Centre for l'revenlive Medicl.n e
Univtrsity of Guelph Professor Animal Health Trust
Guelph f'00d Animal lleallh Research Program Lanwades Park. Kentford
Ontario. NIG 2Wl Ohio Staie University Newmarke1
Ganada 1680 Madison Avenue Suffolk. CB8 7UU
Wooster, OH 44691 1:ngland
Provost A. Deceased USA
Spencer 8 T, BVSc, BCom, MMed.Vet
Prozesky L, BVSc. BVSc(Hons), Schneider O J, BVSc Department of Production Animal
MMedVet(l'ath) 17 Firsl Avenue Studies
Section Head: Pathology Wellington. 76SS Faculty ofVeterlna.ry Science
Department or J>araclinical Scltnces South A[;ic.a Universlry of Pretoria
Fac.ulty of Veterinary Science Private Bag X04
University of Pretoria Schutte A P, BVSc, MMedVet(Oyn), Onderstepoort, 0110
Private Bag X04 SpecDoc(Ghcnt) South Africa
Onderstepoort, 0110 PO Box727
South Afcica Oansbaal, 7220 St George ·r 0, BVSc. MVSc, DVSc
SouthMrka Virus Consultants ln1ernatlonal
Qulrle M , FIMI.S, BA 15 Tamaox SJrcet
Research Scienris1 Scott G R. OBI!, BSc. MS. PhD Cbapel Hill
Moredun Research lns1hute Cenirc forTroplcaJVcterinary Medicine Queensland, 4069
Pentlands Science Park Univcrsiry or l,d;nburgll A115tralia
Bush loan Eas1erbusb Veterinary Cenrre
Penicuik Roslin, m125 9RG Steele AO, 8Sc, BSc(Med)Hons,
Midlothfan, EH26 OPZ Scotland MSc(Med). PhD
Scotland Virology Oepanment
Shahlrudln S, DVM Diarrhoeal Pathogens Research Unit
Randolph T F, BA, MS, PhD Senior Pn1hologls1 MEDUNSA
Agricultural Economist Veterinary Research lns1ltui-e PO Box 173
International Uvestocl< Research Ministry of Agriculture Malaysia MF.DUNSA, 0204
Institute S9 Jalan Sulra.11 Azian Shah SouthMrioa
PO Box30709 lpoh, 31400
Nairobi, 001 00 Malaysia Stewart CC, BVM&S, BVSc(HonsJ, MSc
Kenya 22 MaraisSt, eet
Sheweo P E. 6Sc, DVM, MSc, PhD Baileys Muckleneuk
Re.Id H W, BVM'&S, PhD. MRCVS, Professor Pre1oria, 0181
bipTVM Depanment or Pa thobiology South Mrlca
ln1ema1Jonal Research Centre Ontario Veterinary College
Morcdun Research lnstitule Univen;ity or Guelph Stoltsz W H, 8VSc
Pentland Science Park Guelph Oepartmcni ofVeter\nary'ITopical
Penlcuik Ontario. NlG 2Wl Diseases
Midlothian. EH26 OPZE Canada l'aculty of Veterinary Science
Scotland University of Pretoria
Sbl(ap V, MSc. PhD Private Bag X04
Rlcht I A, DVM. PhD Veterinary Services Ondersrcpoort, 0 110
Veterinary Medical Officer Klmron Veterinary lnstltu1e South Africa
United States Depar1me.n1 of Agric1~turc PO Bo.x 12
National Animal Disease·Cente.r Bet Dagon, 50250 Stuckenberg 8 R. Ph.D
PO Box 70 Israel Department of Arthropoda
Ames, IA 50010 Natal Museum
USA Slater J D, BVSc, PhD Prlva1e Bag X9070
Deparcmeni of Clinical Veterinary Pietermarit2hurg, 3200
Rossiter P .8 , OVetMe.d. MSc. PhD. Medlcine South Africa
MRCVS Univetslty of Cambridge
Regional uves,ock Co-1ndlna1or MadingleyRoad Sumption l<J, PhD. MA. VetMB, MRCVS
Emergency and Rehabilitation Division Cambridge. CB3 OES Dcpanmeot ofTroplcal Animal l{eallh
Food and Agriculture Organization England Facuhy of Medicine and Veterinary
oflhe United Nations Medicine
POBoX34070 University of Edinburgh
Nairobi, 00 l 00 Easter Bush
Kenya Edinburgh, EH25 9RG
Scotland

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xx/

Swanepoel R, BVSc, DTVM, PhD, MRt'VS ThomsonJ Van den Bossche P. DVM, PhD
National Institute for Communfcable Dean: College of Veterinary Medicine Veterinary Departmcnr
Diseases Iowa Sllltc University Prince Leopold lns1ilu1e of Tropical
Private Bag X04 Amos Medicine
Sandringham, 2131 Iowa Na1ionalestrnat 155
South Africa USA Anlwerpen. 2000
Belgi11m
llaylo.r OJ, MA, PhD, VctMB, Oipf Tlkoo S K, DVll:l, MVSc, PhD
ECVPH, MRCVS Senior Scien1is1. Program Manager - And
Proressor of Veterinary 8acterlol01,,y and Vectored Vaccines Exrroordinary Professor
Public Health Vaccine and lnfectious Disea$C Ot!partmcm of Veterinary Tropical
Veterinary Schoof Organi1,ation Dise.iscs
Glasgow Unlverslry University of Saskatchewan facul1y orVc1cri1.1arySc.icnce
Bcarsden Road 120 Veterinary Road University of l'rctoria.
Bearsdcn Sas.l<atoon, S7N 5E3 Private flag X(),1
Glasgow, G61 IQH Canada Onderstepoort> 0110
Scotland South Africa
TI money J F, BSc. MVB, MS, PhD, DSc.
Terpstra C, OVM. PhD MRCVS Van dee Lugt I J, BVS<:(Hons).
Section ofBiochcmistry University of Kentucky BSc~fod(Hons). MMcdVct(Path). PhD
Central Olergeneskundlgc fnsrhut Gluck £Quine Research Center Department or l'atbobiol0gy
Houiribweg Lexington. KY 40546-0099 Faculty ofVcte.rinary Medicine
Lelystad, 822 1 USA Universlly or Ut(ccht
The Netherlands PO Box80158
Timoney P I, MVB, MS, Ph 0. FRCVS Utrcch1, 3508 TD
1blaucourt P, OVM, PhD Chairman and Director 'lne Nc1herlands
Head of lhe Bacteriology Unit Depanmen1 of Veterinary Science
Programme desantt animale CHt,\D· University of Kentucky Van dcrWult M L. BSc(Agric), llSc{Hons),
llMVI' Gluck Equine Rese.arch Ccmer 108 MSc. PhD
TA-30/G l.exington, KY 40546·0099 Spccinlis1 Scicntis1
Campus de Balllarguc1 USA PO Box 40015
Monrpellicr ccdex 5 Garsfontein East. 0060
34398 Ttid uud C I V. Deceased S0u1h Africa
France
1\lmbull PC fl, 8Sc, MS, PhD Van Oruncn Uulo,-van den Hurk S, BSc,
Thoen C O, DVM, PhD ArjempturTechnoiogy Ltd MSc. PhD
Department of Microbiology. Science Park Senior Scicntls1. Program Manager -
Immunology and Preoemivc Medicine Osll Nudeic Acid Technologies
College ofVererinnry·Mcdicinc Porton Down V;1r.f"ine. ;ind lnfcc1ious Disease
Iowa.State Unlvcrsily of Science and Salisbury, SP•I OJQ Organization
Tuchnology UK University of Saskatchewan
2180 Veterinary Mediciroe Complex 120\lcrerinary Road
Ames, 1W SOOJ 1· 1250 Tustin R C. B\/Sc, MMedVct(Palh) Saska1oon. S7N 5E3
USA Professor Emcri1us Canada
Department ofVe1erinary Tropical
Thomp&on P N. BVSc. MMedVel(Mc<fl Diseases Van Kaldercn A, llVS.cCHons)
Department ofProdu<:tlon Anion:~ Faculty ofVetcrinary Science 7 1 PacificStrcct
Studies University of Pretoria Maori Hill
Faculty orVcterinru:y Science Prlvnte Bag X04 Dunedi n
University of Pretoria Onders1cpoort, 01 IO New Zealand
Private Bag X04 SoUlhAfrica
Onderstepoon. o110 Van Heerden J, BVSc. DlpMedVet,
South Africa Umphenou r NW. DVM BSc(Hon) (Nal). MMe<;!Vet(Mcd)
Resident Veterinarian 16 Oalham Rond
Thomson GR, BVSc, MSc, PhD Ashford Stud - Coolmon, America Kimberley. 8301
Main Epidemiologist 5095 Fmnkfon Road South Africa
Pan-African Progr~nimc ror the Comrol PORoxR2.3
ofl!pizoo1ios Versailles, KY 40383
Inter-African Bureau for Animal USA
Resour<:l?s of lhe African Union (AU·
IBAR)
PO Box 30786
Nairobi, 00100
Kenya

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Van Olt$chot IT, DVM, PhD VlsmerH F.PhD Wllllamson SM. BVetMed, MRCVS. PhD
Read ofDepartment Chier Specialist Scientist Veterinary Investigation Officer
Dcpanmeoi ofMammalian Virology PROM.EC Unh VeterlnarySurvelll,tllce
Institute for Animal Science and Health Medical Research Council Veterinary Laboratories Agency
p·o Box65 PO Box 19070 Rougham HUI
Lelystad, 8200 All Tygerberg, 7505 Bury St Edmunds
The Netherlands Souch Africa Suffolk, IP33 2RX
And u~
WaUgren P, OVM, PhD
Professor of Veterinary Vaccinology Stace Veterinarian WIison 1' M, DVM, PhD, Dip ACVP
Utrecht University Department ofRuminant and Porcine Liaison Officer
PO Box80163 Diseases i\rmed Forces Medical Intelligence
Utrecht, NL-3508TD National Veterinary tns11lute, SVA Center
The Netherlands Uppsala, 751 89 Vecerinary Services
Sweden Emergency Programs
Van Reelh K, OVM, PhD USOA
And
Professor APH IS
Labora\ory ofVeterinary Virology Consuflanc Professor 1607 Pol\er Sireet
Faculty orvc1erlnary Medtcihc Depanment of LargeAnlmal Clinical Fort Detrick, MD 21702-5004
Ghent University Sciences USA
133 Saltsburylaan Swedish llnivcl'Sity or Agricultural
Merel.bcke, 9820 S<:iences SLU Yeruharn I, DrMedVet, PhD
Belgium Department or Farm Animals
Werling D , DrMcdVet, PhD. MRCVS Kom School of Veterinary Med.lclne
Van Rens.burg J BI, BSc(AgricJ, BVSc, Department or Pathology and Infectious Hebrew Universiry or Jerusa.lem
MMedVet(PathJ Diseases PO 8ox 12
90 West Street Royal Veterinary College Rehovot, 76100
Pretoria North, 0182 University of London Israel '
South Africa Hawkshead Lane
Hatfield York D I'. 11Sc. MSc PhD
Van Vuuren M. BVSc. MMedVet(Micro) Hert$, Al.9 7TA Director
Associate Professor UK Molecular Diasnos1ic Services
Department ofVcterinnry Tropical PO Box903
Diseases West D M, BVSc, PhD, fACVSc. Westville, 3630
Faculty of Veterinary Science Professor SouthAfrico
University of Pretoria Institute orvetcrlnary. Animal and
Private Bag X0-1 Biomedical Sciences 7.aria LT
Onderstcpoort, 0110 C.ollegc of Sciences Department ofVeterinary Microbiology
South Africa Massey University and Parasitology
Private Bag 11 222 Faculty of Veterinary Medicine
Venter G I, BSc.O·lons), MSc Pahnerston North Universlty of Maidugurl
Senior Veterinary R~>searcher New Zea.land PMI! 1069 .
Division of b'ntomology Maiduguri
ARC·Onderstepoort Veterinary Institute Williams M C, BVSC, MMedVet(Path) Nigeria
Private Bag XOS Associate Professor
Ondemepoort, 0110 Department of Paraclinical Sc.fences
South Africa Faculty orVeterinary Science
University of Pretoria
Vermunt ) I, DVM. BAgSc. MSc, FACVSc Private Bag.X04
Registered Veierina.ry Specialist- Cattle Ondcmepoor(, 0110
Medicine South i\[rica
Awapuni Veterinary Services Ltd
P0Box4000 Wl!Uamson M M, RVSe. MVS, MACVSc,
Palmerston North, 5315 PhD
New Zealand Veterinary Pathologisc
Gibbles Veterinary Pathology
Verwoenl O w. D\,'Sc, t)Sc 1868 Dan.denongRoad
Rese<ll'ch Coordinator Clayton
Faculty of Veterinary Science Melbourne
University of Pretoria Victoria, 3 168
Privat.e Ba.g X04 Auscralia
Onders1epoor1, OLIO
South Africa

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xxiil

Introduction
This is the second edi1ion or the 1.ex1book fnfac1ious Diseases latter three animal species and zebras is 'cquid'. The book.
of Uvestock wi1h SJ1et'i(I/ Reference 10 So111hern Africa which however, not only includes diseases of livestock but also,
was published in 19<J4 and i~ now out Qfprim.Whilc 1he focus where relevant 10 livestock. 1hose of wildlife.
of the first edition was. as its title indicates. more on lhe dis- The country, South Africa, is mentioned relatively fre-
eases of farm animals in Africa south of the Sahara Deserl, que.ritly in the book. -Si.nee publica1ion or the first edition in
the book did include many impor1an1 diseases not then pres- 1994 there have been many political and other changes In the
ent on the contincm which it was thought did constiune a coumry including the creation of new provinces and alloca-
veayreal or potemlal threat 10 Afril:a. The contents of1ho sec- tion of new names, for example, 10 many towns and several
ond edition have been expanded significantly to include 1he rivers. The former Transvaal Province has been divided into
majority of the impor1ant infoctiou.~ diseases of livestock four new provinces: Gautimg. Nor1h West. Limpopo and
present in the world. Hence 1he shor1ened title, the increase Mpumalanga; the fonner Cape Province now comprises the
in the number of pages and 1he set of 1hree volumes. Western Cape. Eas1ern Cape and Northern Cape provinces;
However, in some section$ and chap1ers there is still empha- Natal has become KwaZulu-Natal: and theOrangii Free State
sis on diseases, their vectors. if any, and theirconuol perrnin- is now the Fret?State. We have used these names as well as the
lng to sub-Saharan Africa because of their particular socio- new names of the towns to which th~se have been allo'cated
economic importance in this region. In addition to the 28 protozoa! and nine rickensial
The production of the first edi1i1m was inspired by the diseases in Vnlumc I. chapters are Included on the main
three outstanding editions of the textbook wriuen by Dr vector~ of diseases (e.g. ticks, 1se1sc Dies, Muscidae,
Michlel Wilhelm Henning titled Animal Diseases i11 So111h Tabanidae, C11ficoides midges and mosquitoes), the main
Africa, the first or which appeared in 1932 and the third in principlesfmethods of control of Infectious diseases, and
1956. These are regrcllably now ou1 of print and are vinually infectious diseases of particular imt>ortance at the wildlife/
unobtainable, but much reference is made to them in the lives.tock interfare. Volume 2 contains 83 diseases caused by
present work. viruses and virus-like diseases. and Volume 3, 80 diseases
The demand for the first edition. of which 2 000 sets were cau.scd by bacteria (including mycoplasmas), fungi and
printed. far outstripped 1he sur>ply within a maner of sever<ll algae, as well as a small number of conditions of unknown or
years, and we are still receiving enq uirtes about its availabili1y uncertain aetiology.
almost on a dally basis. It pr<lved. and cominues 10 be, most Each disease is dealt \\1th in terms of its introduction,
popular amongst panicularly veterinarians, undergraduate aetiology, epidemioloi,,y, pathogenesis, clinical signs, pathol-
and postgraduate students in 1hc fields ofveteri'naay and ani- ogy, diagnosis. differemial diagnosis and control. A compre-
mal science, and veterinary libraries. h.ensive list of references is provided for each disease. To facil-
Although fully aware of the daunting task it would iake 10 itate readabiliry. the reforenC8$ ;ire numbered in.the text.
produce the second edition, the editors decided to revise the l'.ach of the volumes contains a list of contents and a
book" because of the excellent reviews the first edition had comprehensive index which pertain 10 all three volumes.
received in several well-known international joumals, the Funhcnnore, the three volwnes are so paginated lhat page 1
two prizes that were awarded to it, and the high national and commences in Volume land the last page (page 2159) of the
International demand for the book. set is in Volume 3.
The first edltion of the book won 1hc following prit.es: We believe !hat, as the firsledition proved to be. this text·
• Bill Venter LJ1ermy A111ard in \996. The purpose of this book \\1U be invaluable to undergraduates and postgraduates
annual South African award is 10 recognir.e 1>u1standing in facull.ies teaching veterinary medicine, and that veterinary
contributions 10 research by university personoel and 10 rescarcbers, veterinarians in fann µra~tke, st0ck breeders
promote the standard o(specialiZcd works published by and ail those charged with the promotion of animal health
them. and the prevention of livestoc.k and wildlife diseases should
• Malbrant-Fe1mcc111 l'rize in 1998. whkh is an award of the also find it a v-aluable addition 10 !heir reference librari.es.
Acadl!mie Vl!terinairedc France. Although every effort was made to cornplete the projecr
in as ~ho.n a time as possible. it proved to be an even more
For the second editiOD we invited 197 scientists and special- formidable and daunting 1ask than the first editio.n, which
ists in their respective disciplines from 24 countries to con- took seven years from inception 10 public.1tion. We sincerely
tribute by writing chapters on subjects in their 6elds of hope I hat !his book will help promote 1101 only the health of
expertise. livestock but also the control of their infectious diseases.
The definition or liveswck in the book encompasses ca1-
t1c, sheep, goats, pigs. horscs, donkeys and mules. It will be TIie editors
noted that the collective noun that is frequently used for the

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x:cv

Preface
The first edi1ion or 1hi$ book, now ou1 of print, was pub- pie where animal husbandry and practices played a role in
Ushed under the title /11feclio11s Diseases of Uvestock 111i1h 1he emergence or a ·new' disease. It is important to note
Special Reference 10 So11them Africa in 199'1 ;(Sa se1 of c,vo that some of these emerging/re-emerging animal diseases
volumes. It was very welt r11c.eivcd by veterinary and animal may also have a significant bearing on public beaJth, e.g.
husbandry professionals. receiving glowing book reviews llifl Valley fever, tuberculosis and BSE.
in several wellknown in1emational journals as well as 1wo The World Organization for Animal Health (OIE) cur-
prizes - the Malbra11t-Few11e11 Pri;.e in 1998 from the rently classjfies animal diseases as Ust i\ or List B diseases
AcademieVeterinaire de France, and the Bill Venter Literary based on lhe n·aturc of their spread, the socioeconomic or
Award in 1996, which is awarded for omsianding coniribu- public health consequences they may have, and their
tions by university personnel in South Africa. This new sec- effects on imernational trade in animals and animal prod-
ond edition has been expanded 10 include the majority or ucts. The introduction (emergence) of an OIE List A disease
infectious diseases pn?s«:111 in form animals across the can have a devastating effect on a country with a sizable
world, this necessitating a new title and expansion 10 a se1 export trade in an imals :ind animal products. It is esrimat·
of three volumes. cd that economic losses due 10 los1 trade are about ten
The focus of 1he book is on infectious diseases of limes those incurrecl clnring dt'pop11lation (stamping-out)
importance to lives1ock and wildlire. Many or 1hesc dis- and disinfection. Early dctec1ion and prompt regulatory
eases have the po1c111ial of transgressing international measures should therefore be regarded as the most effi-
boundaries and consequently a.m,c1ing international trade cient means to minimiz.e these serious losses.
in animals and their products wilh serious socioeconomic The'objectives of the OIE include ensuring transparen-
conse.q uences for countries in which 1hey occur or into cy in the global animal and zoonoses situation: collecting.
which they are introduced. h is well known that several of analysing and disseminating sciemific veterinary informa-
these disease.~ often pose as 1he bigges1 constraint 10 !lie 1ioni providing expertise and encour.iging international
sustainable socioeconomic development of agricultural solidarity in Lhe comrol of animal diseases; safeguarding
communities in developing countries. In addilio'n, 1he world 1rade by publishing heallh standards for internation-
highly contagious diseases such as fool-and-mouth dis- al trade in animals and animal products. providing a better
ease, rinderpes1 and Afrkan swine fever, and those 1hat guaramce of the safety of food of animal origin: and pio-
have public health lmplica1ions, such as Hift Valley fever moting animal wel[are 1hrough a science-based approach.
and anthrax. may be used as bio-weapons or in agri-bioter- Boo.ks su.ch as th.is one arc of great assistance in th.e real-
rorism. iza1ion o[ many of these objeciivcs.
Globallzalion and 1hr evcr-lnrrrnsing trade of animals Infectious Diseases of Livesrock will remind all those
and their produc1s crea1e a si1uation which facilitates the concerned with livestock diseases of their national and
potential spread of many disease's to countries where they interna tional responsibilities. Many of the diseases,
have been eradica1ed, often at great cost. or countries indeed. constitute a very real threa1 10 the world, whose
where they have not previously occurred. The risk of emer- human population continues to expand at an exponential
gence or re-emergence of di.seases in the developed world rate.
is therefore at an all time high. Good examples to illustrate The editorial team should be commended for the
this point are the recem emergence of foot-and-mouth dis- rem·arkable 1ask of coUaling the contribtHions of world-
ease in the United Kingdom and 01hcr cow11ries. wi1h renowned scientists in their respective fields of expe.rtise,
devastating consequences. Climalic changes such as those 1hus adding a tremendous value to this book.
currently occurring in many paris of the world may also be I lherefore h<1ve much pleasure in recommending this
responsible ror the emergence or vec1or-borne diseases in book 10 all veterln.a rians and other professionals dealing
new geographical areas, as demonstrated by 1he recent with livestock and livestock diseases.
outbreaks of Hifl Valley fever in Saudi Arabia and Yemen,
and of bluetongue in some Mediterranean countries. Dr 8 Val/at
Bovine spongirorm cneephalopathy (BSE) is a good cxom - Director Ge11eml 0/E

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x.x,·i

Universiteit Utrecht
·-------
<1ntervet"'

...~
A RC • LNR

Unlverslly ol Pretoria

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xxvii

Acknowledgements
There are ma.ny people. wmpanies and organizal'ions The dimensions of this secon<l edition of the book
that we would like w thank for their invaluable assistance, proved 10 be more formidable than were thuse of the first
financial backing and en1ln1siastic support Without them. edition. and only the experience and dedication of an
the publication of thi~ book would not have been possible. experienced publisher could have undertaken the task of
The contributions llf :Ill the nlllhurs and co-authors are producing it. We extend our sincc.re thanks to Oxford
greatly appreciated. We know wha1. sacrilices had to be Uni),ersity Press Southern Africa and. in particular, its for-
made by them in their daily lives - both work schedulc•s mer Managing Director. Kare McCallum. for their support,
and domestic - in order 10 produce their chapters. advice. technical ability, patience and perseverance. The
We are particularly gratefu l ro, the financial suppun sterling and friendly assistance of Ingc du Plessis, our tech-
that we received rrom several suurccs. This enabled us to nical and language cdiror, is great'ly appreciated. Ow
proceed with the project. thanks are also due tu Marina l'earson who was respons•
The principal funding was very kindly arranged for us ible for the mammoth task of indexing the book and Ethni
by Prof A W C A Corilcli&.5en, Ocan or the Facu hy of Clarke our proofreader.
Veterinary Medicine, U1rcl'ht Univ1irsity, The Netherlands. The unstinting assis1::1n1.-e given to us by many of our
from: colleagues is greatly appreciated. Prof I Boomkcr and Dr
• lntervet International bv, The Ne1hcrland.s J Crafford redrew several of the maps and figures, and
• Utrecht Unlversi1y. Professors M van Vuuren. 1 G Horak and BL Penz.horn, and
Other funding was obtained from: Dr I Picard and others in the Department of Veterinary
• Merla.I France U>ty) Ltd Tropic.al Diseases answered many questions put 10 them by
• Bayer Animal Heahh Oivision us in their various fields of expertise, which saved us many
• University of Pr1ituria. hours of reference work.
ln addition, Professors M. van Vuuren, G P Allen, B A
The considerable role played by l>r G RThomson in the ini- Allsopp and R SwanepOel, and Ors J Godfroid, M M
tiation of the project and 1he production of the first ediiiOn Henton, DC Hodgins, ) A Lawrence, AL Michel, J A Picard,
on which 1his second t·dition is based, is grawfully I 1\ Ver111ont., D w Verwocrtl and O M West, are thanked for
acknowledged. contributing co the book by writing the Introductions to
We are parricularly indebted to the previous directors. the various sections.
Dr GR Thomson and IJr S Cornelius. and the present act - Charmaine Vermeulen of '(elemmic Education Media,
ing director. Dr f' T Po1gie1er of the ARC·Onders1epoon University of Pretoria, F.ri cH van der Westhulzen and
Vetennary Jnstl1u1e. a nd Prof N P ) Kriek, Dean of the Antoine11e Lourens of Academic Lnformation Services of
Faculty of Veterinary Science. University of Pre1oria, for 1he Faculty of Vetednary Science, llnive~ity of Pretoria,
allowing us to use the services of Martie Viljoen who did and Francie Louering of the Department of Veterinary
most of the word-processing for the book and Rina Tropical Diseases, Uni.versiry of Pretoria, willingly assisted
Serfontcin who lent invnluable secretarial and other sup· u~ when called upon 10 do so.
port for the projec1. Both Manic and Rina went out of their Our most heanfelt ,hanks are due to our families and
way at all times 10 make our task so much easier despite friends for their help, forbearance and underslanding.
their other very onerous duties. Their cooperation and
assistance are greatly appreciated. ProfI A W Goerner
We are especially gratefltl to Dr B Valla1, Director ProfJ;merillls RC Tiisti11
General of the Office lnternationnl des Epizooties (01(:), Faculty ofVe1crinmy Sciem:e
Paris, for writing the PrefaU/. U11i11ersiry of Pretoria
2004

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xxix

Abridged list of abbreviations

A C DIVA differenliate infel'ted from
MH1. Australian Animal I lcahh CaCV canine C'llicivirus vuccinatcd animals
Laboratory CAE caprlnc·anhritis,encephallt is DMl!M Oulbecco1s minimal essential
MV adeno-associatcd virus CAEV caprinc·anhriiis-cnccpha.li1is medium
ACAV American Commiuee on virus DMS0 di111e1hyl sulphoxidc
Arthropod-borne Virus~ CAM chorioallan1olc membrane DNT dermonecrotln toxin
ADCC antibody-dependent, cell· c:AMI' cyclic adcnosine monophos- DP! days p0St•inocula1ion
mediated cytotoxic ph,uc DTH delaycd·t)11t! h}'l)ersenshivity
JU)£ antibody-dependem enhance- CBC complete blood couiu
mcnt CBPP contagious bovine pleuropneu· F.
A£ attacbing•and -effi11::lng (lesions) monia EAEC cntcroaggrr.gative e. ccU
,!\EC amino-ethyl t'arbozolc CCHf' Crimean-Congo haemorrhaegic EAV equine aneritis virus
AE.EC attaching aml effacing/;. roll fever EB elementary body
AG anhrog'}'Posls CCHFV Crimean-Congo haemorrhaeg!c EIIHSV F.uropea.n brown hare ~-yn-
>.GID agar gel !mmunodiffuslnn revcr virus dromeviru~
AHS African horse sickness CCPP contagious caprine pleuro- EBL cn1.ootie bovine leukosis
AHSV African horse sickness virus pneumonia ECE equine coltal exanthema
AIDS acquired hnrnunc deOclency cEUSA competitive enzyme-linked ECF East Crsast fever
syndrome immunosorbeni assay ECV equine coronavirus
AlHV alcelaphlnc hcrpcsvirus CEM conrngious equine metrltis ED equine dermis
ALP alkaline phosphmasc CF complement fi.xation EE equine encephalosis
APC antigen-presenting cells CFT complement fQ<alion t~-st EEV e!fuine em:ephalosls viru.~
ARDRA nmplifiL'li ribosomal ONA CFU colony rormlng units Elili eastern equine encephalitis
res!rlcrion analysis cGMP cyclic b'uanylate monuphos· EEL e<1uinc embryonic lung
ASF African s,vine fever phate EF oedema factor
ASFV Alric.an swh'lc fever vims C:H IK Chikungunya (virus) EFK equine l'oe1al kidney
ASPV add-srnble pitornavirus CID cc,mbined Immunodeficiency llMD cpi1.0otlc: haemorrhagic disease
AST a.,pana1e aminotransferase Cl-ELlSAcompclitive Inhibition enzyme:· 1:1-IDV cpimocic: hnemorrhagic disease
ATP adenosine triphosphute linked lmmunosorbcnt US$8Y virus
Cll:P counter-lmmuno-elcctrophore· EHl.:C cnterohaemorrhagic E.coli
B sis EHV cquid herpesvirus
BACTEC radiometric culture system CIN eef.,ulodin, lrgasan. novobiocin El.A enzyme immunoassay
BANV Ban1.i virus CJD Crcuizfeldt·Jacob disease ElAV equine inftl('tious anaemia virus
BAV bovine adenovirus CK cn:atJnine kinase Ell:C emeroinvaslve /;. coli
BCG Bacillus Calmctlc·Gut'rin Cl.A cascous lymphadenitis ELISA cnzyme-linl:ed immunosorbent
BCV bovine coronovirus CLO 01mb,vlobac1er-like orgnnisrn ,ISS3)'
BO Border disease; Borna disease CM! cell,mcdiated immunity ELOSA enzyme-linked oligonucleotlde
BOV Border disease virus: llt1ma Cll,fV ·i,1omegaJovirus sorbent assay
disease virus CNF C}1oto.tic necrotizl ng fact.or F.MC e11cephalomyocarditis
BECV bovine e.meric calicivirus COPY canine oral papilloma,irus EMC:V encephalomyocardltls virus
BE!' bovine ephemeral fever CPO C>~Opat hie dose ENT\/ enzootic nasal tumour virus
BEFV bovine ephemeral fe,er ,·irus CPE cytopathie effect EPEC entcropathogenic E. roli
BH.K baby hamsier kidney CllPV couonrnil rabbit papillomavirus £PM equine proto1.oal rnyeloen•
BHM bovine herpes mammillitis CSF classit~tl swine.fever <"t'phalltls
BHV bovine herpesvirus C5rY classical s,vine fever ,irus Eql'V <'(fuinc p~plllon\avin,s
BfV bovine'immunodeficicncy virus CTL cytoto•ic1:fymphocy1e ERhV equine rhinovlrus
BLV bovine leukaemia virus cn,p cyio10~ic T-lymphOl.ytC precur· EspB E. coli secue1ed protein 1l
BoTV bovine torovirus sors ET clectro,phoretic types
BPF bovine petci:hial fever (Onduri cru community rnberculln uoits ETEC emcrotc>xigcnic E. ccli
disease) cwo chronic wasting disease ET\I C(tuinc torovirus
BPV bovi.ne papillnmavlrus EVA equine viral arteritis
BRSV bovine respirmory S)'ncytial D
vin1s OAEC diffusely adherent I;. coli F
BRV bovine roravirus DALY disability adjusted life year FA nuoresccnt antibody
BSE bovlnc sponglrorm DBHJ direct bilirubin FAQ Food and Agricultural
encephnlopalhy Dr1D Department for International Organization
BSL bi()l;afety level l)cvelopment FBS foetal bovine scrum
BT bhietonguc DFMO dlOuorommhylornithrine FCV feline calicivirus
B1V bluetonguc virus 01 defective imerfcrlng FDC follicular dcndritic cells
BVD bovine viral dimrhoea DISC defe<;tivc infectious single cycle FEK roeinl equine kidney
BVDV bovine viral diarrhoea virus O!SH disinfection. lsolation. submis• FFl fatal familial inson,nia
sion. h)'gienic (procedures) l'PP fatal flbrinou:s pneumonia

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l'HA mamemous haemagglutinin assay Ml.V modined live virus
PIV feline immunodeficiency virus HP ilidircct immunoperoxidase MmmSC Mycoplasma mycoidcs subsp.
!'MD foot-and-mouth disease ILCA International Uvestock Centre M>'c.oides Small Colony
PMDV root-and-mouth disease virus for Africa mMl'SA major merozoite/piroplasm
l'l'A fluorescent polarization assay ll.RI llllcmatlonal Uvcstock surface antigen
PPLC fast perfom1ance liquid Research Jnstinnc MMTV mouse mammary tumour virus
chromotography INF interferon MOMP major outer membrane protein
PPSR false positive serological reac- INH isonicotinic·hydrazlde MOTT mycobacteria other than
lions iNOS inducible nitric oxlde symhase tubercle bacilli
FSE reline sponglform encephalo- IOllP ion-regulated protein MPMV Mason-Pnzer monkey virus
pathy; focal symmetrical IPD infectious pustular MPS mononuclear phagocytk sys-
enoephalomalacia balanoposchitis tern
IPMA immunoperoxldase monolayer MPS mycoplasmal pneumonla or
H assay swine
HA haemagglutin)o IPV infectious pusmlar vulvo- MIH.S mare reproductive .loss syn-
HADEN haernadsorbing. entcric vaginitis drornc
HAl haemagglulinalion inhlbltloo (RP iron-regulated proleins MRM mechanically rccove.red meat
AC hog cholera ISCOM immune-stimulating complex MllT milk ring test
HCV bog·cholcra vims ISH ;,, sir,, bybridiza1ion MSP maJor surface protein
HE human granulocytic ehrlichio· ITEME infectious thrornboembolic mtDN/\ mitochondrial DNA
sis; hydranencephaly rncningoenccphnlitis MV maedl-visna
HEP high egg passage MW maedi-visna virus
HERV huma1\ endogenous retrovirus I MW molecular weight
HeV Hendra vim,< JE Japanese encepbolitis
HEV haemagglutinating JS jaa1,,slck1e N
cncephalomyelhis virus JSRV jaagsickle sheep retrovirus NIii Newbury agem I
HEYM Herrold's egg-yolk medium TV Jena virus NAHMS Nationnl Animal Health
Hl haemagglutination inhibition; Monitoring System
humoral lmmunily L NCDV nwna1al calf dia,rhoea virus
MN human immunodeficiency virus 1/1 lnlex agglutination NCR non-coding region
HN haemagglutinin-neuraminidase LAM lipoarabinomannan ND Newcastle disease
liPLC high-pressure liquid I/IT latency-assoclaled transcripts NE necrotic enteritis
chromatography I.CR long control region NLV Norwalk-like virus
HP$ Hantaviru.s pulmonary syn- I.CV large cell variant NO nitric oxide
drome LOIi lactate dehydrogenasc NP nucleoprotcin
l:IPV human papillomaviru., LOV l3ctate dehydrogcnase elevating NSl.l Nairobi sheep disease
HRPO horserndish p,:roxidasc virus NSDV Nairobi sheep disease virus
HRSV human respiratory syncyrial 1.£E locus·for enterocyte effacement NS! non-struttural protein
virus LEP low egg passage NS!> non-strucniral viral protein
HTLV human T-cell leukaemia virus LI' lethal factor NT neutralization
HuCV human calicivirus I.I loupingUI NTEC necrotoxigenic F.. ooli
LIV louping iU vlrus
I I.OS lipooligosacchandc 0
IMT immunoabsorbenl aggl111ina· LPS lipopolysaccharide OID ovine intcrdigilal dennatids
,ion as,;ay tcs.t LSD lumpy skin disease 011'. Office International des
ICDV icosal1edral cytoplasmic I.SOY lumpy skin disease virus Epi1..ooties
deoxyribovirus tr heat labile OM.AGOD ovine mouth and gum
ICP infected cell polypeptides 1:rn long terminal repeat obscure disease
ICTV International Committee for OMP outer membrane. protein
the T3Xonomy of Viruses M OPPY ovine progressive pneumonia
ID immunodiffusion MAb monoclonal antib.ody virus
lF.C intestinal cell line MAT microscopic agglutination test 0-PS 0 -polysaccharlde
IEUSA indirec1 enzyme-linked MBM meat and bone meal ORF open reading rrame
immunoabsorbcnt assay Mccp Myooplasma caprloolum subsp. OSCPII oligosaccharld!!-sp.ecific cap-
JEM hnmunoclectton microscopy ciiprip11e11mo11iae rure plate hybridization
IETS lmern.adonal limbryo lransfer MCI' malignanc caLarrhal fever Osp/\ outer surface protein A
Soclery MO mucosa! disease OvUV ovlne hcrpesvlrus
lF immunofluorescence Ml)BK Madin-Darby bovine kidney OVJ Onderstcpoon Veterinary
ll'A lndirecl fluorescent antibody MOCK Madin-Darby canine kidney Institute
IPAT fmmunoOuorcscenl anribody MHC major hiscocompatibillry com-
test plcx p
u-, fndln?ct fluorescent antibody MIC minimal Inhibitory conccnlra· PA prolective antigen
test tlon PAAT Programme Against African
IHC lmmunohistochcmical Mll'T micro·lmmunoOuorcsoence test 'l'rypanosomiasis
!HT ind.irect bacmagglu.tinntlon test MIPLD mouse intrapcritoncal lethal PACll Pan-African Programme for llte
UFA indirect immunofluorescence doses Control or Epizootics

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xxxi

PAM
PAP
pig alvec,: ... ::•a
peroxid,c,c . • _..
....·· RAPD-PCR rnndom amplified polymor-
phlc l)NA-polymcmse chain
SPI'
SPG
s1>etiflt pathogen-free
sucro~-potassium phosphate-
'PAR progrcssivi, ·,i•w', ·~11• •Ii.$ rc.~clion ghoama1e medium
PARC 1".m-Arrimn ll111C?t ' \'It>! RB roiiculatc body SI'S sanhary and phyto-sanltary
Campaign RBT nose lle11W1l test SRH single radial hacmolysis
PAS perlodic-acid-:-...l111i ROt rccc1>tor-dcs1roying enzyme SRM specified risk material
PBMC periphc.ml bloo<l 111cmonudear ROI> nlbostimal Oa1abase Project SRSV stlloll round structured Virus
cells RE restric1io1, enzyme; re1iculo, SRV small round virus
PCMV porcine cyuun'*btl,\•in1•, endoihelial ss Schweiger-Seidel
PCll
PCV
polymerase ,lo.i,. ' ,. ... '·· ..
packed cell •,vi• • •• 1>-111,. :J IC
RE,\ remirlion enzyme analysis
rc1>lic.1tivc rorrn
ssnr sheep si:raple brain pool
RI' ST he~t siablc: swine testis
clrovlru,: prima• , ali,·h hu:. RPLP re·nric1io11 fiagmc111 lenglh STb heJt-s1ablc entcrotoxl.n
PONS porclnc derrna, .11111 polyrnorphism S1'Ec; Shil(11 toxin -producing B. coli
ncphropatli)' (11: I1la hi,) ~l'" ' RFVL Rockefeller Foundation Virus SUN scrUJn urea n1trogen
dromc l..al>omtories (SU)-1'M surface transmenibrane
PE proHferntiv,:. c.·i ,•:trhr RllOV rabbi! haurnorrhagic disease svo swine vesicular disease
PBO porcioe cmlt tui<. diurd·w ..·u
4
virus SVDV swl11e vesicular disease virus
PEOV porcine cud('011" ,11t1ahn1.•t1 RI regional lleilis SVN ~•rum virus ncuualization
virus RIA rndit,~immuno-assay svs seminal vt:Slculitis syndrome
PEG polythylcnc t;l} . [\I( rabbll kidney
PENV porcine.en1cri<" ~~··· •1t1fa1!,. R·LPS rough lipopol~ccharide 1'
PESV porcine enw, k r. ''11virus JINA ribonucleic acid· TAS trypicasc-arginlnc-scrfne
PFGE pulstd ncld ok..:. :,lt.orcsis llNI reactive nllrogen in1cnncdia1cs TBA 1mchc'Obronchial aspirate
PFIJ
PKE.
plaque-f,;,nn111: ...
proliferative ha:·.. ,,.,,i.auk
RP rinderpcst TBE rick-borne encephalitis
RPV rinderpest vin1s TBF rick-hornc fever
enteropa1hy RSSE Russiun springcsummer TBILI 1otal hilirubln
Pl parainOuc111.11 L; 1, encephalitis TC 1issuc cuhurc•adaptcd
PIA porcine.iutc~du .. ...d1•,1on1a10- RSV l'(!Spimtory syncy1lal virus TCH thio1>hcnc-2-carboxyllc acid
sis HT reverse transcr'iprnsu hydmzidc
PIM polymorphic imn,,,nodoml- llT-PCH reverse transcriprnsc-poly- TCID !issue cuhurc infectious doses
mmt molecule rnernse chain reaclion TllM transmission electron
Pl.D phospholipase I) RTX replfms in structural toxJn microscopy
Pl.SO pseudo-lumpy skin disease llVF Rifl \/:Jley fovcr TEME thrumboembollc mcningo-
PMWS pos1-wcani111l 11111hi,sys1crnic RVl'V Hifl Valley fever virus enccphulltis
wasting syndrome TFC:i\ transfromier conservation area
PNP prolircrative and nccrotiijng s TGf. trdnsmissible gastroenteriwa
pneumonia $AOC Soulhern Africa Development 'l'GEV 1ransmissiblc gas11oenteritis
Po1V porcine torovlrus Community virus
PPO purified pm1ein derivati\'C SAF scr.1pie associated robrlls TME ihmmbotic menlngoencephmi-
PPEM potemially palht,geni.c SA-OMW South African ovinc macdl- tis
environmcnrnl mycobacteria vlsna vin,s TME 1ransmissible mink
PPLO pJeuropncun,onia-li~e u,i:an· SAT slow (tube) agglu1i11mion tes1 cnccphalo~thy
ism SUI: sporadk bovine TNF tumour necrosis factor
PPR .Peste des petits ruminants cnccphalomyelitis TPGYJ' trypticase-pcpione.glucose-
PPRV pcs1c des petits n11nlnan1s virus SOL sporadic bovine lcukosis yeast extract broth ,vith tt")'.PSl.n
PR pseudorobies SBO spccmed bovine offals TSI! transmissible spongiform
PRCV porcine respira1or)' coronavlrus SCIO severe combined immunodcfi. cnccphalopau,y
PRDC porcine ccspiralory disease ciency 'l'YM trypricase-yeast extraot-maltose
complex scv small cell variant
PRRS porcine rcrrc,t.luctivc and so swine d).'l'entcry u
respiratory syndrume SOA Sabouraud's dex1rose agar UllKO undifferentiated bovine
PRRSV porcine reproductive and sos sodium dod.ecyl sulphate respiratory disease
respiratory ,yndrome virus SI swine infiue.n1.a
PRV porcine ro1avims SIN Sindhis (virus) V
PRV pseudorob1cs viru• Sip ~mple incubalion period VAT v:nfoblc antigen types
PV parasilophorous vacuole SISS segrcga1ion. isola1lon, VliE Vcne-1,uclan equine encephalitis
PVM parasilophorous vacunlar s11bclivisio11, stress reduction VF$ vesic,tlar exanrhema of s\\llne
membrane SL.A swine leuko~e a111igons VESV vcsiculor exa111hcma or swine
SL..ll S1. Louis cncephillilis virus
Q S·LPS smt>olh lipopolysaccharide VIA virn.~ inrectlon-associated
QBC quamitatlw bully emu SMEOI stllll>inb, mumrnilication. VIA,\ virion infection.a.%oclated
embryonic death. infertility antigen
R SMS San Miguel sea lion VN virus 11c111rafization
RA rheumatoid arihri1is SMSV San Miguel sea iion ,\/irus VJlff virus neutralization tesl
RAPO random nniplificd polyme:rn.se s.o.s. stagna1lon, ovcr·abum!ancc vu• virus-like particles
DNA and stress VP viral protein

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xuli

VRG vaccinia-rabies recombinant w y

(vaccine) WEF. wcStern equine encephalllls VARU Yale Arbovlrus Research Unit
VSG variabre surface glycoproicins WHO World Health Organization VLD years lived wirh disabUlry
vs vesicular stoma rills WNV Wesr NUe virus vu. years or life lost
VSV vesicular stomarit.is virus WIil World Reference Laboratory
WSL Wesselsbron disease z
WSLV Wcsselsbron virus ZN Ziehl-Ncelsen

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SECTION ONE
Aspects Influencing
the Occurrence of
Infectious Diseases

.J
§
}

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 1
Vectors: Ticks

RA l NORVALt AND I G HORAK

Introduction greater kudu :Tmgelaphus srrepsiteros) and sable antelope

(Hipporrag11sniger) of 8. decolorarus: giraffe, African buffalo
Ticks are of great economic importance as ,·ec1ors or several and eland of adult Hyalomma margi11m11m rufipe. and H.
diseases of domestic livesrock and of commercially farmed rr1111camm; caracal (Caracal caracal) and rnow1taiJ1 reed·
wildlife in sub-Saharan Africa. Globally. 690 species and buck (Red1111ca J11il1omf11/a) of adult /. r11bic1md11s. African
subspecies of ixodid ticks are recognized. and slightly more buffalo. eland, male nyala (Tmgelaphus a11gasii), greater
than 200 of these are present in theAfroi.ropical region.2 ; In kudu and sable antelope of all stages of de\'elopment of R.
addition. 37 of the 179.species or subspecies of argasld ticks appendicu/atus: zebra (Equusspp.) and eland ofall deniJop-
occur here.27 mental stages of R. e11ensi everrsi; zebra, black rh'noceros
Only a small number of licks are major vectors or dis- (Diceros bicomis). eland and gemsbok (Oryx gaze/Ill) of Rhi·
eases or causes of toxicoses in this region. These are Am· picephalus pu/chellus: large carnivores, zebra and wild suids
blyomma hebraeum. Amblyomma 11nriegntum, Boophilus or adult Rllipiceplu1l11ssim11s; impala and greater kudu of all
decoloraws, Boophilus microp/11s, Hyalomma dromednrli, stages of R. zom/Jezie11sis: and warthog (Phacocl1oems afri-
llyalommn mmcawm, b:odes ru/Jic1111dus. Rhipicep'1alus cai1t1s) of ticks or the 0. m,;JU.batalporci nus complex. Ger-
appe11dic11laws. Rhipicepha/useverrsi evensf, Rhipic11pha/11s bilJine rodems are hosts of the immature stages of H.
zambeziensis and argasid ticks of rhe Ornirhodoros mou- 1n.mcarum and murid rodents the preferred hosts ofthose of
/Jtualporclnus complex.. However, several other species also R. simus. Scrub hare (Lupus sa:mcilis) are preferred hosts of
serve as vectors. but to a lesser degree or in a more geo- all parasiric stages of Rhipicepha/u,t 111nrb11rro11i, as well as
graphically localized conrei--r. the Immature stages of H. margin(l(ttm ruftpes and H. mm·
II is customary to consider domestic animals as the pre· carnm. They are a.lso good hosts of the immature stages of,t
ferred hosts of those tick species that transmit diseases 10 hebme11m, R. appendic11/arus,R. e11errsi e11err-si, R. pulchell11s
them. However. the only ticks that nearly exclusively pru:a· and R. zambe::ie11sis. Rock elephant shrews CE/epha11ru/us
sitize these animals are the introduced Asiaric blue tick. Boo- my11rt1$) are excellent hosts of the immature stages of/, m·
philus microplus. of caule; th e shin}' Hya!omma. H)'alomma bic1111d11s and of R. warbttrtoni. Helmeted guinea fowl (Nu-
detriwm. of ~onh Africa: and the cosmopolitan kennel rick. mida me/et;gris) and leopard 1onoise (Geochelonl! pnrdalis)
Rhipicepllalus sa11g11i11e11s. of domestic dogs. The vast ma- are good hosts oi the immature stages of A. hebraeum, and
jOri!}' of indigenous ricks in the sub-Saharan region are ground-frequenting birds of those of H. mnrginnmm
parasites of wildlife, and indeed a large number of species rufipe.<.
would be unable 10 C'Omplete their life cycles if there were no The diseases of major economic imporranae affecting
\\ild h6sts a\•ailable. 1' Iany of the tick species deemed to be ca ttle and transmitted by these and other ticks are hean-
parasites of domestic cartle, sheep. goats, horses and pigs water. babesiosis. anaplasmosis, tlie theilerioses caused by
are frequently more abundant or prevalem on equivalently various strains of T/1ei/er/a parva, an d tropical theileriosls.
sized or even smaller wild animals. Thus giraffe (Girajfa Of lesser imponance in cattle are the general ly non-patho -
camelopard,1lis), African buffalo (Sy11ce1 us ca![erJ and eland genic mlh.J Llrnileriu"e". spi.rochaet0sb, I.Jeuign anaplasmo-
(Tnurorragus or:vx) are excellent hosts of all stages of devel - sis, benign babesiosis and bovine ehrlichiosis, Sheep and
opment of A. lrebrae11111 and A. unriegawm; impala (Aepy- goats are susceptible to the organisms causing heanwa1er,
ceros melampu.s), eland. bushbuck (Tragelaplrus scripms). anaplasmosis. theileriosis, spirochaetosis, and co the vims

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 >1rno, o,,: Aspects influencing the occurrence of infectious disea~es

causing :-;airobi sheep disease. Hor.;es. mules and donkeys be collected from the ,·egetation. In contrast. the ·waiters'
are affected by equine piroplasmosis as well as by spiro- quest for hosts from the ,,ege1a1ion and include Haema-
chaetosis, and pigs by porcine babesiosis and African swine physa/is and /xodes spp., and a large number oi Rhfpiceph-
fever. In addition to transmitting infectious diseases to livc- alus spp.
smck, some rick species are also associated \dth toxicoses
~uch as ,weating ~ickness, paralysis (e.g. KaJOO paralysis. Amblyomma h.ebraeum
spring lamb paralysis) and brown ear-tlck 1oxicosis, or with South African bont tick. Suid-Afrikaanse bontbosluis 1Afrik.)
bovine dermawphilosis. Several \\ild ruminant species are
susceptible to Ehr/iclzia rwninantimn, the causal orgnnism Identification Adult A. l1ebrae11m are large, conspicuous
of heartwater or cowdriosl~. or can act as carrie~ or chis ticks with long mouthpans, brightly coloured patterns on
organism (see Chapter 40: Fleartwater). Some are also the scumm of both males and females, Oat eyes. and brown
susceptible to certain 111ei/eria spp .. while zebra are and ,~hice banded leg$. The males have yellow-coloured fes-
susceptible 10 IJabesia cat,Ql/i and Tfieilerio equi, the cause toons (Figures 1.1 and 1.2).
of equine piroplasmosis. and wild suids to Bflbesia
trtwtnmm1i. the cause of porcine babesiosis, and 10 Hosts The preferred hosts of the adults are the larger do-
infection with the virus of African S\1ine fever. mestic and v.~Jd ungulates, 10.1• IS3. l,3 to which they attach in
lxodid ticks are also important vectors of several organ- clusters in the groin and axillae as well as on the dewlap.
isms causing disease in humans in sub-Saharan Africa. belly. perineum and peri-anal region.8 • 216 The lar,ae and
These are Rickell,fin conori. the cause oi tick-bite fever or nymphs feed on a wide variery oflarge and small mammals,
tick typhus; Coxiella lmmerll. the cause of Q-fever; and the birds and tortoises.5 ~ t io. 175• w;, 215• 2!13 They are very rarel~r
virus causing Crimean-Congo haemorrhagic fever. In addi- found on murid rodems. even in habitats where large num-
tion. argasid ticks of the Omirhodoros moubmn complex can bers of A ltebraeum larvae are present. 19 113 On domestic
rransmit Borrelia durroni, the cause of African relapsing livestock n~mphs anach most commonly to :he feet. and
fever, to humans. larvae 10 the face. the dewlap, neck and legs. 8 On the larger
The known important tick vectors of the lh'es1ock dis- ground-frequenting birds the nymph!> and larvae attach
eases and tau$es of toxicoses are listed in Table J. I . There- mainly on the head ai1d neck. 110
after descriptions of the identification, hosts, life cycle,
seasonal occurrence. distribution and disease transmission Life cycle Amb(vommn hebraeum has a three-host life
of each tick vector specfes, and the control of some. are cycle. with farvae. nymphs and adults feeding on separate
given . .'\ section on the control of ticks and tick-be>rne dis· hosts. On completion of feeding, engorged 1icks leaYe the
eases follows: this covers the concept of endemic stability, h.ost and seek sheltered microhabitats in which 10 moult or
the history of rick and tick-borne disease control in southern 10 lay eggs. The de\Telopmental periods off the host can be
Africa. recent research relevant to tick control. biological long. In the Eastern Cape Province of Somh Africa. lar\'ae
control. tick and tick-borne disease models. future control may take five months to hatch from eggs laid by female ticks
methods, integrated contr()I, and the control of ticks on that detached from hosts during March or April.230 and
\\1ldlife. nymphs may remain inactive for two 10 three months after
moulting. 168 However, the life cycle usuallr lasts one year
T ic k vectors under field conditions. but mar for the above reasons ex·
rend for longer. The pattern of seasonal occurrence is de·
Most ticks can only be identified 10 species level under a ste- pendent on dim ate and ,·aries considerably throughout the
reoscopic microscope and a conventional light microscope distribution range of the tick. In general. adulcs 1end 10 be
may e\•en be necessary for the idemification of the imma- most numerous during the warm , wet summer months, lar-
ture stages of some. There are, however, a number of el- vae du.ring the colder, dry, late autumn and winter months,
ememarr observations that can be made to assist with the and nymphs during the winter and spring months. How-
recognition of tid,'S to generic level. These are geographical ever. varying numbers of all stages of development can often
region or acmal locality in or at which the ticks were col- be found on hosts throughout the year. 8• 89• 168• uo In the
lected. host and site of auachment, season. and any mor- warmer. moist. lowveld regions of the KwaZulu-.Natal, Mpu-
phological features of the ticks that are discernible with the malangaand Limpopo provinces in the north-east of South
naked eye. Africa as well as in southern Zimbabwe, I.he life cycle seems
Free-living adult ticks can also be characterized as ·hunt- to be continuous \,'ith little indication of a definire seasonal
ers' or ·waiters'. 'Hunters' actively seek hosts by scuttling pattern of abundance for the va.rious life slages.9 ·•· 95• t 79
along the ground when stimulated bya host's presence or by
pheromones produced by ticks already attached 10 a hose. Distribution J\mblyomma lzebrae11111 is exclusively a
Ticks chat follow chis strategy are the .>\mb/yomma and Hy· southern African tick. It occurs endemically in the coastal
olomma spp.. and R. warb11rto11i. These ticks will \'ery rarely regions oft he Eastern Cape and KwaZulu-Natal pro1inces in

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 Vectors: Tic~s 5

I
V

--
--

Figure 1. 1 Amblyomm;i hel1taeumr;,a.e

1Sv co~rtesv o! J.B. Walker. OVJ. Ondersii!poortJ

'
JI
•

/J
, figure 1. 2 Am/Jfyommoheb1seumfema'e
rBvcourtes1 o; J.B Walker. OVI On~~ri,,epoo·:,

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6 ~cno~ost: :\$pec1s influe11clng the occurrence or infectiou$ dl,eases

Table 1.1 l,ck vec:ors of diseases of domest,c and w Id animals ,n sub-Saharan Africa

DISEASE ANIMALS AffECTEO I VECTORS MElHOOOF REFEAENCC..S

!CAUS.ATIVE ORGANISM) TRANSMISSION
Heartwater or t0'.1:!nosis Cattle. she.;>. g:)ats. .v,ld Amb/wn,ma nel)laeum lS, IS. ro· Z. G IJS
{Ehr/ichia /Cowdria/ ruminantium'i antelope AmliiJ,omma vatiega111m TS 34
Amblyomma pomoosum TS 160
Amblyomma lep1dum TS 216
AmblVO/'fl/118 spl). TS 216
B:ivine ehtlicn,os,s Cattle Rhipicepflafus appe,?dteulews TS '48 170
tEhr/ichia bo~1s1 f/h1{)icepalus 1amb:Ziens1s TS 257
Ovine eh1lich1osis
(Eh1//ch1a o~ma'
Sheep Rhipicephsfus averts• eve:rs1
Rh1p1cephalus ou:sa
,lS 162
288
Anaplasmosis Cattle, eland Boophi/us deco/o13tus TS. IS 22:
\Attaplasma marginale) Booph1/us microp'us TS. IS 22;
Hyqfomma marg,ttawm rufipss IS 721
Rhipicephal~s evertsi fJV8'1Si IS 121
Rh,pi,,-ephalus s1m1.1s TS. IS 121
Benign anaplasmosis Cattle Rhlpicephafus s;mus TS l2J
(Anaptesma cenvale)

Ovine anaplasmosis Sheep, goats Rhipicephalus bu;se TS 229

(AMpl11Sma OVJS)

Bovire ballesiosis or African Cattle Boophilus deco/o,-an,s TO 223

reawater 8ooph1/us mrcrop!us TO. TS? i5.235
(Babes,;; bigemir.a} Rt,,p:cepha/us ever1s e,1l'ns, TO 15
f/l11pice,ohatus bu1se TS, TO 75 288
&,me baoeslosis or Asiatic Canle Ooophilus m,crop'us TO ,.: 220
redwa1e,
(88/Jes,a liOwSJ

Bsnign babesiosis Cattle Hyillomma margmawm ruf,pes TO 81, 27'

(Babesia ocr:ut,ansl

EQu,ne p1rop1asmos1s Horses. <ion.keys. mu,es. zebra Hyelomma rronca;urr TO .!9

(ikoes,a caba/11] f/h1p,ceplla/us everis, e,ertsi TS 51. 270
Rhfpicepha/us bu/Si/ TS so
R/11pi<:llphslus rur5nicus TS.TO 60
(as fl sangumeusi
E~uine piroc1as"'10S1s liGrSllS oo,,ke·1s. mu,es. ,ebra Rh,p,cephelus e,e:rs, e,e,w TS 270
(Tneilena et11Jil" • Rhip1cepha/vs evij,.SI m,meuus TS 222
Rh1pteepholus butSa TS 75.2SS
Rhipiceph81us wranici;s 60
{as fl. S8f19Uln8C:S/
Hyolomma delr,tum TS 75
las H. anacolic,m anar~!icum',
Porcine babesiosis Pigs. war<1>og. bushp1g Rh,picephalus simus TO 50
(Babes,e ua1.11rp.;innil Rh1p1cepha/us t1Jf~n1Ct1s TO 134
East Coast 'e-,ar Cau:e Flh,p,cepl)s/us eppend1cula1us TS ·o.~s
(Tneill!'1e pa/V8stfams} {syn. Rhi{Jlcephs/1.1s zambez,ens,s TS ·29
Theilari, perva parva'
9mbabv,e the1lerios1s or ~ttle F/h1picepllalus eppendiculatus TS ·.;s 1s0 . 2a1
January disease Rl11p,c11phalus zamlJe,, ens,s TS '29
I Theiler1a parv11sua1ns)

Comee: or buffalo disease F/hip1cepflelus awrend1cu!ar1JS TS !64

{ TheNet1a pan,esva,nsJ F/h1picepha/us 1amcez.eM1s TS ·29 181
Ji
Rhlp,cephalus duitcn,
'

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 Vec[ors: Ticks 7

Table 1.1 (cominusd)

DISEASE ANIMALS AFFW'EO VECTORS METHOD OF REFERENCES

(CAUSATIVE ORGANISM) TRANSMISSION
Trop,cal 1he1leriosis Cattle Hyalomma demcum iS IS
IThei/eria annulatdi Hyalomma cromedam TS S3
Ben gn bovine mellenosls Cante Amblyom,na heb1ae11m TS .IS. 127 27g
(The,Jer,e mutans. i ve//fera) Amb/yommo variegawm TS 282 283
Benign bovine tlteilerios1s Cattle. eiand Rh,p,cephalus append,culaius TS i26. 281, 305
l TheiJer,a 1auro1tcg1 J Rhip,cepha/us Mmbez1ens1s TS 129

Clvme theilenos,s Sheep Rhipicephalus evens, evarrsi TS 115

(Thei/sria sep;Jrara)" • • Rhipicephatus everrsl m,mez,cus TS 165
RhipicephQ/us bursa TS ;55
Borrellosis or sp1rocilae,osis Canle, sheep. g~ts. ho,ses. Boophi/us deco/otatus TO 269,271
(Bo,Telia 1hei/eri1 donkeys, mutes f/ODphilus microplus TO ?
Rhip1cepha/11s l!Yl'.n,1 e•,e.ns, Tli.TO 2r
African swine fever Pigs, warthog. bushp,g Om,rhodoros moubarecomplex TS. TO. S 2'7, 2i8. 2i9
(Asfa1V1rideel

Na,~b1 sl1eep disease Sheep. goats Amblyomma vaf/egaclJ'TI TS 36

(alld Kise~ye sheep disease?) Rl1ipicephal11s appel/d1aJfaW$ TS. TO 22.Jt Jg
(B11lf)'8vilidee) Rhip1cepha/us pu/chel/us TS. iO 209

Ccimean-Congo haemorrhagic Canle. shee;i Hyalomma merginarum ruflpes TS 267

fever Hyalomma 1ru11ca1um TS 2~3
(Bunr-avmlfaeJ Rh1picephetus ever1s, evsr1s1 TS171 257
Rhlp,cephalus e,errsl mimet1cus TS(?) 243
Bov,ne dennatophilosis Cattle Amblyomma variegecl/lli 24, 145. 178
IOermatophilus congolensis)
Sweating sickness Cattle Hyalomma 1ru11carum :z. 1~ 161
llo~cos,sl
Kacoo para1,s;s Calves. sheep, goats. antelope fxOdes rub,cundus as. ~a. 28$
(tOXCOStS)

Spring lamb paralysis Lambs F/h1picepha/us evsrts, evens, 80.8,

(tox,coStS!

Paralysis Sheep Flh1p1cepha/us bursa 288

(tOXCOS.$)

Pa:alvs1s Calves lambs Rnipicephalus lunula11.1s 128. 197, 273

(toxcos,sl (as R. mcuspis)
Paralysis Cattle Rhip,cephalus preeie.tlatus 202
(tOJtcosisl (as Fl. slmus)

Paralysis Calves. lam!:$ Rhipicephatus simus 185

(tox:cos,sJ

Brown t,ck paralysis Goat kids Rhipicephafus 1•.,;;rtwmni S9

(toxcosisl (as R prows gn>upl

Bro,·m.ear tick 1ox,cosis Cante. an1ekpe flhipicepha/us eppendicul;;IIJs 133. 27~

TS = Trenss.adial, IS= ln1rastadlal, TO= Transo,arial, S = Sexually transmlned between tic~s

Rarely uaosovarial transmission of E111l!ch18 ruminanrium by Amblyomma hebraeum may octur 13
8.Jbes,a equ1\vas redescribed as Theileue equi by MeNhom and Sc.'lein In 1998.1s. It is S1Jggested that equine p,roplasmosis be useo ,\nen
referring :o !he disease caused by tins organism ar.8 bv Babes,a cebe/li
The Soullt Afn~an Theiler/a ovis desCiilled by Jansan and Ne1ti1·; i$ considered by Uilenbarg278 co be svnarymous with Theiler,e sepa:a;a
a parasite originally desc11bed Icom sheep n TanZlln;a wi Theileria separate does no1 inlet: goats

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8 ,a;no,i oxc: Aspecrs influencing 1he occurrence oiinr°""ilous d[seases

South Africa, and in southern Mozambique. Its di$tribution tea.ts of heifers or cows are heat<ilr infested. Over-rigorous con-
also extends inland through the lowveld areas of Swaziland. trol is likely 10 have the negative effect ordisrupting the narural
the Mpumalanga. Limpopo and North \Vest provinces of cransmission of hcanwater to young animals and thus reduc-
South Africa, and eastern Botswana to southern and western inglevels of naturally acquired immunity, consequently affect-
Zimbabwe as well as parts of the Zimbabwean h ighveld.20• ing endemic stability to the disease in domestic ruminant
115 . 213 - 21 • · 273 · 290 · 292 11 is absem frem the highveld regions of populations. 110. 1n
South Africa, possibly because these are 100 cold and tree- Regular acaricidal treatment of domestic livestock, and more
less. but may suf\<ive locally in these regions in wooded particularly canle. can result in the virtual disappearance of all
habitats such as north-racing gullies, ra\'lnes and valleys. It developmental stages of A. hebraeum on sympalric wild ani-
was originally chought tha t the highveld of Zimbabwe would mals up to and including the size of greater kudu asweU as the
be unsuitable for A hebraewn because of its cooler tern- disappearance of free-living larvae on the vegetation. 102. 2 15
peracures. i;s However. the Climex model. indicating ecocli· Larger \\1ld animals such as girn.tfe, African buffalo and eland,
rnatic suitability, 2 i;.1 demonstrated that conditions on 1hc which are excellent hoslS of aU stages of development of the
Zimbabwehighveld a re favourable for its Sl1f\1val. lllll Recent tick,'0.1, raa. 175 arc- probably capable of maintaining popula·
surveys conducted there confirm this predktion.20· 213• 21• In tions ofA /zebraeum 011 a farm, even though che domestic live-
central Botswana its distribution is limited by increasing siock on the property are being created \\ith an acaricide. The
aridiry, and in northern Zimbabwe and central i\lozam- residual populanon of ticks on \,ildlile on such mtxed cauJe
bique it may be restricted by interspecific competition wirh and wildlife farms may serve as an important source oi tick
Amblyommtl L'ariegawm. rm, 260 Ambl)•omma hebraeum is infestation. and consequently heanwarcr infection. 21 0.. ~11 ror
associated with wooded habitats. including coastal bush. the domestic srock on the farm and thus assist In maintaining
riparian woodland. th ornveld and mopani woodland. It immunity.
does not occur in e-x1ensive open, treeless areas.
Amblyomma vnriegatum
Disease transm.issfon Amblyomma hebraeum is the mai n Tropjcat bont tick. tropiese bontbosluis (r\frik. )
vecror in southern Africa of Ehrlichia r111ninamium, the
cause of heartwater or cowdriosis in domestic and wild ru- Identi:fication Adult A variegmum have long mouthparts
minants. Laboratory studies have demonstrated that A he- and banded legs like A. hebraeum, but the colour patterns
braeum can carry higher infections and also a greater 011 the scutum are different and they have beady eyes. In ad-
number of strains of £ r11mi1umri11m than A variegarwn, dition. the festoons of che males are dark brown in colour
the other major ,·ector of this disease. u 1 142 In addition, (Figures 1.3 an d I .4).
field obsef\·ations seem to indicate that outbreaks of the dis-
ease arc more common!}' associated with the former than Hosts and life cycle Amb/yomma tl(lriegacum has a host
the latter tick and that these outbreaks ace also more range similar 10 that of A /zebraeum.86· m,. 216• 273 It ls also a
severe. 1118· 2 13 BesidescauJe. a number of\\ild ruminams.210• three-hos: tick, but differs from A. l1ebme11m in that the imma-
2 11 212 ture stages have a more clearly defined pauem of seasonal oc-
• as well as other animals such as scrub hare, helmeted
guinea fowl and leopard tortoise can act as asympt0matic currence. In 7,ambia adults are most abundant in the wet
carriers of E. rwnina11rium. Other organisms cransmia;ed by season (October to February). larvae from :Vlarch 10 Julr, and
A. hebraewn are Theileria n 1uta11s and Theill!ria Pelifera, the nymphs from Mav to September.206• :ioa This pauem of sea-
cause of benign bovine theilerioses. The occurrence of foot sonal abundance can arise from morphogenetic diapause in
abscesses in goats in the Eastern Cape Pto...-ince, Sou rh tl1e females, resulling in a delay in oviposition.205 Similar ob-
Africa, is significantly related to the seasonal abundance of sef\'ations of onlr one generation per annum have been made
adult A. hebraeum and Rhipicephalus glabrogutawm. 138 in other regions which have a single rain)' season.?16 In Zlm·
The a nachmenrsites orthcse licks around and between 1he babwe adults can be present tl1roughou1 the year. with hea1ier
hooves afford enrrancc 10 secondary bacterial infections infestation~ in the wanner months (September ro May). and
leading to abscessatlon. with nymphs present only from June 10 September. r7s

Control Adult A hebmeum ha,·e long feeding periods and Distribution Amblyomma variegawm is \\idelydistributed
consequently good cont:rol of the tick can be achieved by the through West, Central and East Africa and in somhem Africa
treatmem of cattle with conventional acaricides nr two-weekl)' e.xtends h10 north-eastern Botswana. the Caprivl Scrip of
imef\,als. As the ticks auach predominantly to the w1dersides Namibia, and northern and central Mozambique. 2.'3· 292
of canle, control c-.in be achieved by localized acarlcide appli- Since 1985A mriegnwm has spread from the north-western
cation. It should be noted that in endemic hearrwa1er areas regions of Zimbabwe to the central and ea.stem highveld.213•
214
control ofA hebraertm is on!}' necessary when large clumps of Its spread southwards appears 10 be limited by inter-
adults, forming potential sites for strike by the screw-worm fly specific competition ,,~ch A hebraewn.260 \\ith which it
(Chrysomya bezziana). occur on cattle, or \\'hen the udders or shares similar habitats, hosts and sites of auachmem.ns. 216

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 Vec1ors: T!cb 9

Figure 1.3 Am/Jlyomme

V1Jrregatum mare
\ (By ccur.esv of J.B. Walker,
OVI. Onderstepoonl

Figure 1.4 Ambf.yomma

r varfegawm female
IBv courtesy of J.B. Walker.
OVI, o~.derstepoonl

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lO ,ecnm, o,.-i:: Aspccrs influencing the occurrence <1f infecriou$diseases

Disease transmission Ehrlichia r111ninanri11m. the eause of bers of tic:<S presem.301 • 31lZ ln Ethiopia. north of the equator.
heanwater or cowd.riosis; 111eileria muro11s -and T. 11elifera. adults peak during lhe rainy season in May to June.202
the cause of benign bovine theilerioses: Ehr/ichia bo11is. the
tause of bovine ehrlichiosis; and the virus causing Nairobi Disease transml.ss.ion Ehrlichia r111ni11a11ti11m, lhe cause
sheep disease. Bovine dermatophilosis (Dermatoplii/us con· of heartwater or cowdriosis.
golen.<i.f) is associared with the presence of the rick.

Control The control oft\. variegarum wi.lh acaricides is Amblyomma spp.

Similar to that described for t\. hebmeum. However, the A number of other African AmbTyomma spp. have also been
more clearly defined seasonality of.-!. 11ariega11111i facilirates implicated in the transmission of heartwac.er 10 domestic live-
strategic control of the adul ts du.ring the wet season. stock (see Chapter 40: HearrwaterJ. Except lU1der certain un-
usual circumstances the ability of several of d1ese tick species
Amblyomma pomposum to transmit heanwa1er to domestic ruminants is limited
Angolan bom tick because their adults. "·hid1 appear to be the mosc efficiem
vectors, by preference do not feed on these animals.216
Jdentification Amblyomma pompasum doselyresemblesA The males and females or.all these ticks aze ornate,29?
11arlegmum but Is more heavily punci:ate (Figures l.5 and l.6). havt> long mouthpans. and all have three-host life cycles.u6
These ticks include;
Life cycle, hosts and distribution This tick has a three- • A. astrfon, of which the principal hosts of the adults are
host life cycle similar to those ofA hebmeum and A uariega- caule, sheep and African buffalo. Tt is distributed mainly
rum and has a similar host range.216· i 42 The adults most in 1he Central Afri<.'an Republic. the northern and west·
frequently auach on the undersides of tattle.2' 12 Jr is present em regions of the Democratic Republic of the Congo,
in Angola, parts of western Zambia. and in the southern re- and north-western Angola;
gion of the Democratic Republic of the Congo. lllZ However. • 11. r:ohaeren;, which infestsAf.rican buffalo, and domestic
unlike other southern African vectors of heam,·arer, A. pom- cattle in regions in which buffalo ate, or used 10 be. com-
posum occurs in wet highland areasln savanna and forest.58• mon. It has been recorded chiefly in the eastern region of
86 273
• z~z. In Angola, nymphs and adults are most abundant the Democratic Republic of the Congo, western Uganda
during the rainy season, a lthough large variations may occur and cenrral Ethiopia;
depending on the region and climate.2'•1 • :!. gemma. for which domestic cattle and large, wild her-
bivores are lhe preferred hosts of the adults. It is main!}'
Disease transmission Ehrlic/1ia rumina11ti11m, the cause distributed in eastern Ethiopia, 11onhern and southern
of heanwater or cowdriosis. Somal!a. Kenya and north-eastern Tanzania;
• A. ma,moreum, of which all s tages of de\'elopmem. and
most particularly the adults. feed on tortolses and large
Amblyomma. lepidum varanids. Adults very rarely parasitize domestic livestock
East African bont rick but the immature stages, especially larvae, are freq uentl~
encountered on thcseanimals.111is tick is ,,idespread In
Identification Amblyomma lepid11m closely resembles A. Zimbabwe and South Africa and is probably more preva-
hebraeum, but has beady eyes and the males have \'ariably lent in )..iozambique. Bo tswana and ::--amibia than cur-
coloured light and dark festoons (Figures 1.7 and 1.8). rem records seem co indicate;
• A. sparsum. of which the adults prefer large reptiles. but
Life cycle, hosts and distribution This is a three-host tick black rhinoceros and African buffalo may also be impor-
and the adults prefer canle as hosrs, although camels tant hosts. It rarely infests domestic hosts. l\•!ost records
(Camelus dromedarius) may also harbour quite large infes- of this tick's occurrence are from southern and western
tations. 216• 28; · 302 Other domestic animals can also be in- Kenya. northern and central Tanzania and north-1\'eSt·
fested as well as a number of wlld ungulates. Adults are em Zimbabwe; and lastly
predominantly found attached on the vennal surface of the • A. tholloni. of which the preferred hosts of the adults are
host from the lower dewlap and a."'illae 10 the escut· African elepham (Loxodonta africmm) and hippopota·
cjieon.302 fl ls widespread in eastern Sudan, Ethiopia. somh- mus (HiJl/lllJmtnrnu~< amphihhi.,). 115 Although larvae and
em Somalia, eastern Uganda, Kenya and the nonhern region nymphs have been collected from cattle, sheep and
of central Tamania. 292 In Tanzania. which lies south of I.be goats. domes1ic stock are not common hos~. 1; 5 This tick
equator, adult ticks are most abundam berween October and has been recorded in the Ivory Coast and adjacent coun-
February, beginning either shortly before or shortly after the tries to the west. and throughout Central Africa to the
onset of the rainy season. However, there may be consider- eas1em coastal regions from Kenya in the north 10 north-
able variation in the riming of tl1e peak as well as in the num- eastern Somh Africa in the south.

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 Vectoni: Ticks 11

Disease transmlsslon Eilirlichia rum/11anti11m. the cause mately 70 engorged females can represent a total parasitic
of heartwater or cowdriosis. popularion exceeding 10 000 ticks.~5

Boophilus decoloratus Distribution This dck Is distributed throughout most of

African blue cick. blouboslu is (Afrik.) the moister regions of South Africa. except for areas where it
has been replaced by 8. m;croph,s, and also occurs in cold
tdenUficallon Boophilus tlecolomms aduhs a re small, mouniainous regions such as the Drakensberg range and
inconspicuous ticks with shon momhpans and slender parts of Lesotho. It is absent from those parts of Sou ch Africa
legs. The males. which are usually considerably smal ler 1hat recei\'e an a\·erage annual rainfall of less than 380 mm.
than lhe females, are brownish -yellow in colour and the including the western Free State, and the western regions of
darker coloured intestine is visible through the lightly the Nonhern Cape Province.1 a, 112• 272 In the semi-arid to
sclerotized scutum (Fi1,rure 1.9). They can usually be found arid territory of :--:amibia it Is present only in localized areas
paired \Vith the females. The engorged females are blueish- in lhe north, and in Botswana it is restricted to the higher
brown {Figure 1.10) and can be seen aftached particularly rainfall eastern border areas and a few scanered northern
10 the face, neck, shoulders and escutcheon of cattle. localities. 290 Boophi/11s decolorams is also present in south-
ern :Vto1.amblque, Zimbabwe, particularly the eastern re-
Hosts Boophl/us decolorarw, prefers carcle as hosts. In ad- gions. Angola. much of Zambia, Malawi, south-western a11d
dition to cattle it frequently parasitiz.es horses. donkeys. northern Tanzania. Bunmdi, Uganda, western Kenya and in
sheep and goats.52. 111 147• 287· 302 Conua11 to earlier obser- the wetter highlands and sub-highlands of Ethiopia. 1~-. 151
202 • 287• Joi EL is also found in Central and WestAfrica.52 a,;. 152
\'acions it is commonly present in large numbers on several
wild ungulate species, inclucling Surchell's zebra (Equus II occurs in various vegecation types, including coastal mo-
IJ11rchellil), impala, bushbuck, greacer kudu, eland and sable saic, grassland, Cape shrubland, bushland. woodland and in
antelope.90 · '35. 97• 106, 147• 308 Rather unusually. African buf- u ndifferentiated montane vegecaEion.
falo do not appear lO be good hosls. 106, 139· t-11'. 177
Disease transmission Babesi<I bigemina, che cause of
Life cycle 80011/lilus decolornuis has a one-host life cycle, bovine babesiosis or African redwacer: Annplasma margi-
in which moulting from larva to nymph and nymph to 11ale, thecauseofbo1ineanaplasmosis; and Borre/fa rheileri,
adult takes place on the host .b The time spent on the host. the cause of borreliosis or spirochaetosis. Eland can be
from auachment of the unfed lal"\'a u11ril decachmentofthe asymptomatic carriers of A. margina/eand could thus serve
engorged female. Is approximately three weeks,6 and that as a reservoir of infection for domestic cattle.166 Boophilus
spent off the host. from pre-o\'iposition until tarval batch- decoloratus i< also suspected of being a vector of Babesia
ing and maturatio n. approximately five weeks. Because of trau1r11,11111i. the cause of porcine babesiosis. However. the
this short life cycle the 1ick is able tQ pass through more small number~ encountered on wanhog, even in regions
than one generation annually. and ls usually prel;ent in where large numbers of this tick are present,90 ma.l.e this
varying numbers throughout the year. In souchern Africa seem unlikely· under field condi tions.
low \1;mer temperatures :,ynchroni1.e egg development
and larval hatching.ns Consequently large numbers of lar- Control Because 8. decolora111s spends approximately 21
vae are present on the vegetation when lhe weather warms days on its hosts to complete its parasitic life cycle it can
in spring.i~3 Waves of larvae then occur throughouc the effectively be controlled by acaricide treatment of canle at
summer and into the cool months of Mar and lune. In three-weekly intervals. Bos indicus and Sanga-t)i,e cattle
S0utl1 Africa the largesl numbers oi ticks are usual!~· develop a considerably better resistance to this tic.k rhan Bos
251
present on cattle during the summer and autumn 10 early f(l11ms cattle.2 31• 232 • and consequently require fewer
winter months. 8• r,.;. 230 • ?3G but a spring peak in abundance acaricide treatments.
may also occur.9• 2.% In Zam b ia a spring peak may be fol-
lowed se\·eral months la ter by a considerably larger la ce Boophilus microplus
summer or winter pcak. 139 · 2 '16• ~0, In Nigeria, north of the Asian blue tick. A~ialiese/ pantropiese bloubosluis CAfrik.)
equator. 8. decolorarm is most abundam on cattle during
autumn.•': while in the Gambia peak burdens are present ldentification Adults of 8. mlcroplus are slightly larger
on th,P<P animal~ during 1hP. rainy ~ea~on.15i On \,ild her- than those or B. de.-olorn111s, and are slightlr more red in col-
bivores in South Africa the high.est burdens are usually re- our, but are otherwise very similar in general appearance.
corded in spring and. unless these animals are stressed
because of disease or drought or other conditions. their late Hosts In sub-Saharan Africa domestic cattle are virrnally
summer and aucumn burdens are low.9" · 9 ' Because of its the only hosts and other livestock species and \,ild ungu-
one-host life cycle all stages of de1-elopmem occur on the host la tes are rarely parasitized. and then only if cattle infosted
at lhesame clme and the presence on a single host of approx!- with chis lickare present.""· tor. 14; · ~02

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12 »cno, o~,: Aspects influencing the occurrence of infec1ious disease;

Figure 1.5 Amblyommapomposummale

(By COlJrtesy of J 8. Walker, OVI. Onderstepoon)

Figure 1.6 Amblyomma pomposum female

1Bv courtesy of J B Walker, OVI. Onderstepoort)

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 Vectors: Ticks 13

Figure 1.7 Amblyomme Jepidum male

(By cQurt&$Y of ~.B Walke, OVI. Oncterstepoon}

Figure 1.8 AmoJyomm2 lepidum female

(By courtesy 0 1 J.B Wal~er. OVI. Onderstepoortl

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14 =io, oSI!! A.<pecls influencing the occurrence oiinfec[ious ,diseases

Figure 1.9 Boophifusdecoloratusmale

Figure 1.10 Boophilus decolorerusfemale (engorged)

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 Vectors: Ticks 15

Life cycle B00phi111s microplus has a one-host life cycle. Identification Adulrs of fl. truncatum are medium-si1.ed
and like 8. decolormus is able to complete more than one tick:. with long mouthparrs and dark-brown bodies. beady
generation in a year. In Zimbabwe it appears 10 be present in eyes. and long. red and white banded legs. The posterior
variable numbers throughout the year.l"'; ln somh-eastem surface of the scutum in males is characterized by a depres•
KwaZulu-Natal. South Africa, it is most abundant on cattle sion comaining numerous large punctations, otherwise it is
from mid- to late sunm\er.1 comparath·ely smooth (Figures 1.11 and I, 12).

Distribution HoogstraaJ86 posmlated that B. microp/11swas Hosts The preferred hosts of the adults are large ungu-
introduced into East and South Africa from ~ladagasC'ar, lates, both domestic and wiid.86 • ~t, 17' 1• 26 iThey allach in the
where it had originally arrived with catlle from somhem Asia. tail switch. around the anus. on the lower perineum. and
In South Africa it is now establi$hed in scattered areas along on the legs, including around the feet. 139 The immature
the southern and easwrn coasts of the Western and Eastern stages feed on hares and on certain rodents. particularly
Cape provinces and or KwaZulu-Natal. lt is also present in the gerbils.19, 103. 1~4
coastal regions of Mozambique, Tam.anfa and Renya. In the
interior of the subcontinent it is round in scattered localities Life cycle Hyalo111mt1 1r1mcat11111 has a two-host life cycle.
in the Mpmnalanga and Limpopo provinces of South Africa, which normally lakes a year to com piece under fie,d condi-
in both the south and north of eastern Zimbabwe. in pans of tions fn South Africa. Adults occur in the greatest numbers
the Eastern and Central provinces of Zambia, throughout in the !are wet summer months and the immature stages in
tlfalawi and to the east and north of Lake Mala,~1 in Tan- che d ry aucum1110 spring momhs.119· 1®· ll~. 174 • ?34
zania.112· i-1;. :?a7• 302 There is evidence that where favourable
moist and warm climatic conditions exist it competes with Dis tribution This tick is adapted to di) climates and fs
and is able co replace l11e indigenous B. decolorams. 1~;. ll!!I. 2>1 absem in Lesotho and . with the exception of the Eastern
Boophilus microplus spread into Zimbabwe in the 1970s, Cape. the eastern half of the Free Stare. south-eastern
when dipping ,.,_as disrupted during the pre-Independence Gauteng and ~l pumalanga. and southern Kwa:Z1Jlu-!\atal. it
war. and replaced B. decoloraws in several areas. 11; Long• is present chroughout Somh Africa, 7.imbabwe and much of
term sur\"eys conducted in eastern Zambia revealed chat 8. Moiambique. 11.?. 1• 4 , 2• 0 It is also present in south-e~1ern
decoloratus had almosr completelr been replaced by B. and north-western Botswana; central and northern
microplus in the l O years between 1972 and 1982. ll and ics Namibia; somhern Angola; western, southern. central and
westward spread appears to be continuing. 116 The spread of eastern 7.ambia: cenrral and southern Mala1\i: south-west·
8. microplus in Zimbabwe and Zambia has been accompanied ern and north-eascern Tanzania; southern. central and
by heavy mortality in cattle due to B. bovisinfeclion. ofwhich it western Kenya: and eastern Uganda.81 • 206. 237• 102 It also
is the sole vector in sub-Saharan Afric;a. 116· 18:? occurs in many coumries in north -eastern, Central and
West Africa.116• 152• 2<12, 273 Ac a local level the abunctance of
Disease transmission Babesia bovis, the cause of bovine H. trw1cawm is influenced by the abundance of hares.
babesiosis or Asiatic red water: 8a/1esia big('mina. the cause of which are the preferred hosts of the immature stages.
b01<ine babesiosis or African redwater:Anaplasma matginale.
the cause of bo,ine anaplasmosis: and Borre/ia rheileri. the Disease transmission Babesia ca/Jalli. the cause of equine
cause of borreliosis or ~pirochae1osis. Because this tick trans- piroplasmosi;. Because the immature stages of H. mmcll·
mits both B. bovis and B. lJigemina it poses a greater potenrial wm do not feed on horses the transmission of
threat 10 livestock production than B. decoloracus. B. caballiby this tick has of necessity 10 take place transova-
rially."'9 H,valomma mmcmwn also transmits the \irus caus·
Control The conrrol of B. microp/us is similar to that of B. ing Crimean-Congo haemorrhagic fe,·er in humans. and the
decolomms. Bos indicus·type canle develop a considerably toxin causing sweating sickness in carde and especially in
better resistance to this tick than Bos taurus·C)'Pe cattle and young calves. Onli rertain st.rains of Ji. mmcarum produce
consequently the number of acaricide treaunems required this toxin and then it l& only female ticks that do so. t4 The
can be reduced on these animals. 259 • 28• However. because long mouthpatts of the ticks. as well as their tendency to
8. microplc1s feeds virtually exclusively on cattle, the whole form clusters. can cause tissue damage resulting in second-
parasitic population is exposed to acaricidal imervention at ary bacterial infections and abscessation. The attachment of
any panfo~1lar location as t.here are no ahernam-e hosts. Be· adulr licks co tl1e imerdigital clefts and fedocks of lambs al-
cause of this and !ts slighlly shorter life cycle it would appe,ar most always causes lameness. 124
as if B. microp/11sisselec1ed for resistance to acaricidesmore
rapid!)' rhan is 8. decolora.ws. Control Hand-dressing of the preferred auachment sires
with an acaricide \\ill assist in controlling the adults. le is im-
Hyalomma truncatum practical to control the immature stages because of their
Small smooth bom-legged tick. sweeisiektebosluis (Afrik.J preference for hares and rodents.

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16 ro:no~ I>~'£ Aspects influencing the occurrence of infectious diseases

{l

Figure 1.11 Hya/omma croncatummate

/)
Figure 1.12 Hyalomma truncawm !err.ale ·~~/
, '

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Hyalomma margi11<1t11m rufipes Hyal<>mma detritum
Coarse bont-legged tick. groot bontpootbosluis (t\frik.) The shin} Hy11lomm(1

Identification Adult H marginal/Im rujipes an: large Identification J\duh fl. demrl111m arc mediu111-si1.ed tick,,
ticks \\ith dark-brown bodfos. long mou1hparts a ,culllm ,,<ith long mouth pans. a \ ery dark-brown. \mooth, ~hiny
that is heavily puncmte. beady eyes and long. red scmum. and long rnddi,h or yellowish-brown le-gs \\ hich
and white banded legs. The~ differ from those of JI. may have paler band,.w,
1rw,ct1111111 in that the whole scutum is puncrate and in the
males it is more circular than the rather elongate shape of Hosts Domestic canli.' and horses arc the most common
the latter 1ick. llowcvcr, i1 is \'inuall~ Impossible 10 host~. but shol'P, goats and camels may also be infosted 116
disiinguish bc11wlln the two species by means of the naked All stages of de,elopmem f111.1d on the same hOl>t species.
eye.. J\duhs attach on 1hc inner 1highs, udder. ~crocum and
perineum of canle. 18
Hosts :\duh~ para.silizc domestic a1id wild un1:,'Ula1es.
showing a preference for the larger spedes."1 17~ 28~ Thev Life cycle and distribution /f_l'(l/ommn dl!rrirum ha~ a
attach mainly in the hairless pcri-anal region and on the two-host life c~·dl! with the adults feeding in summer and
lower perineum and genitalia.40 The immature ~,ages are thll larl'aC and nymphs in auuu,rn.~0 The detached en·
parasitic on hares. particularly scrub hare;;, as well as on 11orged nymphs undergo a winier diapausc and moult 10
grou11d-frequen1ing birds. 103• 17 '· 21 ' adults the following summl'r.18·\«;The lite cycle is often asso-
ciated \,;th barns, stable~ and sheds. and livestock become
Life cycle Hyalonmm ma,gi11a111111 rufipe.< has a two-hoi<t infc..~tcd when rhey are housed in Lhese s1ruc1ures. This Tl)'·
life cycle. which under field condition~ in South ,\frica 1akes a/Q111111a I, present along the Mcdltemmean coast ofAfrica as
a year to complete. The aduh11 are mo,1 numcrou~ in the far as Algeria and \ lorocco in the west.'"' It is also present in
eady part of the wel season and the immature smges in the sub-Saharan Africa in norih-cemral Sudan. \\1lich it may
dryseason 11>1. 139· 1•·1 have invaded from the Red Sea coast or ,;a 1he 1\ile River
Valley.~''
Distribu tion This tick is widely dis1ribuiecl in ,,outhrrn
-\frica. and appear~ 10 be absent only from the winter rdin· Disease 1ran~mission Th,•iil'rit1 n,11111/ata, the cause of
fall areas of tl,e Wes1ern Cape Provine<.'. the mountainou~ tropical theileriosis. in those rt1glons 1,1 which If. de1ri1wn is
areas of Lesotho and .KwaZulu-Natal where snow falls 111 the vecwr of T llmmlaca. tropical theilcriosis frequently
winter, and some humid. subtropical habitat~ along the occurs on small farms on which the Jive$tock are close!~
east coast and in the nonh-cast. It is also present in the associ:11cd wth barns and .tables where they acqwre infes·
drier regions of the sou1hem and north-western provinces tations wilh 1he tkk. It also 1ransmits Theileria equi. the
of Mozambique, sou1h-ea~rcrn and nor1h-western cause of equine piropla,mosis.
Botswana. central and northern Namibia. somhern An·
gala. mos1 of Zimbabwe. southern and weMern Zambia. Hy alomma dromedarii
centre! and north-ea,tcrn Tanzania. central and :,outhern The camel 11.iv,lomma
Kenya. western Burundi. eastern Uganda. and in Somalia.
Sudan. Ethiopia and in West ,\frica. 1 s,; 1 ·2 · 119- i:;~ 1-.; ~l•i. ldentificmion ·\dult II. dmmerlal'ii arc large ticks with
Parasitism of birds b~· the immature stages
206 240. 2a; Jo:i long mouthparn,. The $C\1tum of 1111: mak b characteriz11d
undoubtedly contributes to the extemive distribution of by posterior groo"e~ and ridges. rhe colour of these 1ick-~
th is species. varies frc>m ycllow-bro\\n to nearly blac:k. The legs are paler
than the ~curnm and ma~ be ringed by paler band'>.a,;
Disease transmission Ba/Je.i/a occulu111s. the cause of b1•-
nign bovine babesiosis: A11aplasnia ma,glnale, the cause of HosLS rhe preferred hosts arc camels, but cattle. sheep,
bo,<ine anaplasmosis: and the \irus causing Crimean-Congo goats and horse\ ma~ all.o bl! infested.87 :\dults attach 011 the
haemorrhagic fe\·cr. Secondary bacterial Infection oftlck a1- inner thighs, uddt.!r and scrotum of camels. The larval'.? and
tachment sites can lead 10 the forma1ion of abscesses in the nymphs feed on small burrowing animals and hares. but
cattle. the nrmph\ inn~ al5o ,nfost rnmeb. c:1 ttl11 and hor'\e~.

Control Control can be accomplished by localized acari· Life cycle a1,d distribution flyalomma dromedarii has a
cide treatmem. H.w,lomma 111argi11amm ruftrm is difficult twn-ora thrcc-htn:t lifc cych:. The larvae may feed and moult
Lo comrol 01, ca11le herds even by regular aearicide applica· to nymplb on small mammal or leporid ho~ts and the aduhs
tion as engorged n~1,1phs arc cominuouslr being brought in feed on large herbivores. Or the larvae may feed on small
from other areas by birds. mammal hos,s. drop off and moult Lo 1wmph~. which <'an

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18 ,;rr-r,o, tl'-•: Aspl><:ts inUuencing the oc,urrenc-e of infoctious dlsc2S<.>s

lhen either a11ach to other ~mall mammal hosts or feed on the Contro l The control of/. r11/1icu11d11s include~ acaricidal
»ame large animals as the adu.lt~.8'l The life cycle appears 10 trea1mem 01 ~heep and goats prior 10 the onset of Lhe
be cominuous throughou1 the year. 'l11is rick is common cooler momhs of the yt'ar. These animals should also be
\\'here,·er cameb occur in the l'ar, ~liddle and ,car Eabt. 11 b moved from moumain camps with a southern aspect 10
also present in ~Jamitania in \\'est Africa and in '.\lorocco. ,\1- other grnrinl( from lau.• summer 10 spring. The underbrush
geria. Tunisia and Libya in Nonh Africa and is well-adapted to of trees or shrubs, which give prorection 1101 only to the
an arid and e\'en desen en\'ironme111.:;3. 6b In nonh-ea:;ti-rn frce-lhing s1age!- of the tick but abo 10 the small mammal
and f:ast ,\fric.i It occurs in Sudan. Eritrea. northt'rn, eastern hosts of :he immature stage.~. can he remo\'cd." !-'2 :::·;
and southern Ethiopia. nonhern 1-:enya and 11onh-eas1ern This can be ac,omplishcd by prunmg. or fire. or by allow-
Uganda. 6<l. 1• 1 :?<Ji. w. ing goat~ to bro\\'w during the summer momhs. c:aule and
~hecp c.in be used before winter 10 kct•p 1be grass cm•er
Disease transmission Theileria <1111111/nta, the cause of shore. thereby creating conditions unsuitable for tick sur-
tropical theileriosi$. Vil'al or for questing for hosu,.n. 73• i,,
Fire can aho be used
for this purpO)C.
Txodes r11 bicu 11dus
Karoo-paralysis tick, Karoo-verl1;1mmingsboslub (·\fril...) Rhipicepha/11s appe11dic11lnrus
Brown ear tick. bruinoorh<)Sluis (t\lrik. )
ldemification J\dult /. rubicruuhi.s are sma.11. reddish·
brown. eyeless ticks with long mouthpan". I heir legs appt::ar Identification Adults or R. apJ)l!11tlic-11/<1111s are nwdi11m-
10 be grouped ameriorly {Figu res 1.13 and 1. 14). Male~ may si1.c(f brown tick~ \\ith shon mouth pans. 11w leg, of the males
sometimes be attached to females. increase rnarkc.odly in size from the first to the fourth pair and
engorged male,i have a $lender t'ltudal procc~~ (Figu~ 1.15
Hosts Sheep and goats arc the preferred dumestic hn~ts of and 1.16). lliese licks are frequently seer\ qucsiing for hoses on
the adults, on which they anach tu the ventral parts or the blades or grass during the early pun of th<.' 1,e1 st..>aSon.
body. neckline and upper hair of the leg;.5 · ~,:; \\"ild hosrs
include t·aracal and mountain r('cdbuck. iu; The preferred Hosts This species has a wide host range. Ad Lits parasltfze
hosts of the immature stages are rock elephant shrews and medium-sized 10 large ruminants whlle the immature
red rock rabbits (Pronolagus rupestrisi. 70· 1' 10• 1o.;. ;o;.1 siage,- foed on most ruminam species and a vnriet} of ocher
mammals. Including cquids. carnivores and hares.1™'· ~ea,
308
Life cycle l.xorii!s mbic1111d11$ has a three-host life cycle that Among domestic animah. cattle arc the preferred ho~ts
takes 1wo years to complete. Lan-ae are active In aummn and can become \'CT) ht.'ll\~ly parasitized \\ith all stages of
and wimcr. nymphs during wimer and spring of the same dcvclopmem.8 · t"H Several \\1ld ruminant spel'ie,. such as
year as the lan·ae, and the adults in late summer. autumn or impaJa. Mrican buffalo. eland, male nyala, bushbuck.
early wimer ofche following year."' 1o:;, z,· greater kudu nnd ~able anwlope can also be hea\~ly
infe,,wd.' 1 t.u . ·""1 F~male 11y11ln harbour fe\\ adult ricks bu1
Dis tribution This tick is associa1ed with the southern arl· good hosts of the immawre stages.~' 1\dultsauach 111 Lhe
slopes of hill~ or mountainous 1crrain and occur,. in Karoo highe~l number, to the cars of lhC'ir hosts but are al,o l'ound
and Karroid vegetation t~'Pes in the ~emi-arid interior of on other part> of the body.a. 1~·, The llTOponion that a11aches
Soulh Africa. 11 is present in the Western and ;\on hem Cape to the ears \'arie;; between host species and is also depend-
prO\inces. 1hrough much of the interior c,i the Eastern Cape. em on the a\'ailabili1~ of atrnC'hmem sites on 1he ear,;. 1.ar\'ae
in the central ru1d soulhern pans of the Free State, and in and nymphs may anach to the ears a~ well as 10 olher pans
southern and western Gauteng in the Heidelberi: and of the bod)• such as the head, legs, neck and dewlap."
Bronkhorsrspruil di~mcts.99· 112· in
Life cycle Rhlpi<"eµlra/11.< appe11dic11/11111s h a three-host
Disease tra nsmiss ion !;aroo paralr>is (toxicosis) is caused lick with a strictly sea,,onal, single annual lifo cycle in south
by the engorging females and occurs most commonly in late em :\fric:.. Adults are most abundam during the rainy period
summer, autumn a11d early winter when these ticks :tre in summer I December to Aprill. larvae in the late summer
presenL The colder the mean minimum atmospheric tem- and cool period afterthe rains (April to \ugust , and nymphs
perau,res during the lWO momhs preceding 1ick ac1i\'ity. rhe in 1hc winter :ind earl)•sprlng tlune m Octoher'i .8· ~, o, 1·H . ~07
earlier tMs activity commences. 68 Paralysed animals will The pancrn of seasonal occurrence is regulated by the unfed
usually recover if t11e anached adul:s are removed or killed adults. which enter diapnuse and do not cniage m hos1-
using rapid-acting a1..-aricides. but ~evere lo&se~ in sheep. ~<!11king umil the rains start.2n Jn c:oumrics c:l<>se w the
goats and even young calves can occur if prevemiv;; mea- equator more than one life cycle can be complewd rumuallr
sures are not taken. 112• 252 Wild animals such as springbok and no clear pancrn of season.al abundance ma) he
(/Uuidorcas m11rsupialis) can also be affected/" C\'ident. 11' · 15 "

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 Vector~~ Ticks 19

"

> Figure 1.13 lxodes rub;cunt111s male

Figure 1.14 lxoces ruoicundus fernale

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20 >1'CltO.~ o,;t: Aspects Influencing the occurrence of Infectious disoases

Figure 1.15 Rhipireph,>lus appendicu/atusmate

Figure 1.16 Rhip;cephal!JS append1cu/atus female

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 vec10rs: Ticks 2l

Distribution Rhipicepha/11s (1ppe11dic11lmm b an eastern. occur on hotses. and these may require treatment 1,ith
ce111ral and southern ,\frican tlck. 130· wa ils distribution ex- acaricides during the summer. Control on other domestic
tends from southern Sudan. Uganda. sou ch-western Kenyn. livescock species is unnecessary.
eastern Democratic Republic of the Congo. Rwanda and Regular acariddal treatment or cattle on a mb,ed cattle
Burundi 10 northern. north -eastern. central and sou1h- and game ranch in Central Province, Zambia. re$uhed in a
wcstt:nt T.u1:tw1ia. Fu1'lher ~uull1 fl b cunflnt:d to Ila, wetter significant rc<luc:llon in all ~tagltS uf dcvcloprnelll of R, up·
areas, which include the highlands ofMala\\1, Zambia, Zim- pendic11/a111s on sympatric impala and on the vegetation of
babwe and the Angonia and Chimoio provinces of Mozam- the ranch.JO!l Should infestarlon 1vith adult R. appe11dic11/a-
bique. The extent of its distribution in the coastal areas of 1us become a problem Orl wildlife in small reserves. control
Mo1ambique is unknown , It is also present in eastern can be achieved by various methods based on the principle
Botswana and ln Swaziland. ht South Africa it occurs in the or self-application of acaricidc.57• 2•19
Limpopo, l'\orth West, Gameng and ~lpumalanga prov-
inces, along the cast coast of KwaZulu-Xatal and nearly Rhif)icephalus d u ttoni
throughout the southern Eastern Cape Pro11nce. Angolan brown eanick
This tick survives best in woodland and woodland savan-
nah regions with good vegetat ion cover. It tends 10 disap- Identification, llfe cycle, hoses and distribution Rliipi·
pear if overgrazing occurs and it does not surviw• on open cephalus d1111011i is an ear tick that closely resembles R. np-
plains. Its distribution may vary with rainfall. It was inuo- p1mtliculams in appcarance.250 It is a three-host species of
duced into the south-eastern IOwl"eld o: Zimbabwe during which the adults parasitize panicularly domestic cattle and
the commencement of a wet cycle in 1973. and by 1982 it African buffalo. 5' · z;3 It occurs in the coastal and adjacent in-
was estimated that more thon J million ha of the lowvcld land regions of Angola. particularly in tl1e south. Its sourh·
was infested.181 The tick started to disappetir from this re· ern and northern distributions eitend to the north-western
gion towards the end of a dry cycle in 1983 and by 1985 it border of Namibia and the western coast of the Democratic
could no longer be found. 1116 In eastern Zambia. R. appen- Republic oi the Congo respecri1·ely.2AA
(iic:11/aws spread southwards between 1972 and 1982 and
then westwards during the 1980s and 1990,. 11 Disease transmis sion In '\ngola R. d1wo11i is considered
co be the vector of strains or Theii,,rin pnmt, causing Corti-
Disease transmission Rhipi cepha/11s appe11dic11/at11s is dor disease.3 1
thC! chief vector of strains of Theileria pari,a that cause East
Coast fel"er. orher strains of 7'. pari,a that cause Zimbabwe Rllipicephalus u,mbezie11sis
theileriosis: and yet others that cause Corridor disease. This Lowveld brown ear tick, laeveldse bruinoorbosluis 'Afrlk.J
tick also transmits Theiieria raurorragi. the cause of benign
bo\ine theileriosis: Ehrlichia bouis, the cause oi bovine ehr- Identification lthipicep/zn/11s za111bnie11sis and R. app1m-
Jichiosis: the ,•irus causing Nairobi (and Kisenye?) sheep dis- (fic11ln111s are morphologically very similar. The major d if-
ease: and when large numbers of ticks are pr~ent they may ference between 1he adults of 1he two species is that those
produce sufficient toxin 10 cause bro1,~t ear tick toxicosis. of R. u1mbezle11sis have a more conspic11ouslr punctate
Infection with T. raurotragi may cause severe or e,·en fatal scucum.291
disease in eland (see Chapter 33: Theileria taurotragi infec·
tion). Exotic Bos uwrus cattle sttffer serious production Hosts and Ufr cycle It;. host range. attachment sites. life
losses and can become secondarily infesrcd \\1th the larvae cycle and pattern of seasonal abundance are also simila r to
of the screw-worm nr if heavily Infested "~th adult ticks. In those of R. appell(lit:ulmus. 109 • 198· 2811 However, on impala
small wildlife parks situated in high rainfall regions large in- the vast majority of adult ticksatcach around the muzzle as
festations \,ith R. appe11dicular11s can constitute a serious opposed tO the ears. uo and a larger proponion of nymphs
problem irl several antelope species. 133 au ach to the lower legs of greater kudu than do those of
R. r1ppe11dicu/t1111s. 19"
Control Exotic cattle shou ld receive acaricide pro.tection
throughout the summer months. Zebu (Bo~ i11dic11s) cattle Distribution The distribution of R. zambe.ziensis differs
and indigenous Sang::i breeds become fairly re~istant to the from that of R. appendicu/arus. 130· inu It replaces the laner
=
tick a.nd so require fewer ac:ruicidc trcau:ncnt$. 193• 231. In tick in the hot , dryrhcr valley ~ystems of south-eastern Af·
the absence of thellerioses. adequate control of adults can rica (l'.g. Luangwa, Kafuc. Zambezi, Limpopo and Sabi ril .
usually be accomplished by localized application of acari- ersJ which separate rhe major highland areas. in contrast
cide,, to the ears or head. Larvae and n>mphs are consider- to R. appenafc11i<u11s. ic occur~ in the fairly dry em·iron-
abtr less damaging to caule than adults, and do not require ments of northern Na111ibia and in the lowland areas of the
treatment if adequate control or the adulcs is achieved. Mozambique interior. ll is also present in southern Angola,
Heavy infestations with adult R. f/pp!!11dic11/nru:s can also western and ~ouch -eastern Zambia. and in Tanzania north

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22 <1,~,..,",~'~ ,\spccts influencing 1he occurrence ofinfocllou, diseases

of Lake .:Slalawi.:!8l<The dis1ribmion, of I?. zt:1mbezie11sisa11cl than one generation per year. 14'1 All stages ma} be present
R. nµpe11tl/c11/n111$ overlap where 1here are gradual I ransi- on hosts 1hroughout the rear, but their abundance can Vtll')
tions between wet and dJy areas. This occurs in parts of the from season to season.
Ea.stern and Southern provinces of Zambia bordering the
Zambezi Valley. in eastern Botswana, ;md in the North Dis tribution Oftlw 60-odt.l llhipicephalt1s spp. prl.:S('nt in
\Ve,.r. Limpopo and Mpumalanga pro1•intes o( South sub-Saharan At'rica. R. evensi evercsi is.the mos{ widcsprcnd.
Africi,.9 ;;· no. 2118 In these situations some imerspecilic It is most common in the eastern half of the continent, from
hybridi?.atlon may occur.310 Rhipicep/1t1/11s :a1111/p;:Je11sis is Eritrea and Sudan in the north to South J\frica in the
absem from semi-deseri and desert area,.. south. 211& It mlerates a wide range of climatic conciiuons and
in southern o\frica the main factor limiting its di.stribution in
Disease tran sm ission Strains of Theiler/a pariY1 causing the west is increasing aridity, with the critical rainfall lel'el
Corridor disease: strains of Theiler/a pcm•a causing Zim- being abom 250 co 280 mm per annum. 112
babwe thelleriosis; 11,eilerin uwrotmgi. the cause of benign
bo1ine theileriosis: and Ehrlichitl b1:wis, the cause of h<wine Disease transmission ,111apias111a 111argin<1/e. 1he cause of
ehrlichiosis. Although R. .w111be;:ie11Jis can. 111 the labora- bovine anaplasmosis; 8c1besia Mhalli and Tl:eill!ria equi.
tory, 1ransntit $trains or T. pama as~ociated with Zimbabwe ihe cause of equine piropla~mosis: n1eilerit1 jeµamw, the
theilenosis, n 9 11 ,snot associated with outbreaks of !he dis- cause of ovine theilerlosis: Borrelia 1heileri. the cause of
ease in the field. 181 However, the presence of J; parm group borreliosis or spirochaetosis; and the toxin causing spring
antibodies in cattle in buffalo-free areas, in which the tick lamb paralysis. This tick can also experimentally tram,mit
occurs in southern Zimbabwe, suggests lhai it does transmit Habesia big"mfr/(/. the cause of bovine babesiosis.! 5 ln the
subdinical infection. 181 eastern higlweld regions of the Mpumalanga and Free
State pro1·inc:es in South Africa, the synchronom moul!ing
Control Rhipicephalus ;:am/1ezic11si$ differs from during spring of Ol"er-ll'intered nymph~ gives rise w large
R. appe1uiicular11s in tha1 irs populations ne\'er build up ro numbers 01 adults. 111 and these may infes1 newly born
numbers that could be expected ro caui.e :.ignificant pro- lambs. As the female::. engnrge !her secrete a roxin which
ducrion losses in ca1tle. For this reason there is li11le. if any. cause~ paralysl,."'' Locally this is known a~ spring lamb
need 10 direct specific comrol mea~urcs against this tick paralysis bC'cause of the season of its occurrence and the
species. other than to protect cattl<' from infecti<m with !he age of the animab alfected.
T. part,a group of org;inisms where contact with buffalo
occurs. Con trol Adult R. el'errsi evl!rt,iis easily t'Olllrotled by local·
ized application of acaricide to the peri-anal area. Usuall)
no specific comrol measures are taken againM the lmmawre
Rf1ipice11ha/11s evertsi evertsi stages. but these could be controlled by the topical applica-
Red-legged tid, rooipootbosluis (Afrik.l tion or acaricides 10 the inner surface of the ear., and to the
ear canal;. The danger of piroplasmosL~ in Thoroughbred
Identification t\dult R. e1,ertsi e1•ensi arc medium-si7.cd racehorses 01ten warrants the expense of 1remment, in
tlcks that are easil}' recognized by the dark-brown colour of " 'hich case both the immarnre and adul! ticks should be
their ht;avilv punctate scum, be:ad} eves and or:1111,;e w red controlled.
legs {Figures J. l7 and l.18).
Rhipicepha/11s evertsi mim eticus
Hosts The preferred hosts or the adulrs arc domestic and Namibian red-legged 1ick. \/amibicse rooipootho~luis
wild equid, l hot5es. donl!.e}'s, mules and zebras) although (Afrik.J
cattle. sheep. goats and a number or ancelope species. par-
ticularl)' eland, are also freqt1ently parasitized.~"· \I; ioo. HI Identification Rllfplcephalus e1,ertsi mimellrus looki; like
The adults. attach in the peri-anal and inner thigh regions R. ei,err.si e,,errsi. but has red- and ivory-coloured banrlcd
of their ungulnte hostsY Lan·,1e and nymphs feed deep in legs '>imilar 10 1ho~e of certain Hyalomma spp. However. the
the ex1ernal ear canals.a and are round most commonly on structureofits capitulum and itsshoner mout.hparts readily
the same u11guk1te hosts as the aduhs.. a, well as on distinguish i1 from the Hyalomma spp. 2 so. 29t'
bares, 109. 171
Hosts, li fe cycle, distrib ut ion and control Rliipiceplwl11s
Li[e cycle R/1il)iccpho/11s e1•errsi ewnsi hai. a two-host life e1•er1si mimer,i:u.< would seem 10 have the same host prefer-
cycle in which the larvae moult ro n 1111phs on the first host, ences, predilection attachment sites and lifo t.·yclc as R.
the nymphs engorge. de1ach. fall oil' and moult, and the e11er1si ewr,$;,2t13 The adults are mo~t numerous from :\'o-
adult~ attach 10 the second host. With the possible exception vcmber to :Slay. and the immature stages in February and
of some or the colder regions ihe tick can complete more ~larch and from \lay to September. t~ This tick occurs in

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 Vectors: Tick, 23

Figure 1.17 Rn1a•ceotte1Us averts, ewms, mate

Figure 1.18 'lh101cepflil/u, everw evg,rslfemale

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24 ,.-,:n," r ": Aspects inOuencinit the occurrence of inft·criou, disCl!St'S

western Bot~wa na, ,entrnJ and northern :-; amibia and wards. main!~ in 1he eaMern half of the continent. to 1'wa-
southern and \,'estern Angola.~1111- 2911 Its control Is the same 7ulu-:-Smal. South Africa.2~
as that of R. ei,ertsi e1,er1si.
Disease transmission Babesia ll'alltm(llmi {su~pected).
Disease transmission T//eilerin eq11i. the cause of equine the cause of porcine babesiosls. If R. /11111tlaw., indeed is a
pi, oplosn1o;;is; anu Tl1tti/e1 la ,epamctt. the cause oi ovine vector of IJ. 1ra11mum11i. infection would hove co be trans•
theileriosil.. mitted tran~ovariall) from thC/ aduhs of one genennicm to
the acluh, of the next generation because the immature
Rllipicepllalus bursa stages do not feed on suids but on rodents and hares . .-\n-
Identification Rhipi<:epllnlus bursa i, a large light-brown 01her POS>ibility would be the migration of infected adult
species. The scutum is covered With numerous fine puncia- ticks from one host co anotl,cr. This tick also produces a
rions. l n engorged males. the light tan body wall expands toxin causing paralysis in calves and lambs.
laterally and posteriorly beyond the darker ~cutal margins .
•
Rliipiceplmlus p11/cliell11s
Hosts All stages of development feed on cartle, sheep, Zebra tick. ycllo\,-backcd tick
goats and camels.:!88 .\dult ticks attach main!) on the inner
surface of lhe ears of sheep but can also a11ach on the dorsa l lden1ification Adult R. p11ld1el/11s are mecUum-su.ed to
outer surface. as well as in the perinea! and inguinal regions large ticks with a characteristic dark brown-a11d-h ory pat-
of these animals. 21111 tern on 1he ~cu tum of the male, while the ~cu tum of th<? fo.
male is predominan1ly ,..,ory-coloured with anterior dark
Life cycle Rhipu:ephalus bur.~<J has a cwo-host life cycle markings (l'igures I. I 9 and I .20).
1ha1 takes a year to compleu1. Adult ticks are most abW1dant
on sheep during summer in the northern hemisphere (June Hosts The adult~ prefer cattle. on which the} may occur in
and JulrJ. 2ffll large numbers. as well as camels. sheep arid goats as
hosts.ZI'~ CIB7 283 Thcy attach primaril) on the ear~ and the
Oislribullon The Medi1errane(l11, Adriatic and Aagean ba- underside ortJ1e body. including theches1, belly. gen!tal and
sins. and in Libya. Tu nisia. Algeria and Morocco 111 ~orth Ar- pcri-a nal areas. Their prererrcd wild ho~t~ are ,.ebra. black
rlca. Rhi11kep//al11s /J11rsa is also prese111 in some coumries rhinocero~. 11land and gl'msbok. The immature Slage~ feed
in southern Europe as well as in 1he '\ear Easi.2118 on these ~nimal~ as 1,ell :is on harc,.2fl, 2s- iau

Disease trruisnussion Bahesia bigt>mi11a, the cause of Life cycle and distribution Rhipicepilalus p11ichell11s h:.is a
bovine babesiosis or African redwa1er; Babesla cn/1111/i lhrec·hos1 life cycle. The adulls appear to be most active
(experimenmlly) and Tlleilerio equi, the cause of equine during Lill.' m,ny 5easo11.21>2 It is one of the most common
piroplasmosis: l:.hrlicl1i11 ot•inn experimentally\, the t-ause of ticks pre1'ent in the Horn of Africa. as well a~ in and tO the
ovine chrlichiosis: A11apl<1snu1 ovi;, the cause of o,ine east of chc Rifl \'alle) from Eritrea in 1he nonh to north-
anaplasmosis: Theiler/a separma. the cause of ovinc eastern Tanzania in the south. 280
theileriosis: and the transmission or a toxin causing paraly$i~
in sheep. Ois~se transmission rl1eileria ta11rorragi. the cause of
henign bo\'ine theileriosis; and the vims cau;ing ,airohi
Control Acaricide can be applied to the ears of sheep dur- ~hel'p disease.
ing !he summer months.
Control Strategic control of the adults can beach ieved by
Rhipicephalus /1mulat11s acaricidal trl'ntment during the rainy season.
ldentiflcatlon, hosts and life cycle R/Jipicepha/11s lw111/a-
11ts is a small dark-brown tick. Its preferred domestic hosts Rhipiceµha Ius simus
are cattle, shet!p. pigs and dog~. and its wild hosts are war- Glossy bro1, n tkk. blinkbruinbosluis (Af'rik.
thog. bushpig (Powmochoerus ~p.J and African buf-
falo..!l!ll. 289 On cattle it nttacl1es on !he legs. including 1hc Identification Adult R. simrrs arc large. dark-bro1,,1 ticks.
f~et. and in the tail ~witch l'J The hoStb of the immature! The ,c-u111m ha,; a glo,.,-y appearance and there arc fouf"
stages arc ccmain rodent species ,md hares.288 It is a three- longitudlnnl rows of Jargt) punc1acions on the scutum or rbe
host tick. and che adults appear to be most acti\·e during the male.
rainy sea~on. 29 · lS!l
Hosts TI1e pr('ferred domestic hosts oi the adults are L,mle.
Distribution [t has an extensi\·e distribu tion range. from horses and dogs. 8 " J08 The adults attach in the tail brush and
Senegal in the west to Somalia in the east and thence somh- around the feet of cattle and horsel. and around the head and

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 Vec10~ Ticks 25

Figure 1.19 Rhipicephalus pulche/lusmale

•
Figure 1.20 ilhipicephalus pu/chellus femare

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26 '·'·"', o": ,\,peels inl1uencln~ 1hc oc,·u,wnrc ofinfo,·1iou, di~easc<

neck ofdogs.8 IJ!I. !Ill! ~lonogas1ric wild animal~. ~uch as 1he wanhog.~0 - isH The hOMS of the immaturl' stage\ include
larger wild carnivores, zebra. rhinoceros. warthog and bush- hcdgehug•. shre\\'S, ge rhib. mu rid rode ms and hares. :?AA
pig. arc particuhirli•good hosts of1he aduhs. ~3 · 9• 1111 18:; The
immacure s1ages are parasitic on murid rodents. 1 ' 1~~. :rn, Life cycle and dis1ribution Rhi11iceplwl11$ 111rt111irns Is a
Adule abundance appears to be limited by lhe availabilit~· of three-hos, tick oi which the adults general!~ ari.: mos1 nu-
hos1s for the immamrc smgcs. 100· 16'1 merous during 1hc la1c rainy 10 early drv se;isons. 1 !I. 201
In sub-Saharan •\frica 1he majority of records originate
Life cycle and dis1ribu1ion Rhip1ceplmlus simm is a from Lhe l.'asu?rn pnns of 1heco111ine111 from Sudan. E1hiop1a
1hree-hos1 tick of which 1he aduhs are mo~1 abundam and Somalia in 1he north 10 South Africa in the ,outh. HO\\·
during the wam,. we1 ~ummer months. and the immmure ever. ,c\'cral collections have also been made in northern
stages during 1he cooler. dry autumn 10 ,pring seasons. w. Namibia. Thi, 1kk occurs 111 several of 1he .\leditcrranean
93 139
· · nn It i5 widely clistribmed in the moderate 10 high cuumrie,. c1;, well as in 1heir immediatc neighboun. ,md also
rainfall areas of Angola, Zambia. Z:mbabwe. :O.lalawi and in Russia. lr1dia and Paki,tan.2/ll<
)vlozambique in the north co South ~\1rica in the south.im
but rarelr occurs in large numher5. In the north ·e.i,1 of the Oise11se transmission 8r1besir1 tol1nlli and Tlwileri(I eq11i
continen1 it is replaced by Rhipicephalus praewxuum and (both suspected' the caust• of equine piropla~mo,is: and
In the north-west by Tl/1ipicephal11s m11hsammae. horh Bt1/Jes1t1 mw1mn1111/ rexperfmen1ally1. 1he cause of porcine
ricks that are very similar in appearance to R si11111s and babt-siosh.
,,ith which it ha~ been confused-in the pa,;t.~tll! Bcc:auw // t11rn11ic11s has frequemly hcs•n confused with
R. sn11guinl!11s. reports of the transmission of equine piro-
Disease transmission t\11apltJsnw mm·gmaw. the cause of plasrnosis b) R. .,a11guim-'11scould have referred to R. mm11i·
bovine anaplasmosis: J111nplas111n cemmle. lhc cause of bt•· 0·11s.';o Howe\er auernpts 10 transmit B. calJal/i and r. e,111i
njgn anaplasmosis: Babesifl 1rawmm111i. the cause of por· !O hors<'S iranssrndiully from nymphs 10 adults \\ith a Somh
cine babesiosis: and u tllXin cam,ing paraly~i~ in calves and African slrain of R. 11m111icu;, have been unsucce,~ful.-ZZ
lambs. fran,ovarlal mmsmi,~ion of /l. 1m111mmmi b~ adult
Although 1Tanss1adial 1ransmis,!on of.\. ma'1(inale w u. rw·c,111~·11~ seems the mosi likely possibility in 1hc field. ,:i.,
cattle b possible. 221 this hardly seem~ likely lO uccur in the because the Immature stages of this 1kk do r.Qt normally
field as the immature stages reed virtually esclusivel)' on ro- feed on pigs.
de ms. lmrasrndial transmission by infct·ted adult Licks. nnd
more panicularly males. wandering from one host to an· Rhipiceplu1lus spp.
other ~eems a more likely route of ir:fection.zzi Infection of llos1s, life cycle, distribu1ion and disease 1ransmission
suids wi1h B. 1m111ma1111i would sc.ern to occur by the f1fllie· rwo l?hipiceplw/11s spp .. mhcr L111111 those already men-
sin being passed transovariall~· from one generation of adult 1icmed. h'.lH· be<.'n implicawd as 1he cause of livestock
ticks ro 1he next generation of adult ticks. which then il:ed on paralysis toxico,is). On,· is /111iph'epl1alu, ,earlJur1011/
susceptible animnls. 00 (previous!} recordC'd as an undcscribed species of the
/1. f)rt11,11.fgroup <Jr as fl. 111111cmms). The aduhs of /1. war/Jur-
Control Except as a measure 10 prevem anapla,mosis in toni prefer dome~1ic gom~ a.~ host~ and the Immature ~tages
susceptible canle and occasional!v 10 pre,·em paralysis in rock clcpha111 shrews.'iv, ;o 1lowcver. all smges of develop·
cal,es and lambs. acaricidal control ol' thb \[)ecie$ i, prob- ml'nt may be encountered cm ~crub har~ and Cape hares
ably not justified. 185 !w/J1ts c,1µe11si,).qa On very young Angora goa1 kids adult
ticks attach to the head and ears. while 011 older go.us they
Rliipiceplw/11s turanicus nuach h!'re a$ well as on the neck and the briskei: 1 Rltipi·
ldentifi.c ation Rhipiceplmlus 1w·t111i.'11.< is a mudcrote· cephalus 1<•ari.111r1011i is a three-host species of which tht'
si1.ed. reddish-bto\\11 lick. which even under a swreoscopk adults.are most abunda111 on goats from September to Feb-
microscope is difficult to diffen::miate from the kennel 1ick rua~, and the larvae and nymphs un elepham shrews from
Rhipicepl1a/11s sa11g11i11eus. wilh which it has in 1he pa,1 been December to July and from t\pril to October respectively.'"'
confused. io1 The latter 1ick ,·inuall\" exclusivelr parasilizes ;u In :,omh Africa. i1 is found only in the Free 5tate and
domestic dogs.188 /1:onhern Cape pro\'lnces.= /\caricidecan be applied to 1hc
heads ;ind ear~ of u.•I'\· yotmg goar kjds to pre,·e111 paralrsi:<
Hosts The domestic animals from which adult It. 111ra11i The other tick is R. prae1exu1111s (previously re,orded as
cus have mos1 frequent!} been colk>cced in ,ub-Saharon Af- U. s/m11s1, \\'hose adult~ and immature \tages ha,·e a host
rica an~ caule. sheep and dogs. \\ilh no collections from pigs preference similar to that or R. sim11$.iu3 On ca11le the adults
and only a few collections from horses. Its favoured wild anach mainly to the cail brush and the feet. Rl1ipkephal11s
ho~t.S are 1he larger camivor~. hares and some of 1he larger pme11w1a111s i, a three-host tick. and 11~ adults are most
ground-feeding birds. wilh a few collections from :.:cbra and abunda11tduring the rain~ season.icN1This tick is "idespread

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 Vccrors: Ticks 2i

.,
~~ ~ '

,} ."'

.. -,
.'~ '""\
\ ,-'
'
·t "'
I

Figure 1.21 An adult !1ck of tl1e 0'1VihOrftXas

mo~ba;il comple~
II
'1

Figure 1.22 J.;\'ll,Cc ,

hecV) ,nfestat,:m o;
Boop/11/us deco,orin~s

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28 steno,'".: AspectS innuencing 1he occurrence of 111fec11011sdisea~C!S

in nonb-eastern Africa from Egypt to Tanzania. 208• 2811 i',ot withouc specific control measures against tlck.s and tick-
only can It cause paralysis. but Ii can also transmit the virus bome diseases. It is now known that the indigenous Sanga
causing l\:airobi sheep disease. 1!12 and Zebu breeds can develop resistance to ticks. 17 195• 231 •
232• 233, 2-:;i and will also acquire immunity to rick-borne dis·

Ornithodoros moubata complex eases if e.~posed 10 them at an early age. 176• ~ 4 Simarions in
Cyoless tampan which cattle acquire immunity to tick-borne diseases
through natural exposure as calves are called endemicall}
Identification lne C)•eless tampans belonging 10 this stable. 26 .\lost losses due to cick-borne diseases occur in en-
complex lack the sclerotiied dorsal scutum tl1a1 character- demically unstable situations where there are insufficient
izes the hard ticks. and their integument has a mammilated infected ticks to ensure that all calves receive challenge. En-
appearance (Figure 1.21). Male and female ticks can only be demic instability exists in areas that are only rnarginalirsuit-
differemiated with certainty under a stereoscopic micro· able for the survival of the tick vectors or where tick
scope. populations have been suppressed but not eradicated
through the use of acaricides. Tick-borne diseases also
Hosts and life cycle Ticks of rhe 0. moubam complex are cause losses if 1hey are introduced. 10gether with the.fr 1·ec-
parasites of warthog. and complere their life cycle in wart· 1ors. to new regions and spread through susceptible live-
hog burrows. The adult female tick lays a batch of eggs in a stock populations. or it suscep1ible a111ma1s, especlallr
sheltered localiry'in the burrow after each blood meal. After exotic breeds. are moved 10 endemic areas.
hatching the larvae do not feed. and they moull to the fir$!
nymph stage after one or rwo days. The first n~mph stage Historical perspective
seeks a warthog host on which it feeds briefly before moult- Control measures against ticks and tick-borne diseases were
ing 10 the second nymph stage in the burrow. This process is only instimted on a large scale fn southern Africa following
repeated for each of the four or five nymph stages, with lhe rhe introduction or East Coast fever {caused by s1rains of T.
la.st nymph stage moulting to an aduh. The feeding periods par11a) from East Africa. Prior 10 that, tick-borne diseas~
of the vllrious nymph stages and the adults are of short du- had not been reported as problematic In Indigenous cattle.
ration (approximately 30 minutes), and feeding takes place although losses caused by babe.siosis and heartwater had
when the warthog hosts are ar res1 in Lheir burrows. The been e.~perienced in imported exotic caule.128 One of the
nymphs can survive for two years 1\ithout feeding and the most serious tick-borne disease problems of the nine1eemh
adults even longer. cemury was the spread of heartwater through the coastal
areas of the Eastern Cape Province because of the spread of
OistTibution The distribution of ticks of the O. moubata the vector A. hebraeum, introduced by cattle which had
complex that parasitlie warthog follows that of their wart- been taken to KwaZulu-Natal for winter grazing.135 The
hog hosts. which still occur in large areas of southern Africa other was the occurrence ofbabesiosis in cattle imported for
(see Chapter 98: African swine fever). Ho\\·ever, a~ the den- rel.tocking following the rinderpest pandemic of 1896. 128
sity of human habitation increases, these areas are decreas- Eas1 Coast fever. which wa{; introduced to '.\lozam-
ing in size and in number. and wildlife reserves are blque. Zimbabwe and South Africa in 19~1/02. caused
becoming the foci of warthog distribution. Locally the ticks mortality of around 95 per cen1 in the herds of susceptible
are spread via their nymphal stages, which are commonly canle (indigenous and exotic) to whfch it spread. 1,;. It
found on warthog foraging outside their burrows.92• 9 l. 277 caused serious economic losses, affecting agriculture,
mining and commerce. all of which stlll used ox-drawn
Disease transmission The vin1s causing African swine transport. Considerable effort was therefore devoted 10 the
fever is spread from male to female ticks by means of the developmenL of control measures. Early attempts at vae-
spermatophorc during mating. 21 ~ IL is transmitted transsta· cinacion by the t'mincm microbiologist Robert Koch and
diall)' and tmnsovarially by the tlcks.21 S. 219 later by Sir Arnold Theiler were largely unsuccessful. How-
ever. once it was shown by Lounsbury that t11e disease was
Control \:o control measures need be taken against ticks transmitted b}' the brown ear tick, R. nppendic11/a111s, 137
In warthog burrows, but to prevent their spread warthog more successful control measures based on tick control,
that are to be moved should be sprayed with an acaricide quarantine proced1ires. pasture spel.ling. slaugh1er. and
p.rior to translocation fn order to ldll campans that may be dipping in arsenic solutions were instiwted. Dipping
pr<!Scm on cheu: hides. proved co be 1he mosr practical and effective measure. It
was widely adopted and later made compulsory. As a result
the disease was brought under control fairly rapidly and
Control of ticks and tick-borne diseases
then progressively eradicated. Comple1e eradicarion of
Cattle have been present in southern Africa for more than a East Coast fever from somhern Africa was considered to
thousand years.83 and until the cwendeth century survived have been achie\'ed oy 1960.

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 Afterthe eradication of East Coast fever two options were rendered the environmem less suitable for tick suni, al. fre-
optm to ~outhem ,\Mean countries. Tht'r could either con- quently resul.ed in the locati1.ed eradication 01 licks.1!i7
trol the remalr\ing major tick-borne disca,e:, (babeslosi~. However. this ,ituatlon came to an end in the mid-1 'liOs
~naplasmosis and heanwatcr) by vaccim11ion and move to- with the disruption oi dipping in these areas during the pre-
wards reduced tickconirol and endemic stability which had independence war. Large cpi(lemic~ of tick-borne diseases
pre\"lously existed. or they could continue 10 control the dis- oc,·urred and lo,se<. over a five-year period amounted to
ease.~ by iniensi\"e dipping. In South Africa c-ompulso1ydip- more than a million head. which was one-third of t:ic cattle
ping was first abolbhed on commercial caule fanns artc,I the owned by traditional fam1ers. 125· 170 Thesc losse, caused the
choice of comrol strategies was left ro individual farmers. In Zimbabwr vetcrtni111· authorities to re-assess the national
1hccommunal gra1.lng regions-0ft:he Eas1em Cape Pro\·incc poliC) on tick and tick-borne disease conuol. ,-J ..\her inde-
the management of the dipping service wa,. handed over to pendence in 1980, dipping was reintroduced in the trad.i-
the iormer homeland adminii.tration~ during the 1970s. Al- tionaJ fanning area, but wa,; not strictly enforced. From
though the service was maintained hy these administra- 1985 a similar approach was adopted for conunerc1al iann-
tions. enforcement of the dipping programme \\'3$ gradually ers, who were only required to ha,·e tick-free can le if the ani-
relaxed and abotlshcd after Independence of the Tran~k«i mals were to be mcl\'ed off their prnperties.
and Ciskei in 1975 and 1981 respectively. \\'hen these re- hucnSi\'C dipping i; still practised in Swaziland. but i<
gions were reincorporated imo South t\frica In 1994 the Pro- not a~ strictly enforced tb In the past. In otlicr ,ountrie:. in
vincial Departmems of .~gricullure assumed responsibility the region, including :'\amib1a. Botswaha and Angola. imen-
for the dipping senice and supplied both personnel and s,vc dipping is not 111del~ practised or enforced. IL has been
chemicals. Despite the relaxation of enforcement 98 per more extensi\·elru,ed in )!alawi. Zambia and Moz.arnbique
cell! oflivestock owners Interviewed continued to p;make in for the control or theileriosis and heamvater. but not
all dipping events. with disease control gi\•cn as chi' ma111 d1roughom each of these coumries.
reason ior panicipation. Ho Compulsory dipping has now :,.;o mo\·ement away from inumsive tick control can be
been abolished throughout South Africa. However, the made unles~ I! i~ pos~lblc to control tick-borne di,cases of
provinces of KwaZulu-i\'aial and i'>lpumalanga stlll provide cattle (and sometimes those of other lives rock) by immuni-
personnel and chemicals in the communal grazing area~ wtion. and then 10 manage tla: di=ses 10 ensure con·
and report good attendance at each dipping C\"elll. In :'\cmh tinuing, immunity. \!anngcmem mar involve regular immu-
West Pro,ince cattle owners utilizing a communal grazing nization or cr.lves or simply rhe main1cnance of endemic
area have been encouraged to parricipatc in a research and stability through tick exposure. The requirements for tick-
extension pmject dtlring which they were taught skills en- borne disease t:omrol are therefore reliable sources of vac-
abling them to make bcuer informed decisions on the level cines, and knowledge 01 the epidemiology of the diseases .. \t
of tick control they wi~hcd 10 achieve. 1~2 presem all rnccim:s against tick-borne diseases co main live
The dipping policy in South Africa hru. had mixed resul!s. organism$ and for thi~ reason either have a very short shelf-
Some progres~ive fanners took advantage of che availabilily life (unfrozen blood \"aCcinesJ or must be stored at \·ery low
of vaccine~ against babe;iosis, anaplasmosb and heamv:11er. temperatures and transported In liquid nitrogen or on dry
used them effectively. and were able to adopt Oex.ible dipping Ice (fro;r.en-blood vaccine~ or infected-tick stabltatesJ. ,·ac-
regimes. Others. for reasons that were probably only panially cines against anaplasmosis. babr_siosis and hmmwatcr are
under<tood. practised liule or nQ tick control and contained currently available from Onders1epoort Biological Products.
tick-borne diseases through the maintenance oicmlcmic $ta· South Africa. nnd can be supplied to most pan· of somhern
bllity. The majority. however. cominued to 1rcat cattle ar Africa.
regular intenals with acaric'ides. Few 11t·tually succeeded
in eracUcating ticks or tick-borne disease,;, H. 4B. 4 ~ and Recent r esearch
frequently 1he effect in endemic areas was w maimai.n insta· Debate in southern .\frica on the role of acaricide, for the
i>ility where stability would otherwise have cxhted. In th~e control or 1ick-borne diseases was most intense in :he early
circumsta.!'lces Josi.es due to tick-borne diseases commonly 1980s. By then there was considerable ovidenc<: from epide-
occurred if \'attines were not regularly used. miological studies that endemic stability was more wide-
In Zimbabwe compulsory dipping continued to be en- spread than pre\iousl) believed and that incen~i\'e dipping
forced after the l!radication or East Coa,;t ftver. Howe\'er. it frcq11Emtly comributt'd 10 pmblem~ ,,~th tick-borne di,-
should bti noted that Zimbabwe theilc:riosi~ caused by 1 1
case. ·'· ''' ' · ,a. "°· 1
7 81 1112 Lt wa.sor"ued bv. lllOJY\'jJeoplc r.hat
0
.
strains of T. par11a persisted and it was partly for the colllrol strict tick con1rol wa~ necessary 10 prevcm large production
of this disease that compulsory dipping was continued. For losses. bm no reali,;tit: 111formation on the losse, cau;;ed by Af.
20 years after the eradication of East Coast fe\'er. tick-borne rican ticks was available althous:h data existed on the effect of
diseases were cffccth ely com rolled in %imbabwe by dip- B. microp/11,f on the productivity of canle in J\w.trnlia.25~ 265
1>ing. Control was particularly good in the communal fann- In addition mo~ t1uanciunlve infonnation on the resistance
ing areas. where dipping. together with overgrat.ing. which o(diITercm breeds of cattle to Africa11 ticks was also required.

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It was in response to 1hese needs Lhac expcrimems ha\'e been Because much of Lhe information on the effect< of ticks
conducted in several African and ocher counrrie~. 011 1heir caule ho,ts has been obtained u,ing experimental
In Zimbabwe a projec1 to de1ermine the effect~ of.-\. //e- single sp~cies infe,1a1Jons. it is of llmiwd \'l!iue under field
bmewn infestation on live-weigh1 gain indicated that larvae conditions in .\frica since in most regions of rhe cominem
and nymphl> caused no de1ec1able lo~se~ \\'hereas adults animals are inleswd with sc-reral species ,imuit,meously. ~~
did. 1he loss per female 1ick complc:lng engorgement being The ,:,ffect> of rick~ arr also ofcen linked wlrh :hose of ric-k-
approxima1ely 10 g. 192 Estima1ed losses in li,·e,weigln gain borne disl'ases and should be considered in conjimnion
for A mrfoga111111 appear 10 be ton~iderahl~ more severe. with the Jamir. Furthormore. cattle herds contain animabof
amounting w 4fi co 61 g per engorged female.z• 'while 1hose different ages. ~e.xc,, and types, and experiment• w dewr-
for A le11itl11111 amounled co G g. 1M In the lmnreld of Zim· mine 1he eifec;s of tirks should be conducted on entire
babwe the cffec1s of infestation with adult ti. hebmeum on herds ra1her than on individual or selecr.:-0 age gr<>upi. of
the milk production of Sanga and Sanga , Brahman cows animals.·i
was measured over an l 1-week penod. Although the rela- Two projects supported b~· the Food and \griC'uhure
tionship between tick numbers and rcduc1ion in milk ~~eld Organi;,.a1ion 01 the Unired ='!ations have inw~tigated the
or calf live-weight gain during the obsen:acion period was economic impact ot ticks on cattle in za111bi11 wi1h
poor, provided teatS had no1 been dama~ed. an average loss rhe aim of obtaining data on the cost-effec1i\'eness of tlcJ..
in live-weight gain of approxima1ely6 g per calf for eve~ en- control in canlrt herd, maimained under managemem and
gorged female tick was recorded. 193 environmental c-0ndl11ons slmilar to 1hosc in 1he tradi-
On a mixed bushveld farm in M.pumalanga Province. ri.onal s(!ctor.' 2 ··o~ In rhe first the tick-free herd performed
South Africa. 1he weanitJg wl'ights of talYe~ or which the signilicantly beuer 1han the tkk-infcsted herd but the
dams were predominantly infe>1ed with B. dectJ/ornw.s. were value of additional production in the form of in<7reased
reduced by approximately 8 g for each engorged female milJ.. yield. calf gro1\1h race and fertilit) wa~ less than the
present on the cows.~ 11 In Australia mid-lactation Holstein- co~t of 1he acaricidf.' u,ed. In the second project greau•r
Frieslan eows were infested ,u weekly inter\'al~ wilh gradu- produc1hi1y wa~ recorded for a herd spt·ared \\ith an aca-
ally inneasing numbers of 8. microp/11; lar\'ae for a period of ricide once a \\eek in 1he wet season and a, cwo-wecklr
15 week.~. 117 By week 15 uninfestcd rontrol cows produced intcr\'ab in 1he d~ ,ea,on, compared to that of a herd
2,86 litres more milk and 0.1.:J kg more bu11erfa1 per day and sprayed e,·cry two weeks in 1he we1 sea,on 0111,, coc1obl•r 10
had gained 10.6 kg more live-weigh1 than rhe infested cows. ~larch) and a herd recci,~ng no treatment ac all. \\'hen the
ll was esrimatcd 1hni each engorging female tick was re- costs of com rol were taken inlO account, the \1'ct season
sponsible for a Joss of 8,9 ml in daily milk production a11d creaunent. which \1as designed to have 1he ::ire:ne...t impact
1,0 gin body-weight gain OYer rhe trial period on ..\. 1'nri('g11111111. but also reduced the number, or
In Zimbabwe larvae and nymphs of U. app1mrlic11l,1111s Hyalommn spp. and R. nµ11e11dicula111s. gaqi the bC$i result
caused no cle1ec1able losses in li\'e-weight g;lin ofSanga and followed by 1reauncnl throughout the year. Xcither 01 1he
European breed cattle. whereas adult licb caused losses latter acaricidal treatment regimes pre,·ented the tram·
which amounted 10 approximately 4 g for eve~ female 1ha1 ml$sicm of Ea;t Coas1 fe1er when this wa~ introduced Into
complt,tecl feeding. 19" Under field comlltions 1he \\'eigh1 the area towards the end of the trlal. 1~
losses caused by the adults were large in Bo.,· uwrus caul.e On 1hrce caHle ranches in Zimbabwe regular weekly
(aho111 21i kg nvPronP ~11mm<'r) M 1h1• ;111imJ1l~ h:icl lnw rPsl~- 1r<'~1m,•111 rrnvNI 10 ht> ernnnmknlly j11~1!ffahlr In 1ha1 11 re-
tance to the ticks. Losses in Sangr. caule which had high duced :I. hebmeum populations to imperceptible levels ruid
resistance, however, were small and frequently insignifi• elimmated monality due to heanwmer.21 On many farms
cam.1.,. Despite ~everc ear damage in some animals. infes- lhi, approach wnuld not be economlcall1 viablt• and farm-
tations avernging 20 <'ngorging female R. appendiculatusper ers should select the best control options iortheir partlcular
day had no significant effect on rhe milk }ield of 8. rrmrus circumstances.
dairy cows nor that of crossbred H. 1n11m.<i.. Sanga caule kepi Tick res!~tance in \'ari(1us breeds of caule has been com-
on higher and lower planes of nutri1ion. re,pecth•cly. under pared in several countries in sub-Sahnran .Vrica. These
commerdal management on 1he Zimbabwe high\·eld . 191 In· compari~ons have demonstrated that 5anga and Zt>bu cattle
fesmtion \\i1h adult R. appe11<iicu/a111sofSa11ga cowi, receiv- are con\iderably more resistant to the common I} occurring
ing nutritional supplemem on lhe Zimbabwe hlghveid 1ick species than arc• B. ta11r11s cattle. and that crossbred
o.ffccccd 1heir milk yield, a$ judged hr the weights of their cattle show intermediate resbtance. 194· ,.11. 1 !. 2u 208- 2~ 1
calws before and after suddlng. hm had no signlficanc effect Thus. as in ,\uqralin. the potential exists for th,· exploita1ion
on the live-weight gains or 1he cnlvo?s compared with those or gene1k resisiance 10 ticks in 1ick conirol programmes in
of tick-frei: cow:.. 196 Jn Zambia tick infestation,. comprising ~ub-Saharan \frica.
mainl~ :\. vatiegamm and R. oppell(/iollm11.•. on Sanga cows rwo m:1\' chemical$ for tl1e conrrol or 1ic:k~ hal'e recently
caused a significwn decrease In lhe amount oi milk been tesred on caule. The one is a bemm} I phenvl urea com-
produced. 203 pound. known a~ fluazuron, and is an acarine grow1h-

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 Vt.-ct<\r~: Tick, 31

regulator. mhibiling Lhe formation of chiLi11. 2.1 IL 1s admims- wasp release and compared to a control group of cattle 1;,
tered in a pour-on formulation. and female 8. microplm In South .-\fric.1 th<ll,e wasps mainly parasitize the n)mph\ of
presem on animals at the timC? of treaunC?nt laid onlr a few ticks feeding on hare,:10 !hey c,1n be found around the res1-
non-viable eggs. while immawre Licks died as cherwere un- ing place., of the hares and pnrasi1ize panicularly the
able 10 moult 10 t.he next stage. Protection after application n}mphs of Nyalomma ,pp.. .1o which br pr<'ference feed on
lasted several weeks.2 ' The effic;i,cv of 1111~ or related com- these anhnaJ, •ij. i o-1
pounds needs to be tcs1.ed against other tick ~pocies w en- The infe~ctil·e juveniles of nema1odc species that are
\Ure thciniabilhy in \frfca. pathogimic for in~ect< can alsCI he pathC1gcnic for tick!.. In
The ,econd C'Qmpound is a phenrlpyrazone agem Guadeloupe the infective ju,eniles of \1trious species of
known a, fipronil. This chemical ha~ excelltc!m t.hcrapeutic S1ei11er11e111a and l lererorha/)diris had no effot·t on an) of 1h;:,
and per$h,umt activity against flea, on both dog, and cat~. free-living stag(:~ of A 1•ririega111111 or 8. micn,r,/11$ original·
frials 1,11h pour-on formulations of fl pron ii 011 C'.Htle indi· ing from 1he i~land. whereas an Imported ,train of 8. a111111-
catc that it is eifec1h·c against both B00phil11s 111111ulmirs lm11$ was su,r:eptible 10 all or 1hem. ,s:¼ The mfecm·e
and B. microplus. 37 3$ Therapeutic and' persistent efficacy juveniles of two speci~ of S1ei11l'mema could kill tick~ of all
arc dose related and are Increased b~ repeat trea1mems. ~pecics tested within the genera . \mli(vomma. Di.•mu1rt>111or
and Rhipicepha/11s in die USA. 1J 3 I lowcvcr. it would appear
Biological control 1ha1, unlike in in,ec1~. 1he,e nernatodc?s arc unatile 10 muhi-
Although several agems are e!fec1ive or partially effective for plr in lsodid ticks. The ricks are killed hr bacteria of the gen•
the biological comrol of licks it is unllkel> that many or these era Xenorlwl1dus and Phororha/Jd11s that lia, c a >)inbimic
will be eommerciall) 1·iable. 1'e1·enheless. under uarural 3S$OClation \\ith the nematode,. and <'ut<'rthc 1id.swith the
conditions rhcse agents may play a ~ignllicant tole in reduc- Infective 1uveniles.;,.'l'l
ing tick numbers. Entomopathog.enic fungi have been used in ,evl'ral
Domestic chickens are opportunistic predators of 1!ck,. coumrie~ for the co111rol of agricultural pest~ and uidr u,e
The indigenous: ,·arieiics in particulnr. if allowed to ~cav- for the control oi ticks b currently being inve~1iga1ed at 111ri-
cnge arnoogs1 ca1tle. can consll me considerable numbers ous localit.ies. In !..en) a aqueous ron nula1ions of 1hc spores
or ticks, i.'l>pecially if the canle are penned close to dwell- oftl1e fun~i Ber1111 1eri<1 i>(l..<>'inna and Ml!Wrhi::.i11111a11isoplim!
ings in the late afternoQn and during rhe earl> moming. 55• induced a high momllirv and reduct."<! fecundit} and egg
M 6S Red· and yellow-billed oxpeckers .B11phag11se1y1hro- hatchabilit)' in R. appendiculmus focd ing on cartle. 11 a·n1ese
rhy11c/111s and Buphngm· rtfricm111sl. which arc virtually formulations also caused monalhies In all life stages of ·I.
obligatory predators of ixodid ticks. take large numbers of ,,arfeg11111111 and R. <1ppe11tlic11/111us in the l'egetarion. The
these parasites from both domestic cattle and from $CYeral compamti11e case with "·hich thC' :.pores of th,,se fungi can
wildlife species.- t J ?:.II The red·billed birds favour 8, be produced and anificially disst>mi11a1ed makes chem
tlecoloraws and R. appe11dic11/a111s as food i1;:,ms. ; promising potemial agents for the: control of ticks.
whereas yellow-billed oxpeckers prefer the latter rick and Various species of tropical pa,turc legume~ i11 the genu~
Amblyomma spp. 286 ,\s an aid to tick control on b()th wild .'icy/osn1111t,•.(can trap and kill larv,wof 8. mkropllts."2 ·z,, Z••
life and domestic cattle. red-billed oxpeckers ha\·e been I Iowever. more ticks an, llke[y ro sculc and oviposi1 closer to
reintroduced to several regions in which the, original!}' the larger rnssock,, of gra~s plams thon to tl11: ~terns ot the
nrcurrnd in South .\frira. hut ftom whkh thPy hatl in rPrP!ll lt•gume,;100 '>imilarly. qursting lan-:ic "'" mor,• llkrl~ tn :,,.
rimes disappeared. cend a grass tUssock thnn the stem of a legume imd thus high
Chalcid wa.~ps of the g!!nos l.todiplwgtr.s are obligatory legume to gra~ ratios are needed to t•nsure a useful l!.'vel of
parasitoid, of L~odid tick~ and most ,pecies \\ill 01iposit and tick comrol. A number of grasses ai~o have ami-tick
develop only in the nymphal stage of the tick.~ 10 Se\'eral effects."· r s·nic grasses .\Jelinis mimaljlora and Bmcl1i<1ri<l
wa,,p h1r1•ae can successful ly develop in a singl!.' engorged brizmuha both h,1,·c a lethal effect on the free-li1~ng lar1'a<'
nymph. which ls killed during this process.~, 111 Two of the of B. mi.-ro1,l11s, bm .,/. 111i11111ijlora also has a repellent ef-
senm described species of these wasps occur in Africa. fect, thus making ir likely that larvae will avoid it. 1
namely Jxodiphagus hookeriand lxocfiphag,1s 1heileme.lfl. •· • A combination of biological and acaricidal control has
ln Kenya adult/. hookeri were rel,ms,•d over a period of one been 5ucccssfully incprporate<i in ph<'romonc-acaricide-
year into a field in which there were ca11I~ naturally infested impreg11ated plastic tail tag detO}'~.J. !~O Thest> t:11!$ are more
wiih , l,. n1rii!g"r11m and R. (lpp,•nd1culm11s. is.• In ac!dilion the specificall) designed for the comm! of adult , \. ht>bmew11 and
field was ,eeded with nrmphs of both tick species. During ,1. v<1rll!gm11m, or which the feC'tling mate;. exude a phero-
the period of wasp release 51 per cem of A l't1riega111m mone that is ;>articularly attractive 10 otl1er males. females
nymph~ but no R. ((/Jpe11dic11la111.~ nymph, collected from and nymph~ of the ,amc ~pccie:.. !lie incorporation into sul·h
the cattle were infest<'d with the para,itoid. There w.is also a rag, of ~qualine. a naturally abundant mamm.ilinn sl.in secre-
significanr decrease In rhe nu mbers ofA 1/flriegntum Infest- tion that has a long-distance 1ick ~ttractant effect on Amb(l'·
ing the animals compared to the numbers of tick~ prior to om11U1 america111u11 may pro\idc dcfinile advuntages.300

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32 ,Kno, ,,.,.. Aspects inOu~ncinti the occurrence of infectious di$eases

Tick aJJd tick-borne disease models The discow~ of a major gene. which infers resisiance to
A practical means of designJng cos1-effecti\·e rick control R. microp/us. now presents new prospects for rick control in
smncgies is 10 use models.WI. ~ilZ An ecological mode,! was both beefand dairy artimals. This gene was found in a dosed
originally developed in Australia for the one-host Lick. R mi- herd of the Adaptour line of cattle 1hat result~ from the
croplus.Z•I 11 has subsequeml)1 been modified for 1hree-host crossing of Hereford and Shorthorn animals. brel'ds that are
ticks and given tile name T31 lQST. ts·• Ecologicnl parameter nom1all) hiihly su~ceptiblc to lnfest.11ion, llomozygous
\·aJues and the damage coefficient with respec-t 10 live-weight carriers of rhb gene ma)' acquire a ~table 101al or nearly total
gain for R. npvr11dicu/1w1s ha\'e been fitted to T3ll0ST,&1 and resis1ance to 8. mirrop/11s in early life.'"
it can now be u~ed to simulate population d~11amin and con- ~I any laboratories rhroughour the world are working on
trol strategics for this tick. These simulations allow the best the isolation and charac1eri1.a1ion of immunizing amigens
strategies 10 bt! idenrlfied prior 10 the C<>stly and time- present ln Br11Jesifl, Anap/asmn, £ibrlichi<1 and Theilerin
consuming process of field-testing. For example. il has been spp. 140· ;?n!J, 22• ft seems that it is onl)' a mailer of 1ime be-
estimated that the most cost-efl'ec1iv1; strateg}· for the control fore some of these antigens are produced in commercial
of R. appendic11lr1111..< on Zehu cattle at Lake Mcilwaine in the quantities. either by the application ofbioted111ology or by
highveld of Zimbabwe is intcn:,ivc dipping ror four months chemical synthesis. When these new genem1i1>n ,-accines
du.ring the wcr season (December to~ !ard11.sl The reason is are developed the)•Could revolutloniieour approach 10 the
that the adults. which cause losses in live-weight gain, are control of ticks and tick-borne diseases and hal'I, profound
most abundant at this time. Extension of the dipping period effects on callle managemem and production througlt-0u1
could result in increased dipping cos~ progressively exceed- the tropical and ~ub1ropical regions of the world
ing the \'alue of increased live-weight gain. Shortening the
dipping period could cause the \·alueofdecreased li,•e-Weight Integrated control
gain 10 progressively exceed the cost of dipping. Climate. In conclusion, the control of ticks on domestic animals lies
geostatistics and satellite imagery ha\·e been usc.>d as predic- for 1he presem in the integrated use of available technolo-
rh·e tooJ,. for the di~tTibution and for the sea.,onal abundance gies such as chemicals. rhc management of acaritide re-
ofanumbcrofeconomicallyimponant ticks."1 tit ur..22.s. 221 sisLance, ho,t rl·,istance. tick vaccines. and tick-home
As more data become available models will be refined and disease \'accine& as it s-eems unlike!) tha1 an) one of these is
will undoubtedly become increasingly imponan t tools for the robust enough 10 stand alonc.2 '15 The concept implies 1he
design of Lick control strategics. rational use of araricides in combinatio11 wirh other ~trate-
Knowledge of the epidemiology of tick-home diseases i~ gies to cosH!lfec11\·ely minimize production lo~,es cau~ed
steadily increasing and modelling is particularly useful in this by ticks and tick-bome diseases. Such a programme could
respect.32·:<.'l. :1u 5 Models have been dt'veloped for F n,mi11an- include any number of the rnmponcnts lls1ed below.
r/11m transmission and Its epidemiology. •I. l'l'l and for B. bo11/s
transmission. 1' 3 • tH. 2 ' 6 It should ultimately be possible to Strategic control . \caricide is applied during strategic
use epidemiological models to determine Lhe best strategics times of the year in order to comrol seasonal pt.'ak~ in tick
ror tbe comm! or ticl;-bome disease!' fn dilft-rent situations. abundance. It is aimed mainly at adult ticb 10 decrea~e
numbers to levels at which economic damage is less than
Future control methods 1he cost of control.
II i~ likrly that ronvi>mior~al li\'C \'l!rdni>~ 1, ill hP <urpa~~Pd
by a DC\\' generation of ~ 0n1he1ic produclS. It is now well es- Opportunistic control Acaricide Is applied 0111\· \\'hen lick
tablished thm cau le and 01her hostS can he immunized numbers are considered 10 exceed the economic thl'C\,ho!d.
against certain tick species h)' exposure 10 tkk antigens. a1td This slracegy is effective when animals posse>~ a high level of
considerable progress has been made in 1he characteriza- tmmU11ity 10 ticks or in regions where tiGl<s are on!>· sometimes
tion of these antigens.2-q.i ,\t pre.sen! the onJy tick against a problem because offavourable climatic condition~.
which 1·accines are a\·ailahle commercially is B. micropl.m.
thls field was first pioneered b)' the Australians who reali:ted Selective control Only those animals \\~lh rick burdens
that a ·concealed' antigen. which is located on the surface ol' considered 10 be above the economic threshold are treated.
the diges1 ct'lls that line the gut of 8. microplus. could be This is more easily achie\·ed wi1h ,mall herds of caulc and
u.sed to immunize animals against infestation with this should reduce both treatment costs and delar 1he appear-
tick. iw. 297 2'l<I Recombinant antigens deri\'cd from tnrn~- ance of acaricidc re~b1ancc. !ndi\•idual animals, which con
formed cells were then incorporated in a vaccine commer- tinuously harbour very high tick burdens, should be culled
cially kno"~' as TfckGard. 296• 2 ~n Vacclnalion crials against 8. as the) are like!\· ro pass this trait on 10 their offspring.
micropi/1s have now aJ,o be.:n conducted on cattle in Cuba
and Brazll \\ilh a commercially available recombinant anti- Alternation \caricides from different chemical groups or
gen preparation known as Ga"ac.23 ; .:.ia Both vaccines arP combination fonnula1ions of acaricides are applied airer,
also repor1ed to be highly effec1ive against 8. c1111111/mus.as6 nately, The inclu,,ion of acarine growth regulators In this

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 \'e<'tor.: Ticks 33

regime may funher help delay the de,·elopment of acarlcide Predators Pro11ded non-toxic acaricides are used, ox-
resistance. peckers can be reintroduced imo those regions in which
they originall~ occurred. Where catt le are penned close co
Managemenc of acaricide resistance When resistance is rural dwelllll8$ at night indigenous ch ickens can be encour-
already pTesenc. animals m us t not be treated \\ith an acari- aged co roosr close to the enclosures as this will facilira1e tick
cidc from the chemical group ro which resistance is e,idem prO!datlon.
before mo,ing them 10 clean or rested pasture. as this will
result in the new pas Lure becoming conraminated with re- Immunization against t icks Recombinam vacdnes are
sis1an1 tick~ only. An acaricide from a completely different commerciallr available for the comrol of 8. mirropl11s with
group should be u~ed in such an eve111. promising results against oll1er species. The likelihood exist$
1hat more concealed inmmni7.ing amigens will be discovered
Grilling strategies Avoid camp~ chat are heavily infested and that vaccine~ againbt a multitude of tick species will ensue.
with licks during certain seasons. Thus sheep and goats
must 1101 be graz.ed in camps on the som:hernnspecc of hills l mmwtlzation against tick-borne diseases Immuniza-
or mountains in the Karoo. Sou1hAfrica. during autumn and tion can currently be accomplished by using blood vaccines
,,~mer. The Karoo paralysis tick, /. r11bic1111([11s, is most comaining virulent or auenua ted organism, or using ln -
abundant In this ryp~ ofhabicm a nd is prcse111 only from late recced tick s,abilate$. These rather crude vaccines \\ill un-
summer to spring. Goars and canle can be grtl7.ed in sheep doubcedly be surpassed by recombinant amigens produced
camps during summer to denude shrubs and keep the grass in transformed cell~. or by chemically synthesiZ£>d amigens.
short . .lhus climinacing fa\'ourable breeding or questing sites
for the rick. Tick control on \\i ldlife
With iick species 1ha1 take a year lO complete thei r lif<' 11ck comrol on wildlifo in nature reserves encompassing
cycles, pastures can be rested from the end of the season of more than 20 000 ha should not be necessary. particular!\' if
adult rick abundance until 1he commenceme111 of the a sustainable number or large carnivores are pre~ent 10 re-
following year·s sea~on of adult abundance. Pro~ided no move stressed. weak. sick or injured animals. Problems \\ith
hosis suitable for the immatu re scages are allowed on m ticks usually oc-cur in small \\ildlife reserves or on game
the pasture~ during the imervening momhs, most or fam1s, e~pedally rhose under I ooo ha in size, on which it is
thew ticks should have d ie<.! an d the numbers of adulls not praclical also to keep the larger wild carnivores. Healthy
present when rhe lirnstock are reintroduced should be wild anrelopes and l'(Juids in Africa normally harbour tick
minimal. burdens in exce~ or ,everal thousand?'· 55 97 Howe,·er.
In much of southern Africa it is common practice to bum with the exception of large animals such a..~ glrnlfe. African
natural pastures once ever)' two o r three i•ears during spring buffalo and eland, which ma>· harbour large burdens of
to control shrub invasion and remove dead plant material. adult tick:.. the majoriry of ticks parasitic on wildlife are usu-
These <leliberare fire, coincid" with the pos1-wimer syn- ally larvae and nrmphs." qo, l0-1 Severe aduh tick infesta-
chronous hatching of 8. deco/ormus larvae and with their tions on wildlife are therefore frequently an indication of
ques1it1g on the vegetation and consequently many are de- some underl~ing management problem. Thi~ rr.igh1 he
strored. However, the short-term effect on lick abundance gross oversiocking.88 i:u. 184 or overstocking with particular
does 001 compensate for the damage inflicted on some species such as eland, which are large animals that are
vegetat ion t}'l)CS by regular burning. highly susceptible 10 rick infes1arion. 11l0, ni It could also be
A high stock density may initially favour the propaga- due to the introduc1lon of \\ildlife specles no1 Indigenous lO
tior. of licks because of t he .1bundance of hosts. However, a region and hence 1101 adapted to the local tick species, ln
should $tock number~ become 100 large the habhat may be the laner evem these animals may not only acquire enor-
damaged to such an extenr that it becomes unfavourable for mou tick burdens bm could al,o succumb 10 c!ise3$es such
1.he survival of some tick species. Human perrurbacion of the as hearrwater or one of tl1e theilerioses.
environmem by chopping down tre~ for firewood or Before control measures are instiluted rhe underl~'ing
removing them in order to esrablish pastures can have the cause of the problem should be addressed. One of the first
same effect. steps in this direction would be 1.0 remove any animals rha1
are vlsuallr,-ery heavily tick infesced as well a~ any diseast'<I
Host resistance Sanga or Zebu cattle can he farmed with nr wenk animals. The lattrr animals am 11nrmally <i>v('rely
minimal tick control, ranging from none in dry areas 10 stressed and consequent!) harbour large tick burdens chat
strategic wet-season control in higher rainfall areas, ca n lead 10 contamination of their surroundings Thereafter
whereas B. 1a11rus canle require intensive tick control fonhe sw cking density should be examined and corrected for each
greater pan or the year in all but d ry regions. The amoum of wildlife species in the reserve or on lhe game fa.rm. Wildlife
tomrol required for crossbred canle will be somewhere species that hi.$toricall~ did 1101 occur in the region could
between these two extremes. also be removed.

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34 •EC11<J>1 o,;r: ,\spects innucnclng the occurrence of infectious diseases

Pro1·ided the habitat is suitable, the introduction of spe- plied to animals by constructing a single entrance co a
cies such as blue wildebeest could be contemplated. These watering poim, thus forcing rhem ro step on a pressure-
animals have an innate resistance to ticks and remarkably few plate or interrupt a light beam that will acth·a,e a jetting or
adult ricks are found on the:m in their natural habiia1.!IO· 96 pour•on device.74 • is; Domestic cattle lhat ate regularly
The inrroduction of oxpeckers can also be considered. This rreated with an acarlcide. can also be used to conrrol ticks on
would only be feasible if the reserve or farm was locaied in a a small "ildlife reserve. These animals can be introduced in
suitable habitat and was large enough 10 accommodate the 1he wet summer months when adult ticks ate mos1 abun-
birds. or if the surrounding cattle farmers use only acaricides dant and can be herded during the day into those parts of
rhat are not toxic to the oxpeckers. Although fire may kill large the reserve where ricks are deemed to be most numerous.
numbers of free-living licks present on the vegetation al t11e None"" wildlife should be introduced on LO a wildlife reserve
time. its long-term effects seem to be limited. particularly if or a game farm without having first been thoroughly treated
animals are not kept av.,ay from the burnt area once there is a with a reliable. broad-spectrum acaricide. This will reduce
Ousb of newvegeration. 15~· 263 the risk of introducing ticks into a region in which they did
Should chemical tick control be' required. acaricide can not previously occur. as well as eliminating the s1ress of pre-
be administered by means of a selfamedicating applicator existing tick burdens on the new inlroducnions.
coupled to a bin containing licks or food concentrates to The regular tteaunent of domestic animals on mixed
which the animals are anracted. 57• 249 In South Africa several cattle and game farms can resull in a slgnlflcam reductlon In
wildlife species are more inclined to take supplementary tick numbers on the \\~ldlife on the farm. Those tick species
feed in such bins during the winter momhs because of the that use carde as hosts for all their developmemal stages. or
seasonal shonage of graziog. 88 Consequently mainly imma- chat prefer cattle as hosts for their adults are particularly
ture ticks will be killed. Acaricide ean be alllomatically ap- affected by this procedure. 102· 309

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C6re d'Ivoire. .ncltl Traptm, 2•1, 2111-405. lr.:mi.misslon orCcuWrtn ri,;,,lmmtlum b}' Amt,lyomma llebraellmc
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\he disease. Sl!l!l!t11ttniil Hepori ofth• 0/rr<tnr of \tt1,r/na,y Smtlcl!s twd 1-4 UttU1DL"(Jl0UT, J.D. I. MAUlllR.86. A.• 198-L Sweat[ng.skkness: a
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U 88.kt,,:\1 £:-.~. D,t.,, Cl!\':)f!}.:, O.:.f., CUAM, c .• }tA.i>Otft', M, If Jltv..~Pr. J,f\.A,, 1998. 1t11,rjiue;o Global Plm1~1il'C, /\llgusl 28-Scprembcr I 1991,. Kn.tgor
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12 RINA~. LE.W- 1920. SO\Hhcm Rhodesia. Rtportof1heChfefVetcrinary hrbro.eum and htanwotcr on thrc~ runches In Zllnbab\·1t. Zlmba/Jwe
Surgeon for rite Ywr 1919. Veterinary Jo:,mnl. 29, l li-131

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 Vectors: licks 35

:u BUG\',U:l. t.•• 19.ss,. La ·1nu?.tdie. de KiWn)•f. du mouton du~ a un viru~ on l.hc 1!:C"Onomtc 1mp:,ct of 1.ic.k., on SanR3 canle! in l.:tmbia.
filtrable t'I 1JOU'.l51lme pnrdos tiqucs. 8111/it1i11Agricol,d1, Con~'O IMg,,. 46. E.i:par11n~1/lnl 1111d Appl«·,!Al:m'(J/<>ro•, 21. 3 19,
115>-J.162. ,13 fJFT:,~ • .!,.L., ~OR\' ,1.. R.A.I., ,o;,-;ow. l .. [tETJft.. T.f., ;l,\,\H.\.'\' ._ \1. ~ f'UMJl,)(,t",
23 •uu.. M "~,1~n"u.. s .. O\'f'l(C'S"O. o." Jtr.s,1;. Ii.A.. i996. Suppression of M,J .• 1.996. The c.•p1dcmiofog)' o(heart\\;ncr; c..astobhshmcnt ,1nd
Hoopl,ilus microplu$ populations whh tluaY.uron- nn ac.arim.~ 1trowth mrunienance of cndr1111~ st~hlllly. />nrm:110/oro• 7ixl«J•, IZ, 4QZ-IO:I.
wguhuor .,tus!falia11 VrwrinmyJorm,ol, 'N, ~68-170. ,44 m>v(»..,\.f.. 1!!1'9 E.pidtndQlo.gy otndcanm)I of hU\'lnf.l l.)11b1.'!IIO'-i' in
24 lllllltUl>f~E.. ,u.. 1,es. llc:trt,,'r1wr inv.ttfo\. th(' C:iribbeMl. Pc,m,cl({tfl1J:O' South .\frk:rt.Jmminl o/rltt S01111i A[riro11 Veuri1t<ll)' .\.<1nri<11fon ;o.
:'odny 1. li>--l77 357-362.
,:; ,u.cuVt. <:. ,933. Th~lnf,-ctlon or,·a.rlou• tick ,~cl~ '-'ith Bol,;.,.s,o -15 n~, o!'s. A,1,, 19!1 RhiJ>ktpiUilt~tJP/1MtUruliu11s: rau-re nnd ,·l"'C1nr of
bif,'i'mfnn, tb uansmU.~on 1U1cl ldcuuiffcuion. Parn,itoloJ:> RCl·m.r-eh. 1.;, dlsea~, In Africa Jo11mol ofthd,011111 ,tfrk(m l"r1m,11r,;·,wmimfo11
324-3.~0. 52. 315-322.
26 CAUD\.\~ L, ... 19n~ Vaccination a1t3rnst bovine b.1bcsiosis. tr1: >11u..ut, q."' t:Vtk\· tt.• t98t hpfdemlulogy nod conaol of b,nint"
.ib oc \ O!i .•
LH •• PISO, A.I, I, Mo.tJL\'m', r.r... (cd.S), lmmumty JO Blood Pllffl.,(/(1'1 tifMm, bnbcsio.st~ in the ~1.11nl mltUa.mJ.s.. /1t: wmn!J1l!AD. c.,u. &. r.1l1""'· 1.11.
ar:tl.·\nfm<1lf. ~C\,.· York: PINturn Pt~!'. (cd,). Tkk B«i/o~· tmtl C:,,nuol Prneeedll~ of,m /memmlumi/
41 ~uc.,,. , ••1.., H~'Y. J.•P. AQ.•\.M, F, e, ,,oRn.• P.(." 1958 'nleTlcl".SOflll<t Clmftrtnc~. 2; 29 Jnnual} 1981. HhOdL"- Unl,"t'"h)', Cr,1ham~11>wn,
.Vorld (Mari,la. txodida/. w,m,11cla1,,r.. ~nocd Mages. lfoscs, $otuhAfri<"a. 1;~.
Dislriburion, Pnns.: Ors,om Nliuons. ,17 OE vos..l.J. Ac ,orr.1rrr.R. ..., .. 1983. The ~mJtt of dckt'.ontrol on the
~ CIJFFOltt>, c.>I., 11.UX, 1-1'. ~ IJOO<;>TJ•ML H•• 197&. S.asonal Md A.'J!lOn.ll epidemtoJo,",y of bo\inc b~bcsio~i,. o,ukrsr1,11)(J()tl Joum.JJI of l'rttruu:ry
abunda11ce of ticks <fxodid ..) on hare, fl..crfX)rid~el tn Kenr•-J()unw!of 1/1!5,arrh, 30. 3-~.
·11"1/ail En1omo/"I:)·, 13, 4()...1,. 18 Pt \'OS.A,J, & ,nus# 1.A,.1981 Obs('rv;n:ons:on 1hc m1n~mt~s.1nno(
2-9 0>1.Boa.,-r~ f., 199-S. Tht lifo C)!clt! of Rhipi~plttll11s lwwlntuJ ~<"Uma.nn. T1wlleria11m111n.$ an ~outh ,.\fnco. 011dtnr11poort Joumnl of \'ez,"'tinnt)
1907 (Acarina: lxodidae) undet lnbnrnH>r')' cc,ndilion..,. \,i1h noa-s on R,'i~t,rch, 48. HI.
IU>ccolog)' In Zimb~hl<C. E.xJ1'1rlm,•uwl tmtl 111111/11-d ,lr,am/ogy, I, ,19 p[ \\'Ml r,,r.. 1990, Tran~m-arlat ~TMffllission or /Jliheua ,wQO;t; by
JI ;-32S. ll)'<llomrna 1runcan11n. Oodc.,Nt{!J)OQ11),mmato/Vtti"rlno,.,, R,~rr11.s;.
30 coou:, •...,,.. 193.i. A SN1tth for tick parosltts In S<>u<h Afrle11. ~100
01tdentcpot1tt Journal of Vtufinn,y Scicnuond \nimttl Jntftr.tll)'. 3~ So Of \\'AM IJ.T.. t.dfal'.Z~JU!t\01.L\\L L.M. II roll;ilfil<P:. r t , t,S:: Tht
23-12. tr.Ut~,·arlul ua.n~mh,s10'1 of &lb,_t{(11rm11mnn,,J by Rh1pf«pltt,l1u
31 U,\ GM.C.\. 11,M, 4' S,ERR"-"'0· f',M,11.. 1971. Con1rlbUlft'.IO p.Lrlo e,;i\Jdo dn J.lntlJ.$'to domouc P•.S> 01fd1•~rtPQOrt /qumalo/\'ro.•rmary Rts1-Y1rr,1,
1h,·Jl~rtc.w dnccrin• m•hgna dos bo,1nos, em Angola. /\cw l'~wm<lria. 59,21S-22J
\"fll•IJ LlsbM. i. HI. 51 OE:\\ \.U. n.r.. • t1nm1nt--H. t-1 , 1987. The 1ran<s,,tdlal trJn.,tnisiJonoi
3z 0,\1.L\\Tr?- ,~ r . 1~7. Modellint: tlck·barnt" dl"t!illUS in tfc4 /n: lk,1be1fo eal,af!/b~ RJ1111fc«1fJltillu~ ,werr.,;f ~nm,d. Onffer.m·J,tXHl /mlft1t1! (Jj
,.1rn1111..,T, Jl.w•• {Cd ,), TicJ.-.,: ,mrl Tfc:J.:•lxmtc l)f,f«.SC$.. Procerdlr:1;10} <m Vt1tvrimll')' Rt,i:c1rth, 54, 6S5-6S6.
ln/mmrfo111JI ll'orkf/1011. Ii-21 Febru~ry 198(;. l'iy•tl&lJ· limbobwc,
so 1>1vto,u, 0.0.. 1\1:,. Th~ lnddonci, or1iclcsor floo1,M1<u,pocle-.011 c•nle.
Canbtno.: •...C.IAR. sh..,p 4nd goai< In '/fgorl~. Trop:rat ,\11imt1/ l/,a/1/i m11/ PIOdtu·lim:. -
l3 i)AU wn;r. \1.,., \'OUS(';. A,$ •• '\l<\J-fO~Cl'. o. h. &, $Ul uutn. ll.W.• 1g86. 35-39.
<'.o,np.ir:u"" cpidcmiologi of1lck-borne distil><" or tat<le "11h 53 n'ouvea\. rM \'A~ t>Ut ....·unr "·· JAC.()Utcr. P. '"Jl'lst.lJ.\.'~ J. • a>9-;
•mpha.<i, on modelllni:.Jm uow1tu. >1.1 .. red.I Prom.'lilng, oftltt$i.rth Det•cr!on of T/1~/J,rln a11nu/Q,a b) th• l'CR In ,tro \C•n: bod!dac
lmm,nrfona/ C,,11gr,-ssofPnra1ftology. c:~nbem, ·\ustnll•n .o\cadomi· of t<>llecicd rrom (';!tile in M,1url1•nlll. F.x11<•rimMtal nttd 1111p11,11
Schmet:. /\ooroloro·,21 27~291
:l4 o,UlJ~1:v. R.. 1930, ~atu.rat 1r;:m~mlssion orh~;inwntrt of ihecp b)'
&11 t:M'l\t/FM, ~.,1.. P~'C'V~ T.'\:." umui:. 1.1.,., 1988. Thr tfm.'Clopnwn1,1! ,-u(c:~).
AmM,w1mma varlegarum (Fnbtici11, 1794), Pams/101"1{>·, 22. 2ff0-.267.
ofAmbl;'Ommaltttbrik•um a.ndA.mbtyommtt mtll'mort·tmt on Lhl' leop:uJ.
:t.li OAVDM2\. R. ~ Huosos. ;.R., 1931. Nairobi .sheep disease.. Pmasuor~·~ 23, tonoiie. (;(><l(',)t•lo11ct p1.,,t1ul/J. Oudrrsti'J»ort Jour,wl ufVt•r.;rr,'.lJ.t'f
~7..52.; R1'fror.!1. 5S. 11-13
:,6 l'>"uusn. f', s ffUD50S, pt. 193,l, Nnlrobi "hl.!ep disease nntumJ nnd .SS OSU,\'E'ft, J:. ro•JRl(, LI, I. ~()).. O.J .. 1997. f'.r~lial!on uf llvbH>('k tit"k!t b~
t;1.--p..:nmcm~l tr.:msmi!sion b)· ti<'1'..: other thao Rf1!poCilpJ.alus thi(kcn:. ol, a ti1,;;.~ comrol m~lhad in o1 U!!Outte•poc,1 urb.tn
«11pmdlculn11tt. Paramology, 26, 496-50!\. erwironm1:nt. Oml~rJt1•1JO(Jff Journnl r,f\:tt'1imuv Rt':tt'1m·h. 6t 273-276.
:rr .OMI:.', R.B., ,\UftD;~, LIL. GfOk{.1,., JS;., UU~'TFR., f~\, I, J&........ ,t"-. P.. 1998 36 1>8£'ftR, J:. ,:011tut.. 1..1 & KOK. 0.1. 1.991;1. l tC'k dt\'cr~hy. "bund.incc and
Thorap,,utlc and persls1ent ullic.u:y uffiprontl ai:,,in<t B00pl1i/u, ~1::n.sonnJ dynum,~ 1n .1 r~~1urct•poor urbnn l!O\ironmcn1 1n I.bl• Ftt't:
mlrrop/w i,\curl: lxodldacJ on c:.1111~. Vcwl11nry Pnrruftol"I:)·. N, State Pro\inct'. Ondt•T$1r:1NJOtt fournal o/Vettri,100· R~$i•11rch, 6.r,.
161-?76. 30~316.
ss Q,WL\ R.1$., CLtit<GL f.E•. nu:m.Jt, ,.~. 61 Jl;A.NNJS, P•. 1999. fa11.lu.ulon o{ I\ S7 uu~('.As, 1.M.1 ~ms!($. °'··
1992. Tick comrol on ,~:ind 1aumltlJKfJJ ory•xl
pou,,on rnrmulntion or rip:onil ,i;,1inM Bnn11ltl/ut nmwlniu;; 1Aarl: and buffalo (S111«r«,< ,nffi•I'/ wnh tlume<hnn l'!l. pour•on <hro1gh ~
lxadida(·• und<tt na1uml Sou<h 'roxo,s field condil!ons. E.xperimcnwl ,m,/ Dwican App1btor.Jou..rnal ()/tf:e SQmt, A{rim11 Vi:·unruu,· VJoci11ttQr1,
Wplltd /\rorolo,!tl'.13. 351-36,l. 63.7-10
39 DA\11;.S. r..c:. * ,1wAJ:"1}1A.. F.. J98,?. QUJlhi::utvc !tud!~ of the tronsmls:stoo 511 m.nt. ·'" ., ..~.sru~. c;. t9ou. lx<idid tkb (Ac:,irino.1,odidoc uf c,niral
of X3irobi Yi~ep dl<c:isc ,;n,J by flh/piccphn/11.<npptr:diculatw Africa. Vol, I. C'renenll !nuoducuon. Ce.nus .•\mblyv11mu, Koch, IS.H.
(lllodo,dcll.. lxOdidM).Joumal o/Comp1ira1fc't Patltolol!,"!·, 92. l>-20 Ammtei dit ,\1!1..~ Hoy(II tit• I.Afriqur C,:nrrttlr. Trn111r(11. ~(gu11tl'
4u p,.-..,!!-, A.J., l9'86. Bil.illuij,4JJh~ ufllu: l~utliVh"t,1'\nl tH~mt::iupu:rJ Am1f1ft.1. s;,1~/u lJ Sc:frttc.'t'S.7A(J/QR1qu.,':,1, Xo. l-45,
<.:halc,doldea. Ellcynidar). p:,rasites <>I ticks (Mm.1,odid;icJ, "i:h note< 5~ rn11, .,, " ,,,.,-,.,,. Ci., 1961>. 1.,odfd tich tAc•rl110. l:<11dldoc• ul <:eru~
on 1be,r biolog)', Tijd,cllrlft tJ(>tJr E1110111ol0/II•· 129. 181-190. Africa. \'oL Ill Gtnu, fl/1iµlc,,pl1ulus Koch. 1844.,lrm<tl« 1/u Mose,
41 P.lf R\AR05.. ,. r. ,, ~ .,,.AMt, D.lf,, 1.ga9~ i\~o de grnminC"as forra,ge1r~!I, cm Royal de t. :-tt,;q,,._•Centrdl,· l't"n•uff'n, Re/.gtqt1(' -\nm,t~·. \int' m n ,
klr'\ih fnfesr3mes do campatu dos l>ovinos. 8oo1111i1,,~ m:cropliu. Scwnce-J 7.oQf(lg,qu«'J. Xo. u-.
Pts<1mso \-°c:>U·rirumn BrasiW~m. 9, 17-21.
,;o lMGi.. "·· 19~3. Die Ubeni•g<r d,-, Pfctdcplropla>mO',<•. illf• \hl•r1•llun0
4:/ DL C.U'TTlO. ,., .. JA1'>1t,:)..A,U,. MINJ,\1.m'. ti- 01 GIULIO, G.o,, .PE&.\US". :\ und 8!.ttlngte.. 4,rlurfur \t/l$n•1iJt:lr.aftllclw m:tl P'r11lm1<l11• (u~J,l,,/trmtl,..,
11~aA.,1, tu;, ettt7.YU~. H.(,,tl, a: Sf\-YM~G\\t, f'., 1997. Long·u:rm studk~ nl. 209-240.

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36 .reno~ n-,1: Aspect~ influcnc:lng 1M occurrence (If lnfoc1in11• diseases

GJ f<'tRhl>.\•PtS;\..... 19.99, Geot.t"t~n<"'S ~nd tt:ncu~ s..-ns1ng using ~C>AA Rl11plc.-plln/1<< ~,~rm ,., ..mi XeumM, 1897. during n:,plctJon .tnd
\\ I lRR satt"llitc imrtJtc.11· as prt.·tlicmc tnnl!\ in rick: d.Isuibmlttn :ind aipltbon 1Jw p.1tho•n11.'ChJnism ofthb to'<teo,u.. /11: \\1TITCHh..o\O, G.B. 11
hobtt~tsultablllty c~1lmo1lnn, for B<>opltllu,microp/11.< 1\rurl: h<><lid,teJ ~IJJSO,. J.tl. tt.l!'i 'f/r1· 8fofo10•1md Omttol. Proct't.'tilngiof an
In Soulh ·\mcrica. \ t·u•rlnftry Pt1rnsttolog)·.8l. 73-32. lntt•~mutonnl Cmt{t.•tt'IUt'. 2'i-2.9 lanuacy 1981 Rhod~ Unlwr,lty
62 rat..~,\~Ol;t.~RU\'AlCAO:I\, M., Cf!U1.·\'1\ZQUO, C•• SOlASO•'t'fRCAM, J. £i GrahoirJ.101,i,. "<nuh Mric;1, r,.13.
,~HO.\•\.'At.QUtil, z.., 1999. Anti•tkk cff<.-"<'1..(j iJ( s:ylo~'flmllt.• lwmlll., ~rat 8() (')Ol lU~. R..,. BlilUUJf ',IIOUT, t.r)., 1986. StUt.fkt.l nn lht :tb1ht~ Of ditft'rtmC
St) l<IMlntlir1 ho.1,m.(<1 on plou t."Xpcthnwualty lnf~cd ,,,th &x,pl1ilu$ )traJns or ?OJ>U111uo11-.. or fcmdle R;,;pif~{l1mtu~ ~,,•rt.sf M \'Yl~i Ac..irin..
m:cn,pluslar'\'ac in ~,1orc:l.os. Mc.•xko . E.r11c.•rw:,w:a/ and .o\;)plltd t.xodida.e· to produce pnrn!)~s in ~htkp. Om.f1·r.<tt'ptJ<Ul Juw·,111/ of
.,.,,,rntog, 23. 171-175. Vt1t•rt1Ulf}' f«'lt'Orrh, 53. 19-2·L

63 Q.G,lL l9o3, A ncw,,rn~.rn tick p.;ua.>Jte, lluntff~llus tht~lc.•rm•

f'1l1DU.1t, 81 G.Ri\Y, }.~. f.. Of\·(')~. ,.,.. t~81 Stud le~ on n bo\:fne 8nl1t.<>sfa m1n.,mincd b)
~p.nov. Ontlcr::;1.:poo11 /(m111altif\fcrcnnaf),· RQearcJ: 26, 61-Q$ • plme. ii>t1lomma t1iari:111,m,m rr,JipesKoch. 18·14, Ondr.rsrrf)(Y)r: Jot,mal oJ
6.1 Fl.0\1), ft.a.. ~tntU:R~r. R.\\. la \1;\\\\'ALI), t.,,f l-98';' \fodctUJn~ lid. \'t'Tt'rinmy R~'.Jt.·o,d,. 48. llS--223.
manJgt•maL Ill. Dc,li;ulni; 00111ml ,1ro1<1gie<for Rl:rpltepl1nlu, 8.2. 1.u.\la •u, • 4.,01 m 11., 1978. lnflu1.1nc~ of inf1;.•\Mticm r.ite on tick
rtp1>~mdlculnmsust.ng1311os-r. In: '\1m1~1 fl.\\1., (ed.. 1/rkluml parnl~~i,. In ,hcep tndu<•'<I b)' /!11/11/0,11/111/1,s ""'"' ,,,rr;sl '-t•umann,
n1·k·L'Om~ DJ.w,u<S. PrrX,..,<Yll11&,ofm1 lntmrt11/on11/ l\ork$IIOJI, 17-21 1897 r,urhwr,-·Ptmt.dt(l/u1:,."1,.; 183-191.
Frbrual) 1986. :-:yanga. Zlr11bal>wo. Canberra, AC!AH 1937 The htstof)· and dt.'\ctopmem of iht.~ thtl!t! indi~~nou,.
t3 U,\'ft\'t:\" R.i.:•.
/;; >OUfttf L.J. i.. un,w;, 1.G,. 19u,. Ttek-induml pJrttl),iS of !he springbok c~lll• br.cd!< 1>f Z1nihah1w. In• r.M.,T. 1., (('d I f>ro.·et..trr,g, ofu
S<1111hA[ricar1 Jounial11/11·r1dlif.· H-""· 11.
t:ll-ll3 U"ork.~11op<ll ,\1(1t(IJHJ( ll~/.t."111d1 S;nrl1J11, 26-2i h•bn,..U) l t•8f.
66 fOUSUF LJ. ~ HOR.U:. 1.G.. 1091, Tht Si."a\or,3J ,u:ti,1~·oi .ldult 1..xod1d t1tks
7.imbabwe
on .\ngor:t g4~b fn tht.suulh \\·~tern Otting, Free S1~tt,, Jour,ta/ u/ tlit' B.: JWS,\.'., .... \l,, DtPltllU o.a .. \.\:C>O• .\..0•• 0llHt\',1110, I Ii. l9~l. Pr~d.tUO!'I
South Africn.11 \'Nerinm,•,-\s..totltttflm. 62. lOt"-106. on ll\·estock ud,., b\' chlck~ns.. l'l•l4'rinti?· Pa.rt1.filO!o~,. 38. 199-204.
67 The hfo cyrfo of lx<>dt:> rublrundus ,\rnri:
fOUA1£, LJ, & 1TOk-\J;. 1.c:;.. 199.1 8,; HASS\.:,; , \I,. DWtou,. on • '-1U,-.;\•1x1, Q',M 19tl~ lnfhwnr~ of 1.!xposure
IJCodidne! ru,d iis adap1ation IQ a ho1 dry emi,onrnent. E.<p,11111,mal penod and m;magl.'ll1cni mmhods on the rm.,:t.voncss or chickens 0<
m11f Ar,11/ied Acaro/011>·, 18, 23-35 pn!<f:nort- of uck., fnf~tmg can1<?. Vt:t.trf,ia.ry Pnmsuology,.43. 301-~09
68 J<>Uftl>, 1,1 .. 1<0R.'ll-, 1.c;. ., ~o,. 1>.J.. 199ij, ~patlal and 1empo1~l vnlf:ulon, i6 1100<,STl\.\.\t.. 11 .. 1s;6 .-!f,tcan lxodo(dM. I ol. /, Tl<b of1ir, Sur/1111 [IV/rh
ln the commencet11l•nt nfwa...,mal acthit·y in L'1e 1'aroo para.lyr,.b t1C'k. spocialrtffltrf1.C'•" rq f.qttllloria Prorlnc.-nntl rd1h p,,..llminnry ,,.,,ft11-so/
/zod,,s rublettrulu,. 0,ul,mtp,HJrl /011m<1/of\'e1rrl1wry H,,.• .,,r/,,
&.,. 1he i:w~rrra Uoophdu,. ~farg_aropus muf Hr,1tomm:1. Re~:irch .Repon
~05--308. X:0.1003050. 2s.o- Ocp.1nmcni of the Na:\"y, Uurcau of \1l"Chcme and
69 fOUJIIII;, LJ., HOR.~ 1.c &, M.\.RAh...... 1988. °l.n undC!\CT'fb.cd RJliptetp/tnhu Su.rg4'r.. \\"3mlng1nn O.C
,pecie-:Mocia1ed 1,i1h R,ld poral)..,i< or ,lnE,(lr• &~"· Joumol of1/re 87 tt<>l><;STM" H .., .,",u' >1 s .. 19;6. Tho 11tk, 1,nd<1id<a of Ei:;'P•·
~0111l1Afr/m11 l'r,,rin(lry,l,Gt,c/111/011. :;9, l7-l9. /oumal ofrlu: fg_,ptton Pu/11/c fll/irllh -'ls.,octni/Qn, 33. .; 1-85.
70 ~nu,ul!. L.),, HOAA>:. I.C. ~ VA~ on: IICTVtill. J.j .. 1992. Tht• rel..itiVl· host 88 IIOM~ :.<... 1980. The C(ltllnll ol para<lt<-. In ~nh·lo1>c in ,111:,U game
sr:1tu4; of rock eltph3nt Shh:ws El,•p11rmculll.>t myuru.3 and NtuU;!qun rock n·!l,Cl\~. /bu,,,nltJ/'1lt1Sou1hA[rfca11 Ve1c1iut11)•Ass,Jt."Jntum• .i~, 17-t9.
mice AtrJrom)'t n<mr(JJ/11t11t$Jt ft)r uconumknllr imporutnt tickti Small
89 t1nKAt<:, !.G.. l'JtlJ. Plma~tl"S nf domestic :-md \,·Ud ttnlm:al, fn SUuL"t AiriCl,
·Vr/Cb11 Joumnlo/Zbol&io·,27. 108-1 H.
X'\'. Thl' """;,,.~nal prt"\~encc o{ ~:·ctop;,:m~itL~ cu1 fmp,llti and artle In rhe
it f'OURIE. 1.1 .. IIOAAK) l,G. I. VA~ z:n.. J.M •. 1991 Siu~so(:m~chm,·01 :ind non.ham lr~n,,·.Ull. Ottdtrsh.'(XHJrt /()Ufl'uH 0/l',·1,•rfomJ Rrs4.'tlrtir, ,19,
tmra,,.'p(Kltic inlcnauon dco~1llt"~ tti the brown p~~ uck 85-!13.
!<Jupl«J11ullu$ pw1ttat11Jt on Ant;ora g()81$.. E.x.JRrim~•mal un,t .1p;'1irt.l
90 HOR.\~. 1 G 1998. The rtlanons..l-iipsbem~cn uc,ks. ho1t~.ind the
.-lcnro!og;;. 12. 243-z,.;9.
cnvr(onmtnt In 1h(• Kruxcr ~3donaf Par}.... In r.oos:,.. t.. • KOtK~uu,. M.
;2 roua•t. 1.1•• to>:, o., .. non.,.-. t,G. • ,·.,, zn... , ..,,,.. 1995, An e\'nluatfon of 1l'.d!.\ ·n1e s«ond ln:t1mmio11al Co11f11t-untc D11 ncf.:.·lxJ"-1.' PmhfJJ!,..11H. o:

,1rat~1~ and threshold e<1111rol mcu.,u,-,s a~:l.fn.st th•"~'"° p~roly,,s the 1/o.\:.Vt't't"r Jm,•tf<Utr: n G!ohol Pcts/N!Ctltv. ,\u.,1~1 ~8...September
uc:k. J;,;od('$. rubicmutu,1i (Acuri: ixodfdae), m Soul.h Africq. F..tJH!r;nwnwl 1 J995. Kruger Xodonlll r•ark. South Africa: p~t'Cdlngs•nd ob>traCLs. 2,
and ,;pp/Ir,/-ium,logy, 19, 14?•153. 113--126
OJ .. li:'RU(;H.. L. $!\..,"\11\!\'. ~. « \'.\.~ OHtu;o.;cl,. F, l'9~8.
i1 11)1Jklf, l.J ....v.:. 91 IIOH,o:.. I.(,., \~,ll0M!i1\f'"i', M•• KRECE>.., It.( I. D()(.))l .. f.l:11 J.. 199:?.. Arthropod
Conrrol ofJ;aroo pam!ysh 1lck, through •"$•:ationmunogome111 In , pords!I"' of ',jlnngbok g(•insbok.~udu\. i:,mff<:, 3nd Burch,11'• and
cool\:s. L ,. "ornsa111J>. ,,. ,~l,}. nr,~Sttond lmefnnt10,mlCnnfMt11ei H.:irun:J.nn·sn•b:.a., in th~ E.cos.~, and tlttrdup ~::i1ur1.· Re\f!n~. ~amibia..
011 Tl<k·bom, Pa1//ogms Ill 1/1~ flcm-V,,..,,,.. lntcrfao.~ n Clo/xJI Ond,•rg~/~KJfl tnun:11} o/Vcrtrlnary Rt.S.e<J!C1' $9, ?*2Si.
P..apfcti1-.,, Augu>128-Sepiember I I 995, l(ruser :-.aw,>ool Parl., ~u,h
9'2 HOR.\K. :.c; .. 81(.(.i), H.c lfA'lo6....q.,~. T.:S. & H,\;\.S'of'.\, lU,, l9&3 The
Mnca: ptoo,<.'ding., ;ind abmac15. I, p.19.
p1e\.1Jenct.• or ht!'lmmth and arthropod para"-m:.") or wart:iog.
:'4 fOUJUf. t.J,. \1VfE.R. J.~. UORS'.\V.:S, .... 1'0).. r,.1, 19.98. Auton,a1ic Pharodr.0t.·n~t. <lt{lmmi<cts., In South \Ve.~l Afnc:u/Xi1nub1;s. Onder~u~poort
.tpplfc-ator (or pout•(U\ ct>:np()und-.. hr: t.OO~i'i, t. ~ 1U)Tii">Hl11>1 ,,. l"'h} Jo:irn«I of\ ',•1trinMJ' R~$1Vm:h, 50. MS--148.
The Srcond lmrnm1to11at Q:mf,·rl!m-eon Tkk-born" Pnrlms~1m m ihtt
93 m.1K.\~. u, .. U(.)r.)MKC'R1 r. 1>l \'OS. v.• N)·1·<,1 1~1 M... r .. !'81. Para:,hes of
J/~;. \·,..,,u, lnt,rft1c,: 11 (;/11/x1/ p,,,..,,....ti,.-.Augu,1 2&-!><:pi.mher I 1995.
dom.,_11< ond wl!J animals Ill $ou,h tlfrlca. X)(JII, l!elmln<h Dnd
Kruger :-:a1ional Park, :iouth ,\lrlca: pro:><c<dlng, a1ld ab«rncll, I p,57.
arthr1>1>0d p~ta.l1e,,ofwnrthog,. Plrn=l1wru1ac1hlop,crr,<, ,n !he
~ Ff\lt»uc,n• .:.1., 1988. ·1 rnn\J'Jli\.\Jm, nf RabrAin. /n; 1:ucnc . .\t,. ,·t(t,), eastern lr.ln~,·.;ud l..ow'+'eld. 011df!rs11.•poort Jrmmal r(l'n.•rmr,,:.·
&1l1"$f0$1$ in l>oJ111!1rlr A>1/ma/., (ltUI \1011. Bll<:;l Raion. Fis CRC l'f<'"· Rt'Jtmf\~,,I 53.1.;s-1~.
23-:'>2.
'M H()K.U.:. 1,G•• 1JOOMJ.1Jt, f," t l.\~v.su. t.tu, .. 199;j. P3rn1,i1~ a< dometttir
:-6 rn1so1. J.F...1899. Towatd.s:. pl'ml3nr,nt ~Im.Ion (orcontrolll~t\\ttle W1d "ilJ an1m•b ,n Sou1h.:1(rtcn. XX.XIV \nhmpod 1>'1t>•it11> ofnyo:I,.,
Ucks. J11ttmolfa11ol ftJurnal /t,r Para.<f:olo;J·. 29. Sii-i I in nort.h~c.i.... tent Kw,l.Zulu-N:ual. OndcwJrJl()()rt Journal o[\ c,1M1tta,,.·
rr GAU JV,\..~. c.J. & H()J\I\);, u•.• 1~7 1:k,dy ~ and habitat ~dercmunanr, Rm:-arr'1, f,2.171-179.
of Lickrntcstations of wlld un5;.\tln1es 10 South A.inc;) Saurh .-tftu:flfl 95 HOR:U:,. I (.,, ll()(>.-.1J\1.N, J, ,PICK:ETI, ,\.M, N OU \IO,, v., 199!! Pllra\itC'o of
Joumnl o/Wlldlifr RC$((lrc/l, 2i, 6:>-70. dom1.•,nc: •nd wild unim.tb In SoutJ1 Afrk;L :o;x. Ec:opara-.iu.ot.: o(kudu.,.
;a ccnHL , 1967, nru In 1h• !illuth African 7.oologlC.\I ~='*
Collc-c1ion.
XII-Th; genm Booplrllus Cumrc, 1891 Wld ,HtJrgorop:i, !:&1>Ch. 1879
In 1he c:><1trn Trnn"·""I l.o\\~~ld and 1h1• cas1crn Cupe Pro'1nce.
Onder>Jt'JJ()t(JN /oumnl tJf\!eterlnctry Rd~1trth. S..9. 259-2~
Ond,.,srqx,on /0111110/ of \f,•1trl11my RCH,ir<h. 3-1. 81-101>. gt, H08..\1', u;., 1,,
,·o'.'11, v... bRQ.\,•,·. ,,.k., 19-83. P.,r.t,iht\ ot d'>mt~tic and ~,·ild
-;,1 GOTIIE, tt- 1981 Tho 10,lc ph1t.«' of p,ral,-.ls Inducing fomalo animal, in ~nuth A!rl<a :\'\1 Hclmln~rnm!nnhropod p.,ra<llos blue or

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 Wctors: Tick< 37

1nd bfa.c;:,\Udtbt$~t 1.COnmxltnNes lOllrlm.,s and ComuxtraNn ;,.r110t1 btology. J)eJN,rmzr,u t>j 'V(r:cu/Jural T1•dwirul Sun:irt:.i Rvpubl,c <if
01u/J:rs1tvoortJoumnl 01i·111t11muy IC~$.Ctitcl1. 50. 243--2.SS, Sout11. l{ricd. S<.·1,•m·.._, Bu.lforln :'\o. 39;i.
9; 1-fOAA~. t,..:. •• Dt: \"os. ,·.~-OE Ja.EJUr;, a.o,, 19R.-. Paru...,.jr..,..-, or dome,;tlc;: ;md ua uowt..u. n.J.. n r .... 1, 1.s ft' ltrJM>;, r.r... ~9. '111L· ho..,t ~i.:uu~af the
.,,Jd animot, ,n South \fnco :SYII \rlllrop<Xl paras11;,,ol llurchdl', smped mo111e. Hhtt//<lomi~ p111111/lo, in rcla11on 10 the tick ,-e<1or, of
~ro. Eqrms imrclwllr. an the.- ea..,1Nn Tran.·w.i.a.J J.owveld. 011dt!t)/fl/>0qr1 hc:mwatcr m south .-\inn 011dm11•pt)<m Jo11m11/ o/W1m11a!)•
loumnl ofl',•wm:ar,· Rrnoq,c/,, SI. H5-IS·1
"""'""'· 56, ~291
'#·
98 1101t,~ 1 •·fUVl'Ufl, , .•, .. l!:hJL P~rA.~ile,; rJfdOm\.~llc .rod \\lid anhn,1J"'fn 1111 1111. Iii: .• lllt.\".\D "'·' ... otJ\'fR. f,11, 1998, A TC\1cwon the \1$e"Or
:iQuth ..\frit#, XXIX. lwdld tltk> on hare,. in tht Qpc l'to,in<c and 011 Jxt>d,plmguJ. ,,-a'ip, (tfymcnoptccJ: EuC) rtida,cl u:. natur.Jt enc::n:c,,. for
htire~ ~r.d red roe~ rabbla In th• Clr ,,ng~ Frtt ~131< Onlf'7;<1,•pwn tlu.• conuol or uc;;.,., L\ct1n; lxod1dael Srsrt•11m1rc and ~·\pp1a•d A.c11raf1>r,,r; .
/ourn-nt 4_(Vetmmny Rf.,ror(/1, 58, 261-2;0 3. 1!1-28.
99 HOK.\J,,. 1.r.. & 1 ouEtJr, LJ., 19<J.2. Par:bitt,, uf dontc--tk nnd ,,11d ;m!mali- In 115 JA,.... i , . n c. ~ ,.,,r 1 \\ .o. 19~t,. Thi: cxprrimcmal tr,msnU$.>ckJn of
5outh .\frlr-3. )OC{t •..\dul: ucb 011 shrep In the !'.>po Pro\1nco 11nd In tho ni.•lltri<1.orl'shy Hluplc~plwlusn~rtsi. U1uh•r,u•JHJQ11 Jo:<m1Tlof
0=&• Fr«• ~:.,w Omf,,mr1,oor1 Jo11m11/ of\'c1,rin11r,· ll,'!<t1trl1, 59. Vi--rerlnnr:-· Rt"".-rtrrh. ?'t l-6
21>-?83,
Hb JO-,.G(J.\ti, f .. PrltRl, ll.D., ,1CH,1i&nnv:,.1: I' 0.3,,, \tl~hl, f-,t .. ~H;R\\t, R ~. ,$,
100 HORAK I.G.. H)U1Ut LJ., ',()\'l"W£. 11• .\. & \V'lll.L\~1 ..., r .J 1~1 P;irmltcs ,:..ca:r,s. " · ,988. ~prdemioloizy M bQ,ir\c l•ab...i•»i• w1d lltTapl:t>mo-i,
of domestic and wild anlmnl• In Southi\!nc<L XX\ L !ho m=•< or IN :,,,mbl't, TtQp/r,1/ .~1111i:n/ ffm/Jh nnd /'r(ll/111·1/011. 20. 23,1-2 t?,
l'<otUd uc.i..infl-"SUUlcuu. on bird~ :md m'1m1n~J, in the ~Jounu11n /...cbrn
11:" JOS~<f.O:'\ , ~ X,. ~H\1R. D G, '"l'SCHO>,, A,I.•• latft')\. l',h..,;. A"'\',fU J.• 1998
~:nion:,.I ~rk. OndrfSttµQIJn Jtmrmll of\'tt1,irf1111r,• Rt1.,,~n:h, ~d'. 12S-
135, Produ1mo11 cff•ct< of co:tl• 11ck rsoo,,1111111 ml,·ro1llt1>, 1nr..,.1.11ion ar hli;h
)it'ldlng dolt; CO\\• V,;•rfnnr,· f>o"11f10/q~·· ;a, G;-,7
101 IIORA,;1 LG., tACOl' GUIU.AR.)100, A., MOOI ~I\~\ LC, k tl["\'O~. V,, 1gk;.
u8 MMA. t.J•. "'- ·.\\\',\..'-GI. t~~ .. 19<18 Concro.l of lh·6lod ut:b tn A'rl,u.;
Pitr.tsiio;. of domestic and wild animuls is SOuth i\frlra. XXII. lx<>dld tick.,
pu,!loil>di,it.~"" ofhioluga~t <'tJntrol ttSI~ the, cntumogt..nou\. fun~J
on domtstic dogs :1nd on wild c-hmiVore,. Ourt,rrsh'{HJ(1rt J111tr1tt1/ ol
Bt"nuh•ritt bmstmm .ind ,\1'4r,,rlri:lum 1111f.u>1>llnl'. In: <:f>O/'~. ,. ,.
,•cu1rit1tJ1'1 Rl,!Sinrth, M. 573-580
MOrH\11111,>-. M int~,. n,_.S..'c:01111 ltt1&.'n1111io1wl Conft1un,·,: ou Tick•lHJmr
102 110J&A11:. i.c;.,c I\S1c;,1n. >S,'.\1,, 1986. ,, c:om1nmson uf1hc ockhun.lcns-01 l'mllv;:~ms ~, rh1110,1-\.'.,·,·tor /n1,·r[t,(1t,·" Glul111t Pi..•q1>«"1f1·,·. .:\Jgu-.c
,,1Jd ani.mlll.s. in;,. nature r-1.-"'icn.·t rind on un ndj:lcent fa.rm ,,hen· lick 28•Svptombe'r I l!l95 l;su~w ;s.,,llon•I l'vr~. SouthAfrlc,,: prot:o<dlng,
c:om:1:>t i$ prnctl$cd. Jour,ml of1i1t! Stwd, Af,1'ctw l l~i:rln.af}' ,•\sl<>(·1t1m111. :ind nb111r~,c1~ I S-16
s;, 199-203.
119 t:AJ.\."£.ll, !.-1 x .• 1.1m1U$:T. lt,W.. ntmn,r. A,:-.,. (-:.t'lst1,.. , r-;, L. J\I H-OH1, Jt R.,
103 MORAS: u;. «MACl\'OR. ~~. vr ..-.. 193; rhe scrub hon:. a rcJ1;,bh.• •!M· Popul~tlon dynt1n1le1 ol tlclc> on Ankole caule in fiv< ct:"lugknl
ind1co.tor o! th-P prr.sence of ll_mfmnmtt n,;b in ~h<: Cnpc t•ro,inee. 11.h1t..... In 8urtimJi aniJ ~mncglt...._ for lhclr con1rot Pr<'•'•'tWW t i!t,mmilj'
/n11ma/ ofrl:,Sourl11¼frim11 V~tcr/110,y,4.<s<YC(lffttlll, 58. IS-19. M,•dldm•, 6, 19!'<-222.
104 HOM$. 1.c•. )~AC".J\Ok, lo..• .\1. ot t .. Pl·T."J'\, T ".\. r. D- \'05,. \ 1~'/' ~me 120 >,;[ ' " ' · u.u PUlhL"I .... N,D .• WUGH, u, ..... \\'ILL..\f>SL.'i, P•• 1989, \':Jc:C'lnmlon
11,ian und mcmnuthan ho'.'OlS of -~ml,lymnmn- Jr,~nmf'um ..,nd a~1ln,t JJ.Qo11Mfus1111uopluJ• lo1,.'7lli1t,1ion of unllK~ru;-on th:k 1r1u: ct!ll,
nmbl.Htmnu, nwnr.11r.-11m lAC:tr1' l.xOdfd.-.e: Undt~lf71QfJrt Jm,mnt of
and tht:lrlnttrnction ,,i1h tht hi::m immune '}~tem. Jrx1J..,i111r,Ulll mu/
l'r1,ri11my 11,'JVttT<h. :;4, 3qi-lO~. Ap1!ll<YI i,'11"1/ojO·, 7 U-lft.
,.c. ,1001 ,, ,~. L.c. & r.01,atE, t..J .. l!llti' $cm\\.• wUU hD"h of rhe-
103 11C'tfv,i,:. 1:1 U.. GrntSft\, C... & ,,1,,.·M)~. IU,f • 1988. A cllsntH(:..b~~~ mod~) fot
JO,.:t;,
1\arQo p.ualy<ht Uc~. /l'()((,•1 r,Jl.Jictmdus Ncumnnn !904 1Acarj:
th~ devclo1>n-.cn1 of the ~xudid ticl<. Rllipit'4:ph11lt1StlJ)JNlldk11lmu.s in
lxodfd.,), ()mlmr,poort Jo11r,u1/ nfl'etunt111'J P.-starrli a·t 49-~ I. 1;n<t Co~t fo\·er ronc), \ t•u,J,wn· 1>1ullSiWlog;. 29. J t-51
l06 UOllU: I.G. .. PO t'CltTEJt r.1 .• W.\LKI:R, J.B.• Dt \'O.S.. \ 4r 800\S~.-,.. J•• lg.SJ. 1n KIW,\....'UJ,;,,\, ., .• ,rE\\.\IT (",1., IK'tW:\, ~.M.., IU- \\',\,\I.. n.T. JICE.-C,. \
The L,odtd tid.. burdcn<.Qf \'ariOUS tacg~ mm1n.mt'Spc.1r.te:-. in s;o\llh SCIU)UtA..~. us. 4; J•tTT[\', Ir,: 1~•• l~~- Punlcip.am~· t't:~ate'h ~nu CS1('HSIUn
•\frk'Jn n:ume rtsel'\ t\. ().ndn.</f!p<K)tl /ouru11J af~'~"lt'rimu,· R~11rrl1. :approach to rJd; control in a de-\ '-'lop1nj,!~l'tl:tafSou1h Atrh::,. /11:Ct'M'J\'"~
1

50,221-~28. t ,. mn 1tc.11111,. ,1. ed'- Tir.· Stl(-Onf/ lnu,rnmlonnl Ctm/1.·rt•tttttm1

107 HOR.U:.1.0.•. .!rolll.PP}'\ . .... J..StGHI, M.\1.& tU!tmus, ~ L, 1986,. P.1r=a,qu,-~ of ,,t
Tirk,bor1tt· Patlm,1;:oH tit(• llo.H, Vtttor lntt•rf,lct~. ,, G!()bi1I Pt1.-,p«11:·,.
dome,tic :ind \\ild .utln1al, in S0u1h Alric~ XX.I .\Mluopod p.iruslte,, of .\ub'U."'l ~8-Scpa•mbor 1 199l. Krugtr X.stlon:..1 r1.1rL. South --'frka.
\·aal ribbol bonteboka.nd scrub h:ues 1.11 Lht' \\t,t\.'rn Capt' Pr0\1Mct'. ;uvc.·l"t"<Jrng., and .:.h,u.icp,. 2. 481.-&~l.
O,ukn·tt1µoo1·1 /(1u.r,ull ofVDurlt1tu')' lltt!4'Vtrrlt. 33. 18";- tft; 12) 11:fk:.A,....... ,1 .. IIU"IUI\. LJ. •• Pmm A:\f_"\. \J3 •• Cl,\)POOI~ V L- ",\.\Jfa,lj. ):., ..
108 HOll.U. I.(;_, SPlQ;CM. t\,M . ._ OMi\.-\C,-:, L..l.-0•• 1999, nuctu;itlon~ in lht c.1.,1.FH.t.. l'JSS, Lntontop-."'!thof,!~nic nt"mmodc, n.:-. •1 pou.•nwtl hful~k:d
lburu!:inte <>l 8"<Jp/zi/r"d~<Qlorat11S3J\d thM, Rhipici:pl1u/1;.< ~p('CI.-S on control mt.•thCld for t1<:l..."- 4n11rthoJ"tl1t.• .\cu• \~r.l,: A(l:dtmJ·ofkf1n1.~
rcget;t.don durln.g l l COJ)W('Uti\'e yea.~.,,, . . V."f\llRO\'..\. .)1, & L\IUJD,\. )I 8,.i9, 355-16.
red."} Thr Third IIJfl'flUtli01ull C-OtJjl+nmCi'OII Tic.ll rmof Tirk•l>or,u•
12.:. ,.:o,:,. 0.1.,. FOURJr. l.J,. l~l- Thl" rulL• of llyulommn ticks Ln fO()t
fla.thogc1ns: into tht• ?ht Ct'IJll.lf)', ,\uitUSl 30..~pten1ber3 1999. Hf¢1
lnfosfaUons :lnd 1l'IIIPQr:'U'\· l!'inlt'nc>). or sltl'-t:I> u, ;s~mt,;irfd ,~gion of
Tatro ~tounrun,. SlovJXliL :m,1n,c1>, p.l 1a
South A(ri<'A. OtUlt•fjtt•lHN>t( /oumnl of\ ·cr~ri,uu')' R~arth, 62• .?01-206
109 fl(')RlJ,: J,G,., .srzcun, A,.M 8-R.\.\C~ LE~O,, l 1fSl,JIQH' . R.L... l,\(;X U.t.. M r. ,.
125 U\\HI ~n I,.\,, t()(.C,I:\, c.;. ..,. A -.:Qlt\'AL It,.\ 1.~ 1980 The etli'Ch o!\\:.&rOr:
U'\'.:s. A,C.. 19~ P11r-J~th o! dOllle$tiC und wild .:m:nw!"' in ~uth \frka..
the conuoJ or dtfic.ssc,of tl\'\·s.totk in Rhotles1Cl (2Jmf>.3b;,.,.·.:,J. \t•rtrmnry
xxxm. t,;odld tioks.<in scrub hartlS 11, the nonh-oa,wrn r~1011, or Rt!t'U!'f1. l0i.82..s.3
Xonhcm and EMtt:m TnutS\~ll\l and of Kw.1Zu1u•.X;\tal. Ondt!t$r...•poon
/Ounml 11/ \ ·l'rtrilJtll)' Rt·st,.v,rrlt. 02. 123 ·131 1.26 L,\WRC'tt 1.... &. ;\' u·.J,.T.\'"Zrr r•.,;-,1 . 1980. bc,tt.uu.m o(" non,p1nhottNur
71uritl!rtll of cattle rr.in,m.ined by RIJtp;i>eJJlmltL< tlJ1Jh.YJdteulams...
no 110AA~ 1.G. & \\'tUL,~tt-.1:.1 •. 1936. P.arasi:c-s ofdom~1.1c: and \\1hf aninrnt~
/Jmbt1b1<'t' , ctcri,1my Joumat. 11 . 27-JS.
In Somh Africa. X'"\11f The,crowncd gwn1:t1 fowl \'11mfd(l 1rn•l,'>J.gn·s). an
ini,s:t0n.uic ltu!..i uf hnrn.ituu, iMxl/U ti\J,,:, Omlt•r,wrJtmrt Ju11w,1t <Jf 1;;:1 t.,.\WUt:"liCI', '·"MWM·,;,;u , .....1. 6 "J(tf(\'/U,. 11,A,I, i981 hol,l'ion of
\~ircrtnnn• Hescort1'. SJ. l 19-J.22.. ntrJll-ntt mt,,n11s111 /.1mbabwc. /.l,11lxdm'r" Vi•tcnrrt11)' Jrmmat t2. 2;-!iO,
UI HOI\.\J;. I.(, .. WtU 1 \M'i. t.f. 6 VA.'- ~CIWA"\V't'J(, r.c. 1991. P.arei;;1(e,: of 1:,,8 U\\IU:St:f , ..,. (I ~or~VM ft. \.I, 1_9;a. :\ hblOI) uf1icba.nd titk·bomC'

dume.ilc and wild nnimals in South Mrlca. ;o.,,. r.,odld tic;..,, on ,lwtp ,n U.15,•a"c"' of r.m1c tn Zlmb4b\\C Rhode.!\lo. IU1CN1t,~ia.n \ 1•1-.rm1m)' Jot1rr.ut.
the nortf, •..,s,em Orange free Srnte ~nd in 1he ta\1ern Cope Pro,incc. L0.28-10,
Ondt!rst.epoort /oumul <>/Vrummtry Ht.'Scv11'1t, 58, J t.5--12!¼. no.\.,. 1983, Rltlpfrnplmlus
,::,, L\\\'Rl'.:N('I. I,,\. \Ok\ \I~ H. A,I N Ult f-MU
J.9';'"6, fkk5,.,. mht..,.;1nd tn5t.'<b
11.2 unwo.z.. C-1.. \\'AU.t-k, t.at,t. , t\•1u.. 1t.M .. :nmlJ~~f<·t~~ ~ ;1 \'Cccor CJf hQ,1nr TI1cUcrluc. Tro11fcol ;\ntmaJ flMllh
Infesting dom•stic animal, in S<.>u•h Arfico. P~n I, l>,:$crip1i11ns and ,md Pmdt1rtfon, 15. 39-l!

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38 >".cno, osr: A~1>ec1s influcndng lhl' occurr,:,nce or infec1fo us discas""

UO U:S.S.\AO, P•• L't-::PJ.ATTl!..~ll1R, R•• S(HW,..L. R...A.1,, liiUSDUIT. ,; •• DOI.AS~ T.T... lxodid t,ol;s 011 cru~c on o llhadesi~n higlweld tsrm. Rlux.lN/tm
C.flOZL II .• \\'AUa•.k, j,t\ .• IR\"IN. A.I). It P[Rk\", a.o.. 1990, GtO(rnphicaJ V~tttmary }iJumoT. 8. 2--6.
inform:ulon ~1·~tcm" fc,r smd~·ing th~ cpfd~rnioh~m· 01 nul~ di~a~ ISQ M.\T1101., 5., \1fL T7.lll. r,,<, ,. &- HOMS.:, 1.r;,. l~T Sues of a.ttaclum:ni iUld
cawt'<I b)' TJte/1,rlct p<11w V,rc,ina')· R,'tJotd. 126, 2~26:?. den,iry a.,...,.,men1 o( ,~odld 11ek> f,\carl: lxodid&el on imp~la ( 1cp;~em<
131 uw,.<.., • .. 1981. Th~p,uholo!l) o(Rl1ipicrpT,olust1ppMdirulltl1,s m~lampi,s":, E.xper11m•1m,t mulA1mll1."tl Aam>la.&)'1 21. 179-19?.
infestJtJon nf ci:snU Ttwrou,,g,uot)~. lrn \'r11m~1U..\ll, e.1. ~ ,:u-a.-t0s. u .. 1s1 ,umn)~r.. J.C. At.ou1a. \t,H,. 19SS;. tlrft l:todfrl tlrt:s uj Ug,mdu t~·tlu.•r
1.•tb, f/d.' 8/ofQ,O' um/ CumruL Ptrx':Mllllt,~ oftm lm,·omt(Qm,(l u•itlt SJ>tfltJ Purtimmt :tJ llgnuda IH't'fm.Jt•oftlr,·,, Pt,•1,,•m \tiown
C..onfru11u 27-2~ Ja.nunry 1981, Rhodr:,. Uni\NSU}. Cir.,hJ.m~U)\\"n, DiJrrlh111/011. Colleg~ Parl.. MD: E111ornologlrn!Socleey uf ,lmorlc,
:>outhAftica:. !9!)-:?0,1
1S2: '.\\i\lTIOU, lt.C. JAS,':EH-.1...., COR.11.. s .. Y/lVl, f.A,, 1,A ,nn v..s. I. \'E.RIIUl,...U..., ,
132 UWl~ t..A.1!1.!9- \"aih1l,i.slu,epd4&(,;se. Repon 01 th.t\'t-tl'n~ 1997. Sea-.onn.1 prL"\a.l1tncc- or tic~ Jntl titk,transmitk'Ci h!lmop.i.r.uitn tn
O.panrru·m. !:•ny,,, !or 19~7. pp. 4:;, 51 tr.ldUlon...dl~· m,..n.)~d :Xd.1n1.~ rau1e whh n•!t•tt,nct• to ~ua1esic tJck
&J..1 LIGJrrfOOT,C.J,& "lootW.U. fL:\,I,. 1981, Tick prob!t'.JlU m wddlift H1 c:,,nrrol In rho Gambr.i. llw/101/11ml v,1,ri11ary IIJ11,,111<>/c_t::,·. 11. 3-1~!~8.
1.frnbab\\'c, 1 TI1e cfft'<1s n( urk 1,arasjr~m on wUd ungul:u~ Saud, 153 ",uu'o,. 11 .. "'""' "· "-.~o,u. s.. ,99~. Po1bog~nl<'ity of 1- l;ola,o,; or
.',frifu11 /01m1<1/ 11/\\'ildlife l!<Wllt'('/1, I 1.41-4$ cntomoph:sgou~ nt•m,,todt!i ISreincrru.•rnnctd;,e ond Hr1trorhohdiudac
13.; L()PU•R.£90~j(., ,,. ,c. or WN\I o.T., 199.; J,c~ \·enors or&lb..51(1 for Lhe ticks .4mbl)'Oittmh 1•t1rlrf(<2111m tFabric:fnS>i. &tt>pl1ifuJ m'icroplm
rrauwumm in domesuc pi~ an SoUlh Afn(3 (n: f;1:h1h lmtmnho11nJ :Cn.nt-~ttinl1 and Boophi/W m11wllfltU tSa)•>. &.·,~rinwnwl tmd .~pplltd
Co~g=c,; Parasitology. {l<'1ob~r I0-14. U~4. \l>,.1ro<1, Vol I 2.31 Amrnlogy. 17.11.11-338
Imm.Turk~ 15-.; MA\,,·,u.D. G.r•• D.\U,WST/' "'·'· i, Mff>II IQl.l'. R,\\ .• c980. A ,J.\l(::U '1J)J>roa.ch

135 wu:-.~&Ul<Y. c.ft .• l8.99, The bnnt 1lc:k. •.;mh/;-nmma lh'lm.u~um !,;.ock. h> ,Jt,,
to co1nle dck ..:mmol ln: PrtJc.,,,1JJings a[ f<mrth Blttn11:dl Co,:,f.,•r...'llct,
Slmulo.titJlt ~ttl)'O/;,\"'11nll11, l:Sr1sb.Jn,·
JiJe hi~toryand h~birs.Ag,iru/tura,' /ot1ffl(l(. Capt Towu 15, -;~a-,74:i.
IS,$ '.\;CCUU..OC.:tl IL, 1'-.-\\A\l.+ W,}T\J'\('-"R. \/A, R., ~HD-,\ 0 Q.J • .c. ~ffl.;\.. lf' t:..M.~,
u& t.OU!\SBU1'\". c P•. f.900. Tkk hu-anwa1~r exp<-r1merHi... .-\J(n'culwrnl /tmnu,I
l!,68 . .\ study nf the ttr ..• hlilUl'y ur the 11ek Rluµ;rPplwlut
oftlrrC41,...o/Ci,oq,I Uo1••, l6. f,82..;;a7.
n11prndu:,,tmus - the .m:.111 ,·1:c,or uf Eu.st c:aa..;,t (to,·er .. "·Hh rcfl.!r~nce
\37 LOu~,;1n.rJ1.\'. c.1• .• 1904. Tran.s.mls-:.iun of Afmi,.rn co.1.>1 (t'\·er.•~ri,utwml ,nu, beh~,;our undei field «mo111on< and "irh regard 10 its conrrol In
/o:m1al. C.,pe Imm. 2~. 428-432. ~ukum.sJand Tana:rmt.. Bull,:rm of Eµi::.(J()tic Diuusao,r tl/ru:n. 16.
l38 MACJ\OR. ~.Y,.. m: f t,/ HOHA>... l,G •• 1~;. Foo· ;ib~(',;.,..., int<>3'-''" u:1Mlnn 1-:"i'-5'00.
to 1he ~·.·u,on.Ll ,1bundanc.: ul' aduh '\m/,IJ'tmmm fo•l1meum rtnd aduh \!',6 ,,01utOA!\, u ,. ~Ot,."1:\, 11•• 1~ft. Rt:dlscnpt:on or JJ.1besfr. ,oqul l..~n·ernn.
Rlt1plc,;p/ialmgl11/m,.mm11111111.~c3ri, lxodld,10, /o:m1n/ ofth~ Soutlt !<IOI a, n1t•1lma .,,,,, Mehlhorn, Sche,n 1998. l'nm.mo/111:)• R•"""'h.
84 .
."ifncm: \'1!t!rlnar,· ·!SllA-u11/011•.;a, 113--l lE. :G;-r;5.
l3$ ~c.,r.uoo. r , C"OLHO. M,H .. \l.f\080t.ll ,. M H,, ,w. \:0,. ,uMu. t\ )9i':" 1.57 Ma~nu. T.• t')!t7", \'t)(tdingi.t-:n11lgl.c en dlc bt!hc:t!r \-31\ bu!iolul~·
F.Lo!agt.obcudle, c,! l><oditl tlcb (Acor,. lxodldac• in /..imbl• Ill bc.metting,opwHd. BSt 1Hun< proj~. Unh'et>il}'OfrheO...-n~• Frc-c
St."it$Olu.l act11.-ir,v .1.n<l ,;1tachment~hi.·~an t<i!ltlc. \\ith nott.'~on 01ht.•r S1aw
t10)1, 1.fUllt•t1n oflmomo/ogu:nl /ks,mrch. 6i, t ~ t-l 7 ::s..
l58 ,u..;..uuu.. J.1 ~ "UH.\AL., n.A,t •• 1982.. f!ai;:tor" 1unu~udng.1h~ !ip,1ti.1l
UO ,:~H \.'-:, ",,M .• .UL"-<>l'J•, U., t{t)('.A',!, t...~t., 1•;\UU'H. Cj.Jt. ~ JO;\'.G}JA.'.:, F,, 1~99• d!J.1ribu:ion of Rl11pict1plmlll.) ,1p11t•udkul11w$ in t;),J\! R«rcutfon.11 P~k.
Vacdn~ >Uategles h.1r (.Qu,Vlrin ruminmulmu infcc:dol\s ,.tnd thuir Zlml>Ab'•'"· So111/L-1/rln111 /u11m1111>/ W/1,11/fi• R,'14~1r<lt. I?, IIB-123.
appllc.,.tian 1<1 u1hor chrllchlal in!crrlo:i,. />a,as/rologv To,/ay, 15. lG9 M\\'AN<,I, I ~ .. llA."-<t.\'\ ,, "'·· ~AV,\; f:. 11.1,, r~'SU:OlA,'\, s.. 19i7, 11ll' fmpac: of
290-l!l>:, /xtJtl/ph~tguJ hook~11. u lid: p:m,..,1wid... 011 1tmblJ'01111nn taric,s:tilum
Ltt M,\JL,:\, 'S ).!,. t'El't.R, T,f., Sr.MU, $,M,, S-lMUl. ll,H .. 'CJR\'\l, l't.A..I t- tV,HHl.l, (A<,ri: ltodid>el 10 o lll!ltl uiol In t:enra. l!Xp,!t/1111:111,u a111' App/led
A..L 1~5 l.ab(,r-ator)' rented Amblyomnu, hi.ilurumm .:md .4.mblytmmtr, 1\<'11111101(1·, 21 117-126.
'"""l:l'lmn dl!,cr 1n their ••i:sccp1lbill;y m lnt,'Cllon \\1lh C,,wtfrln ,r,o Nrn, \,.o., 19-1.;. Die oc>rdrat;ans \'on hanw::i1c:,r dcur t1mbl,1'0mmn
rumim1,r11um. /;pldtmidletgydntl /11/f!f.tiQn, l l3. :·MS-303. 1>0mpo.t11m l)OmtZ. 1909. South Afrlt:lm Scitnt/Sl, 1. 83.
14:! \U.lt\N,, \I. ,•1!11:.tt, f,f',•• $1..\IU, i,M,.~l\11\t, b,lf, '\'.(Ht.\'Al, ft A..I U H,\UhJil, 16, -'S r1. \\.O,. 19:>4 Hynlc,mmt1 m111s1t11t.t Schulte; 3 n.-ctor of swe~tulg
.\.f .• 199H Acomp:,rh,on o( ,u~Ct!J1tJblJltv or Ambl_\'t>tmnn helm1ttt1m and stckl\~$$. /or,r,u1l oftltr SomIt tlfrtam Vcu•rlm1r,• .wr,lir:o: A.ssQ.c1a1if'Jn. 25,
Amhlyt>mm() ,·ori~«rum to tnfoctlon \,ich 11\'t Cowdritt rumir,tmtittrn 19-20.
,.1r:u.n.... , ln: coox~. 1 ~ noi·H:il-lllJ.>, M., ted\ n,<"~nd Jm~rn11rlt111nl 1h: ,nt.r \\ .n 1p56 i\ rnn,nhd;uion nr our k11nwlwlst n! ,h_. nan,mi,,!(,n
C:<Juftun, ~ un Tick-./NJrm: l'tllhqgtn$ a., 1lw U<>,l· Ven.Ur /111~1,v~o
ur1,ck•Wn11.~ dl<.caSl-t- Omkr,~·poorr /fJWtudaf\ilf-:rinary R~rrl1, 27,
(;fobi1I l''1tTpfClit l' Augu,t 28-Scpttmbcr '. 19~.5. ~rus:c1 .'.:,uic:mal Purk,
11.5-163.
Sou1hAfnta; prucc.'<llng,, und ob>ttaca. 2 32~322.
141 \l.rm,r1 , o.f., 1~69 Elo,ine bab<$[Qslo:astud> off~c101s concerned In
163 , .. ,,:1.
\,,,,,1 19$; Tht•fl-.•rio..,1,, gonderiosb aud c.,,auxt<mno~: ~ tC!\-it'!\\
Ondi.'Tlltp<tqrt Jor,mnl of\.'Nt+riuar;• Urscarth. 2i, z75--;,9,
u·..n,11ti ..1un.•~111u11t uf'fi<lpla1/ ,ll<llirh1cui1d /'11/«litololl,I', ~1. 1-1-1.
Rcwll'\\1 or h,'(.."(.'Jtl dC\tcJoprltl!tU~ in the pt0(01.0Q}OID or
16.t St:.rrt. \'.,O., 11)6,.
14.; !'tUIU~'£\ . n.F.,. HU~. ''·"·· 1912, f.pi10(>tt01oglc:at f.t.L'UU\. in the control o! 1kk-bome di-\ca.,.~, . St•t'.Ond me1tt11J: oftltt• F,\~OIF. &JNrl pmwlon
bc1\"10C b4bt......io!ioiS. AtJJ.troll(m Vcwrinary /011rntd. ~a, 292'-298
tirl-lJ<>m,dlA'tU.., o/tt,,.,rnrk. Ollm. Working paper \ o I.
tis ,u" r1,u, 11., U\$t.J\f.. ~~, ~LAR.1. n.,., 1u,u.L,.,.., T. 1992- Scudft-!o- or tht" ro1 ..~ t6S t\flfTL \,,.o.. 19;~ Tut l.'XJ~rlml-n,~l tramtnl!>SJ.on o! T1teilt!rta o:•ts by
of .-Vnt,trc:um,w trtrh"8tUmn in tht! tran--;rnh>lon of tkrmatopl1llt,$ RltfpiuphnlUJ dt't'r:Si mi11w1fr.us :md R. bursa. OndctJtt'IXIOrl Jo:,r,u,/ of
amgol,•mt.S. Jn. i:1\.-.t, ft .. rt r,H. , . • HURA~. 1 ,ltd<., Trd.; \ «1or8iologJ' \ i•tt'tittnf) Re;,-,,rl'I,. 39, 83· 85.
- l/f1/Jml am/\ ,r.•ritUlfJ'.~,p,-cr,. Berlin Hehlell,,eri: Sprln~<1.Verfag.
166 NGf.R.\;\'\.\'A, l,J,'.\ \'L~TI-.R, LIL. l'LN'7,JIOli~. ti.I.. M>I." ~•• '1W,\:O.UA. 1, ~
14\i \f\:~n.:,.\ Pt• '\O~.\.."i01.A, "'VA;\. AV1 I\Pf\,.. \\. 1~97, Tkk.<c,,n:rol b) -.;\-osct S..'-t \998. l.'h.zracumz;iuon or ,tuoµ/asma isohm.~ from dand
.. ·nn:tJl.,;c.e.lc c;.ttde fiamH.•A tn 1h,• '-'~nlr.ll £.ta.,ch~n\Dp,4• PrO\inc~ Soudt t 1cmrumisw fl/)'J. • P.11tHJJ;t:Oici1) m wutc -.uu;I ~htti> und O~A profile-$
r,J
:Un,-a. Jor,rtinl ri:t• Satttlt Afrlcr.m \ltrirmnr,·..,\J,,fM1ntion, 68• ..;,5-48, an.ll}-si,<. \"tt<r/n.,,10, l'ura.uwlogy. ;~. 109-122.
u- \t,\$0:,r. \.. & SOR\"AJ.. "··'·'·· 198<>. The tlch C'>f ltmb:1bwe. 1. l'hcgcmu" 161 ,o~VAt. ••,.1 • I!!'"" T11:k problem, ,n r,l3rJon 10 Lmd utUl,.auon ln
Boop/1/lw. 7J111bab11., \~·r,ri,inry)oumnl. 11. Jo-43. Rhodeslo. ,'lltodmart l'emloo,:,• /011m,r/, 8. 33-~8.
1.;8 ''"' r,.:o.\, *'·"··
1567. -lll~lenol\~, in Rhode.sia; ! A s.Lud~· oi~Ji.'l~n()<"1lr 166 soK1·.,i. • , ,. 1917. f,:ology of the 1icl. Amblyom,11,r lr,·l11,r..,,111 Koch in
*cimons oi. er l\\O :staMlJV, /our,ull oftlu SowhAfrirtm \,~,uri,mr,· ,he \!&>l•,m Caiw 1'1'1,int• ofSoulh Afric•.1. Oi.iribu1ion and 1><0>00"1
Mtt11a1/.~nno11, 38. 93-!0Z. atthi1y. /o:m:al of/'11m,/10/ugy. 63, 734-739.
1.,:tt ,tAf'0'.'1,;. 11 ., , SC,h\"AI.. It.A.I., l'Ji'7 Th~ .. c,uon;il ocC'Ururn:.e nf i.duh 16' sn•v.u. ...,.1.. 1979 Th~ llmt1ing o!frt:1orho,1 .,..,lnbilll'· for th•

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 Vectors: Ticks 39

,mmotutc siogos<>n popularlon growth fn economlc31l)' important 191 SOR\'AL ft.A.I .. ~UTHtnff'. M.W•. <il8SOS, J.0,, l.'.t"HR. J.D., TKt>RNE. L~. &
lxodid ll<k\ /mm111/ of Pamsirnlogy, 62, 28s-28, t!u..t..~•v.uc.t:,.\.. 1997. Th~ cfftclb of 1ho bf'O\\'lt ~ar-tkk. RJ:ipfcrpha/tls
170 ~QJ\\'At.. K-\..l., :979,, lic.k inf~tations; and nck•bomc dl.Sf:~ in ttpf)<'lldlculnt,«. on mill. producuon In dnltyc;iule. ,\/<dim/ ar.d
Zlmbabwe Rhodesl3 /011rnnl 0/1/:c S<,111/r African \l~rrrinn,yAuo<illllort, v,1m,u11)' Emo111ol1>gy. 11. ISS-158.
50. 289-?92. 192 sutc,,u. ttA.t, sunn.Mr. R..w... !OAO-t..,.$£.x, o.r..,(,';1J).so~. J.D. ttia:RA. J.o .
1q89, The ellec1 of !he bone tick IAmb!)'011111ui /i,.btn411m1on the weight
m ,o~,·.,1.. "·'·'·· !!NII, The tk~•ofl'lmb,bwe Ill. R/1/p/c,p/m/usa'l:m:
t1Ytlsl. 7.1mlmlJt<1"" Vc;.erim1ry /Qr1rn1,/, t2.3 l-35. g41ir: uC Afni..andct Ske:~. ~ ·,:tt•,;1u.uy l'ti.1Wit(Jle,gy, JS. 329-3~ l.
193 SOkYAL. K.,\,1,, SUTTIJI..Jb'f, R,W JOKGl::>.~LN, O.G. ~ );.f!M.Jt. J,n.. 199;- The
112 .SOR\'AL. R.A..1 .. 1981 Henrcwuccr In Tribal 'l'ru:,t Lands in southem
7jmbab\.;c,, ZlmbttbUce) V,111.•rhmty /OUrtUlf. 12. S6-Si ...fTect.~ oflhc bont ck-1.. Amblyqmma hebmoum, on milk prodcct,on of
S:mga ~nd Sang• x Brahman c:ml<: Medifu/ olld Vtttriflt>'t)' Er.romr>/010·,
173 SORVAL. R.A.t .• 19B1. Are·ll.S$t'S~m~m of U'tt' mrc ofdipping in the conuol I I. 143-147,
or tlek.·bomt dJsc»e in 7lmb3.b\\l'c.•. In! wnn1ili.t\.O. G..B.. .t Gm!o~. f.D.,
194 ~'014\'At.. lL.A,I ,, '-UTll1 H~l. ,u.. . . KERJI, J,l>.. 1996. lnfoMttliOn!, ohhc.bom
lods!. Tick Biolo1Q•n11d Cb11110L /'roc,,1•d/11g,oft1• /ntemnrlonnl
tlckAmlllyomm,i lr.>bm.!11111 :Ac.,rl: lxodldoc) on dill'ere111 breeds of
rA1tfqm,r,. 21-29 l•nu•ry 1981, Rhrnle> Unlversil}-. G111h~m>10M1,
South Aftl.ca, 87-00 canle m 7.imbabw<!. f.xperi111e11ca/ 1md t111pl/e,/ Acarolog:,\ 20, , 9 - .
19j !l:OJt\rAL tu...l., SUTlfEJtil. R.\\'., ruRl.1, I,, G1Jl$0N, ),1). &. ~fftA, J.O,, 1981,
174 NOR\:AL, R.A.r .. 1982. The UekSofZirnba.bw1!. JV. Thcgcnusfl)'diommtr.
litnhflblt'C VtU'rlnary/ournnl. 13, 2-10. ' Th~ l'ffet'I of 1ho brown car-tld Rlll111<'epl1t1/11S "PPCl'dicu/otu, on tit•
growth of Sanga •nd Europtt•n bl'Cl>d caulc V.Nrlnmy Pnra.</•olog,• 30
175 ~QRVAL tLU.. 11}113 The ti<!<$ of 7,lmbabwe. \'ll, the genll5".~mblyommo 149-1~.
7./mbabw,, \'cterinmy /011mal. I •I. 3-18
196 SOM\' AL H.1U., StITlftRS'I . H...\\ •• l,;.UMXI. J., 0:RN. J,0.'1 GfS:SOS, J.D.. 1997.
17G :tOR\' u.. "-"'·''1' 1s8J Arguments 4f;4l115t imcn.,hie dlppini- Z:mbabu ..- I he.- c.ffttb "l lh~ brown t::ir lick. RluµillpltotuJ upfM)nillculnru.t, on milk
\fer,rinory J~umal. 14. IS.ZS. production of S:!np cantc. .".f,•dieal a,,;t \'t1,,rfnnry Emomo!og:,:. 11
t:-7 t934- Resina.ncc to lkl!')phllw ,t«,oturatus ln thl· African
:,,:())WAL ft.A,1,, 148-154.
bultllo (S}ll<,VTt« w/J'"11, Zlml.,11,u., V~t<rinn,y Jo11rn11/, 15. 3-1-35. 197 :mM\'A~ " ·" 1. " naEL.£. x .. 1983 1"hi.• ii~sorllmbnb\\:c. , . Rhrpr«JJhnlt,s
178 ~O.A\IAt. R.A..t .• ag.86.Amblyommn 11t1.l'1ttJV1tum ;:md :.,cutl! bo\i.nt' /111111/111:,suno Rltlp:rrpl,n/111 rrtwsp,s, 7.1mbl'lb«>.' Ve1umn(}' Jo11rn11/. I3.
donna10philo~fs. Pmn.,110/"I{)' Tl)l/n,1·.? 2J J..h.17.

1;9 SOR\'A.I... R..i\,t., .\.~DH£w. 11.a. g, M~l.l'l.l ,K, ,1.1,. 199\. ScJ.sonal occurrt.nce or 198 >ORVOL IC.A.I .. \\'AI.J(ll]l, 1.a,,. e.OLRl)k;<f, , .. 198~. Th<' ecoloin· ol
the bon1 tick c,tml1/yommh hebrnc11111) 111 !he southern lo,weld of Rh/11/o'J)hn/1<; .om/J<:lrns/1 and Rhl111c,,pltn/11s «J>f1'/lldiculiuus
Zlmbabwe. Ex/)/!rimemnl 1111rltlJ)Jlli<"'- ,1"1tOlo11J', I 3. 8J ·96, tAcanna: L"X:ldoei \\ilh pmicul>r rcfcronce to Zimbabwe
011<1i•rslcJJOOrt Jovm(I/ o/Vt1e111mry R,~.re,,,,h. 49.181-190.
Epidc,miology
1150 ~Ok\'M.... R.A..1.. flVAl, .8.11.. L"VRLl\'Cfl. , .•,."" UH()\\'S , .• 1984
1
o(tick,bome dlse;ises of ca1tlc 111 Zimbabwe. JI. Anaplasmosis. Trop;cal 199 0 C:\U.1\faL\:\',C,I , • .MU)U!,. u.•.. l•f1J1R,·r.~.- P(M.RY, n.o .. 19:96.
.~11/11111/ f/ralrh nnd P1(>(J,miq11, 16. 6.l-70. lm1~tlgc1ung th~ cptdcmtolot;> of hear1w;t1trr (Cbwdrlrr mmfnanm,m}
in(t.'tt!On h~· ml'.sn~ of_. lr..\nSml\'licm d)'n:tmic~ model P,1ra.'f(tolt>lD'·
Epfdemlology
l8J '\UA\li\L. M..\.t .. UVA7., D.11 .. JA\\'Rl:.~Cll, l,1\, ~ BRO\\'X, ,\.. l98S,
111, 49-f>I.
Mtlek-bomc di="'• o( cuttle In Zimbabwe. Ill 71,ciltria pan·o group.
rrop1wl Anlmol Heal1h (Jlld Pro(fuctlQ11, 17, 19-23, 200 PAL'lth. 0.H. llUI\A."<011'\\'A, f .R., KOC\S, lri.M. Jt J'IJtOWX. \\'.C,. 1999.
:.tolecuwr ba<i, ro, ,11cc!n~ de,clop,nc111 ng31n,1 lhe eh,llehilll
182 ~OR\'At... fl.A. I., 1'1\'A.l'rt tl.lL, t.AWRJJXCI'_. I.A. -11 lli\HU.C:t>UN.f• 1 ., 198J.
p.:ithng~.nA~uipla1m,> nu,rginntr. PtmWtofogy Today. JS, 281-286
Epidcmfolof_zy o(tid·bomc dlscasc of ca11h1 In Zunbabwe. I. l\abesfo,1s.
Tropical ,1mm11/ /1001111111111 Pmdt1r1100, IS, 87-94 ZOI PE(.;R.\.\I, R.0.,. C:UJ:JOJU), C. 'it... \\'i\0:Ut. l.S. t. KEIAAS-& J.11 .. 198;
Clurificauon of tho Rl11plttt>l1nfosst,11gut1w1u gmup L\cm i""do1de\1,
183 ,oRvAL. u.1. t.. U.WM.L "-',.l!., I.A,. 1979. TI1e conuol of heam,"attr In h:odfd3t'1 . I. fl. ,iu/«1111;. X~m.tnn. 1908 i:u1d R. lurttn.icu.c Porr.emntse\',
7Jmbabwe Rhode,la. Z/111/Jobw., R/10</~in ..\gr1a1/1uml fouma/. ,6. 1936. Sysremt,Tic Pom,lroh>gy, 10. 3-26.
161-llu
20~ PUffl..\.'11.! R..G., ll~'tl ltAi,\L. tt, I<\\ ,\..'iS,l'..I, U,\'.., 1981. Tid.'$. (Acari:
18.; ,'()lt\'AL R,i\.. l, & UGllTroOr. t.J., 1982. Tick prob!tmS ln \\11dli!1: in hodoi<Je,J01 E1h1op1:i. I. Disiributlon. et'Oiogy and hos: rcl~lfoM!tips or
/.lml>ol>wc. fa<ior< influencing 1ho occunonto •nd Qbundance of
sp,,cit'> micung liw,1ock. Bu/Ir/in o/EnromolDfJMI Rr.l<'OTclL n .
R/1/11/rxp/1al,,. ap1*1111i,·,itams. z;,11b!lb1t•· Vtm/111:.,y fountAI, 13, 11-20. 33!hSS9.
185 ~OR'V.,\1.. " ·"·'· t \tAWS. c.A .. 1981. The! ricks ofZimbnbwc. IL The life :.!03 PU,Fti\;\1, J\.G., IA''l!'o, A,(),, OOt;,Tl'RWIJK, G,P,M,, ~U.OR:.-.:, t,;.J .. LnlCHE,. J..
cyclt', dhtribuuon ,md lh"bl.a. uf Hltlpftcplui/uys:lmd KiKh. I~.; C':Hut<1rn, M,. 111):l.E, I,., Cut7\V~\, l(.C.,tl.. ~tWML lt..T. 4c CHfZH-U~. f .. 1991.
7.imh<ibwc \'"t•t,•tinal)' Journal. t2. :?.-9. S1udits on the ,..:1momlcs ohic~s in Zambia. Expcrimemal anti Applied
186 <OI\V~L " · "'· "•llAAY, • 0 .. 1990, lniroduct,on.$Jlr~d :i.nd <Ubst,qucm Ararology, 12, !l-26,
disappearance of tlt.e l>1own ear-iici<. Rh1f1ic,pha/1tsnpw1U11,11kt1U$, 20.1 Pf.<11iAM, Jt.O.. LL\:CHE. J,,. CJIIZ\'UAA, H,C.,R,, S\ffHtiRST, 11..W,, fl.0\'1). SL.a,,
from the somhcm iow,·cld of7Jml.labwc. Ex1wrlmc111(1/ am/ Apf/1/M ,;E.nA. J,t>.. • ,t('.);OS.:I K. l',J .. 198~. £tT~t or1ick cont ml on li\·C\,('igh; gain
,lt'Orology.9.103-J I I. of caul• in centrnl 7.umblu ,\/e11/c,ll 1111d v,w1rt,>ry Emomr>log;,·. 3.
187 ~ORVAL. It.A.I,. Pt JI.RY,. (I. D ., KMUSKA, R,L. &KUNt>LRT, Ko, 1991. The use of 313-320,
climate da1• inrerpoladon in ~rimnting thr dbirtbution of .~rnbl}·omm,, 205 P£GAA.,'\I. n.e.. \1\\'ASf.. Lt .• 1.f\'~OVIC:, n. ~ 1¢-XCfl'Ji\;\, F., 198-8..
wiriogawm in Africa. Pr~i-emi,~ VcwniUII)' ,".frd:Cln... 1 t. 365-366 ~orphogcnouc dlap~uw in ,1mblyo11111111 mrlog1111,m (A.:.,rl: I.U>dldarl,
188 -.0R\1AL. JI,,\ , •• PlllR\', I\ , n.. \U!L.T't.f. R. ~1.l .• ).'1\US~. JU.. "° 80011-t, -r.u., tSg4, ;\fetllra/11111/ Vtr,rinary Enromology, :i. 301-307,
Facto~:ufectin1t the dlStnhuuon~ of thv UC'ks- ..\n1l}'(lm11:tt lrnbraeum and :?06 Pt(.ftA.\l, k,G.. Pf.JISt'V, 8..U.. ~lUSfSI, f .L.. &: )o(W.i\~.\UMO. 8,. l98-6. f..<OI~· and
.4.. tYJn'egamm m 7.Jmbabwe; unp11cauo:\S ,,t reduced .:scancldc U$.3g(". phmol~· 01 ue~ m /..amb1:t: ,tra~>n.11 dynamiC\ on C'ilnJc. E..rp,n·m,muU
Exr><rfmcmal n11tlApp/,roMt110/ogy. 18. 383-40,. mu/ t11111U«1 •!t'urofO!(I'. 2, 2HS
189 :-:o'Rv.u. fl \.t. .1i<.~11c.1Rt. ,.1H198.1 ln1ersptcilkC'1')mpe11uon between 207 ,w.
rE<;RA,,., •..,,. .... 8.P, A• >CllEl,... Hf., 1984 Seasonal dyn;mlcs or the
Bo<)philtJ$ tle<olaron,s and lkJ<,philu., mkrop1,,.,;n southern Af:ic:a. hr: l)MISitlc ond non·p;).fllSHic >l•f.e> of c.udc 1iw 1n Z-\mbio. /11
GIUmnf>, o.A. t, "°"""'"' CJ!., (eds). Acaroloio· ;'/, Vol:?., Chichester. f.R1...,.m1:.. o.., &. uow,~"i, c.c., {e<b). AcarofOi;O• VT. ,·ut 2. Chichcs;er::
Ellis llor,,QOCI LJmi1ed, 1242-1246. Ems llorwood Umnrd, 1163•1188
190 ~ORVAL tc.~f .• M>~L~::.Hl~t. D,fj.,AUN\, !-" It BUiUUUCE. M,I,, t9S6 ~8 rtORM1, H,G.. WAU:J:fl, J,b,. ('J t»()JU>, C~M, tic IO!Jlt/\~:,., J.C., 1987
Effic.tt}' or phe1omone-acarlrlde•impregn1ted tr1ll•mg decoys (ot Comporison o/ populations of the /111/pfrl"phaltissimu•group. R, llrn1,s,
conuolling the boru ti~k+Amltfy'1mma hclJraeum (Acari; lxodfdae), on R. prru.•tc,;ttm.s. and R mullsamM !Acarl: lxodidaeJ Jnmnnl of,\fed1cnl
c;a1dc 1n Zimbabwe. l:.xf)erlrne,,ra/ and Applied Arara/Qgy, 20. 31-16, E,,romo/og,,, 24. ~ 2

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40 »c11,s '"" Aspects influencing 1h11 occurrence of infeclious diseases

:!09 Pt.lllorusi. o.. 19.;o. Ll ~stro-cnteritc <.'morra:¢c.t <fl'llt pecore. 227 R:.,..:sno1.ru. ~.i.. t99i.Ab1ortc nnd hfotfc: dc,cnnJn:.uu.s ofthe seawnal
&;,eriIMntl di ironsmis<lonc col'11'1/pfc,pl:alus pulrl:ellru' Bot1111i11a dyn:unk~ af 1he tfck Hhlplet!µlutlu.<iJ/1/>t'lldlculnh,s in Seruth Africa.
ilvlfa s«te1a ltal/a11111/I Mwllcillt1" lg/,n~ l'rtJp/c,1/<. IO. 164-170. .\JetllMI a11d, t1tt.1ri,u1ry1 £momoto~·. 1 t. 25-37

210 r!'ff.R T,f., .,~t)tJISON, }l.(',., '6'1RR.UlC;f M J. OMAHAS, '!i,..•,t, 1998, 228 MXDOLl'll, s.L J. 1t0<;[as 0.1. ,'l93. 'fowa.ids new populatlon rmxM, as
O~monsu.a.tion oft! rnrrie.r St.UC (or CO&A•dria ruminnnu1,m m wild tools.for the c.o'ntrol or Afrtc,an tic~ :md ri,1k,bo.m e dise~.. /Ji: <':ooxs
rumlnanis ttom Mr!<Ul./011m11/ ofWl/dllf, D/.1,.·=· 34, S6;-,75. L. &sou1<u1u,. M, ted~}. 77J1..1 s«oml /11tttmmi1>1wl Cnnfrtr1rc,1 ou

:OJ Pu-tft, r.r ,\."t,ftt~O~, b,Co liUKHIOOl, M,t, Plil\K"I.. 1$,D,4' \t.'\tt.-\;\, ),M,,
TkJ..-1.Jum~ /imlt(JJ:!:NJ. ul ilW Hu~J· \r'U..1«11 lnJ<t1fu1,~ II <ilub«I f'r:1~'1Nt.'lfrfl.
1999. Susccptlbillty •nd c;11Ticr,-wtu• ur impale . .,.blc ond ,.,_.,be for >\u-gu~ :?S,,,Si'pH.Yftlh<'r I 199:>. Krugur-Nallunal Pu~.suu\h .:\fric.:i;
CouTlno rumfnn111/um !n(ocllon (he,trtwaier •Joumal oj /•aro,:/tology, pn·Jt·eNling~nnd ab!nrncts. t. 2.o-35.
6.5.~63--172. ~~ RA-STEOAO:.n• .t;.s;.. 1g,33, 7ur Fntt;t! der Ubcnr%-c.rdc.'f
;ti2 •r.rn·K , .... \l,,Ut,\.~. S.M .. t~AMHf:r. A..F., ~C)K\'\l R..\.S. ~ UURhll)(!ti.. M,J .. S<:h:slp11op!4smosen ,n Azer!>a1d$Cha11 Cl rnnska11koslet> ,lrrhu• fm
19911. Th~ abllil}' or t<lrrier c:nttlc to <ciw a, ,~sel'\ o:1"$ o( Cb11 drirJ 11·mem<1111/1/lcl1< ,md />mktis<ltr 1'ierl1rilkrm1r~. 6i 1-i;. l 80.
n,mlmmrlum 1nf\..~tlon for ,\mhlyQ1mn,1 ft1tbrt1<'tm: tiel." In: cooN~ 1 & :,JO KU:1"',, .. 1982. Dynamic• or tick pop11lutlon, Uc•rl: Lxodldn~I in di,•
R<tnlMnU>. M. 'Cd~}. '11rrSecmul Jmenmt!Gnal Corifer~11uon 1'fck·hom~ e;ss~em ::ape Province of Somh Afrtcn. Jounut I of .\{rdfg1/ F.momol~gJ 1,
Pmirog.:n.t "' 1;,, 1/,nr•Vtcror lntcrftu:~a (ifq"'1/ AtrJJh.-Clit·~. r\o.gus1 19. 679-,00.
28-Sepr.mbet I 1995, Rruger National Par!,;. Sou1h Afrka: proceedings 2.11 ttl;CHA\', "" o,.um. J,• u~. o,A., 1991'). Resi.M.anrrnf Br.i.hman ~
and Qbw,am. ! , 185-188. Simmcr.t'11er cnn!~ 10 <outhern African 1tcks. 011dmtc/Jl)Ort Jo:m,nl Q/
2-Q Pl'itiH. T.t.. ru:RY. ls.t),, 0°C\l.l,M-1lf.AS. ("..I. \IH)Ln. c.;..... ~UU~aJ\. w .• \'v1erJ,:t;ry1 /rei~nr(ir, 57, i-12,
,tAU%a,tA. w • m11t.1t1DGt. \f,f. &->t.\lJA"''· ,_,,, •• 19~. The dt"'trlbullon of 2:J2 RECH.\\.,.. .&. ~o:rii:h.1.F-\\,-,.tra, ;it.\\,, 1991. Relatfh· r~i"'t.anc~ot si~ eatde
hci>.rt\\'ll~r in lh• hlg)wt•ld orzimi,,.bw~. 19U(H~,. Ond,m,poorr
hrtt..-d, tn \htl tick RooµJuluJ thx"tlformtti in South ,\!rlca. Ond1.•N1"'poon
Jo•mal o/l'r;urf,:ary lksmtc/1, GS. 177-187
Jo11rn11/ of V.•t,r/11111')' lli>Kwc/r, 53. 181-18G.
2W f'IFtl!R. T .r . PUUtl, n.o.. 0 1
CAU.AOIL.\X1 c., .. )lf'i.l)U.Y. G.F.. SJ1U~11l.\, \\'.. 23,:1 Rll11 w. v. t. ,ut1,Mc1<<:. ".n.. 1986. 11ck popuJ,uwns orcwo breeds of
M,U')7.lM"· w.. aunmu,;,:, '-S,I, IQ> M.\H,,~. ,.M,, 199!. Oil'tributloi,.,, of th~ r:!ule under fidd rouditJo:1S, \\ith n note on blood componl'nt,5; rchued
, 1·cc-ton orheam,JtCr. Amb{vommn Jitlmtrnm :md Am11/.-.·tm1ma
10 ho$\ rtsi<"ta.ntc. ,,,. MUrR. J,Jt. t, HAik, J.A .• (\!d.1'l. .Worpito!ogy.
11t1rlrg111111n i.\cari: T,odldae). in Zlmbabw•. E.tp,rlm,1110/ and App/led
Physin!ag:,. wtd B<•ltn1·/oort1/ Biology of11cl:J. ChJchcs1cc Ellis
.~rmo!ogy. 22 1:t>-740. IIOt'\\'OOd.
ill$ rirr~~·- T.S, ,- HOIIA<, 1.<J .. 1987. Th• clleci of dippini on porn.We n,nt
~4 R.&C>l,\\', \,. £ftol'R.8UG. ~.[. &c.lJ LLfK, U.A,, l987- D~-nanucs or•..\frkan
frce,lt\i.ng povuI0tiou5 ofAmblyommu Jzrbrn~um on a. fMm and on ttn r:cl< ,\Cart: ixodoidoal popul~tions ;no nnrnl'lll Cr\-·Congo
ndJacem mature mo-rvt-. 011d1•r.<1q,o,,rr /c>:m:a1 o/\·turinMJ.· Rt1C(Urli. hmnorrh:lgk ft.wt fncu,-,.Juumul nfMrdia1I Entomolti~· 24 . .575--58.1.
54. ;;29-533.
233 nm,."·' 19~0. Thdifc <-yde of t.ltl/1P.)'lr> b1g1,mfr>11 !S1111thand Kilbourne,
216 PCTN()', l ••~ .. IIORA~. 1.C. & R.l!Cll\\', v.. 193'". 11,~<-colotn· of tl\l!Arr:can
1893) io the tid. \t.'ctor Boophl/us m1cropl11.s cCane-Mrh'I!. tiustra/Jtm
,·cc1ors nr ht.Jrf\,-nwr. ,,ith pnniculnr rcfr.r~nrc io ,\mb1..,-,mma
Joumnlof .\J::_rin,lmral Rtsenrr.J:. 15. 802-821.
lt~bm..:um ,md ..tuthl•NftJYIW '·""tci•mum. 01tdmt~ptJCJr: !our,wl of
\'r:1.•rfmuv Rt.\"Mrth. 5-1. 381-.'J9S. :t36 ROatRTWX, ,,.D .• 1fN51. A !'our y('ur Sllld)r or thesciOOnal Ounu.atidfh lr'I
the ,xcuttenco ol the blue nck./JoQp/11/11, d,<:olom111.< Koch In ,he
2:r; Pl,OWRIGHT, w•• PJ.=R.R\', ,~..·, &UR.Sil(;, A•• 1974 ~cxual u.inm,i ..~on or
African s,,1nc ftver ,,ru~ 111 t.he udc Ornhl:odoros moubam 1,ortinus
C.Oi!.Silll fdijJOn~ oflhe~3)tem Cape. /n;W1UlEU£..\O, G,D, $<Gl-..~O:'\, 1-l',.
Cedsl I ,ckBiolo,ow:cf CcmtrQ/. Ptvc«,U11&, IJ{u11111,um:11wtnl
Wal,on. R,shil'dt ill l'ttori11n,ySc/mcu. I,, I06-ll3.
C,mferr:1~.27 -2!> J,nwiry !!1111, Rhod<:S Unl\ernt)', Gra1:mm<11m.
:?18 rtoh'R1GHT, \\,, P't:MMV; r..T, 1!r PMRCL ''·"- 1,;o. TralhO\'at1al fnfc·cdnn SOulh ,\lhtll. 199--204.
\\1th .\Jncan .,\,inc- fever virus in the arga5m tick.. Ornlrhodorqs rlwubmn
-Z37 k001UQ1TF.7., ,1 •. \l,'5~.,nn.c.14. 0,\ FO~SE<.1', .,.H .. RA,\J~~~.f .. '\t.;\Clt\OQ, H.,
1>0rrm111-. \\'alton. Rtt1Mrdi ill \'ernruuuyS:;ien,:.t~ 11. 582-584
L..Uli,rcJ.\. \'."OE u rur.."\-rc-. r., 19~ Effect ofvacdnanon,,ith: n
ll9 PUJ\\'JUt.HT, \\,, PJ:;RR\, ~., .. VlilM(",J ... '""· & P.\tu::l:.ff, , •• 19;0, Experimental rec:ombtnaru Bmtui ~lnttRen prcpamtiQn on nu.tura.l in(e~t.'ltiun .. or
infecuon of thl"' aigai.ld tick. Omi1ho<lQm.t. muubuta 1,urcinu.t. wilh Boop/1//t1smterqph1s 1J1 grar.lng ~QII)• and b~rpur-eand crossbre<l <4Hl~
,\ll'ican ,win~ fetor \'iru;.Arth/11 jllr/;ffilmte Vin11forsrl11mg. 31, 33-50. in Bru,l 1<11:rin,. 13 18().1-1808.
220 •01c,1m·~....., .. >97,'- Thu IICC·C'Yrlcor 111,oow /1Q1•/$Md 111,1,.,,,a ~8 NQnJdQUJ 1,, ,1., PF~1Cfti:'T, \1,1., ~:OU111S.. .\,fi.. U 'll'\ltTA. \'., l.OR£N7.0
111,i,mittli in de~ and ln ca11le in Suu1h Africa Pl>.!> 'fhl?<is, l!and 111 \fl.S. t.. lllllUCR.\. J\., C.01\00\'l.s, C:,, MNCl!F.4 P,,A., R,\~10~ t ., WTO. A-
.·\fnl<:lan, u,wmuy. C.\~.~. )1,. PAU.'\7,UE!A. 0,. TRIG\H..AO, :\,. l l.,- Q:O..\RT J\., ttl Stitt.RA, Lt.
22\ J)(>tG-t~'laR, l :1 .. 1981. Tfck 1ranflmi.S.!({un of an~pl3.,.mOS!>.m South Africa. m ._. •Lf, n. , .. 19gs. C:Qnt<olof 0,1<Jp/1il11• tmcropl"s p,;pulnuons,n
lit: WlllTOIC.\t'I. c;.a, 1,, musos.1,n., (ed.$). Tfcl..· Bfo/010· (Intl Comrol. gmzingcnule ,"3ccinatNt \\ilh ~ re-corn bi.mint Bm66 DJU.ig1.m
Proccedl11g$ of 011 httm1t1/io1111l Cd11fct.mu, 27-29 Jnnu.u, 1981 Rb~d,,s prtp;aJ11t.fon. \ 'ttrrtmiry Pum.mtJIOgJ', ST 339-349.
lJniu!rsi(1. Grnh:lnistown. South.-\lrlca 53,.56. 23'9 ~,11su.:1o1., ~::,n.-v r ~ Vf,M'J:.K, 1•• 1998 The dC\'l'lopmcnt o(
222:. POYG.11:rt:n. r.T.. t>E WAAL. o:r.. & ros-xerr. u. \992, Tr3:nsmi~ion :sort cmomopathogmk nc:mntodcs wlt.hin the tick 8o,ophllm ammJ.nms~-ln.
d!agnos!sor c,qulne bulx.-sfo,ls in l>ou1h AlrlC1l . .\lemlirias do /11>1/uuo c:ocrv~. 1. • R0TJ!$Jtlt.D. M, (ed,:}. nu.- s«ond ImcmnrlQn«I C"njn'"·n«<m
Oswtlldc, Cruz. Riu 1/1..• Jtml!ira. 87 (Suppl 1111, 133-1 .J:?.. 'f7ct,bante Pathob,."4..'l{I ac 1hc HlJst~, 1a:1or /mer/ace: n (;ioool P~upc.'Clll't*,
,IU!ll!St 2$-~iombcr I 19S5. Kruger Na11onal l':uU,o~th Afrtca:
Z23 r,nmfTE.R. F.T. 1r-u.s. 11.1 •• 1977. Light a.nd clt.:ttron m1cro~C'Opi:
J>mce<."C.ing~ o.nd a~t.mc,s. 1, 2-t.
obsol'\•ouons on 1hc d"cl<>pmont of /l«bw111 bi1;'<'mill11 m 411\'~e.
n)'mphac and non-1<ple10 rc:n:llo,, o( JJ<>op/1il11td?r0lorm11s. 2110 S,,\.,,'TOs 01.,~. r. \.1'.. 1993. !')omcdatn concerning the dC.l."S
011d~r.,tcJJOl)rc /ourntll of\/ecerinnry f/tsef1.tch. •\~. ?13-232 (1\corn>a-l,odo,deai pn><ently known ,n Mo,.ambique. Gan:ia dl!(>rr11.
~,ril'S Zuoloi:iw. lisi10.,. 18 (19llll, 27-tll.
22.4 PQTt:lf.TI:11, f,T, k VA.~ RJ'..,S1.\UJUj, , .. , 1g8;. T ck trun,mi~..Jun of
i ·\nrtpla,tma n.'1llm1e. OndctrlU!f",)Ofl }t)umnl of t,.111''rinary R~i«m:h, 54. 241 ~HOlTl. \t, \1,. ~1•1-CS.:tn. A,)t •. LO:'o.l&4\HO, P.t. 6 ~'°,$U~·. ~.1 1Y91, !he
;...; eil'ec:1 oflkk lnf~tatfon on the protluttivit)· of COM of 1hrrt breed, of
ctttdc. OttdcNllpo(>rt /OtlnU1lufV.rtt'rlnmy lli"SC'mch. 58. -1-;.;.
~ Pllb"JOS "'· ,t.Jt Jt>S~F!JA~. 1-.. 1a99. Prntc,c.."tii.-e hnmune miKhanfsm< ,o
1ld" and tirk·bnrne di>e.u~, or rumin'1!lu. Pnro1irol1J8>' Todtij•, 15, :142 5EltAA:<O. ,. ,1.H .. 196:t, Con~ldL'fll.~Ot'l- ><>brc o.mtlrfologia. «<thv~,a e
255-2,aa. biOlQgi, doK ,,odideo, do< g~nero,,11111,1,.,mmn e Dertr,1Jtw11or
,s P..\SDC)l-PH. S,E.. 1!.f:93, C1inunc. S':m.•!Uttt im::igtry 3.nd 1hc,C'2.sonal a.s,innl2d<>S cm Angola. H,l'lm, lit Cil11rins l'uterf11arl1u. ;a i81-20-l
or
nbund:mcc the tlcl< R/111,iaq,/111/tu 11ppo111/.u/111u, in wu1hem Afi'Jc:,; ~.;j SHEPHl:.IU), .... ,., S\\'A , , ..vcrn1.• R.. c.:oR~f..l, A.f. & MATIIE1•• 0..1989.
a ne'\\' p.cm,pcetl,·e ,\/rtlla'I/ and Vez.:rinary Emomology, 7, 243-258. F,'Pcrlmenial .iudie.on lhc rc,piicnt!nn and U-lllsntlilalon or

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 Vectors: Tic~ 41

Cd mean-Congo hcmorrh:sgtc ((n'"t'r \1N't ln ,ome ·\mean tlck --fkde,,. I 199,, Kruger '-~tlonJI P;,rk. !i<>ulh Afrk;i: piu«t'<ling,, .u1d .b,ir~m I,
,\mtrlcan Journal of1rop!cal ,\Jt.ttl((uw,mrl /lyl{k•n.:. .;0. 3.26--3.lt. JlJ....1-1.
2,~, s1101tr ,.J.,..
,ul\\At-. H..\.t., 19111. n,c ~~nal nahia} or kllfp,r1.•µl1nflu -'63 .SUTil61t.:,. r. R.,\\ JO,h, R J, $. ~... HMOntU~O. JLI,. 1981. Trop!(cl Jc::,wnts:
hpptmflcu/(1111, '-c11m,1111 1901 (M,1rina: lxodid•~J ,n the hl~hvcld of of the jl('nu.\ St,,:lo.qu11/h.•J.·1mmobih:tc :'JU.d klll rnttlc tick...: Vtu,u~
l.1mbabw,• llhod<':<io1 /oumnl off~lfOJ/t(JltJW, 6,, i,-84. fl ~nr/<111), 295, J:?1>-l2J.

u; • ,on\·,r\L
--tlOkf. x..J. ft.'"'··
1~1. l<l?'Rulntlon of ~~asonal otcurrttrtc.·e fn :?t;J SUTIU.kSf. R.W. s. }t,\,\'\\,\t I>, C. t .. 1965,.,.\ C()JnJ)Ulttii'.(.'dsy.s.u.ma for
1h~ tic-icRhrp:t·,,pholushpf.N1ulicr1taws ~tum-ann l9GI. Troµ1rnlAtwnal matthht8 climatt·'!t in -tcotoRy .Agritultu.n.• £CW)'St~m, nnd Em'iron11h•m.
Hraltli mid µ,,nftictfort. I!. l!l-2f,. 1a.ia1-m
.: 16 ~,urK, A,n.• 191\3, Babt"Jfil /Jo:,iS! l'Ompmcr ,s,imu:ation or thc:' tt'Jiltlonship :i.6$ ~Ut1Ulb1, 1\.\\,. '1 \l\\"\J..D, t. .... WU\. In 4 lo,if f.t:'I \'.\, 0.,, .. 19,8,1, The
bl's,,•e;:n aht• tft~ 1.·,•t1or. paro.,.i1c. nnd bo,into ho,t. Fxp,.,1fmt•n:ol cflect '1f cant-: rid. 8oop111JUJ mtan11lus1 on the gro,,1h or Bol wdtcu1),
P11ra,/10I0f!;t. 56, 2i-4fl. 11. mun,., -ttetr:,. \r1i1111hn11 Journul n/A.gtin,lrrunl ltistlltclt. :H,117•327

u: ,:-.NA..'-', A•• fOtJRH L.J .• t::QJt: o.J. ~ ,mu.Ai,... 1.u.• tS9-1. Ven.teal m.1,vauon or :66 '-Uli-nttST, n.w. wn..o,. 1 .,. sanr,. n. ji ~1 AA.. J.u.. 1988. :\ sun·c-! ut 1ht
:.duJt fX()(ltt mbftmu.lw;, the K~noo p.-m,h~f.; tfck \ari· b,Odld.u.• .1hllit} 1,f tropltal 1-l'gurn~ in th~ genw, sr,·/0#1111/reJ tQ tr-1.p li1t,ue o! thC'
E.rp,,nrr.rnrnl anti ,lppllt•tt A(nmlf>/C', 18 JOl-110. cattle :ltk, Boo(lltl/us mlrropr,., l~vdldaol. ltmra/11111 Joum"l of
B.r1Nn11wn:a: A1,rr/ru/rrm., ?6. 473-ti9.
u.8 SOW}io:,. c;..,. .J:..\AYA. a...P•• 1990. Comj)::iruon of r<.>S-is13nre in 1hr~...
l.67 \\\' \XEPO!>L f .. , ~nttJ'TlUJt!t, p:,, ~HE.fnlf k1•, A.I., !\lC!G!W\"JtA'I ~~!..
breeds <>f catd• ngwnsi Afl'lcan lxodid ticks i;x,,..r:ma111nr e11:d,1µpll<'d
'\'bl. "·'· & JUPP. P,<> 1933. (:rimcan•Congn h1.•mnrrh.igk [O\'tl ln
Acaroroz,·. 20. 2?.1-230.
South Mrica,,lmcrr<V111 /1111rnnl u{Tro11f(,i/ ,\/<',I/cw, ,mtl H.1~,nt 3:?.
249 sose.x.smxr.. O.E., ,UJ.,.\S, $.A., S(Jft\',\l. R..A.I fr- MUlJUIW.t, ,, , •• ~996..\ ~I!• 1~07-1 us.
medtcaung ~ppllca,or for control or ucl.> on d~or. .',/rdlf./11 '""'
\tr1r.t1r.ory Em(Jnwtogy, 10. J.19-151
268 1'TCltEU.Jt 1 1988. ht\Jd)' of th; c!Trct of 1lck ,nrc,1:ulon on llwwo1~!11
g;tJl'I or cat:teo in thl" Sudmt Tmpit./J.I Pt'.sl Mm1(Qt1.1nu.•m. 34. 16>-J G;.
?SP sou>A DL\S. ,,.. 195<>. Sub'1dlos para u c,,1ud<1 do• L,od,dto• de '\ngola.
-::t,9 nu nu,.,•• 190s. Trlliism1>:oi11t,n and lnorulabJlit)' of Splritlum 1Ji(•l/1...n
P.-c1wn11. 2. 127~280 09H-J948Hllcv11111 J>£1. 1-154,.
11.a\'Cnm . Pr,wvl/11,:s of rl,e ROJirl liorn•r_v. ,U.
50'1-.';1!6 •
.2,51 S.V!Cl..£1"1' \ \1.. Df a.L!.JU;, 0., t..'i"IIU~. C II. IL~llOLTl, '.\t,:'\l. lq~
~;o nu 1UJ1 A ,,os. rraJ\"-tnh,.,fon of~quinr piropl.a11,mosf~bl :f\~\. In
Rc:1;,~Stanc1t or ~gunt. Borumi!Til Q.Od Hc.rerord caitl~ co ncks m .a
SCJu:h r\lrica. Jor,rnul ojti~mpnmtitv PmlwluJO' and Th,1rop,>uru:s. Ht
Bushn,ld reg,on of South Amca, Omlfr$1<f)()(Jrt /011mttlt1fl ,wm:111,· 283-29:?
/lr<ectrr/:, 56. ?-1!'1-250
r,1 n-rr 11.r1t. , .. 19oi..), Trnnsrnir...~nn d<'."~ splrtJJ4,.•s ~, dt•s. plt'Qpla-.:mcs par
252 ,,•1Cliil:-rT, \.M, 1, ur,~t. u,. t.988. A ~Uf\\.j'Oi K..uoo tfd, p~ral)-..l\ tnSOuth ,1ur~,cnt<S t><Jl<.'<r> dt' tlques. 8111/,•tlfl dr S/><"(~111,1,. 1'1>1/,0/Q/(f, f.\Mlq11,.
Afrfci\ o,,,Jrrstqpoort Jo,,rnul o/Vr1eri11nry R,,.search, 55. 89--~\:?.. 2.2ll3-~9l
The
233 t;:pir.nTI \ M JIOR.U: 1,f,,, \'.\"'',it.UH~,>. A Dfl"-"Cl..1L.-t,.O, 199.J ;:.;:,: t10)1 rK. t... 1949. Zoolu~c.11 S\m-i.~ uf 1h"-' Unto.r, oi· South.~fric.a Titk
,/Tea of veld-buming M the -nu! abund,nce offrtt,ll\1ng t><>dtd sunc,\': P.ln 11-··Hiuribuoon of BIJOpl:fflll ,l'a/pol.llK>phliln) d,YO!aru::u.
lkk, 4) do,cnnined b\ dr.,i:,;amplin;i. 0111l,•r,1tpaor: /,;111nn/ of th4,.1 btuo tide. Omk•rac;UJor1 Joumo I "f\l'ct11Yitu1n•-&i~11«. 22. 25:'.- :?68.
1
\ tterinruy llt'.-<arrh. ~- 28S--29:t
l?7J 1n1r111'~ G,, 19~2. 11:v lxodairltfl Pmaslcf"5 ofVur,1bmu,s ,n .Afr,,a south nJ
2311 :,Pie.Mn, ,.M ..... ,1At.AS, ,.a., l!J78. Ccne1l0 lncompatibfli~ b~"''°''n rllL' S<,lmm (flln'opit111 Rf!g1cm.i Projeci S ~)~. Rcp~H1 co 1he t>irct'.tur ,,r
/J()(Jph//tu tfrto/1,mius ,Koch. 18441 and lloophl/us mitrop/11, \'etenn,1,cy ~"'IC~. Ondl't:tltpootL
CancJ1111ni !1188 und h)-br!dstcrilll}· ol ·\u,:rallon >nd Soulh .\frii.-.m
:u.: 111l.t\:A,. $,.t .k ,1,"o' t , t98t bola1ion nnd 1mn~ntl,-slnn 11Can
/k,c>p,1Jltu 1111,,,,,1/u., \C3.rlna: J.xodidacrl Ondf!IJ!f!JH.Kr1 Jour1 u:l of
unidtnt1fit:d Ba(,-'jm ~p. info(ti\'l° fur c.anlc flmh~rsr~poo, r /oJ,mnl r1/
I ~!dmra,;, /f~.11>,m:h. IS, 149-IS3.
\ ~lc'ttnllt)' .rl~·,rrrJt .l3.. 155-158.
:a53 ~T.\Mr.,. s .. 1q59, l tck pnral~'Si~ in the Kam,~ Jre.ts o! S<,uih ,\frlc-o
l7:1 11h"IM \~, ,.r, ..1, ~uTx.. w.a., 1958, Rhipktph;di~e tlc:k tOXfc:o,i.c hi c~ttlr·
011d,,.1,•prxm /011,1111/ oft ·4117/na,)' R,•s Mrrlr, ll, 169-22;,
lb po..1il>J~::swa\'2dng~t.~Ul'I Ccrt:a.ln dbH'::l:tt.~, Jm,r,u,/ o/1/1t• ~qt11J1
:i~& iT,\~1PA. "' • nu ton. I\,, ,958. Pnratr~tlri nf "to,·k du\! 10 tht.· l,;,1rf>U -~fnwrr i ,r,rl,rnr; l/i'1ffn1I Aurx:imio11 29, 39-SO
p,,roly~, uci,. •lnxl,n mblcwu/111, Xeu.J. /i/Jut/1 A{rt<an /0,11,:it/ oj f.l.." RO.\tUU>. 1,N .. 19:'8. ·\nli-t[c~ gr.»-n'i n_', :hl'
2·1 b 11111\U'Sf.'>S, J;,C.. J40.\,
Sdr1J('~. s... ;t.j 1-?-i6. ba,r-s for du\'rlopsn~ pr"'nic-.:il rrop.fcaf tick ctmtwl p;:ck.11R~ rrupfrul
~~; <ifl>L ,~ \,'.It ..l'f9..J Tr.-.n~mission nr F.hrllrltta ll0111J. h~ RIH'pict/>lttllu5 ·tnimn/ f/ealllt am/ PNJduct/011, to. I 79-182.
~pp, ln South-,\l'r1rn. Juumul ()[site SVwh !l/i'l1.".t1•1 ttec,,,J.Jfflt:,· .'U:ioclntlou. m 1110,1~~. G.,.. J985- Thv 1.lpid~tn1ology nf ,,Mean S\\iOC' fcwr rhr roh• of
65. 159. tr\!c~h\1ng, host~ m Alric.a. Onders,tp,wrt Jourml/ o/Vt•H•r1mu; Rt:1•"11ti:h.
:tj8 S.11J'ttlRfffl)~. ,.~t.
6' ''" 'A.Ob.. tc.. 1!)65, lloOS:tingoe'h3viournnd hflSt ~2 :!(H-209
j.(•ltclion <>f O'<l)l."C;.e~ tA\'t.'5: BuphJ.Wn,tL' in :..tortnti WilOllfr R~~f\l:l'.
8ots\,·,an,a: ond ea-.trm Coprivi. South \\e~tAfnca. So111J: Ainw.11 /uurntJI
:no mu ""t-RG, c... 1~1. nn:ilC"nJ.1 .spcde., u, do1m•..1lr ll\~10clr-. /,:: IM.\1,,
\,o, cu.sst~Gll.\" '.\t t• ""vou,<,..\.~. (td~}. Ath•m,c~ In tlri.TComral <>/
o{Zoolo~·. 20. 2,:-2 10. J'hol/mom. The 11.,guo, Boston, Lqndon: Mortinusl\ijhotl Vubh>h•<>.
:i.;!i MIi m ttS:1. ft.W•. 198i. ,~ 1he Australian J'Ht~l ma.rugem~nt up-proat:h to :?'79 wu,n .. K<., c.. l.93$, -\\iqul~irions nou,·lllle<i 4.1.uu, t., connai",11cc du r61~
lki..control rr!t.\":m, co \frrc:i? ,,,~wmn11un. <,.a &r..-reso~. I.IL eds' ,·~cteur de :iquc, du scrue ~ml>/yt,mmr1 fbt1>i.lld,1eJ. fwm~ d'tlc,·11gut If,•
Tick &'OIOJt.·mut Comrol. ProCCt'tlf11Jl~ oftm l11tfmatfo1wl Co1tfi-r,•,1r<. ,\/.',/,'('/11; Wtimrr,/ivdN P1>J• fttJP1<YIIAA, Jli. 61~L
21-29 JanU3J) 1981,Rhodc-., Unh~t5it). Cruh.\mt-tuwn. South ..\fric:.1
~ u1u~tffKG. c. "A~'t>RIA't-~ M.r.. 197,i-l(,u•mmu.umu Jf!Jfflrmu, "P· n.
19..135.
(~p1,rw,>;1, T:icllerhda~ . • nc" blond pom,ite <If dome:<dc ,t,...,p 1n
~, S\nu, a;.1. N.w.. IO!t. Tllt'drnamic:, C>fhyhml 1..onl,sbcmt:Dn 11r\: {,\earl) r:m1.anl:i. Rentt• tl'P.l,•1·t1,C!,( ,•r d4! \ltf,i«im· Vtt.!ri1111it~ des Pny,
.sr,ec:ies. hm:rnt1tit,1wl /outn11/ of Pnr11slt()ln~·1I;, 921-9.26 ltvplcmix. 27, 4;19-165.
::.61 ~u·n.1f.JtST. PU\ "'OAUWJT1.~ ?-1.J. J.97'9. l'topres.s. in the ~f\'CIOJ)ITil!nt oia :'!81 Ullf.-;a.nFK'i, (.., Pii:ftl(. ~.\I. U\\'kf.~tf. J,.\ .. t>I! \f0S. \,J., V.W.SG. rt..\\',&
populuoon mod,l fonhe cmtle 1ielr. &>oplrllu, microp/r,,, ln: ... 1,,,u.n•.LU!.. 19b4 C;au;.aJ o.\~n~ ot hO\inl' ,hci1niD.sb 1n 1'11'1Uth~m
Pro.wdi"I!' o.ftltt fourth /nrm,a11ouar C:t>ugr,.s of. lc./lrologv. ,\u)!Uil ,\frh:--1. Tmpf(Ol.~1tjmul 1la1l1h mul Pr1X111rtfnn, \,;, 127-140
197~. Saa!Mdcn. i\llMrt.l.
23:: UU,.[Nl:fU«:;, C. KOR~'.\\ f. &J•I D6A$t:,. v .. 1974. ~mt expe..rimtnJ.> on tht
25::.~VlHl!RST. R,W., flC:ffO, tt:.a.. MAY\\"AI r,, O,f. f;,\'(i0\1J\!lriS. A.r, 199,8 1r,msm1sston of 11,,.,/,.ria muums tllwil"•r. 1900> 1md TJ1,tilrrft1 part'lt
~todclllng for rn.mngonu,m orAfiian licks 111. ~xs. ,_ ~ ~<>111"u1.0. Cflwller. 19041 h)' tlw tl<ks .~ml.>l)'Olllf/111 twrf"l/QJIUrJ CFabritii..,; 1;9~
,1. (&di). Tile Suond ltt!enwrlo11nl (',on/11w:c·i· (JI: Ticl..··bomt Pmi:o,:eui .'.Ind llhlp(~plmlusopp,•ndfcutmus ~eumann. 1901 jn Ugandn.
al 111, HMl· 1',>'IOT /111,r[nwn G/1>/Jo//>,rspt'l'til... August 28-S•pi,mb<>r rw,..'mm~ditftt zmd P1un;1iloftJgfe'. 27 3:?3w:.t28

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42 s t = OM.: Aspects lnHuencing the occurrence of infectious diseases

283 Utu:.XimtG. c. & !!CllllL'UOt:R. u.u.c.. 19;6.. Studll'S OJ\ Thdleriid3e \'3ccimtion .-.ga{rht 84,QpJ:ilu.s mlcroplus-:Jnd ~·Jia bot'/$ u.~in~
(Sporotoa) in Tanz;,nht. l. Tkk trnnsmissfon of Hmwumul'nus L"e'/£furw.. rocomblnrui1 .nrfgens.. .Wcmorin, ,to Jrw/11110 Orwaldo Cm:.. Rio d.-
Tmrll'l1111editin 1111d Partuilol<1gi•, 27. 106-H l. Jw,rlro. a; 'Suppl 11ft, 289-"..9~.
r.o.• 1978. Resisnt.nce tO BOQpl,i/u$
28.1 UTf.CH. -= .•.\,~.. \\TH,\IHOX, It.It. & li..'DU\, 2.99 wn.LW$t~. 1•.• Al OIXC:. G.11.. ~tr:1a XX-A. M.\'. ft Kl MP. n.H .. 1989.
mfm,pfw iCanes,rinl) In dlll'e<cnt bl't'1'ds of eo~ule.,IU.ttt'll/tllnfoum«I of lmmunolog,clll con1<01 of o porasillc ru1hropod. ldQn1lfteation of o
,tgrlculwrnl RMMtl'II, 29, 885-89$. prot1.."Cti\1t antfgt-11 lrom &:>oph/lus mk:roplw.. J(,u1nal oflm.nwnology.
285 ''"-" Rt.'<sau,,c;, s.• 192a. Tork para!~ ln ~http, Fanning in !,outlo ,ifric,i 143. 13'1>-t.351.
.,1.,cb. 661~2 3,00 ,,,i.sos. LJ.• suntutn n.w.. 1990. Oviposi.I:ion sitCj of Boopililll!
286 VA.'< ~,=e.,.
v.o .. 1ss1. Th~ tcd·billcd oxpockor and 11, n:la1!on 10 stock mictopl1JJ (C.1nc!itrinr (Ac:srina; txodldaeJ 1n !JrJ•lnsnnthtt .and g.ra...~
In Kenya. EtN Afrk<111Agrlc11/li1rt1//011n1t,I, 17, 1-11. p.i<tures. /1J1,r,wl of tilt' ArlS"alitm Euu>nu>logit:dl St.w:irty~ :?9. J01-10S.
~i \\'Aua;R. J.n.• 1.91,i. Tiu: lxollitl 1)'rl:s ofKNtjrt .,t Rtwl,•J,•OftJrdSl!III JOI ll'.O>HN, <;.H .. 19.a. Field ,ooor>tudies or oplwo"c 6.rn c«lst '"''"· \1.
w,ou,:li:dG,t10/ tlt~ir Um.ts amt Dl,s.trlbution. r.ondoru Com.monweaJth Ttu: Ott\:.rrenttt of Atflblyomm,, ,,,.,,;egmum 1md A. lepi.d,.mz in the Ei"lSt
ln(titute or Entomology CotlStfe,·cr,on,,s. 811/Tt1111oj'F.pi:lomic n,,..,,.~111 ,\from, 16. 183-203.
288 WAUrf,R. 1.s.. ,'Euuss.1.L & BORA<, 1.c;,. 2000. TJi~C.mlJ 1\hiplcephalus n,~ /xq,//d 11('/.;s o{Thtt.11/100. ,4 S:udyof
3<>'l 11'0~1\S,G.tt ~ \\'\lJ;fR, 1,8., 1967,
(/otan, f>'Oliilf111:1: n G1111/e 10 1lie Browr, Ti<ks of1/oe W,irld. Urute<l the Zoo11,ogroph;-ofllu• t:todid~ofa11 /!4$1 AJf(t;(m Counr,y. London:
Kingdom: C'..ambr1dsc Umve<S11y Press. ' Common•,,-e"1th I11>1Jrn1e of En1omology.
:t:89 Wf\.l.KEA, J,B,. ~lltA~::t", 1,1., J'F(';RA..\1, R1V, & CUfll(ifW, r..M.. 1988, 303 VODCR, ,,,. .. ~Tl.\'tX.S. A w, $ CKQUC:11, ,u:.. 19~9. S<.1ul.llin~: :t muurally
Clarification of the st3tU5Qf Rllipfctpioo/111 mct11pis Ul>nit>" 19011 ond obundnno mamll1'1ll1n sklnS<.'(;rcuon nnd long dh<31t« uck nn,ncum1
ll111p1rep/1alus /111111/nwsl'/cunumn. 1907 llxo<loodca. lxodida.e l. L~rati; lxodidMl, /1J11mnl OJ Medlen/ /i111,,molo11Y, Jt;, 52o-S29.
S;~fCmMiC Pnm~/101',gy. 12, l:,~-186.
JO• ,ou:<G . .1.s., ,,e,. Toe epodemlology <>f thelleno,,s In Ea,1Afri~•· In·
290 \\'At.nA, J.l;I .• '\1JlHUTZ. n.~ JON~. C,E... ,918. NOlt.'1 on thr Ttcl.'$0/ lfW1~. ,,n.. C\J",:\l:O.<it1 \N, M,P, &'l'OUSq, A,S., ,ed.\1,.-\dt'C'lt/~i In tht
1

Bouu,01,a Ccnnan ,\g4,nt,· forT<'ChnlJ:al Coop<ratlon. Wt•,1 <Jcrm,111. Control ofno,·1/rriom. The Hague. 8osroo, l.ondon ~fininu• Xojholl'
:9.1 \\'AlJQ;Jt, t.8•• .S01C'\'AL. tl:•.A.l.. & C()~WJN, M.•J>.~ t98t. R11/ptc.:plt4114 P\lblL<h•r,
::ttflJbe::l~nsis sp. nov.• D n~w Ud. from casttrn a.nd ~mhern Afrlca.. aos iuu,c.. A.>- ,~a;. ModeUing 1hccplde111iofo8)'of 1hcll~rl~-slY. l11:
tog~11hcr \\ith a rc..•tlt:scriptlon a( Hltiplcc.•plt11ltn apJNJ1dfet1laru~ >UTHl'l!ST, ~.w., red.l, Ti,k,and Tick-borne /)4,•as<.s. Proc.•rr/fngsof011
~tumanr... 1901 (Ac;ufntt: b:odllfae). 011dr~tt!ptl<>rt Jtmninl of\'curn.,mry lnt•marlfJ,101 WorkJloop. 1,-21 Fobn1oo; 1986, Xyanga, 7.iml!•b"•·
R~•t!llrt'II, ~8. 87-104, C,nbem: ACIAR
2.9:.! ,,,·.uJ...U. r..a. '6'01 \\'AGll, A.• 198';. Th, lick \'CCtOIS of Coudrla 306 \'C;)U~G. ,\,:,,., GM001Y.....:Hut:i. J,(.,•• ~ 1,uU,k. c..u... Ji:.\~HAf, C. "· 4- q,\c;c:, 0.,\.
rumlnanrfom lxoclofdeJ: lxodld:te, GcnusnmiJlyommaJ ~nd thvlr 197,. l~la\lon of• n,,.;/ma species from eland (Taurotn;gr,s or,;,!
dismbution. Ondt-rs.JcJpoori /11unu1l of Ve1e1i,:an• R111,1'(1tch. 5·1, 3SJ.-379 infertl, • (or cn1lle. l.'r/1scloriftftir r,0,,..111111'.di,:,n ,md Para.<1.Jo/ogiq, ?7
::g:s w \I.Kl R. J a. & scuuu:, ~.<:.A,, 1904. Records.or 1hc bom uck. A,nlJ(\'()mma lUi>-194.
lttimumm. from ch,• nnguJatt l()~(liS(", Ch"'''"'' <mgu(nUJ And 1t,f.'
107 l!E(:Eft, u.. HOkA,. '·"· 4< CAUU)WH1., ...... l99H, O)'n3mlr. oHr,e•tMng
leopard :ono1$t'. Gc<i<loe/011< /Jflfdo/Js. Ond,r;,e{){Xlr1 Journal of b(c,dld ti<I:., un ,,game rnnch In the C.ntr.tl Prtl\'ince. Zar:tbl:~
\f,:m,ool)'lloscnrc/1,51.171-173. 011der,1<poot1JoumQ/ of letcrln~7 R..,,111(/1, 6S, ,;9-;9
,o
294 \,'1lLU)S.Ex. "·· 1986. lmmuno!ugkal -appro,tc:h~ thl' con:rrol of dck..-... JOB nl!GI!l\, u.• UQM,,\)., i.e.., t-.AULD\\~U.. ,U:.,,. u..,. \.C., 1998. hodid tick
/11: "II" c11. >t,!.. 1ed ProcewJfng, of,,,~ S1<:it /111,malional Con1trC1,,<0f
on(e,aotion_, u( wild bird> and ,nammols on• gomc ranch in Cenir3l
Parasitology. Au,omlfan Acndomy o(Scl~ncc, C.nborn.
Province, Zambhs. Oml,1f$ltpoort /ourrml o/V1tltrint1ry Rrsear<'h. ll.5,
~95 "'IUAOSl:<, P.. 1997. Va(cines, genetf<:< and thcmicnl~ ,n 1lck ~on 1101: the 113-124.
A11s1mlfan expc1l,nce. '/top/(1>/ Anlm11/ H,a//h nt1d Prod11ctlon,
.3"9 'l.ltG!!.R, t .. HUM)., ,.c... f.AVl.D\\'1.U... c',-1 .. , U\"' ,\,('.. " DUtH~'tA.. ,. nu P.. t~98.
~11pplcin,n110 Vufuntt 29, 91S-94S.
1be eife<t ofrh~mlrnl 1lcl< eontrol on eioule on frtc·llvulg lxodfd 1tcl:,
29'6 wt.Ll.AD.si.~, 1•. "
JO!\ r.r fi\.S-. r.. r999. I mm t.m olOg\' of tbc tic~-hos.t ond 1111 tic!.$ pom,.,llc on wmp31rio omp.,la in 1he Cemrol Pr<"in«
ln1omc1i11u and tit• coni.rol of tich and dcl.·bomc di>=es. P{lfQSi/oloC,)' l'.wnbia. Somli ,lfri,;,,11 /qll1t1Qlof\\11/JH1fe h'-1rrl1. 28, 10-1 ii.
Today. 15. :!.58-262
JIO '/,f\"):0\'?C. D., J'f(;IU'.\h ft~ .. ,o:i:ur.Jr\l\. V. 4, MW,\tr. E.T.. t!,86. Biology or
29; ,,1UA.D:tt.~. P. & Jo:t~\tP, O.lL. t988, Va:cdn(lliVn \\ Ith conet~C'd' tlntlgcn.s Rhlplcephn/11., 11p1><'rorfirul11111und II :11111/Jr.;/rnsl., Jnd pn1due1io11 of•
fo, tid,conuol. PnrosirologyTodny. 4, 196-lll8. tcnllc hybrid under fobomtori,·cond1tion1. F.!<11<-rlme11rnf and Applied
:.198 WTUAO\f.~. P .• t:l!).IP. 0,11,, f.Oij() S, r..s. ir \\"RJC'iH1, t.t,., t99!t. Succ~fuJ Mt1rofogy, 2, 285-2.98.

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 2
Vectors: Tsetse flies

R J PHELPS AND D F LOVEMORE

Introduction Biology of tsetse flies

Blood-sucking muscoid Diptera of the family Glossinidae and Anatomy and sensory sy-stems Detailed studies ha"e
the genus Closiirltl, known as tsetse flies. are restricted almost been made of the anatom/6· 161 and 1he physiologyll3 of
entirel~·toAfrica and some of its off-shore islands such as Zan- 1se1se flies. Leak1u bas recorded the advances in knowledge
zibar. although they are absent from others such as Madagas- of 1hc biology and ecology of tsetse mes over the previou~ 30
car. There is one old record. published in J906. of tsetse flies rears.
from south-west Arabia, 29 and a recent one nearGi1.arin Saudi Tsetse flies (Figure 2. 1) have one pair of membranous. me-
Arabia,61 but they have not been reported there since then. It sothoracic. functional wings, while !lie rnctathoracic "1ngs are
had long been known dtar domestic stock could not be kept modified ro ionn halteres. They can fly at abom 6.3 metres per
where tsetse flie, occurred and that disease wa, associated sccond71 and the power of lift generated by I.heir wings is un-
with theOies. Ho\,·ever, it wascnot until Bruce. working in Zulu- usually Iarge.90 1l1e age of tsetse mes may be estimated from
land in J 895, demonstrated the 1ransmission of pat.hogenic the amount oi wear on the hind margin of their wing~. while.>
trypanosomes of livestock by tsetse flies that lhl! reaso11 for the length of the vein forming the blade of the 'hatchet· cell
li\''l?.stock deaths became known. (ln this work. Zululand refers (Figur e2.2) can be used as an index 10 the size of the fly. 1" The
to the lowveld region north ofI.he Tugela River 1n KwaZulu-Na- energy substrate used in flight is proline.22 and lhe s,ored en-
tal Pi·o\1nce. South Africa.) The role of tse?Se flies as vectors of ergy is in the form of abdominal rat.·n,e amount of proline
human 1rypanosomosis, and the importance of Clossint1 pal- available to the flight mu.~cles in the 1horax is limited. so tl1e Oy
palis in this respect. was also demonsmued b1 Bruce and his cannot make sustained flights. 'Nhen this fuel is exhausted. the
co-workers in 1909. 1• fl)' must rest umil further proline has been synthesiJ.ed in the
The 1se1.Se fly-infesled areas of Africa could not be ex- cells of the abdominal fat bOd),u and transported to the flight
plored easily before lhe ndvem of mechanized transport, as musclos of the thorax.
animal-drawn transport was cxduded by trypanosomosis. The functional mouthparts, or haus1elh11n. of the 1se1se
The presence of tsetse files was thus a major obstacle 10 the Oy consist of the labium. which is t.rough-like in cross sec-
development of much of the condnent. a situation which tion and lenninaces anterior!)' with the demiculate label-
pertains in man~ places even today. There are those. even lum; the labrum. 11ing along the dorsal surface of the labial
now, who look upon the tsetse fly as the guardian of the lrough: and the hypopharyru:, which lies within the tube
natural ecosystems of Africa. and who would like this Oy to formed between the lubium and labrum and which has the
remain until humans have learnt to manage the land tn a salivary glands attached t0 its base. There are no mandibles.
sustainable manner. i\l presem. though, the human popula- and the maxillae are represemed by the siout maxillary pal pi
tioJl of Africa is expanding i:apidly. and the economic situ- {Figure 2.3) between which the baustellum lies when a1 rest.
ation is such that the burgeoning pc>pulation can, for lhe The haustellum. or the haustellum plus the palpi is often
most,part. be accommodated only as peasant farmers. The termed the proboscis. When the fly !s feedlng. tht' haus1el-
importance of livestock co such people is very great. not only lum is lowered. die labcllar teeth cut into the host'sskm and
as a source of food . drauglu power. and money. but also for ihc hausrellum positioned in the wound. Blood ls then
the importanr role that livestock. especially cattle, play in sucked up the central lumen of the haustellum. and saliva
cul1ural alfairs. Under these circumstances it is essemial re containing an anticoagulant Is (eleased imo the wound. The
try 10 eliminate. or at lease control. 1he diseases oflh·estock, labium ts well supplied with sensilla which monitor the
amongst which trypanosomo~ls plays a major role. bloo<I now and qualily.
43

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4-1 ,,rno-. ,M: •\.,peels influencing ili~ occurrence of infet:1lnus disuases

\. 7

,,

'
'
'

.•
!

·.,. ';,,, ·.

'"~" '....
r
Figure 2.1 G/l}Ssma mors,1ans

Temperature of the host" s skin is a primary stimulus for the duced the numbers of mes attracted 10 the ox. but did not
fly 10 start probing for a fecdiligsite, ruid adenosine-5-triphos- nffcct the proportion of flies feeding on the ox after arrival.
phate in the host's blood stimulates the fly to engorge.6!1 Only in areas of very light disease chollenge would the repel ·
Both sexes of the 1se1sc fly arc obligate blood suckers and lem be of use in pro1ecting animals in the field. :--:on-volarile
generally feed at three• to five-day lnren'als. although they substances may al~o be detected by sensilla. as is shown bv
rnay feed more frequently if the oppom111icy occurs. The the initiation of copulato11 responses in the male Cly bi the
hloocl mpnl is 1hP ~nurc:<> nfhoih murhion nncl \\.11rr for 1he romarr of its tihi~I r<'ceprnn. 1\li1h fl'nrnlt> sex pheromone. in
fly. Food reserves in the form of fnt, and the amount of re- the form of a wax on 1he surface of the female's thora.'-. 1~ '
sidual blood in a fly. give an indication of its nmritlonal sta· Well-developed compound eyes arc present in I he rsetse
tus.~00 to which it~ behaviour is related. 11 fly and are important in host loca1ion. I lungry nie<. are par-
Tsetse flies can detect odours by means of sensilla situ· ticularly attracted to mo,ing objects. bu1 a1 shorter ranges
atcd on the antennae; if their antennae (Figure 2.$} are re- they can also rec-ognize stationary ho,ts prO\ided these are
moved. mes are less able to detect their hc>sts than are intact large enough to be resolved by the l')'es, which have an ab-
flies.nu Substances ckairomones) produced in the host's solute resolution of about three degrees.231 The eyes can
breath. such as carbon dioxide and ac:etone. attrac1 tsetse also distinguish between light and dork. a useful attribute
flies.Z35 while a f,mher a11racti\'e component of cattle breath for seeking out shaded microclimatcs when ambiem tem-
is l-ocum-3-ol.24' Kairomones in the urine ofAfrican buffa. peratures are at lethal levels. 107 The spectral sensiti\it)' of
loes and cattle which ottract tsetse flies have been isolated the compound <')"Cs of tsci,,c flies ha6 been in\'CStiga1ed84
and identified as phenols. 174 • 2 1" However, some and it has been sho\\11 that it is the spectral ceflecti1ity of
kairomones are repellent; Lhe odour of humans repels some rhe object wl1ich is detected by the fly. nlue-green and red
species of i:setse mes. notabl}' G. p111/ic(ipes.234 One of the covers on Oy trap~ i11crease their capture rate, while grct·n.
phenob presem in ox urine. 2-mecho;,.yphenol, is repellent yellow and orange covers have 1he opposite effecr.ll--'• Com-
to G. pallidipe.:s and reduces trap catches by 60 per cent.226 parison with a graded series of grey traps indicate-Cl that the
Placing a source of 2-metho>.-yphenol ne:-."1 to a bait ox re· Hies employ some colour information ,md not contrast

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 1mm

Figure 2.2 W,ng of a :setse !i\. w11h tile

ha1ther cell indicated bv st,pp, ng

ological age of flies. as the shedding of eggs from the four

ovariole, follows a regular sequence.30• 205 The rate at which
the eggs mature is a function of1empera1ureso. If the ambi-
ent temperatures are 1,..,,own. an estimate of the chronologi-
cal age or the lly can be made from its physiological age.
Man}' aspects of tsetse fly behaviour have been investi-
gated.11· 12• 6 1 221 n -. 251 :.everal of which have been used
Ansta of antenoa to devise methods for their capture. When a full under-
standing of their beha\iOur has been attained. It may well
lead to improvements in the presen t me,hod<- used for this
M2x11fiar, pal01 putpose.

Life cycle and population dynamics Female tsetse mes

emerge from their puparia about two clan before
males 119 mi aml, aJter expanding and hardening their
wings, they seek a blood meal. In some species. females are
receptive 10 males as soon as they start seeking food and
Haustellum
often mate either when taking their first blood meal. or soon
aft<.>r. Female~ acti\·ely resist mating more than once.120
They are viviparous: ovulation taking place about four days
after the first meal 11·ith the first larva being deposited about
10 days later. At deposition. the larva isa. the end of its third
instar (Figure 2.5), having been fed on a nutritious fluid dur-
11mm I ing its developmcm within the female. Each subsequent
larva follows at an inter\'al of about 10 days. The duration of
Figure 2.3 Lata'al ~sp~t oi tile head of a :se!Se fly with ;he haustellum lan•al develotllnem in the laboratory is temperature-depen-
lowered 10 the feeding pos,tlon dent. and mar be predicted by a Formu la.~3 l'he female feeds
at regular interval~ throughout her life. and if not \\ell nour-
ished, her reproductive perfonnance declines. 70 • 160 It has
infom1atlon aJone. The same effec1s have been obtained been shown in the field that high host density. resulting in
with flat o!oth screens of differem colours:e:? black is le~, at· very well-fed mes. may cause a reduction of the interlan•al
tractive than blue, but more mes se1lle on a black surface period by 1wo dar~. relati\'e 10 that predicted on the basiS of
than on surfaces of other colours. mean ambiem temperature.9 ·1 Females live longer than
Eyes contain pteridine pigments which accumulate with males in the field, and muM live long enough to produce cwo
age. This feawre is now used to detem1ine the age of mes of mature Jan·ae if the population is to remain static. and more
both.l>exes.130 than that to pro,1dc for mortality and population incrense.
The sexes are easily recognized in 1Setse mes as the male The newly deposited larva moults to form the prepupa. hut
h)'j)Opygium is well developed (Figure Vl) . The male clasp- remains within the shed third instar cuticle. which then
ers are concealed \vim in the hypopygium when not in use hardens to form the puparium (Figure 2.6) ,,;thin an hour of
and dissection is necessary to displa} these stnsctures, lan•iposition.267 Lan·ae are deposited in shady places "tjth a
which are important for specific identification. The internal soft substrate. Within the puparium, the prepupa moults to
genitalia of females may be used to derermine the ph}•si - give the pupa. and a final moult within the puparium gh·es

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-16 ~=o, o~,: ASpeels influencing the occurrence of Infectious diseases

the pharate adult, which emerges from the puparium in

due course, the duration of the puparial stages being
rernperarure dependent. The insect does nor feed from the
time it leaves the female fly as a mature lan·a umil the
adult emerges from the pupa, a form of reproduction kno1m
as ad ... notrophlc l'ivipai y. t\t a commmc t~rnptmHurn or
25 •c. males emerge after 30 days and females after 27 days,
while the respecth-e figures at a constnnt temperature of
16 •care 101,8 and 100 days. 179 The integrated mean tern·
perature from a Stewnson screen gives a good representa·
lion of an equi\'alent constant laboratory temperature 3$
far as development within the puparium is concerned. 178
The life cycle of a 1sct~e fly is sho,wn diagramatically in
Figure 2.7. 1mm ,
Male tsetse Oies emerge in equal numbers to females 1
and, after expanding and hardening. seek a blood meal. AJ.
Male
though the males may copulate soon after emergence. they
are no1 fully fertile until they are a few days old. [n the field ,
males may possibly mate more than once but, as spermato·
genesis does 1101 occur after their emergence from the pu·
parium,~• 1heir sperm reserves arc soon depleted and the}'
are general!}' considered unlikely ro mate more than once.
Males feed at similar intervals 10 females. but have greater
energy reserves for flight as they do no1 have 10 nourish lar-
vae.23 In general, the mean life span of males in the field is
about four weeks, while that or females is at least eight
weeks. 119 but these durations may vary from place to place,
depending on prevailing en\~ronmemal fac1ors.
The popula1ion dynamics of tsetse flies have beenJnves-
tiga1ed for many years since the development of the mark-
release-recapture method b}' Jackson. 1 1" This method de·
pends on marking flies with dots of artist's c,ll paint 114 and 1mm
1 1
1hen recapmring them. Capture melhod, range from c:uch· Female
ing flies from a mobile or stationary bait with a hand net 10
the use of variou~ C)'lle~ or craps. manr pauerns of which3"·
91 232 238
• • have been developed since that used by Harris.~8
Figure 2.4 Venua. aspect of tne terminal portion o! meabt omen oi both
However. no known capture method gi\·es an unbiased sexes of G/cssina mars,tans
sample "~th respect to po po lat ion density. sex ratio. hunger
srnge. or age.'43, 1s,. 200. 2<1u
In view of the sampl ing biases, estimates of population Ecology of tsetSe flies Tsetse llie~ are distributed over 11
density of tsetSe flies vary widely. In addition. the assump- million sqi1are k.llome1res in i\fricn between 14 •x and 29
1ions underlying population estimates are often \iolaccd.25<1 •s. The northern Ii mies of latilUde are highest on the west
Consequently. the population ecology of tsetse flies has coast. while the southern limits are grea1es1 on the ca~t
been reviewed.zot. zoz and a new analytical approach to coast, where the 17 •c effective temperature isotherm 21 i
population ecology has been developed. 198 gives a fair representation of the his10rical limits of the
The rate at which tsetse fly populations grow is 1101 1setse fly. These limils are detennined by climate, often
known, alti1ough a mathematical modelling approach has through its effect on vegetation. Ale hough the tsetse flr and
been developed which indicates the limits \\1thin which the its puparium are \\'ell waterproofed. 18· 19 regions situa1ed
population gto\,·th rate must lie under gi\·cn environmental below the 500 mm isohyet do not support 1secse flie~ unle~s
conditions. ba$ed on current knowledge of tSe1se fly biol· there are watercourse~ wi1h some vegetation along them.
ogy.92 It has been shown that no population of tsetse flies Temperature is of great imponance in the ecology of the
can survive if the d:tily mortality of females is greater t.han tserse fly. Not only does it affect imerla.rval and pupa rial du-
4 per cem, <llld in the field possibly 2103 per cen1.9Z This is a rations, but it also influences the ny·~ acliviry. Below 15 •c
very useful criterion for those concerned with tse1se fly tsetse flies are inactive and above 35 •c they seek out refuge
eradication.

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 Vectors: Tsetse Illes -I 7

Posterior respiratory lobe tsetse fly distribution is limited.2 1 T11e larger fly species are

I not subject 10 chis limitacion to the same extent. Tempera-

tures over 40 •c are rapidly lethal to both large and small
species of tsetse !ly1 13 and their puparia.180 Increasing alti-
tude moderates remperamre, and this is reflected in the dis-
tribution of LScLSt' mes. N.:-dr tht, eyuator 1hcy may occui U!J
to an altitude of 1800 m, but in Zimbabwe, which lies fanher
south, Lhey do not occur al altitudes above I 200 m.110
The vegetation rype at a given locality is great!)' influ-
enced by temperature and humidity. Apart from gra..~slands,
which do not support tse1se flies, all forms or woodland,
from savannall to rainforest, can provide a suirnble habitat
for some species of tl1ese flies, but no one "egetation cype is
suitable for all species. Artificially planted plantations can
provide a suitable habitat for tsetse flies,:iao and the chickets
·:::
. (sometimes comprising chiefly IAmana camara) which de-

\- ,
,:..
'?;·.
.>
velop on aband1med agricullural land are often good habl·
Lats for them.

Host p references and locaLlon The distribution and

abundance of some tsetse fly species, especially G. morsi·
tans and G. pallidipes. which are often referred 10 as the
game tsetse flies. are closely related to che numbers and
habits of certain wild animals. The developme111 ofmechod.s
for decennining the animals on which indi\ridual tse1~e flies
1mm
1
have fed. by analysis of their blood meals, has given an in-
Figure 2.5 Th110 1ns1ar larva or a muse fly sight into their feeding preferences. 193• 235 Preference has
been shown for the warthog (Phacochoerus aerhiopicus) and
bushpig (PoU1mochoems porcus), as well as some Bo\~dae
Posterior respiratGry lobe such as the kudu (Tmgelnph11s srrepsi<;eros) and bushbuck
(Tragelaph11s scriptrlS). Other wild animals, includlng el-
ephar11 CU>xodoma nfricana), black rhinoceros (Dlceros bi-
comis) and African buffalo CSyncerus coffer), are also fed on.
but less frequeruly. In the case of the elephant and tl1e :\fri.
can buffalo, this may be due. in pan, to their tendency to
move over large distances. Primates are poor hostS, and
some animals such as impala (Aepyceros melampusl and
1.ebra ( F.qnu.~ lmrr/11zllii) ;rre rarely fed on although che)' mar
beabundantm 1secse-infes1ed country. The blood of impala
is adequate to nourish tset~e flies, tl2 and 1he reason for mes

1
1mm I not feeding on rhis animal may lie in the beha\1iour ofchese
animals. 1-5-1 Behavioural reduction in the aLtack by tsetse
flies has also been sho\,11 for juvenile warthog.224 lt has been
Figure 2.6 Pupanum oi GloSSJna mors11ans shown chat. 2lthough lhe blood proteins of a range of \\~Jd
animals differs, tsetse mes can effect lvely use all Lhe dffferent
rypes of blood. 1&1 In the absence of its preferred hosts. che
sites. such as rot-holes in rrees. animal burrows and deep 1se1se fly can sun·ive on the blood of olher wild animal spe·
fissures in bark. in which they are inactive. The eyes are cies. Nucleated red blood cells from birds or reptile:; have
important in this refuge-seeking behaviour. JO, Thus the also been recorded on occasions, and G. pul11alis in pan-icu·
temperatures must scay within the 16 10 35 •c r-«nge for long lar may feed extensively on reptiles.
enough during the day for tsetse flies to remain active and Domestic animals, particularly cactlell. la.t and donkeys. 9
able to seek food. The~· are inactive at nigh1. For the smaUer can be good hosts for 1secse flies. but sheep and goats are
species of 1secse flies. the puparial duration at low tempera- not. The anal)'sis of tsetse ny blood meals from the northern
tures is so long 1hat the insects· fat reserves become de- region of the :'>iyanga district of Zimbabwe pro\1des
pieced prior co maturation: this is one of the ways in which convincing e,~dence of C. morsiums' ability to s unq\·e

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>18 , CTlr" "'" ,.>;spec!$ innueneing the occurrcn,·e of infectious diseases

•
b

Figure 2.7 Tr.e l1;e qtle oi a tsetse fly

a "blooo-engor,ged female e: nv e~rg,ng from !he pupanum

h "!al\ 'pos,img female t : lateral a!pec1 c' a :se1se fly
t'" lhtrd tnstar larva g = dorsal aspec: or a tsetse 'Iv
d =puoimum conraming the pupa h = some no,;s 01 me m:Jrs,;ans group of tselse files

almost emirelyon domest ic cattle. 193 lt is also known from 10 locate its host.lo. 11 In the case of vision, shape, move-
West Africa that G. ,,alpalls. "'hich feeds on a \\ide range or ment. colour. and light and shade differences are important
hosts. can adapt well to feeding on peri-domestic animals aspecis.85 · 99 ?ll The contraSl between light and shade may
(animals such a,; dogs and pigs which occur around the be important in determining alighting points m\ the host. H
hou,es of peopleJ. 122 As the range at which a moving ox can be seen b) a 1se1se fly
In the laboratory. feedi ng defibnnated blood th rough a is about 140 m.• il seems that odours are responsible for at·
membrane i~u~ed to maintain colonies oftset,e Oies.161 For tracting flies from greater dis1ances. 2:1o
some unknown reason, pig blood fs superior 10 cart le blood Studies have been made on the behaviour of 1se1se lies
for chis purpose. in an odour plume.24 71 87 but ruu
details of how the tsetse
Both vision and odour detection are used by the tsetse fly ny na\;gates in the µlume have 1101 been determined. At lo,,

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 Vector": l ,ct,e llil"- ~9

wind --peeds an odour 1>lume meanders and. even l Om from laboratory. by contamination of Ilic mouthpan" nf certain
lhe source. may !live an mcorrect indication 01 the diren line other species of bning flies.2; 7 :\s l1')1>nnosomc~ survini ror
13
10 the sourct> of the odour. • ijG •\1 imermediate "ind ~pceds only a short time omside the mammalian or tser~e :ly host.
lhe odour plume,> ~,rnightens ar\d gh•c, a better indication of a II) with comaminaied mouthpa rrs ha:. 10 feed on a ~u,-
the direclion of the odour source. but at very high wind cep1iblc host soon afterthc infective meal in order tot rans-
speeds the odour plume disinccgrales rapidly and il> not of mil the µara,ite mechanical!~. 1'1yp1111osoma em11si in
use In indlca1in1? the source or the odour. William, i,;,i ha.~ :'\'orth Africa. i, rhoughr 10 be rr:111smiut'd cmirch me-
modelled difiercm tanics which rnay be used by the fly 10 chanically. and T 1111•11x in ::>outh America and :0,lauriuus is
locate an odour ,ource. and ii b likely that the lli is able 10 assumed lo be tran~mitted in thi~ wa~. In southern Africa,
use a \'3.fiecy of methods. depending on the prevailing con- mechanical transmi~sion of the uypanosomes uormall~
ditions. Glossi11a pallidipe, has been reported 10 cross a transmmed b~ tsetse flies has not been unequl\·ot-dlly
clearing 9;!3 m wide. where there wa,, a conspicuou~ bu,h- proven in the field. 10• ~ 5 On!) some species of 1,.c1se flies,

vision are involved in chis lype ofmovcmem.

..
<:ov1:rcd tlllf on the far side.21 :; l'ossibh scn~es other than and thl'n only In parts 01 their di£trlbution range. uansmlt
trypanosomes which cause disease in humans T. bmcei
gnmbieme in \\'csc -\frica and 7. brn,·ei rhodesien~ein cen-
Tsetse flies as vectors of trypanosomes tral. eaMern aad southern Africa). In the ca,e of li\'l•srnck
Tsetse flies are important main I>· because they are \·cc tors of d isease. all species oi tsNse flies tested can transmit the re-
certain species of protozoan para,itt/~ belonging to tl1e ~pon~ihle crypanowmes. although indi\idual specie~ mai
genus Trypmwsomn. This genu~ i,; ,pli1 into four subgenera. not trnnsmh nil the specie~ or ll')'panoM>mes. and somt• urc:
Of thc:;e, one. Pyc11omo1111s. c;ontains only one ~pecies. T. beucr vcnors than Ollters. Tnc inrcction level or the tsetse
s1ti$, which is of minor importance In causing di,eaw in -\f- fly population \'arie, from place 10 p lace. but is general!~
rit:a. The subgenern D11tto11el/(I, .V111momo11as and T0•p11no· bet\\'een 5 and 1:; per ct•n1.
zoon comain the species that cause most cases of The c·ondilions under which tsetse tlie.$ become mfcctcd
trypanosomosis (naganaJ in ~wd., whlJe all the organism~ when feeding have been reported. m. lbS Within the t~c1sc
causing trypano~omosis (sleeping sickness) in humans be- nv populatior ~om<1 individuals arc rl!fraccory to infcc-
long to rhe subgenus Trypa11uzo011. The indi\idual :.ub~en- tion.156 11tis resistance is maternnll~· inherited and is asso-
era can be recognlzetl on morphological .ground, but the ciated \\ith the presence of rickettsla-Jike organisms in the
species within rhe ,ubgcnus 1'ryp1wazoo11 arc morphol()gi- midgm. in. is- pnnkularly in newly emerged flies. 15' \ppar-
call) idemical and their sepa ration is based o n cp1dcmio- C'ntly lectins are invoh·ed in prOlC'Cling lhv tsetse fly ngainsr
logkal or biochemical criteria. Tl) p<1no$ome, bl!longing 10 infection by nypano,umes and these rickensia-llkc organ-
the Jjfferent subgenera tend 10 devl'lop in different parts of isms affect the level ol the lectin~. 1"' The work on lecrin~ and
the 1secse fly so. when the fir is dissecwd. one can often rec- rickcu,ia-111:e organism~ has so far only been done on labo·
ognt?.e the subgem,~ involved b> It, location 1sec Chapter ratory cultur~ of i,,et~c flies and tryparlo$omes. and the re-
12: African animal trypanosomose.s, Figure 12.21. The de- sults have not i ct been <'l'>nfmnc<l In the field.
velopment of o:,; •\ probes for the identification or uypano- le has been suggeMed that che presence of lr)1ranosomes
somes In tsetse flies•~ enablt!~ rapid and accurate in th•· hau;tellum of the hetsc fl)' imerfere~ witb che ~em.Illa
idcnrlficarlon of some species within the Oy. Trypano,omc., monitoring the r.iw of food imake and leads 10 infected m~
of major importance In sou thern Africa. ~nd lheir usual dc- prob ing wi rh their mouthp::irts foi· a m eal morn ofltm lhon
wlopment sit~ in rho tsetse fli•, arc given in Table 2.1. uninfccled l\ies_1r~ Thb would be an Important ractor in the
Transmission of the parasites after the; have 111\dergone frequ<'ncy oi disease transrnission, but no increas~ in the
a de\'t-lopmental cycle to give th (' mcracyclic infective stage. probing activiri· in infected G. 111orsi1n11s cemra/i$ has a, yet
i~ known as cyclical transmission. Trypanosomc;. can also been found in the field. "'"
be transmiued mechanical!~· by srringe passage and. in the Many wild animal ,;pccies. particularly bush hue~. kudu.
wan hog a nd bushp1g. awsubclinlcal ly infected\\ llh the uy-
pano~omc~ lhat affect humans and livc\trn:k and ari: impor-
Table 2.1 T,vnallGSO/l"e, a' maJot imponance ,n sou,rern Atr,ca and tant reservoirs of lnfcction. 1:!2 Domestic nnimals ~uch a~
the,r usua de>.e:opmer.; s,ces In the tsetse f;y
canle and pjgs ma~ be reser,oir hosts of nypanosomes
SUBGENUS SPECIES SITE OF OEVEL0°MENT which aOc.,ct human~. ; '· '"°
Ourt1mel/11 T
\ T'.tl-,'I. riaus:-e!:urn rdenlification or Glossi11t1 and I.he species in
Nanr.oma~as i CfJngo/ense Haus:e'bm it'ld 111,dg ; southern Africa
; s.tt11ae Haus-..e lum and r:: dgut
rr;panozocr. T bruce, urucei T<etsc flies were formerly treated as mcm hen; of the family
M,dgu: aru: sa11varv g!anas
T /i!uce, gambiense M,dgu: and sahvarv g!anos Muscidae, or a sub(m11ilr of it. They ore now generally
i i11L"Ce' ,hodesiense M,cgu: and salr:ao,r ;lands placed in a separate fomily Glo~sinidae, which ha~ only one
genus. G/ossiiia.

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50 , .. :oo\ .,,t: ·\spec1s Influencing the occurrence nrinlectious disease,

1mm
1
Figure 2.8 ..awal asµsct of a female G,'ossina mc1s,12~s
N1>1e fo1v:aid-l)o1m ng 0robost1s

Flies in the genus Glossiua can be recognized by the fol-

lowing features: lhe proboscis points forward when at rest
1l'lgure 2.8); the wings have a hatchet-shaped discal cell

(Figure 2.2), and the antennae have fea1hered bristles on the

arista (Figure 2.9). Wh en the Oy is at rest, the wings lie onli!
on top of the other (Figure 2.10), a feature which separates
them from other common blood-sucking Diprera such as
horse flies (Tnlw11u,spp.) and srnble flies (Stomcuy.< ~pp.).
There are three grou µs of species within the gem1s Glos-
siiw. whicb may be distinguished largely by their habitat
BristI:. v:lu
preferences. Ho\\'e\•er. feature~ of their genitalia are the de- coml)Ounrl
finitive criteria. Thcsr groups have been given subgcmeric featn!r,o;i
names. bm these are rarely used. Species and synonyms
within the subgenera ht1Ve been listed by Pont. 188
The fusca group (subgenus A11sre11ina) inhabits rainforcs1
or dense riparian forest. The pnlpalis group (subgenus ,\ 'em·
orhina) also inhabits rainforest. but some species extend o,5mm
1 1
along 1he riparian fringes far out into savannah woodland.
The morsiM11s group (subgenus Glo.<Si1ta) is restricted main!)' Figure 2.9 Laie•a asoec1 of the antenna of a tsetse ;ly
10 sa\1!tlllah woodlands. In the wet season the} spread
throughout the woodland, but in the hot. d!y~cMon tht'yarc
often associated \\1th vegera1io11 along the drainage lines. morsir<ws group
The following taxa occur in southern Africa: • Glo.~.~inn amreni n11sre11i Newstead
• G/ossi11t1aus1e11i mos.urizensisTravassos Dia&
/11su1group • Glos$i11a moffitam morsitans \<','t>~twood
• Glos:sinfl sclrwerzi :-!ewstead & E\'llns • Glossina mor.si1n11s ce111ralis Machado
• Gloss/Im ((lbt111iformis:\\'cstwood • Glossi11a pa/lidipes Austen
• Glossi11a na.<l1i Pons
• Glo.~inn bre11ipnlpis Newstead Apart rrom c;. bre1,i11<1lpis. the f11scn group i~ confined to the
nonh of Angola. and ~pecics in Lhis gro\lp are generally of
pa/11alis gi:oup tlttlt! importance In dl$easc transmission in so;11hern Africa.
• Gfo.<si1u1 pa/pa/is pa/palls (Robineau-DesvoidyJ The same is 1rne of G. pallicem 11ewsrettdi in the palpnlis
• Clossi11afi1scipesf11scipes Newstead group.
• Glossinafuscipes q11n11ze11sis Pire~ For the non-specialist, the diJTerentiation between G.
• Glossina juscipes mnrrinii Zump1 pnfpa/i$ and G. fusripes and identification to the subspccific
• Gfossi11a pallicem 11e111suradi {Auscen) level or all the species is difficult. and im·oh·es mounting and

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 Vecl<l1'5: l'set,e me,. SI

...

\ .'
.. .,-~,.• :
...... ,..! ... -._
,.
. ' "
.h'!-..
...
'-\:Y

,,••
"

Figure 2.10 Dorsal aspet: c'

a !en,.ale Glossma mo1s11ans
show,no the pos11,on of toe
.wigs wnen the fly 1s ai rest
1mm Note the forwa1d-poin1,rig
1 proboscis

close cxomination of the mnlc and fomolc gcniio.lio. Infor- ground colour \\iLh black bonds ncross it. Black banding
mation given in chi~ chapter will enable the reader to Iden- occupies most of the swface of the abdomen. Upper sur-
tify the subspecies of e specimen. if th!> is required. after it fnce of all segment~ of the hind tarsi brown or blackish
has been identified 10 a species level and Lhc distribution ....................................................................palpalisgroup-3
maps given below have been consulted. Reference ,hould :'l!ot a~ ahO\<? .......................................................................-1
be made to the keys give11 by ;,1achadoH~ for the subspecies 3 Posterior margin of the hectors straight, or in the form of
ofG. morsira11$, w ;\lachadoH 7 for subspecies in the Jlulpalis a shallow curve ..................................................fusr:ipi!s s.l.
group. and to the key 10 the sub~1,ecies ofG. (IU.{te11l given by Posterior margin of hector~ deeply ~plil by an an1eriorly
Oias,53 should ther\,ish to confin'n their idC'nrifications. poinring triangle ............................................... pa/r,aliss.l.
A simple key lO the six specks ofimpommce in southern -1 Upper surface of abdomen bright ochreous-!a\\11y in col-
Africa i& given below. our with tr.ices of dark trans\•erse bands which are not
sharply defined ...................................................a11ste11l s.l.
Key to tlle species of Giossina in southern Africa Not as abow .......................................................................5
I Large fly, wing length 11 to 13,S nun, medium brown in 5 Last cwo joim-s of from and midd le tarsi without contrast-
colour, anterior cross-vein 1hickened and darkened to ing bro\,n to black tips. Abdomen brownish-yellow on
appear as a spot on the wing• ...........................l;re1!ipalpis the dorsal surface with distinct. comrasting. black 1rans-
Kot as above ....................................................................... 2 verse hands interrupted on the mid-line of the abdomen.
2 Upper surface of the abdomen with a dark grey back- Bands are square-cut a1 the mid-line and often appear

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 ,,,:rlll, 0'1·. Aspe<:ts inOucndng the occurrence ofinrectiou, diSt'a,c•

as lf seen thr<>ugh varnish. Fringing hair.. of the behavioural differences between 1hese two populations.
antenna! nagellum more than a third of the Oagellar The two are, however. inrerfertile, so there is apparemly no
width .....................................................................pallid1j1es question of different subspecies. G/ossiria pallidipe.: h, very
6 Last rwo joints of the front and middle tarsi \\'ilh contras1 - variable in colour pattern. with tlH' darker fom,~. in which
iog brown 10 black tips. Dorsal surface of abdomen the abdomen is almost as black as that of G. palpaliI. coming
brownish-yellow with contrasting bands of black inter- from hea\1ly shaded areas. while the lighter coloured iorms
rupted at the mid-line. I.lands 1end to be rather indis- cornc from less shaded areas. 20
1
tinctly defined posteriorly. are not square-cut at the A new species. G. borgesi. was described by Pircs. "'bm it
mid-line. and do not appear at If ,•amished. Fringing il> regarded by Pott~ l'Kt and Pont 11111 as being a srnonym of G.
hairs of the nagellum only one sel'enth of the ,,nllidipes. The male genital claspers of G. p11l/idlpes In
Oagetlar wid th ................................................morsiums s.l. Mozambique are very variable and range from those
s1ronglrresembling the clasper$ ofG. lo11gipalpi~ a \\'est Af-
The e:-.,ensil'e studies on the genetics of the gt'nus Glossilla rican species) to those typical of G. pallidipes.{' Haeselbarth
ha,•e been ni,icwed by Gooding." The taxonomy based on er oi.611 considered G. pallidipL>s to be a subspecies of G. lon-
nwrphology is generally supported b~ lhe evidence from ge- gip,1lpis. An anatomical feature on the male da,pe~ of G.
netic studies. rn sou1hem Africa. onlrG. ,wsrtmlhasa degree longipalpi$ has been c!escribed11 which may be used to <lif-
of uncertainty in its taxonomic affinities. Even on morpho· feremiace between G. fonglf)alpis and G. pal/idipes. How-
logical grounds it does not fit well in the mOl'$iH111s group. ever. Thakcrsizt, sho\\·ed tha1 no1 all c;. /ongipnlpis had 1he
and evidence from molecular genetics and breeding data feature so it is not always of value in separating these two
suggests that it is only distantly related to the morsiums species.
group. 76 It has been proposed 1ha1 G. n11sre11i be split into
two subspecies.s.1, 55 and that 1he species G. a11s1e11i i1self
Distribution of tsetse Oies in southern Africa
should be placed in a sepanue subgenus Mnd1ndomyia.5"
bur it is coo earl~• 10 sa~· if 1he latter wlll be ,,idely accepted. In this section. the distribution maps and lis1s 01 species in
Differences between 1he Giemsa banding panerns of the 1he \'arious countries made by pre\•ious workers''0 · th.i. 26';
chromosomes of G. pallidip<!S from Uganda and those from have been used as a baseline. Where possible. these data
Zimbabwe have been demonstrated. 138 and 1here are also have been updated from recent publicaiions and the

0 01stribuuon of ine !,anus G'ossma

H1s1oncal limus
ll2J Definite records oi flies
1/21 less tenain :ecoros o, 'ie!
Figure 2.11 O,swbuuon
of lM genus Glossma m
soutnern A!11c.i

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 Veclors: Tseu.~ Oie, 53

'

FUSCA GROUP
• G brevipa/p1$

D G. schwe:z1
• G rabamformls
Figure 2.12 Distnbution of
tsetse fly species in the f11$UI
group in s01JU1ern Africa

PAlPALIS G110UP
0 G patpalis paloa/is

0 G.fuscipes 4u11nzen$it

• G fusc1pes maron//
Figure 2.13 Dis,ribution of
lllD G. palliceta rrewsrearf1 1seise fly species 1n 1he
pa/MIis group lo sou:hern
Africa

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54 >lCllOS oxr: Asix.-cts influencing the occurrence of inrtcrlous dise3$eS

MORSITANSGROUP
O 6 molSI/BflS mors,raos
O G mors1tans central,s
llID G pallid1pes
0 G. avsiiJni
* TypelocalityofG mo/SJUJns

HiS1oncal limits
IE] Oeflo11e rewrtls of ilies
Figure 2.14 Distr1but1on
of the rsetse fly spociss IA'! Less certain ;ecords of f11es
in the morsirans group in
scuthern Africa

personal knowledge or people currently working in the lield, No recent information Is available from Angola, and the cur-
whose assistance is acknowledged 1\"ith gratitude. The dis- rem boundaries of the distribution of tsetse flies d1ere may
tribuclon of the genus Glossina in southern Africa is shown differ considerably from those shown. This is especially true
in Figure 2.1 J and tha t of the species fusca, pa/pa/is, and for the area infested by G. moriita11s c,mrmli.<, a spede~
morsila11s groups within the genus is shown in figures 2.12, whose range has been expanding rapidly in some areas in
2.13 and 2.14 respectively. Indication of the presence ofHies recent years. !he hiStOrical range of this specie5 cover11d
on the distribution map (Figure 2. I l ) docs not imply that large areas of Angola. including the drainage basin of the
they are uniform ly distributed over the area, bu t rather tha t Okavango Ril'er, 126 so much of southern Angola is potential
they are present where s uitable habitat occurs in the region tsetse fly habitat.
concerned. Some ofche areas shown as tsetse fly-free have Cabinda has all the species recorded above for Angola.
been cleared relatively recemlrand, in such localities, pock- apart from G. morsirans cemmlis. In addition. two fusca
ecs of flies ma) still exist and will need to be eliminated as group flies. G. naslti and G. ha11i11gw11i. nre pre,ent.66
they are found.
The polential for use of s atellite imagery in determining Botswana
dis1ribmion of tsetse flies In Africa. as well as predicting The only species pre~enr L, G. morsirans cenrmlis. ils distri-
monalit} rates and popu lrttion dei:sity, has been demon- bution in the OJai1·ango swamp area varies from ~·ear to year.
°''
strated.199· 2 Predictions of the changes in 1selse distribu- according to the extent of seasonal noodlng and the conse-
rion fol1011ingglobal warmi ng have also been made.204 quent movement of game animals. The historical limit, to
this fly's distribution in Botswana~u. ·1, are shown ln Figure
~gola 2.14. After the rinderpest pandemic of 1896 to 189i. it was
The species that have been recorded from Angola. excl uding reduced to a few isolated foci and, a ll11ough the recovery of
Cabinda. are: game anc caule numbers after the pandemic was rapld, the
• f11sca group: G. sclzwea.i, G. u,baniformis 1ser,c fly's recovery was ve1y slow.411 bm eventually it
• pa/pa/is group: G. fi rscipes q11anze11sis. reached i:s pre\iOusly known Umits in many places.
G. pa/pa/is palpalis. G. pallicera 11ewsreadl The pre-rinderpes1 !secs<: fl)' infestation in Botswana
• morsitans group: G. morsi1ans cemmlis along the north-east border with Zimbabwe. and along rhe

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Zambe,d between Livingstone and the Chabe River. has not tion in Zululand. South Africa. G/ossi11a mor£it(lnS appar·
yet been re-established. Flies in these areas may have been entl}' did not reach as far as 7.ululand. Since the rinderpesr
G. mor$/1a11s mor,itans, but there are no known specimens f)andemic. G. morsium~ and G. p//llidi/Je$ han: nor crossed
a,-ailab!e \', hich could be used co confirm thi~. the arid area between the Save a nd Limpopo ri\'ers in
Mozambique.103 :u~ The advance of tsetse nies into the
l\,[alawi south-ea,t of Zimbabwe1ll-l might well have extended co the
The species which have been recorded from \1alawi arc: Mwenezi (\:uaneL,iJ River.\\ hich is on the Limpopo drainage
• fusca group: G. bnwip(l/pis system. hnd no comrol measures been taken. This could pos-
• morsit<t11s group: G. morsiral/$ 111orsira11s. G. pallidfpe,~ bib!y have enabled the,c two species 10 reinvade their former
habitat in southern Mo1.ambique and. in the case of G. mor,i-
\\'hen describing G. palliflipes. i\ustenJ had specimens from tans. the nonh-eastem r~ion of5outh Africa.
:l.1alawi in the material before him, This ~pecies ha~ not been ,\ puparial cas(' of G. f11sciple11ris was found in I!162 in the
recorded from there since then. a1>art from a reference to it riverine forest along the Uanctse River (a tributary of the ln·
by Moloo 162 in 1985. Recem surveys (1sing odour-bailed cc>mmi Riven in southern Mozambique,5-t which is in an
traps have shown this species to be distributed in ~lalawi as area currently free of1se1se !lie,. but is kno\\11 to ha,·e been
indicated in Figure 2.14. infc!~tlld prior m the rinderpcs1 pandemic. This species has
The e:-1.ent oft he dfstrll>mion of the tsetse Ilies In l\Jal.iwl been recorded from the LMngsmnia Mou mains. east of the
varies seasonally. \\~th the permanently 111fes1ed area~ being northern enc ol Lake illalawi.~1; However. the climates of
largely in game and forest reserves. This situation is compa- southern Africa are known 10 have changed in recem geo-
rable co chat which existed with G. Jl(l(/idipes in Zululand. logical times. and the diMril>ution of G. fiisciple11•is might
South Africa. w<•II hn,·e ex:ended much furihcr south thnn at present.
There may h:we been isolawd relic populations of this spe-
Mozambique cies in southern Mozambique up to the time of 1.he rinder-
The species which have been recorded in i\lmambique are: pest pandem:c.
• fusctL group: G. /Jreufpalpis, G.f11scipleurfs Austen
• morsita11sgroup: G. auste11ia11s1e11i. G. (lllSteni mossuri- Namibia
.:e11sis, G. borgesi, G. morsira11~ 111orsi1a11s. The only sp.:cies recorded from Namibia is G. morsirans ce11-
G. pallidipes trt11is. Concrol opetntions since 1964 ha,·c removed this fl)
from much of the region eas1 of the Cuando (K\\'ando River
In thls work. G. /Jorgesi is treated as a ")nonym of G. pallid· and ha\'e limited its ,pread weM of the river.
ipes. follo,•.ing the view of Pous l<>O and Pont. 188 flowever. The historical limits of the fly extended to the Oka\'ango
the variability in morphological fealun:s found in G. pul/id- Ri,-er dramage ;irea. so 1he threat of expansion of the m-
ipes:,o· 187 and In the Glem~a banding pattern~ <>f the chro- fested area in ~amibia is a very real one. \ final solution 10
mosomes of ahis species. uu indicate thnt G. pallidipe.s mar the problem in thi~ par1 of Africa can only be achieved b)
contain subspecies in a manner analogous to(;. morsitans. Namibian co-operation with the authorities in Bo!swana.
It Is pohltible that funher study will ~how G. /Jorgesi to be a Zambia and ·\ngola.
\'3lid ca.-<on.
Glosswa au$U!ll i mo.,suri;:.e11sis is rhe subspecies found in South Africa
miombo woodland with dense undergrowth in the high The specie$ which have been recorded from South .-\frica
rainfall, medium to high ahirude areas along the ;\tozam. are:
bique Zimbabwe border. whereas G. m/$/en/ <wstem is usu· • f11st(I group: G. bre111pafpis
ally found in the drier coastal thickets. The type locality of • 111orsitntl$ b'T'Oup: G. t111st,!lli. G. /Jra11do11i Chubb.
bmh G. borgesiand G. austeni md$.<11ri::e11sis ls the Sitmonga G. mor.<imm morsitmi.<, G. ,,allitlipe.~
hills area50 km north-east of F.spungabera.
The distribution of the ~pecies in ~101.ambiquc i~ based The hlstoricnl distribution of tsetse tlie~ in South .-\frica i~
on the maps of Ford and Katondo,G6 Dia,;; and Takkcn.w' dealt with b}' Fuller."~ The only species recorded \\'!!St of the
However. the caprure in 1990 of specimens of G. paliitlipes Lebombo ;\fountains was G. 111orsiw11s morsitans. whose
in Zimbabwe along the Mozambique border. just south of t},ie locality is in ~Orth West Province (Figure 2. 14 1. All
the \_lazoP. River. 2r inc!k,ues that tht> c!is1rih11t1on of rhis Oy twtsc> Ille'< tfi,appeared from the northern rt>gioM of South
in Mozambique is greater than is indicated in figure 2.14. Africa after the rinderpest pandemic. although reduction in
The rinderpes1 pandemic of 189610 1897 had a profound 1he cxt(mt of the infe\lation began before thar.67 possibly ai,
impact on the rsetse !lies in Mozambique. especially south a result of the ac1hirie~ of hunters moving up from the
of the Save Rh·er. Before the pandemic. G. 7mllidipe., and southern regions.
G. morsiccms e~-iended 10 about rhe latitude oi ~raputo.';,1 GIO/!$i1u1 /Jre11ipt1/pis occurred in 7.ululand and probabl)
and G. 1x1/lldipes presumably linked up with the Oy popula- also in the gorges of the larger rh•ers where tl1ey cross the

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56 .,,mu, "'L' AspccN inOuencing 1he occurrcnceofinfcctious dlscasc~

Lebombo Moumains. 07 G/os.,f11aftw.:iple11ris may have nlso Zambla

occurred in 1hesegorges.st Glo.<si11a /Jm11do11i. which has its The species which have been recorded from Zambia are:
type localit) in Zululand, i, a synonym or G. ausreni. which • fusca group: G. l;re11ipnlpis
also occurs in Zululand and is probably G. m1sre11i m1steni as • 1ml,,t1lis group: (,.fiw:ir1es m11r1inii
it is found in the coastal thickecs. The historical records • morsita11s group: G. morsittms re11tmll.;,
show that G. µ11lli<lip11s was the dominant tsetse ny in Zulu- G. morsiran~ morsir<u1s, C. paT/idipe;s
1.and. bur although !here have been reports of G. moflirtws
from chc area.. Lhese have never been confirmed."' The his- The map gi,·en brFord and Kawndo,GG nnd the list given by
torical distribution oft~e1se me:. In Zululand indicates that it Moloo. 162 indicate chat G. Ju.wipes fuscipes is presem in
must have been an extension of !hat in Mozambique {Figure Zambia but this has not been confirmed. 18 Phelps and
2.l 1J. Al some time, the former distribution of G. palfulipes L.ovemorl'! 81 consider the flies recorded as G. palpa/is fu.;ci·
In Zululand mu~l have been linked to that in l\lo;,.ambique. pl!P 10 belong to the taxon G. fusc:ipes 111anf11li, as indicated
although in post-rinderpest times chere hawi been no G. pal- by Machado. 1:·
lidipes in Mozambique closer than about 650 km to Zulu- 'file boundaries of tsecse fly discribmion in Lambia are
land.60 no1 monitored continuously. and advances and rece.s!>iom,
At 1he lime of the elimi-nation of Glossin,rpal/idipe.~ from occur naturally. so the limits shown ~hould be regarded as
Zululand ln 1953. le was known that both G. brevipalpis and approximati<m~.
G. a11sie11i were ,till present. They we(e confined to riparian Flie~ rrom the isolaced Keembe pocket or G. morsita11s
fringe vege1arion, forest~ and thickets. Canle rarely enrer ce11tr(llis, east of the main distribution zone of 1hi$ species In
lhese densel) shaded areas where the flies were locared. bu1 we~tern Zambia. h,we a different ;illelc frequency for some
minor outbreaks of trypanosomosis have occurred ~ince enzyme si ~lem~ from flies in the main distrlbulion area, 78
195~. and were contained by using curative drugs. 125 A indicadng Huie gene tlow between che two popularions. TI1e
major Ollibreak of 1rypanosomosis occurred in communal durmion of chis isolation ls not known. and the current ad·
area cattle in Zululand in 1990, and the disease was diag- vance,- oflhis subspecle$ from the west may cause 1he,e two
nosed widely in 1.he area between the Umfolosl River and the populations to link up.
Mozambique border125 (see Chapter 12: African a ni mal Recem survey,. using kairomone-baiced traps. h11ve
trypanosomoses). shown G. pa/Jidipes 10 be more common in Zambia than had
Changes In land use, such as lhe establlshmen1 or been pre\iously thought.
game farmi. and forestry projects. as well as large increases Robinson ec a/. 1%. 197 have used multivariate analysis of
in human and cattle number,, have taken place in climatic factor, and remotely sensed vcgc1ation data rn pre-
Zululand since the dis1rlhutlon of tsetse flies ,vas last dict habitat ~uitahle for iseise mes in easiern Zambia. Good
plotted in rhe 1950s. ,\ survey w determine the current agreement ha, been found between these predictions and
djscribution of 1setse flies in Zululand was initiated in the kno,\il distribution of tsetse Dies In the area conc:emed.
1995 and i.s scheduled for completion In 1999. 17.? The It WO$ also found 1ha1 the ~eparatlon of G. morsiram cenrm-
survey results to date are shown 1n Figures 2.11 and 2 .12. iis and G. mor.•iw11s 111or.sira11s distribution in 1he uorlhem
The results show that csetse nie, now occur in some pro,·ince of Zambia ' f igure 2.14) could be accounted for on
areas where !lie} were not recorded at the time thac Du !he basis llf the maximum of the minimum temperature at
Toit did bis work.60 The two species may occur together any given altitude. Areas with a colder minimum tempera-
but in some localities, such as Hluhluwe and Umfolozl. ll!re are inhabited by C. 111orsim11s centralis and chose with a
only Ci. brevipnlpis has been recorded, while in Temhc warmer minimum temperature by G. morsimns morsf1a11s.
Elephant Reserve only G. austeni bas been found. 125 Both
species of tsetse fly were involved in the 1990 otnbreak Zimbabwe
of 1rypanosomt)sis. GIO$sina tmsteni was found 10 be re- The specie• which have been 1•ecorded from Zimbabwe are:
stricted to vet)• densely shaded places while G. lne11i- • f11sC/l group: G. lmwipulpis
palpi.< will range out from denselr wooded areas into more • morsiums group: G. ausreui, G. 111orsi1011! morslta11s,
ope11 woodland panicularly al dusk. Glos.firm 1,re11ip(l/f)it G. ptl/litlipi:s
,hows some nocturnal activiry. especially on moonlight
nights. (;/ossi11a !1re1ripalpis ha, been recorded along the border
, "~th vlM~mhiqnP in rhP C:hipingr dis1rk1. a~ hac; G. aiwtml
Swaziland (probably the high-altitude, moist environmenc subspe·
At presenr, no tsetse flies exist in Swazjland. des). ln recem years, though. neither these specie., nor G.
l listorlcally. G. pt1!/idipesoccurred. at lease seasonally. in pt1/lidipe.; have been reported from this area.
the soulh-easc of I.he countT)1, 60 while G. m1stcni and G. In 1992. G. morsfra11s mcrsitans. (j, pallidipes and G.
brel'ipalpis were probably prescnc occasionally along the armeni were captured in odol1r-bailed traps in the Honde
Swatiland/-:'-1o:zambique border (Figure 2.1 1). Valley. on the eastern slopes of the mountainous eastern

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 \'e~tors: rscr""' flic, 5i

border area of Zimbabwe. very clo~e to the border \\ ith rently experiencing thc.w population pressure,. TI1e situation
~101.ambique.1 '" :-;o tsetse flies had been recorded from this ~how~ no siws of ,1ahi!i1Jng yet and. in the absence of cata·
area pre\ious!y. although relatively large numbers of cases sll'ophic C\·ems. populations \\tjll conrinuc 10 increase for
of uypano~omosis had been recorded sporadically from many year,,, The population nu mben; arc al ready in exces,, of
c:ml<? gra,dng there. any known historical levels in thi, part of ·\frica, so e~-ampll'S
l'he hisrorical distribution of 1sc,1,se 11:es in Zimbabwe' 111· of th<: j)U>I itneractiu11, h<:tWC<:11 ltumall~ ,U 1d lS1.'lst: me, 11ta\'
111 is shown lt1 Figure 2.l I. I he southern rsehe fly 'belt' had beofacadernic imere,t on!~. However.some indication of the
staned 10 breal.. up prior to the rinderpes1 pandcm!c36 and ,ort or int~'rnction~ that may occur are given.
disappeared comp!e1eti after the pandemic. An ad\•nncc of n,e effect of humnns on the cJifferem subgenera of tsetse
r~Nse me~ into the south-eastern corner of Zimbabwe irom Oies differs greatly. due 10 the difference~ in habiiat require·
i\lozambiquc in the vicinity of the Cionarczhou :\'arional ment~ of the tlie~. The descrucrion of the forests of Africa,
Park during the late 1950s and early 1960s wa, controlled part!) ior comcnercial timber and partl} ior seulcmellts ond
and this part of Zimbab1ve is now free of tset~e me~. 191• 19:; agricuhural 12nd. is proceeding rnpidly, and thi~ proce)s re·
There ha,·e been reports From Moza1ribiquc during 1999 moves the hnbi1a1 or the /11sra group nf fli~. lt is no longer
that LSetse llies are beginning tO recover from the earlier possible to leave cleart>d land fallow after il has become too
control operations in the norlh-we$1ern corner of Gaza depleted in nutrients 10 suppon crops. so clearings become
Province and could soon pres1m1 a u,reat to ~outh-eas,em permanent. The human population uiso destroy~ the food
Limbabwe agajn, suppl) of the 1secse tlics rhrough hu nti ng. so they disappear.
rhere are high populations ofbct~e HiC'son the \lozam- l'he/1w:<r grr,up of Ole, nre likely to sunive only in forest re-
bique side of parts of u,e eastern border of Zimbabwe tFig- ~ervc,, in lmure. In soul.hem Africa, (;, br~l'ipalpi,, which is
ure 2.11 1 and occasional cases or trypanosomo,is occur in the rnosr \\'Ide!~ distributed member of the Jitscn group. I,
cattle in adjncem areas in Zimbabwe. these ca~<.'~ arc attrib- confined to ri':erine fore~ts and to the few humid fore,t, that
uted to occasional tly mo, emems and to cattle grar.lng over occur in the region Thh limited distribution makes it a rela-
the border in Mozambique. Only the northern part oi /jn,. 1ively ummponam ,pecies in the comexi of disease trans-
babwC' currently has a resident pop11 lt11mn of both G. 111or,-i- mission in routJ1r.m Africa hul ir is of imponance in
m11s mor;it<ms and G. p<lllidipes. 7.ululartcl, "here G. 1ml//diµe,; ha, been eliminated.
·\!though some intermediate form, m cc:rm; of male The influe!'lce of humans on the pnlpnTis group oi nies is
genii al ~mrcture~oiG. mnrsiumshave been r;:cordcd, crn~- not alW(lYS ad1·er-e, and they <'~n often exist in close contact
ing experiments have confirmed lha1. ai present, G. morsi· with p1:ople and their domestic siot:k. The.w file$. pa.'"ticularl>·
1m1s morsirrms is the only subi.pecics presem in G. ,x,lpalis. feed on a wide varicty of wild and domestic ani·
/Jmbabwe. 182 It is not known whetht:!r in historical time~. mat~, including reptile,, and they ma}' be invol\'ed in di~eaM?
the Infestation in the north-west oft he count11 wa, G. mor· 1ransn1i',~ion even in densely populated area,. 12t·n:cir habi-
;iums 111orsim11s or G. morsi1a11s centralis. fhe latLer species tat is riparian \'egetarion: humnn activities dn not always rt>·
b expanding ii~ range rapidly in i'ambia at present panli :.ult in the elimination or thi> , ,:gctation 1ype. \griculcural
because the c.xren~i\'e control measures needed to contain actil~ties tend robe in area6 further from tht' ,trean:.,, in high
the fly are beyond the resou rces or that coumry. l'ossibly, rainfall :mms. and the tree plantations grown 1herc rm\)
1herefore. C,. morsira11.1 ce111mlis may ~each the /.ambezi themselves c~1end the habitat of the fly. Patches or natural
mver in the 1·ici11icy of Victoria !'.ills and should thb occur. ,·cgc1a1ion protected l>y human, for cultural reasons also
Zimbah\\'e may be invaded from Zambia by a subspecies of occur in many pans of Mrlca and ma)' provide sui1.able habi·
G. morsi((lfl$ not pre\iou~Jy recorded in Zimbabwe. tat t'or pnlpali~ group flies, provided domcsiic animals .ire
readiiy a\"ailablt: a~a food sourte. l\lcml>crsofthe ~ub~pecic,
of G. fuscipi's can be doselr associared \\1th humans and. in
Control
western i.:cnya, they have been able to live in exwn,ive thick-
Environmental modification c1s of Lt111rr11111 ca11wra. which haw dcvelopt'<l as n result of
Effects of humans on 1setse flies in santnnah and r iverine poor land managcmcm.2•1:> Flies in che pa/pa/is group ha\'e a
areas In 1he past there have been many !h1ctuntions in restric1cd distribution in southern ,\frica. but where tht.'} do
human populations in Africa which have been amibutable occur, they may be important vectors of disease in hoth hu-
to disease epidemics. tribal conflic1s or the colonizacion or mans and domestic stock.
Airlc~.65 The advem of western medicine ho!> reduced the 11 ~ on 1he 1110,~iums gr()up or mes that humnn$ have
death rate in human populations in •\frica dramatically and, m ade their greatest impact in southern Africa. c;1os;"i11a mor-
as 1he birth race has remained largely unaltered, there has sitans and G. pal/idi,:ws are widespread here. and ate charac-
been e:q>onential growth of the hunian popula1ion. \\'here terbtic' of the undi~turhed ~al'annah Md 1,·ooded sa,·annah
these high human populations are confined, for example by vegetation types, where wild animals are ptemiful. fhe agri·
lack of a<ldiriortal arable land or territorial boundaries. the cultural activities of humans, and their use of fire. have
human densit) has increased greatly. Southern :\frica Is cur- played an Important part in the development of savannah

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58 ;•.t.,'<>X ' "" Aspeets inlhtcnonN the OC<'ura•m:e of infectious <lisca~es

,·ege1a1ion, ~o In this sense, h~tmans ha,·e been of benefit 10 t·onfined 10 the ,anctuaries and are allowed to increase, 1he~
the tset.Se fly. But humans often emer tsetse ny-in[ested sa- will modify Lhe habitat. These habitat chruiges may make the
'>'annah coun1111 ro underrake agricultural actl\iti.es. If culti· area unsuitable for t~ctse mes. and for many other animal
,-ation in 1he~e area~ h continued on a long-term basis. the ,p~les. as has happened in ~ome East African park:..&. .\
vege1a1lon becomes either cropland or grassland, and at tht similar process i, taking place on the escarpment of the 7-.am-
,;amc time the wild unimals on, killed or move 1w,uy and the bci,, Ri\'Cr \'allc:, in Zimbabwe, where hlllslopc woodland is
flie$ ultimately die out. In 1he past, land was abandoned being <'onverted 10 grassland by high densities of elephanrs.
when its fenility had declined. Such land would. in the
course of lime. re\'ert to sal'an.nah \'egetation and the wild Rinderpest This disease entered Egypt from Arabia in the
animals would return. making the area suitable again tor late 1880s. and a pandemic ofit swept down Africa in a wide
morsimns group flies. Thus the siruacion was one of Jong· varier) of animals. It had reached Lake ~lala,\i by 1892 and
term rises and falls of fir populations. depending on the crossed the 7..ambet.i in 1895 to 1896 (sec Chapter 49:
acthity of human populations. This shifting pattern of agri· Rinderpeso. Thl'rl' are endemic foci of infection remaining
culrure is becoming less frequent ii; sourhern :\frka. and In r:ast Africa now. but none in somhem Africa.
more permancm occupaLion of the land by humans leads w This disease almost eliminated caule in the infected area
permanent removal oflhc 111or.<irm1sgroup of flic;. The den- and s11Yerely depleted the numbers of many wild animal
slt~· of human se1tlement needed to aclueve 1h1s removal of species in sot1thern ,\Inca. Not aJl " ~Id antmals are suscep·
tsetse tlies is not known exac1fy. In West :\frican savannah, tlblc, but African buffalo. kudu. eland (Taworragu$ OIJ'X).
densitie.s of 15 to 39 people per square \dlomem: reduce bushbuck. bushpig and warthog died in large number~.
morsiinnsgroup flies co very low le\·els. and at human popu- Other bo•.ids such as sable ancelope (Hippo1rt1g11s 11iger)
lation levels higher than Lhis no tsetse Illes e:o.i~t. 170 Somc and roan antelope (Hippotragus (!(/Ui1111s), impala and blue
parts of southern Africa ha..:e always had a hlgh human wildebeest ,Co,mocluwres umri1111.~) were less affected in the
popula1ion densir) .2 ll An accoum of the \,·a xing and waning pandemic. while elephant, rhinoceros and hl?popotamus
of the Glosslna population In the )lossurise region of lHippopou11111t:, amphil,,ius) were unaffected.~& JI I The sus·
'.\1ozambique, consequent 10 cha11gcs in human population <.'eprible animals thus include 1he preferred hos~ of G. mor-
densiries in this heavily seuled atea berween 1861 and 1889, ,ir,111s and G. pnllltlipes in particular. 1~·1• 205 This\\ as not. of
was pro,~dcd by Swynnenon.213 cour:;e, known m tl1e time of the rinderpest pandemic. but
In the 19i0~ the cMl war in Zimbabwe pre\'ented control after the dlsea,e had been eraditatcd from somhern .-\rnca,
01 tsetse fiie~ along the north-east border of the countt)'. Stla'\'enson· Hamilton 2 ' poi111ed out lha1. as 1101 all animal
Over a period <>ffi\'e years the fly (in this case G. morsiwns) species were equalJ~,affected. the tsetst> flies ml;st have been
invaded thl' country rapidly. reached ih historical limits, U$,Oclated with those that were mo~t 5uscep1ible. The 1'iew
and e,·en progressed slightly (urthe: in some places. How- held by some people at the time of the pandemic that
C\'er. much of the country that was reinvaded w;is marginal rinderpest \'lrU$ was itself pathogenic to tsetse flies. has
fortsetse flies and their number:, remained ltm, but the Um· been shown co be incorrect.27 The effect of the pandemic
furudzi Safari Area, lying in broken but well-watered coun- was not uniform. and in some localities appreciable num-
try and with a large wild animal population. pro\'ided a good ber, of animals of the susceptiblo :species survh·ed.211
habitat and flies built up to large numbers there This i.nva- The effect on the G. 1110r$/rr111s and G. pt111i,/ipl!f; popula-
sion ga\'e ris~ to cases of trvpanosomosis in caule near 1ions of 1;1is se,·ere, rapid change in the emironmen1 ,,·as
Shamva, an area which had been free of tsetse Oii,,, since the dramatic. In Bot~wana. 1se1~e retracted to four ,mall foci.
rinderpe~t pandetnic in the 189()s. whih: in Zambia, the Kafue tsetse Uy belt ~hrank consider-
ln the Sebungwe and Umniati districts of Zimbabwe, ably and 1serse flies disappeared completely from the WesL·
outbreaks of human t.rypano,,omosis. in 1912 and 193~ re- em PrO\ince.~u., rhere were alSQ chan~ in 'l.amihia and
<;pt:Cti\·ely. caused the authorities ofthe day co eliminate the southern ·\ngola. In Zimbabwe, G. morsiums disappeared
problem by 1110\ing the hurnon population irwolved. 7 ln the from much of it, range. 110 and dbappeared complete!~· in
long term. this son of nc1ion fa\·ours the vector since. in the Somh ,\frica, 6~ In '.\loa11nhique, tsetse flies. particularly G.
absence or any Oy control measures, the abandoned agricul- mnrsirnm and G. /Jllliidip1.1s.s I tli~appeared from the '>Puth·
tural lands re, en 10 suitable t.Setse fly habitat. Si111ilar e\'ents ern pan of their range.h7 In pans of North '.\.lossurise in
have occurred in East Africa.l'-1 ~101.ambique. (;. morsirans numbers did not decline In ,-pite
- Human, al~o affecr the di~tribution of ,~e{'!;e flit!s indi- of heavr mortalitr from rinderpe$t among wlld anlmals. 1 ' 3
rec:.-tly by creating wildlife sanctuaries. If these areas are in As there have be.en similar reductions in G. mor$im11s popu·
tsetse ny-infested counuy. they~-en-easa source of the flies to lat ions after at least four outbreaks of rinderpest in East Af.
domE.'stlc o;iock grazing on adjace111 land. a, hn, occurred In rica. there can be little doubt thnt the cause ,,·as lhe remO\'al
Zululand6Q and is current!)' the case in '.\lala,,i . i;- There is of its food supply. Fifes may haYe sunived in some localities
generally re~lstance to the use of any and-tsetse fly measure bct·ause some herd~ of susceptible animals remained unin·
in the sanctuarie~. However. iflargeelephant populations are fectcd. or because of the presence or no11-susceprible

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 \'ccrors: Tse1>t !lie, 59

animals ~uch a, rhlnoc<'ro~ and hippopotamus (often VelJ' area. par1lcularly kudu. The results from the!ie experimem~
localized in their distribution) \\'hich can pro,ide goo<! led co 1lw wbsequem u,c in 7.lmbabwe of selective hunting
sources or blood for tsetse flies. llhinoceros blood has been of the folu preferred species between parallel fences 10 pre-
round in c;, mor$ita11s and G. J)<lllidipes in Zimbabwe at vent the ad\'ance of tsetse flies. l'he fence immediately adja-
some localltics. 193 and G. bre1 1ipalpi$ i~ kno1rn to feed cent 10 the fly area was more robust and large animab such
rcadily on hippopotarni."1"'' The e;.dusion oi rhinoce,os as elepham and buffalo were kept awai from ii br limited
from the animal ho:,1. list during the campaign to diminate hunting. This approach to animal reduction initialli en-
tsebc flies by game reduction In tht' 1930s may hil\"e played abled the tsetse ad,ances co be contained, but became iner-
a considerable parr in maintaining 1he tsetse illesM and ,ug- fcc1ive during the Ci\'tl wm in the 1970s. Whilst huming
gests a possible mechanism by which flies were able to ,mr- operations ,,ere resumed soon afler the cessation oiho,1lli-
vi\'e the rinderpest pandemic In Zululand. 1ies, including those conducccd for the protection of fence,.
1hese ac1l\i1ies wcr<! shon-U,·ed and bi the 1990s had heen
Animal reduction as a tsetse fly control measure The pre- al>,mdcme,J com1>le1ell' in favour of other techniques.
rinderpest re1reat of G. 111oniu111.~ in the' Limpopo Pr01ince
ofSouthAfrica6; and in north-wesiern 2.imbabwe 36 was m- Vegetation modific.ation Swdies on rhe biolo~ 01 i.secse
trlbmed 10 the acti\ities of big game hunters and the ad- flies have ~hown that in hot weather they depend on the 1·eg-
vance of settlers from the &ou1h. The effect~ or rinderpest ctarion to &hade both their refuge sites and the sites in which
provided further evidence of the impormnce 01 \\11<1 animals larvae are deposited. \'egetation pauems hnve been modi-
to G. morsiums in panicular. /\s the animal populations in fied bi fire i11 ~outhern Africa. and the tJlimination of fire
southern Africa recovered after 1he rinderpest pandemic. ,o may result in :he dc,·elopme111 ofp:merns unsuited to ,ome
did rhose of the tsetse fly. and they began rn spread 10 their tsetse specie.,. 74 · 2 ' The use of this approach ro conrrol
former limits. The flies soon came Into contact with humuns 1setse me~ in East Africa and :t.ambia has 1101 b"'cn generall}
and their dornes1ic stock, and lhc governments of the ,ari- \ucres~ful. a~ it is impo~slbte ro exclude fir,• from larg<' areas
ous countries had rn undcrrake measurt!s 10 reduce the dis· of Africa complereh·. 1• 2.1; t\bo, this method rakes efTert
ea~e prohlems associated with tsetse flic,. The most obvious slowly and there b usually a demand ror quick.result;, where
\\'Oy 10 anac.k the flr ar that time was 10 remO\t' it,; source of i,;et~c fl)' problems occur. The direcr effect~ oi bush ti rt'$ 011
food by reducing the number of wild animab. adult 1se1sc flies and theirpuparia hove been irwestiga1ed in
In Zimbabwe. the number~ of wild animals were reduced Zimbabwe,"5 but no e,~clence has been found that the~e
hy huniing between parallel fence~ 16 km apan. 111~ \ broad fire$ control the Ilks.
spectrum of species was shot. and the method was success- As tsetse Hies do nm orcur in grassland, the clearing of
ful in reclaiming large areas of what is. now $Orne of Zim- woodland to produce grassland can create a barrier to t;,etse
bnbwE!°S most productive fann land. The large-scale flies. bu1 the c!ifficultl is that. lo be effective, the clearing has
desrrnction of wildlife is abhorrem. 111 but at thnL time ii was rn be wide. ahhou~h 1he exact width needed is no1 known: ii
1heonly practicable method ofconrrolling tse1~e Oi~. which apparent!) \'Sties be1we11n species. Barrier cleari11g~ have
threatened disaster 10 1l1e fam1ing community In Ziml>ab- been made in Zimbabwe on 1he eastern border with
we. Broad specrrum b'(lme reduction campaigns against Mozambique 11 t ancl in Zambia.ill!, Zululand.I'° Bmswana 16
tsetse 0). either between rences or not, were undertaken in and l'>-tala\,i. where dearing was combined with seule-
Zambia.'~- ~b5 Botswana1 ij and Zululand •S0u1h \frlcal.'~ ment.110 All hough barrier clearing, have achieved moderate
bu1 the success achieved was generally lltll as good as 1ha1 in success, they are expensh•e rn create and mus1 be main-
Zimbabwe. t\s allema1ivc meiho<h of1~etse co1urol became tained indelinltel~. They arc not widely used at present.
a\'ailable, game reduc1i<1n campaign, ,,·ere abandoned in l\uempts have been made to u,e herbicide~ 10 comrol re-
most countries. generating vegetm.ion.r 18 but 1he chemicals are expens1\'e
1-ollowing 1he demonstration of host preferences of anc,I may have deleterious effects on the envi ronmem.
tsetse flies.~;,~ an experimc.111 was conduc1ed in Zimbabwe Particularl1· In East Arrica. the vegetation requ.;rements
between 1962 and 1964 which showed thar a spectacular de- for cse!'le Ov habnars can be dividrd lnco essenrial and non-
cline in G. mor$itans numbers follmwd the humi1\g out or essential components. The essential componen,i; might
warthog. bushpig. bushbuck and kudu. 4P Elepham and compri~e a small proponion of 1he total area and. if re-
buJfalo were clri,•en out of the fenced experimemal area moved. would re;,uh in 1hc collapse of the fly popuJaclon. In
prior;o the sum of the experiment, because of the dam:igc more arid IXt«s of the mes' range. C!lsemi:tl habi1.1c b con
they do to fences and also because they can be a good source fined to \\,Her c01uses. bu1 in hc1tcr watered areas l'cgcta-
of food 1,·hen prcse111 in an area. No other animal spec:ies Lion pancrns wirh well-developed upper and lower canopies
were hunted . .\s warthog provided about half th~ blood are thought 10 be l!~~enrial for the nle:,' sun~val. In southem
meals of G. mor,irans at the experimenral Site. the effect ol Africa the discrlmlna1i\'e cle.iring approach (remo\'al of
the remov,11 of just warthog was testecl.2 '•3 It was :.hown that essemial habitat onlyl has been auemJ>ted but not
the rues ndju~ted rapidly 10 other preferred ho~,s 111 chis widely used.

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GO m:, 10~ ,,,t: Aspci:1s m!lucncmi: the oc~11rrrnce or mfectious dbt"u~e,

Dbcriminatini clearing was ,,scd berween 19•1 l and 1942 from the Di•. 1his campaign was moumed joint!)' between
10 elimina1e G. morsitnns m.':tr ,\bercorn in 11onh-eastern 1962 and 197.J by S0u1h Africa, Mozambique aild Zim·
Zambia."·' \'egctatio.n was removed only in the shallow val- babwt>.1+' 1 ' 5• 106 1""· 1~5 Ground sprni1ng has been lhe major
leyi, at the headwater:;. of small Mream,.. which formed only a mt.!1hod u~ed 10 eliminate tSetse nles in Zimbabwe from 1960
small proportion or the total t<ell,e fly.infested area. This to 1986. and ha-, also been used extensivel> in both :--:igeria1"'
c;,mpaign was one of rhc few successful am:mp1s at dis· o.nd Botswunn:'8 In Zambia, ground sprnying has been done
criminativeclearing. In Zimbabwe, ex1ensive discriminative with both knapsack i:.o and motorized sprayers carried on
clearing in the south-east of the country \,·as done between u nimog vehicles. ;i. Ground spm}~ng has been conducted in
1956 and 1961 in an attempt to arrest ad\"allCes b) both G. the Capri\; strip region of :S:amibi;'I and e:-'j)erimemally in
morslmm and G. pallidipes into 7.rmbabwe from :-.101.am- Malawi. but large areas have mil been cleared of ise1se flies by
bique. • Yluch of thi~ arid area is ma~ginal coumry for tsetse ibis method in either of these countries.
tlies. 103 but in some places riverine fringe \ egetation and The ~fse of long-acting insecticides leads to e1wfronmen·
patches of hillblope woodland provided reasonable habi1a1 ml contamination. and much concern has been e:1:pressed
and abundant wildlife. The clenring dertainly helped to limit over tsetse fly conrrol schemes based on ground spraying.
the flies. but it did not eliminate them. The coMs were high, /\cute poisoning or birds and other wildlife br sprai~ng ,,i1h
as much of the work Wa$ d()1W by bulldozer!>, and the de- di1;?!drin against tsetse mes has occurred in Bots\\'ana.1!0
struction ot riverine vcge1at1t>11 Is likely 10 b(- perhlanent in Hesrdues or DD J ru1d us meta halites are accumulated b}'
this Vel)' harsh pan of the country. This form of tsetse fl} non-target org,mis.ms in sprayed areas. Following the indi·
c<mtr<>i has not been used recemly. l.'ither in 7.imbabwe or cation of potential adverse elr<.'CtS on animal populations In
elsewhere in southern Afric~1. Zimbab\\'e, 15· a comprehensive four-year smdy 1,"as under-
wken in a section of the Zambezi River Valley in Zimbabwe.
Pesticides The studr area bordered on Lake Kariba. where ground
Application from the ground \Cter the Second World \\'ar, spr.1) ing with DDT and, in a small scale uial. deltarne!.hrin
rhe advent ofsynthetic organic insecticides. and particularly against tsetse flies was being conducted. The re~uhs57
the long-acting organochlorint!.,, led w new methods of showed linle or no elfec1 of DOT on reptile populations,
tsetse C'()ntrol. The flies· hot, dry season habitati, (Le. their some indication of effects on some ant species, and delete-
essential habiiats) can be determined by sampling for adults rious effects on populations of several species of insectivo-
or puparla. \\11hin the essemial habitats refuge sires of the rous birds. ln particular. popula1ions of the while-headed
Oies can be recogt1ized. 11"5• l>S6. i,ri The determination of black chat l Tl1am11olaea amoti). which feeds extensive!)' on
the$e sites is easier in very arid counuy, but e\'en in well- ants. were severely reduced bi acute DDT poisoning. Eggs of
watered woodland Ir ls possible to recogni1,e them. 1f thi•sc terre,mial avian raptors. ~uch as the African goshawk
hot season resting and refuge sites are treured with suitable vtcip,er rcic/1 iro,. sho,, ed marked th inning ofLhe egg shell. a
long-acting insecticide. applied from knap$ilck sprayers b) fac10r known 10 be associated with chronic poboning by
team, mo\'ing through the bush. the me,- will ultimately DDE a metabolite of DDT. In many parts of the smdl' area
come into comact with this insecticide and be killed. This along Lake .Karibo. eggs of the African fish eagle (Hafiaeetus
so-called ground spraying involves the recognition or essen- r·ocffer) showed c,,gg shell thinning. whereas in other pans
1ial tSetsc fly habitats on ae:riai photographs and the subse- very little egg shell thi11ning was evident. ;,.:'o detectable ef-
quem development of access tracks 10 these places. fects of DDT wl'.'re found on soil processes. Residues on
followed by the appticarion of insecticide to them. The in- spra}ed tree 1runks and soil were 1101 long lasting. Xo se\'ere
~ec1icide ls applied in the cool. dry ~eason, ~o that as much adverse effect~ were demonstrated on the fish species stud-
country as possible can be treated before the onset of the ho1 ied. There were indications chat populations of some spe·
weather. Where there is heavy grass cover. i1 may be neces- cies of bats roosting in hollow trees could oe ad\-erscly
~ary to bum 1his before appllcacion 1>f the fn<;ecricide. which affected b~' ground ,prai~ng with DDT.
is decomposed by fire. The most widely used insecticides in A smaller <tudy" on the em~conmemal effects of delta-
this form of tsetse fly control ha\'e been dieldrin and DDT. me1hrin used in ground spraying against 1se1se flies. showed
Howe\·er. in Botswana, Namibia nnd Zimbabwe they were that popt:la1ions of non-1arge1 invcnel>rate species were se-
more recemiy replaced by synthetic pyre1hroids for envi- verely affected in both terreslrial and aquatic habitats. :\'.o
ronmental reasons. The substances u5ed for this purpose major elfects were ~een on other animals. although the large
~ave a residual life of at lea~t four month~ in the dry season. decline in in~ec1 populations may ha,·e caused lnsecri-
Ground spra}ing has now been replaced almost completely vorous species to move to umreated areas.
In these coumries by the odour-baited. insecticide-treated ~one of the species found to be adversely affected by the
target technique, again for environmental reasons. pe$ticides used in ground spraying against tsetse mes were
ln southern Africa, an advance of tse1se mes in south· endemic m the s111dy area. Following elimination of 1he
ea,;1em Zimbabwe and sou1h-we~tern /\·Jozambique wa~ tsebe flies, it would be anticipated thar invasion from sur-
halted by ground ~praying and a large area was reclaimed rounding un~prayetl areas would occuc, as well as recovery

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 Vector~:T,etse Jlie-, 61

of the re•idunl animal populations in rhe sprai NI area. Thi<i in Zimbabwe, Del1amcrhrin and alphacypermctr.rin have
would 1101 happen if Lhe spr.aying wa; done near senled also been used in Botswana.2""
areas. a~ human settlemen t results in major changes in the Uirge-scale aerial spr~ying campaigns were conduc1ed
ell\ironment and many habitats are destroyed. at irre&'lllar imer\'Rl'- in Zimbabwe between 19.5:-l anc.l
,\1101her form of insecticide applicaLion from the ground l 988.:i:i. ,;u, to: tr~, The more recent aerial spra)'ing. spon-
is the u$e ot peHicide dusts. Thi~ approach wn) u,ed in Zu- sored by the Regional T~e,~e and Trypanosomiasis Control
luland. South 1\Irica.w but not elsewhere in ,outhern Africa. Programme, has been successfu l againsi Ci. 111ars/1<111s but
less so again,, G. pn/lidi1;cs. Aerial spraying conducted in
Applicalion from the air The first aerial spra}~ng cam- Zambia has been large!~ ;.ucce~sful. 1: 5 No in all in,ecticide
palgn against tsetse 11les \~as in .lululand between 1945 and work against t<etse llie,. the cleared areas mny be reim acled
1948.60 Initially. a coarse spray uf atomized liquid droplets if they are not develop~'<I and settled by humans or llw op-
was applied. but this resulted in the ionnation of toxic re- erations are not progre~si\·e. In Botswana follo,,ing suc-
sidual dcpo~ib of 1he chemical. This apP,roach ha~ not been cessful prellminn11 aerial 5praying trial~ in 1972. pan~ of th(.>
used elsewhere in southern AfriC<t but has heen tried in \Vest Okavango Delta were sprayed annually from 1973 to 1991 ex-
Africa, where helicopters have been used to apply the spray.' cluding 1988). In geneml, although not exclush·el). the an-
The second approach to the aerial application of lnsecll- nual opera1ions up to 1979 alcerna1ed between 1hr wcswrn
cide was the use of insecticidal smoke or thermal aero,ot.00 and sou1h-easwrn perimeters of the Delta. Thereaftt'r. the
Tl1 this fom1, normally long-a,·ting in$eccicidc, such a~ DD1 operation<; ,,·ere ainu:d at tht· maiu body of the infostation. z,;o
or hexachlorocyclohcxanc (fom1erl} known as ben;,..ene :\(though pestidch, application rotes are very !11\\ In the
hexachloride or BHC) have no residual action. and the i,m1- aerial spray method for tsetse fly control many aqua1ic or·
ronment is simply fumigaced. In Zululand, the applicarions ganism~ an· killed. c;.pecially in shallow water. l'he effect'> of
were made in the early morning and late afternoon to take aerial ~pra~ing on non-target org..'U1isms have been moni·
adt·aniage of the 1emperature in\"Crsion condlcions. which torcd on several occasions 3~- sa 153 and it has been found
pre\'ent the aerosol cloud rising from th~ ground The aero- that generall~. if 1he in~ec1lcidc b applied at the correct
sol was sprayed in the rsef!;c fly habilat from aircraft flying level, and only In one year. no permanent damage to the cn-
just above the tree tops. i\ slight wind is an advantage. as the vironmcm occurs. These conclusions have been ~upponed
aerosol clot1d 1hen drifts through the habitat. bm ~,rong by the St'ientlfic Envlronmemal Monitoring Group.2";
winds disperse this cloud and its applica1io11 ha, to be which is part of the Regional Ts.etse and Trn,Qno,omiasb
stopped until favourable condit ions return. Control Programme. The same group also stated chat 'if
The aerial application of insecticides inlliall~ developed areas without open water are treated with endosulfan (6 to
'°
1
by Du Toi1 has now become more sophistica1ed. but the 2-1 glhaJ. and area,; with ri\'l'rs and Jakes are treated 1,ith dcl-
undedying principles are the same. :\11,opp ha5 re\iewed tamethrin (< 0.3 glha). 1lwnaccording 10 our present know·
insecticidal me1hods used against tsetse 11ie~ and outline:. ledge the side-effects ,ccm 10 be accep1ablc · 6"'
the current approach 10 aerial spraying. The de\'elopmeilt of Other me~hods for generating aerosols ha~c been med in
rotary atomizers which produce a unifom1 aerosol droplet anri-1se1se ny work. Thermal smoke generators have been
size and. In particular. che development ofnigh1 nyingtcch- used, in which a combu~1ible rnLxture of ~odium chlorate,
niques that pcnnh spray aircraf110 operate throughout 1he sugar and DDT r1rc 1gnltcd 10 generate an inscc1icidal
nigh1. have made the sequential applic:ulon of insec1icidc smoke.GU Commercially ov<1il:1blc iiwectic:idal fos: gener;1tor5,
fre>m 1he air a more practical and economical proposition. such as the $\\ingfog. ha,e been employed in 7.irnbabwe and
Such applications need to be made 01 regular intervals, Botswana, and the I ehicle-moumed TlF,\ has been used ex-
based on the time ii cake. for a newly emerged femafo tsetse 1enslvely in lambia. fhe use of these methods ho,·e wnded to
ny 10 deposit her first mature larva. a proces£ which is tem- be discontinued a~ aerial applicmion technique:. impro\'ed.
peramre-dependent and may also depend on how well fed
the fly popularlon is.94 Applkations mu~t continue until all lvermcct in nw use ofin~rmec1in admiJ1istere.d w animab
the puparia which were in the ground a, 1hes1an of the op· as a mean:, or killing isetsc me, has been inves1igated. t-"' Al-
era1ion ha"e gh·en rise co adult mes. ano1hN process which though lies feeding on trenuid animals died. the co~t ofi1er-
Is temperature depende111. Operations are carried out in the mccti n at the dosage levels required to be eifoclil'e is
winter months. to take advantage of the tempt!ralure Inver- considered 10 be un<'conomic.al. II has been suggeMed 1ha1
sion ,comlitions am! gencrally c;almcr weacher. bui this formula1ions of ivcrmectlns other than 1hose used may be
means that five cycles of spraying at 10· to 18-da} intervals more su!cablc but they ha1e not yet been ioves1iga1ed.'.>11
may be needed when the long puparlal period a1 low tem-
peratures is considered. Endos.ulfan is the mos1 widely used insect hom1ones ln~ect growth regulators such as di-
insecticide in aerial spra;•ing at present. but deltamethrin flubenzuron. when applied to :,tdult female t$Ct5c !lie:>,. cause
was found 10 be a suitable alrematlve in thE' last Regional the female to produce lal'\ae which do not puparin1e. 1?1
Tsccse and Trypanosomiasis Control Programme operation Similar cffec1s have been found with insect junmile

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62 ,, 110, '" ,\,pcc1, h1Hucntlng the accummcc of infectious di:,t,a,c,-

hormone analo1,•ucs:;~ 115 u.~ Field studie~ In Zimbabwe (Figures 2. 15 and 2.18J used against G. morsi1a11s and
with the ju,·eniie hormo ne mimic prriprox·yien have dem- G. pa/Udiµes ha,·e a blue outerco\'Crto anract Oiesand a sec-
onsrrated that ii ,·an be used in conjunction with traps. tion of black material within the trap to induce flie~ 10 enter
using the system devel<>p(•d by Hall and Langley.89 to. in ef- 1he traps :ind •cttlc. \\"hen rM<ly to move mam· of the, flies
fect. sterilize the tsetse population. 9 'The technique was de- on the black clorh ny upwards into the white mosquito net
\"eloped lttrther b) i!pplying pyriprox}fen 111 place ol conical canop}. which leads to the retaining de,·ice. It 1~ im-
convention.ti Insecticides In tht• wide!~ used odour-bn.iti:-d portant co have the canopy well illuminated as the up\~ard
target system for killing tsetse Oies.9' Laboratory smdies on flight bin response to Lhc ligh1. In Z11luland. both G. brel'i·
trillumuron. a chitin syn1hesis inhibitor, ia.: show<.'<! that II p11/µis and G. m1$tt'l1i can be caught in a low trap \\ith a blue
induced abonion in tsetse flies at hight:!r doses as well as ol!ler cover and black internal crossed screen, and with two
preventing the emergence of adults from puparia at lower conical canopies in a lateral position. 17:! These two fly spe-
doses. thus elfectivcli• ~terilizing the flies. Triflumuron has cies tend ton~ horizontall y when taking off. rather than ver-
been te~ted on a field scale in 7Jmb:ibwe and the prelimi- tically as is the case with G. pt1llldipes and G. morsi1a11s.
nary lndications were that it would be a sunable candidate Crossed plywood paneb paimed blue and covered with
material for u,e in 1set>-e control by means Qf odour-baiicd sticl..·y material h:we been used 10 catch C. a1isce11i.102 .\
targets. These materials arc a\'ailable should it become un- much imprO\'Cd \·crsion of this t rap has been used in sun cys
102
desirable lo use pyrethrc>ids. for both <J. llllSl<'ni and G. brel'ipalpis in Zululand.
n1e perfornmnce of traps for C. morsirrms and G. 11111/id-
Traps The use of traps in auempts to control tsetse Oies ipes has been grea1ty it11provt>d roUowing the idemificarion
was first developed in Zululand, South Africa in 1930.98 ofkairomones produced b~ hos1 animals. Carbon dioxide is
) Ian~· G. pallidipit.s were killed in this control campaign but k.nown 10 anract ail sp1:cics of t~ctse flie~ on which h ha,,
the flies were not eliminated. Four per cem or more of the been tested but it is not 1ea5ible co use it on a large scale In
female flies in a populatio n must be eliminated each day for 1hc field. h:airomone~ found so far which can be u,,ed in the
a tsetse population LO decline to e>.llnction.~: Despite chc ficld arc m:e1one, methyl ethyl ketone (butanone'. t-octen-
largl' numbers killed in llarris's traps this te, ei of elimina- 3ol toctenoll, -1-methvlphenol, 3-11-propylphenol and 3-m~
tion could not have been attained. Various traps were de- thylphcnol. :'sot all of 1hese ,ubstanccs are e!fecrh·e again~t
:.igned In the 1930s arter 1he work done by Harris. 11:1. :!.M all Iii~. and not all need be included in a bait for 1110 he E-f-
most being based on a visual stimulus to the fly. o.lthough foctlve. The bail used in 7.fmhahwe for G. mor.,irmi..< and G.
Fuller and Mo~sop1i8 had shown clearly the importance of pallldi11es t<1nsbts o! 3-11-prop)'lphenol. oc1enot and -I-me-
odour to G. pt11/idiJJl!s. thyl phenol in 1h11 ratio of 1:6:12 (originally 1:4:8) comalned
Since the 1970s intensive atrempts have been made to in a polythene ,ache1/·14 245 • L47 The sachets a:re made from
improve traps for tset5e flies both in \\"est Africa32 • 1' 1 and in low-density polylhcne Jay-Oat tubing with w;1lls 0.15 mm
Zimbahwei'1 91 • 2:18 In West,\fricil. after the anractlvenessof thick and heat -:ealc."d to contain 1he bait. Sachet~ measure
blue cloth had been discovered. the effectiveness oitraps for 5 cm by:; cm and conrnln :ibout IO ml of bait. Release rates
Oies in the pa/pal/$ group was grcatl~ increa..<ed by using from the :,acheh are a runction of the densit') ofrhe polyeth·
traps covered with blue do1h (Figures 2. 15, 2.18 and 2.19). ylene 1he thickne,s of the ~nche1 walls, 'iUrface area of the
\'arious patterns of traps have been developed in .\frica.42 ~achet and the temperature at which the sachet is held. 22u
Traps may be used 10 moniwr iseise populations and are The relationship ";th temperature i,, exponential and sur-
\\idcly uwd in Africa to assess 1hc success of conrrol opera- face temperatures of ~achets In the sun are up to LO ~c
cions aiainst tsetse flies. If the traps are rreaced \\ith insccti· greater than ambient Lemperaturcs so it is difficult to give
cide. or have a coll.ector system from which the flies canno1 exact rates or bait relea)e. 11 i~ often quoted as total released
escape. the trap~ can be effective In killing large nwnbers of per day a1 a given ambien1 temperature, but the actual re-
flies. Such traps have heen used LO bre~k Lransmission of try- lease ra1e varies great!}' during the day. In addition LO the
panosomo,is in livestock and human,. 1s. 59 · 113• 1·12 bait In the sachet. acetone or butanone is rele.ased from a
In eastern and ,0L1Lhem Africa. bright royal blul' glass bottle with a hole in Llw li<l.22ij Mean release rate>
(phthalogcn blue) cloth has been incorporated lmo 1raps. 1hrough a :!-mm-diame1er hole are about 150 mg/hour and
which originally had a white oUler cover. a colour kno"1110 1hrough aG-mm-diametcr hole about 500 mg/hour.•M Plas-
b(> a11racrive co ~ome 1se1se flie5. Bo:h blue and black c.lmh rlc bonie~ ,,~th appropriaw thickness of the walls and sur-
ilHHllll11Ctl\•t, lO G. mursi1<111sand G. µullidlpes.'" but if gil'en
,..,.
face area can be: used to dispense bucanone ·· and also 10
a choice. the flies prefer to alight on black cloth. 1'01 all ~pe- rt:!placc- ,achet~ for the octenol and phenol components of
cies of flies react in the same wa~ to colour. In Zululand it the bait.
has been round thac G. amteni is strongly amacted to bright The kairomones used in the field in Zimbabwe agamst G.
royal blue cloth but not 10 bl,1clc cloth. ' 24 \\'hen g1,·en the mor~iuwsand G. pa//idip!!sare ineffective again~t the palp,1-
choice. thi~ tsetse Oy alights in roughly equal numbers on /is group oi flies in West 1\frica. apart from G, tachi11oides.
black or blue cloth. Traps such as the Epsilon and 1=3 which responds LO the bail used in Zimbabwe. although

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Figure 2.ts tps1lot1 trap snow,ng the eitra~ce

Figure 2.18 ~3 ,rap •h ·, ~g entrance and collecting appa•a;, s

Figure 2.16 Ta,c:e, wil/1 black centre .ind two n1ming pa~;is

Figure 2.17 Slat~ t!Olll ia,get being sprayed v,1m a;; :ame:Ji:,n

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64 • '"" t•\I; :hpec1s lnOucncinv the ocn:rrcnce of ink-ctious dbcasc,

different proportions of the consliwents are needed. In 3 suitable re<,idual insecuclde. rhe !lies alighting on it or sim-
\\'est Africa G. /011gipalpis (morsiran$ group• and G. medi- ply touching it will be killed. Canle dipped in Dl)T solution
,;orum /usca group) also respond 10 the bait u,ed In Zim- \\'E.'re used to <'liminate ,mall population~ of G pal/ldipes in
bab\\'e,i•n Zululand irl 19-19 ti(; The same merhod was im·estigated m
In Zululand, C. 1111s1eni re.sponds 10 t"drbon dioxide but East ,\frka.zs. bm was 1101 widely accepted. TI1e develop·
not 10 any of thl? mher known kalromont-s. <i/l>.<s/1w lm:vi- ment of synthetic pyrethrold Insecticides. such a, deha-
palpis rc.sponds to carbon dioxide. octenol. ·1-nwchylphcnol methrin. which are good acaricidf>_, as well as insecticides.
und acetone (but n<•t to butanone). 1?~ The dose~ of octenol has re,ived interest in using dipped cattle to elir:1i11a1e lSCt$e
and 4·mt!thylphenol used in Zululand are higher 1han those me, In lightly infested area&. In Zimbabwe, a we.I-developed
u~ed m Zimbabwe and the use of acetone 1s onlr 01 benefi1 dipping sys1em against ticks is opera red in most part,. of the
in the d~ season. ·1 he reason for thb scu~onal t.'ffect has not cmu11~·. ln communal, or traditional areas bordering on
yet been determined. known 1~c-1se II~ areas, where caule numbers are large
Behaviour of tsetse flies in odour _plume~ resulting from enough 10 give good coverage oflikely Oy habitat, a dip wash
release of kairomones has been inve~tigated 1 .!:?J. ,i, and it comaining ddtamethrin i~ being used. Cattle are dipped
has been ~hown that flies mai' react ctilforcntly at different every 14 day~ or. in the absence of dipping facilities. treated
doses of odour. Comparison of synthetic oduu no the odour mon1hl1' "ith pour-on formulations of the dip. Flies aligh1-
from an ox. at the same dose, indicated that further attrac..'t· ing on the c:aule are killed.2rn or knocked d0\\11 ~o as w be
am(s) e"ist in natural ox odour• tor both G. morsiransand G. available 10 1>reda1ors. and th,• reduction in trvpan()somo,is
pallidipes. 1\ similar conclusion was reached for r.. brei•ipa/· cases in areas under 1rca1ment has been dramacic. This ap-
11is, and possibly for C. a11s1<mi. 12° Comparing odour from proach is considered a 1·e~· promising form of l}et,11 fly con·
small to large numbers of caule "ith an equivalent do~e ofth11 trol \,·here cattle dcn~itic~ are high. !rt term5 of area under
synthetic odour used in 7.imbabwe ior G. morsira11.< and G. trea1men1. dchamc1.hrin dipping is 1.he most \\idel> used
pa/lidi1,er' demonstrated a relation,hip between quantities anti-tset~~ mt-asure in Zimbabwe at prcsem.!O. The cos1s
ofodour and llycatch, su that a tenfold increa,e in odour gave are less than tho,c iur an> other meLhod of control. pro\·ided
a 2,3· ro ~.8-fold increase in catch for C. pallidipR~ and 1.4- 1ha[ rhe infrastrncture for dipping procedures is m·ailable.
t'old increa~e for G. morsiums. It wa~ aho sho\,11 that odow· Successe~ 111 using insec1icide-treatcd cattle havl' been
From the cattle w,1> more cllcctive than that from the syn- reported l'rom ~ouchern and eastern \frica.~0 as \\'ell a,
theiic odour. again indicating tha1 further aurac:iant(s) re- from Wes1 Africa. ll has been shown that pour•on applica·
main to be identified in host odour. Skin secretions (sebum) tion~ of insecticides do 1101 reduce the attructivenes, of
of cattle also alfect the beha11our of t~etse lli11s,-·;? especially caulc co G. 11<1/lifiipesand do n0t affect rhe proporrion of flies
after alighting on a stmiona~· cloth screen. that feed on the animals.6 Disease transmission can thus
The way in which tsetse flie~ move through the habitat still occur after tht> caule have been treated. and "ill only
has an important bearing on wherl trap\ should be posi- cease after 1he 1~e1sc population has been reduced b> the in-
tioned so as 10 be most effecti1•e. Tsetse nies use game 1raits secticide and 1hen only if 1here is no invasion of flies into
when folio\, ing odour in thicket 176 and 1end to 11) round the area 1,here can!~ have been 1rea1ed with insecticide.
bu~hcs rather than over them. \'alt:' ha, developed a deter- \'arioth insecticides for use as pour-on, plungl' dipping or
ministic model to inctic:arc how tiies move through three sprarlnfl of canle are now available. 2
vege1arion types in Zimbabwe. 241 When placing trap~ in the In comparatl\'e trials in i.imbabwe with three different
fic:,ld, they should be clearly visible 10 a fly bu1 need nm be in p11ur-ons. the arih,c Jngrcditmls of which wen, alphacyper-
a large clearing. Provided there are breaks in the \'t1getation merhrin. deltanwthrin and c~,fluthrin. respective!}. the
of over a metre wide. a hungry fly ,di! mo\'e up an odour dung of the animal treated with alphacyperm~>thrin. which
trail. Tllert>should be no dis1rac1ing host-sized objcctS, such was dropped about ~c~·en hours.lfter the inseccidde was ap·
as rocks or small dense bushes near the trap. ,md ii the trap plied. was found to be lethal to dung beerles.~42 In the event
is of the ,inglc entrance Epsilon or F3 type. the entrance of this phenomt•non being common to oilier pour-on, and
should face downwind. le ls particular!\' impona111 that the also dip wash!/,. their widt'Spread use for r~etsc ll~ and 1ick
canopr of the trap is well illuminated so that flies mo\'e up control purposes could have considerable en\'inrnmemal
inco the collecting de\'ice. lf a trap is needed in a po~illon implications 10 soil f(•rtility.
that doe, not meet the requirements. a small ;11noum of W'herc 1seL~e flie, are abundam. cattle canno1 he casil~
trimming of the vegetation \'.aJl improve thll quality of the maintained. bul .inificial l<lfgcts can be u,cd agilin~t the
position and resuh in beuer catches. l n wcll-gras$ed areas. it mes [Figures 2. 16 and 2.17). The~-c targ('1S consist of a cloth
will he necessary to make a clearing round the trap to pro- scrc<!n in a fram" mountl'd on a pole. In some \'ersion~ the
tect thl' uap again St fires. cloch i;. !lankcd b} bands of black mosquito netting, which is
not readily 1isible to 1se1se mes. The screen and thl' gauze
Targets The objec1ive, or target, <)fa hungry 1sctse fly is the arc spray¢d \\ith lnsectidde. often with llehamcthrin sus-
hos1 animal on which II will feed. It the target is treated wirh pension concentrate. in \\est Africa. blue,-crecns. which are

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 Wctots:'rsetse Ille, 65

disadvantages oirhe doth and neuing target wa~ that the

neulng soon became torn. It was found 1ha1 an all-blac:k
cloth rnrget 1;; m wide and 1.0 m deep was a more robust
substitute. Tith all-black cloth rnrget was used widely in
Zimbabwe In conjunction with k.airomones and wl1h delta-
me1hrln as the insecticide applied to the targer CFigu ,·e
2.17).
Work in West Africa on the JJnlpalisgroup offlil" showed
thm bright royal blue targets were heller anractams than
black rargets.i,.1 In Somalia, it was shown that a combination
ofblu<: and black cloth targets were more effective than all-
black targets against G. pa/lidipes.2'.!11 Furcher work in \ ,·r.st
Africa showed that the best pauem for G. palpnlis \',a$ a
bright royal blue centre panel fhmked by two black panels,
with both colours being amnctive 10 the ny but the black
' _....·--;"' cloth being the preierred substrate for alighting. enhancing
the tend.enc) for 1his fly 10 set tie on the outer edge~ of any
Figure 2.20 Blue-l:lack odour-ba"ed insecticide-:reate!I target used in targel.
Zimbab\•1e agamst G/Qssin;, n:ors11ans morsH11ns and G peHidipes. A bait
sachet is sho\\111 partlv withdrawn from the pocke: in tll3 b ~~ clot~ and a A detallt!d scudr 1, a, made or colour combinations and
bottle of buianone is on the grcn,md in front of ;he :arge: materials tha, could be use<! to make targets for G. morsi-
cuns and G. pnllidi11e;; in Zimbabwe.2~ 0 Although larger tar-
gets do perform beuer. an acceptable target had a central
good visual atrractants, are used so 1ha1 flies come 10 the tar- black panel O74 m wide, flanked by bright royal blue pan-
get. alight on it and receive a lethal dose of insecticide. Suf- els 0,5 m wide . .-\II panels were I m high. Both G. n:orsiums
ficien l 1arge1:; must be deployed to ensure that flies have a and G. pa/lid:pe~ tend to set Lie near the centre of a rarget
high probability of encountering one. Very good results have and this trend was enhanced b>' the black cenrre panel,
been achJeved u~ing this met.hod In West:\frica. " which isa preferred aligh1ings1imulus for these spccies.:\n
I! has been shown239 that tse~e nles flying in an odour additfonal advantage of chis pauern of 1arge1 is that only
plume are diverted 10 host-like stationary objects and a va- rhe black cloth needs to be treated with insectiC'idc. Sereral
riety of visual cues are involved.ui Differences in the de- alternatives to cloths were identified for use in target con-
gree of response by different species of tse1se Oies to host- Strucrion, but have not ret been accepted for control op·
derived stimuli ha\'e been demonsmlled.t 52 in Zimbabwe, erarioni, in Zimbabwe.
targets are used in conjunct ion with the kairomones dis- In the original field tnal of the odour•baited target sys-
cussed above. The rnrget first used in routine control op- Ulm247 the insecrkide dosage wa~ low and 1arge1s had 10 be
er.nlons consisted of a black cloth cemre panel. flanked by sprayed e"e~ four momh,; to ensure high fly mo:ialit). A
panels of black mosquito nelling. mounted cm a pole and study 227 shov:ed thar a d<>sc of I g. active ingredienr. of del·
pivoted so that the target faced across the pre,·aiiing wind tame1htin per square metre of cloth killed almos1 alt flies
(Figure 2. 16). The combination of an odour and a vfaual at- nlighling on hover a period or a year. Test cloth~ were held
traciam has been veryeffeetive in attracting both G. mor.fi- in full sun and exposed to 1he elcmems in the field. D!fferem
ttms and G. pallicli11es. ancl the black cloLh panel causes qualitlei: of cloth l'aried with respecl to rei;idue longe\'ity
more mes to alight than does bl ue cloth. Hies colliding and also in the extent of f11ding of the black dye under field
with the black netting also contract a lethal dose of pes1i- conditions. Only lhcs11spension concentrale formulauon of
cide. l"hese targets were first shown 10 have potential value dcl1ame1hrin gave the required longcl'il}, although alterna-
by their use in controlling G. morsimn~ and G. pallidipes on tive insecticides may be used.1 In limbabwe, th<' applica-
an island in Lake Kariba. 246 The first large-scale lield trial tion of insecticide is now done at a cemral point and, to carnr
(600 square kilometres) was subsl'queml~ conducted in for possible 1-ariation in cloth quality. lhe in,.ccticlde b ap-
the Zambezi River Valley. where both G. morsit<ms and G. plied 10 give 2 g of active Ingredient on the central black
pallidipes were ~bundam.207 The le\'el oi success achieved doth panel to,i5 m2 J. This blue and black. target is the pat-
in this trial. in which tari;et:. baited with acetone and tern U$Ccl in Zimbabwe at pre~ent (Figure 2.20).
ocrenol and sprayed with delramcthrit: were deplt}yed at lnvesligations showed thac 10 g of the phenols and
three to five per square kilomerre, resulted in a campaign octenol mi.,ture were needed in a sachet if il was to la~t a
against G. marsirans in fhe Umfurudzi area of Zimbabwe. year. It woltld also be necessary for the sachet 10 be in a
This campaign was successful: tsetse flies were eradicated pocket on the blue section of the targcr for this degree oflon-
in 18 months and. complementary t0 1nis. trypanosomosis gevity in fuU sun. 111e bait would ooly last abom seven
in canle in the area disappeared completely. One of the months if held in a pocket on the black clorh.n~ :\ bonle

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66 <rl"'l'>' " . .\open, ,nOuei\ct:lJ; the occurrence oflnfecliou, di~c~sc,

containing 750 ml of arctonc or buumom: \\ith a 2-mm- diffC'r her.wen species. and in different \'ege1a1ion pauern,.
diameter hole in the lid }ields odour for a }Car ,\1 the ,iie oi the 1arge1 in the field. feature» such as good vfs·
The original approach to deploying odour-baltr.d target, lbilit\', absence of distracting object~ and pathway> through
in Zimbab1,e wa,. to aim at a relatiwl} uniform d1s1ributiou the vegetationi• arc imponani. 8$ is the case for odour-
of 1argeri. on the ground at an a,·crage of thrcr 10 five per baited trnp>. It 1:, necessary to have a fire break round each
,4uarc kllu1111:u1:. 1 •· This d1:11,Jty gave a mpld rcllunlun in target . .i11ll llw grou11cl round the target should bl! reason·
the tsetse population. and climinat.on of G. morsiran,< and ably level ,o the target swings easily.
G. 1mllidipes co11ld be achieved i,ithin a year. This approach TI1ere are fac1or:. that adversely aftec1 the use of the
has been used extensively. with Lile moditications hl iarget odour-baited targ1;1 S) ,tcm tor 1sctsc comroi. The prime
colour and longer lasting insecticide and baits. with good re- cause of failure is theft of targets by mral populations. Fail-
sults in Zimbabwe. \lala\\1, 'samibia and 7..ambia. provided ure to ma.kc adequate fire breaks round Hll'f(ets is another
tht' guidelines for the method were iollowed. ·\ modification major cause of failure of the target system \\'ind damage to
of the pattern of deploym<'nt in l.imbab\\e ha~ been fol- tar1tet:. h ano1her ~e,·cre t'llcct in many localities. and targets
lowed in rhe broken country near the Zambezi River \':tile} may become ini,ffe('ti\l' In a matter or wcl!k\ alter being ,et
esc:arpme111. As was the case nt the time 1hat ground ~pray- UJ)- ,\ltcrmtth·e mmerial, to cloth may help ~olve this prob-
ing was in use, targets were deployed only in the es..ential lem. In .ome localities. damage b> "ild animals such a~
hot ~eason habitat or the me~. which wa, g.-nerally asso- elephants. baboon, and hyal'nas can rearh high lt>l'els, al-
ciated with drd.inage lines. These situations can be seen on though in much 01 che undeveloped country in Zimbabwe
aerial photographs, ;md careful planning 10 po~ition the tar- dam~ge levels are low. r)omcsticcattle are also kno\\TI 10 de-
gets in trnnseCt$ along the drainage lines wns done. 1he stroy odour-baiwd 1argClb, A major difficulty in operating
number or targets per square kilometre totnl area w,ts ~till the system i~ to ensure maintenance or the lllrgets. To relo-
about four. but the density in es,;er.tlal habitat wn, higher. cate targets rcqmres that 1heir position be marked on aerial
Thi!> approach has given good resul!:. in elimination of both phowgr;:iphsor map~. and global positioning >}'stem IGPS)
species ofOies where drainage llnes ore wcll de,·clopcd. It has instrume11ts arc l'cry useful in this respect. .\ccess to the area
not always been Stfccessful where drlinagc is indeterminate, being treated i> es)emiol. and the supply of the target.sand
and in '<tiCh areas a regular grid of targeu; ma~ be necessary. bai~~ at the correct time~ is a complex logistical problem in
FolJo"ing the outbreak oft!) panosomosis in Zululand in much or Africa. fhc capacity to operate the odour-baited
1990. the all -black odour-baited target sr.s1em ust!d in Zim- wrgc1 system doe. not exist ill all the 1,e1se-infested coun-
babwe wa~ introduced in a small-~cale trial. Some initial re- trit•s of southern Afr!ca, so the elimination of the t~ct~e fly by
duction in tsetse numbers was noted but the method did 1101 1his means will n<>1 he a shon- 1erm operation.
e!iminau: the flies. 125 Research 1,·ork ~hO\\'ed that G. bri•t•i· Some oi the,e difficuhic,. might be solved Ii it were pos-
pa/pis and G. f1t1sre11i dl!fcred in 1heir rcspon,e 10 black ~iblc lCl convert tree~ to odour-baited target;. This can be
cloch as discussed above in Lhc soc1ion on rrap,. and only G. done qunc easil} for tree stumps, but ll has been ,hown that
brevipa/pfs respondt!d 10 the kairomon~" us.ed in Zimba- it is not ea,y to do for 1rees?1' 1and was more expensive: to do
bwe.12.; \\'ith th<i knowledge gained from eiqJt:rimem~ on than using clorlt tnrgets.
the reaction of the 1wo fly spcdes. a 1arge1 was developed ..\11empt:, have been made 10 encourage local communi-
which. although not lite optimum for either ~pecie•. was the tie< In ~e1~e-infestl:'d areas to run their own N~be control
be~t com1>mmise. 111c target w1i- 1.,5 m long and l m high operati01b. A succe<,~iul uial. u~lng Lraps rathe• than 1arge1~
with a cemre blue panel 75 cm \\'Ide. nanked b>· two 50-cm- and a simplified bai1 consisting ofcau.le urine and acetone.
11~de black paneb. in Glo,tsirl(I brel'ipa/pis tend" to alight tO· wa~ nm at :,:gumman in Kenya. and ~imilar ,ystem~ have
"-ards the edge~ ofa target and the black cloth enhance, this been operated in \\'est Africa. ·9 A community-opera1ed
effect so. for this species. only Lhe black doth would need 10 odour-baited carget S\'Sh!m 1s under invc,tigatlon in eastern
be treated with pesticide. Glo$,l11a au;1(•11i is attract<'d by the Z.1mbia. Preliminal'\ result~ arc encouraging, but it has 1101
blue cloth but 11ill alight on either colour of cloth so. for this yet been dcmomtratcd 1hat this c:ommunil} \\ill operaw the
species. the whole carget would m'Cd to be treated. Tlw bai1 "ystcm for protracted periods.
used in Zimbabwe has also been modified. and b only effec-
til'e for G. /,re1•ipalpi.~. while for G. ausuml only the ,i,u.il at- Target barriers The !ir.1 large-scale trial of odour-baited
traction to bright royal blue cloth will attract the nv. When targets 1c climinaw (;. mor.~i,ans and G. pallidipe., in Zim-
the daily pnttcro oi movcmum and dir.pcrsal of the 1wo fly bobwc2'; demonsmm•d that this could be achieved in 12
species has bee11 determined, it should be po~•ible to devise months if the mean targei den&ity ll'aS three 10 nve per
an odour-baited target syscem which \1ill eliminate both square kilometre, Trap samples from a traverse across the
G. brel'fpa/pis and G. a11sre11i. cemre of the main i1wasion front ~'ielded no flie:;, fun her into
The pattern of deploymem of1he odour-baiied targets in the treated area than 5 km. The wrgetsalso exerted a marked
the field has been modeUed!•ll · 262 on the basis of known reduction of che tsetse population omside rhe treatment
features of tset~e movement. The ceployment needed will area. These results ga,·e riMno 1heroncep1 tha1 r~rge1scould

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be used as a barrier to in\'asion by 1~c1se nles. lnitially, ii \,as on cattle treated with insecticide and that 1ra11,:mis,ion of
coru.idered that a narro11 barrier 11ith a high den~ity ot 1ar- t rypano~omo~I, ocrurrcd.
get~ would gil'e the bos1 rC{.uh. II has ~ince been sho1111 1NI Odour-bailed 1argt'l harriers to ln\'asion b} tsetse flies
tha1 such harriers do not prcvem some tlies crossing them. can be e!fectl,e. but 1hey must be cc1refull~ maintained a1 all
l)sing knowledge of fiy sun·i\'al rates, cxpecuuion of 1hc.- nc1 times. Attempt, art' hemg made to p111 the maintenance uf
daily movement of 1he flies and the prohabtllryofa 11} being target barriers out to comrac1 but, as the barrier, wtll ha\·e to
killed by an odour-baited Larger tn~ated \\ilh deliamcthrin. bo maimained for che foreseeable future. it remains to be
Hargrove9l ha~ computed the probability of me~ c::ros~lng seen how effect ii e thi; will be.
barriers of different wid1h~ and target densities. An eltuc1iw
barrier widLh of eigh1 1imes 1he expectation of net daily Other approaches
mo\'ement of the me~ (ahou1 a kilometre in l.imbabweJ, Chromosomal abnormalities Following the developmeni
when the 1arge1 density was the ~ame as for csetse elhnina- of theso•called ~tcrilc male technique for in~ect control. the
tion in 12 momhs. was proposed. In Zimbabwe, the target principles oi which 1,ere reviewed by Knipllng. 1r the
density would be a mean of four per sq~are kllomene and tnllthod ha,; betn tried against a wide range of in$ect:.. 128 .-\~
1he width .ibom 8 km for G. pt11/idi/1l!.S and po~ihl> Iese; for 11 hac.l been \'ery ~ucce,sf\ll against Lhe screw-worm fly Co-
G. mor$/u111s. The number of rargets in\'oh·ed would be less chliomyfa Jwm/11/1,orax' in :,.;onh .\merlca, It is 1,c,1 surpris-
than that of a barrier of four line~ of iarge:s0,5 km apart with ing that it was considered for the control of Glo~illa spp.
a high density of U1rge1s in the lines. TI1e probability of a Oy \\~lh Lhcir ve11 low ra1e, of reprodurtion. Initial theoretical
crossing the proposed barrier would be about 0,001. As the considerations for using 1his approach agains1 GlossiluJ
barrier exens an effect beyond it~ boundary, r.he Oy number~ were reporced b) Simp~on in 1958.201 The irradiation of pu-
on r.he invasion front of the barrier decline as the barrier is paria as a method for rearing male tsetse Oles with chromo -
approached. The predictions on barrier cffecth·eness do not some abnormalities and thus effectively sterile if mated co
take into account game trails. foo1po1hs and road~ that CT0.\i~ normal femal~s l\',h d1:velopcd br Po1tb. 110 '
the barrlE'r. Such features may reduce the effectil'enes, of The firs, ticld trial of Lhe~1erile male 1echniquu \\'a~ con-
the barrier through transport of flies on vehicles or animals. duc1ed on an island in Lake Kariba between 196i and
A 1rfal ot the type of barrier proposed by tlargrol'e9 ~ was 1969.'6 The test popula1ion was reduced b) 98 per cent in
conducted on a section oft he boundarv between Zimbabwe 1his campaign. and Cll'entually reached the stage where no
and 111ozambiquc in the Mudzi distrin.m fhe barrier was Oies could be caught by any of the available sampling meth -
•10 km l<mg. 8 km wide and target lines were 0,5 km apart od~. In thi!. trial, ,·hromosomal abnormalities were induced
with targets ~paced 0.5 km apart in rhe line. The country Is in male!> by chemical mean!> as !he llics emerged from field-
rt<latil'ely !lat and che cargec dcnsiry In this regular pattern collected puparia. Thh,approach obviated che need for large
was rough!~ four per square kilnme1re. l.<>cally 0\111ed cattle colonies of behe t1ie:; 10 produce puparla for Irradiation at a
ttl!ated wi1h delramethrfn gruz-ed within the barrier and for a known agc. lMs
disiancl' of about 20 km from the international boundary. A fu11her auempt at using the sterile male method, this
The mean dcn,lly of ca11le in the 428 kml trial area was 8 10 1ime based on 1he irradiation of puparia produced by a large
12 per ,quare kJ\ometre. Ille distribution or cattle was not colony of tsetse llie~. was made some yenrs later in Ea,, Af·
uniform and there 1rns no comrol oven~here the,· grazed. nca.264 The area imo \\'hich the treated flies were released
\lonitoring of tsetse me« (main!~, G. morsitnn~ bu1 wifh w:u- i~olated I>~ :t wide mn•! cleared of hu,h which in the
G. µallidipes also presenrJ and trypanosomosis showed Ille~ cven1. was not compll'tcly t1ffecch·e. 1 ho 11}' population in
to occur in low numbers no furtherthan 2 km rrom che inva the area was reduced to a very low level. but \\as noc elim-
sion from and for case:, or trypanosomosis 10 be rare. A 1ri.1I inated. The development of membrane feedir.g tech-
W3$ designed 10 check whether the insecticide-treated cattle niques ~1 ha~ made latge-sc-.tle rearing of tset~e llies more
alone would provide a barrier to mes moving into Zim- practical. and release of male flies sterilized b~ irr-.idiation
babwe. rollo11ing remo,-ru or che targets. there w.is a rapicl from such colonies has re,ulted in the elimination of ,ome
incroa~e in the distance into the barrier that flie" could be fly popular!ons in W.::,1 i\frkn. •J
caught and also a rapid increase in the number of cases of On Zanzibar island. G. au,,1e111 has been eliminated usmg
u-ypanosomosis in the c,utlt•. induding some iatalitie\. the ,terile male approadi. Large-scale (up 10 one million ie·
Following replacement or the- Large1», the £ituation ,oon mnles) rearing of Oie, u•ing membrane feeding technique~
returned 1othe originalstaie. The trial demonstTau,d 1hat, at was done on the mainland oiTan;mnia and puparia \\'Cre ir·
this Jocallt). the densiLy of insec1icidc-trea1ed cattle did not radla1ed. resulling in ctrectl\'ely sterile male flies, Other co11-
pre,,em rues from ;\loza1ttbique emering Zimbabwe. l low 1rol measures such as treating cattle with insecticide;. were
far into /.imbabwe the files would penetrate "·as not deter· also in use, but chi! sterile insect technique W.b the mclhod
mined, as it appeared tl1a1 they might cross 1he 20 km-wide which finally eliminated lh<· ny. The last wild lly to be caught
belt in \\'hich cattle had been 1rea1cd ,,i:h dehamethrin. so was in 1996.1q'
the targets \,ere replaced. ll 1,as clearlrshown that flies fed An alternative 10 rearing flies for sterilizmion is to

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68 ,, ,,"" ,v: Aspect$ influencing 1hc occurrence of infcc1itJ11< tlisc3<.<?s

srerlli1,e males in a wild population bv chemical means (au- Stralegy of trypanosomosis control
tosterilization). The development of efficient odour-baited There arc three main strategies which may be used. indi·
rraps h.is opened the way for lhb approach In Zlmbabll'e. \idually or in combination. to control trypanosomosis in
where a suitable sterilizing device has been developed 1"' Jivesmck. These arc 1he administration oi curati\'e or pro-
and tested ,-uccessfully?4a but the ~·el') toxic nature of the phylactic drugs: the use Ol ll')-panotoleram livestock. and
chemicals used preclude~ its wldesprend appllcatron. The vector comrol or eradication. The choice of the tactics 10 be
1uvenile honnone mimic and the chitin synthesis inhibitor used \\ill depend on existing circumstances ~!any varl ·
mentioned above may provide a safe way to aumsterilize fe- able;., such .~s the species ol fly. vegetation type. topography,
male tset.w flies effectively. These method~ elimi11ate lhc and both human and animal popula1ion densiry, mu.st be
need for the cosLI)' anificially maintained tsetse fly colonies. considered. \lerhods which have been successiul in one lo-
l Lhas been suggested t hal. under some circumstances. the cnlity may 1101 be effective in another. The strategy finally
sterUizaLion approach to tse1se fly control may be best i,,n chosen u.ually depends largely on economics, and, in gen-
but, until more accurate information is available on fly popu· eral, the benefits denved from controlling the disease
lation dynamic~. the use of population models in prediC'ting should be greater than the coM of achie, ing this objecti\'e.
lhe outcome of control measures is unlikely to be accurate. l:xceptions do occur, however. and an example ls the threat-
Other approaches to control bi!sed on lhe genetics or ened invasion of pastoral areas by vecmrs. Here. large e.x·
tsetse flies have been investigated. The possibility or elimi· penditur11 may be required simpl~ to avoid rhe potemially
nating G. morsitm,s by introducil1g G. swy,111erumi into the catastrophic los~e~ that might occur if the im'l!~ion were not
G. morsiian;; habitat, where it would not become esiab· checked. i\ good example oi such a situation in soulhem Af.
llshed. wa!> examined.11 ~ fhe,e two :,pecies crossmate Tica was the control, between 1959 and 1969. of the advanc·
readily in the laboratory and the inwrspC'dfic cro~s issrerile. ing bel~ lly front from :...lo.'.ambique imo $ou1h-eastem
It was hoped that G. morsira11swould be reduced in number, Zimbabwe. which was threatening 10 rcinfcst the Limpopo
as man) female$ would mate 1\ith male G. su.,-11neno11i, Rh·er ,·alley 1"' Game park$ and game or fores: resen·es are
however lhe field experiment was not succci;sful. 11 Other abo ~pecial cases. L!ule or no benefit is gained b) the pMk or
suggestions for control based on genetics-such as releas- reserve b)• cumrolling tsecse flies. but a.djaceni agricultural
ing labor:11ol')'·brcd strains of tsetse flies. homozygous for area,- benefit great!:,. This situation occurred in the earlier
translocations, into natural populations~ 5 - have not been pan of il:i" cemul')' in Zululai1d, South Africa."° and is the
cried in the field. current ~llllatlon in pans of Malawi.
The economic aspects arc 01· prime importance .\lost
Biological control Some animals. ~uch as birds. wasps countries In sou1hern Africa cannot finance attempts ro
cBembex spp.J, robber Clies (AsilidaeJ and spiders (Hersi/ia eliminate flies ,,iLhin lheir borders. ·111ey can only try 10
spp.). are known to prey on tsetse flies on occ:asion. 171 As !es.sen 1he 1mpac.t of trypanosomosis either br comrolling
these animals do 1101 feed only on tserse mes. ther are not the vectors or hy u~ing drngs to control the disease. Should
candidates (or ellminauon of the flies. it be possible for a country 10 eliminate the flies ,,;thin ii!,
A number of insect para$iloid, \\hich parasiti7e on Glo~- border,. a, is the case in Zimbab\\'e, there Is still the
.<i1111 puparia have been discm·ered. 1'°· 171 ReC"ent taxonomic perpe111al threat of reinvasion from tsetse fly-infested
re\'isions have altered the old names of manroflhem. 81 One adjacem countritts :\ucmpcs to prevent rein\'asion b) the
of lhe parMi1olck wa~ released on a large scale in an a11empt u~e nr :irtifiri>,l barrier,, surh a~ 70nP~ nr "lrra<. rlrarrrl of
to control G. mor$iuws in Malawi.' 29 but it was not success- bush. general!~ ha\'e not proved satisfactol')· il1 the long
fu I. possibly because most of lhe parasitOids are not host· run. The U)e of newt>r types of barriers. such as inseC'ticide-
specific. trcatcd, odour-baited targets, or the dipping of StOC'k In
A parasitoid wasp in 1he famih ~lutillidae tChresco- deltamethin, is still being assessed. In an}' event, the
m11tilla glossi1t(le Turner) is well kno\\11 in Zimbabwe3'>. 101 maintenance of barriers is cxpensiwt and external funding
102
and hai; also been recorded in Zambia and Mala,,~. agencies do not readily contribute to such ongoing control
Under natural condition~ it appears to parasitize only pu- tncasures.
paria of the genus Glossina, but in rhelaborat0ryitca11, \o\ith Eradication of the tsetse Oy is, asof I 999. the only perma-
difficuh:y, be reared from Sarcophaga sp. puparia.34. l(l{i It nent ~olution to lhe eJimination of disease caused b~ the try·
has not yet been po,sible t<) culture the\\'asp in the numbers pai1osom('s which they transmit. ~lelhods used 10 kill che
11ccded m a11emp1 biological control. t~etse flies arc based on detailed knowledge qf their biology
A ,irus, widespread in Africa. 1hat infec~ C. palli,iipe~ and ecology. in To achiev,· l.'radica1ion. 1l1e fUe,; must be re-
and cause, hypertrophy of the salivary glands oiboth sexes tnoved up to nalural barriers such as lakes. open g=sland.
and sterility in the moles ha~ been reported. 1'.".t If it is pos- heavily ,cttlcd area, or mountain range~. not merelr to ter-
sible ro de,1se methods by which the dissemination of the ritorial boundarie:.. unle!;:; these happen to run along natu·
,irus could be improved, it has potential for biological con- ral barrier~. Clearly, co-operation between neighbounng
trol purposes. countries iS impcrativ<i in order 10 achieve this objective

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 Vectors: T•~cw me, 69

and. .JS 1.he cos1. is likely lO be higher than most coumries ground spraying. the use o[ targets. and !.he topical applica-
in\'olved can afford. international assistance muM be tion of deltamethrln 10 cmile. The cleared area is protected
soughL by a series of target barriers located along the tsetse front
A prnposal 1.0 pro,ide a permanem solu1ion 1.0 the 1.se1.se and regular application of deltamethrin co all canle depas-
fly problem in ~lalawi, much of:-.'101.ambique. eastern Zam- tured within 30 km of the :\·lozamhique border.
bia and Zlmb:tbwc was prepared In 1983 by Peter Thoma$ Although !he Regional Tse1se and Trypan0,~omiusis c;on-
Assuc1atcs Consulting Services rlimbabwel and the Minis- rrol Programme hM been terminated, ir seems likely that
ter of ,\gricuhure (United Kingdom).''' In the long term, 1.hb collaboration In tsetse control in southem .\frlca will be
comprised the elimlna1ion of the Uy bell common 10 these continued through the escablishme111 of a regional network
four countries. except for the coumlj east of Lake Malawi co-ordina1.ed by Zimbabwe as the Southern African De,·el-
and north of the Zambeii. The region 10 be cleared of m~ <>pment Community':- Centre of Excellence in tsetse control.
extended to namral barriers for almost the whole area ln- This a.rrangemem ha.< still 10 be endorsed b}' the SADC
voh·ed and left only a small stretch along the Zambez.i River ~ linisters of Agricu !lure.
in Mo1.ambique, where an artificial 'barrier would be In 1996. a new grouping of agencies, the Food and .-\gri-
needed. The European Commi&sion (of the European cuhurc Organization of the United Nations, the World
Union) agreed to finance the programme, which was known Health Organization. the International Awmic Energy
as the Regional Tsetse and Trypanosomiasis Control Pro- Agency and the Organi7.ation of African Unity/International
gramme of '.\lalawi. Mozambique, ZMnbia and Zimbab\\'e. Bureau for Animal Resources initiated a programme called
Implementation was divided imo two phases. The first Programme Against African T1ypanosomiasis CP.'\.~T). The
phase began ln 1986 and the second iu 1992. The projeet was stated objectl\·e b to promote integrated trypanosomosis
closed at the end of 1998. control through co-ordinated international action so as to
The first phase involved the follo\\~ngcomponcnts: iden- impro\'e food security and sustainable agriculture and rural
tification of the extem of the problem; the deYelopment of developme11t.
rsctse-control techniques; pro\'ision of training: emergency 111 this chapter. drug treatments, trypanotolcrant lh·e,
eradication operations: and environmental monitoring of stock and attempts to develop .a vaccine against tij-pano-
control activities involving the use of insecticides. The somes are not discus:.ed. although they may be effective
evaluation of the first phase recommended the continuation srrategles for 1rypanosomosis comrol (see Chapter 12: Afri-
of the project imo a second phase. The prime objective o( can animal rrypanosomoses}.
th is phase was to develop a comprehensive strategic plan for Trypanosomosis will cominue 10 be a major constraint 10
an integrated strategy for the eradication of tserse-rransmlt · livestock developmem in much of sou thern :\frica for some
ted trypanosomoses from the common nr belt. The second time to come. Countries such as Malawi. and to a lesser ex-
mam objerti\'e was 10 complete work s1aned in the tlrs1. tent Zimbahwc, ha1·e a relati\·ely high human population
phise. During the second phase. because of the scale of rhe density. so the need IO eliminate tsetse flies is more urgent.
undertaking. the objective of the programme was redefined However, even in sparsely populated cow1tries such as
as integrating strategic tsetse and trypanosomosis control Zambia. the population growth rate is high and. should it re-
to support sustainable development in the countries main so. it is only a matter of time before the need to elimi-
concerned. nat.e nypano~cnnosis becomes u rgent Population growth
It i~ unfornrnatl' that thl' projP.l't wns rPnnir1atPd whE>n it ra.res may change in the f111ure. and furtherimprov1,ments in
was. because all four countries bad achieved considerable measures to control trypanosomosis may be developed but,
successes. In particular, Zimbabwe had cleared tsetse from over a cenmry after the 1895 demonstration b: Bn1ce of the
20 -100 km' in the north-eastern part of the cOUntlj', includ- association between uypanosomes and tsetse flies, 16 there
ing all of the farming land which was O\'emm by flies during is no indication yet of any rapid or easy solution to the del-
the War of Llberalion.79 This result was achie\'ed by a com- eterious effects of uypanosomosis on humans and their do-
bination of control methods, including aerial spra)•ing. mestic animais in southern Africa.

RefeJen ces
,u.s.wP. R.. 19/14 Con1tol pf 1$l!t~c Ilic:$ (Olp,~r:,, Clos>lmd~cJ using 3 Aum.,, e.t.. 1903, .-1 r,14nogrop/• 0/1/:e t.wt.u> Jli'4 tC.11111(;!0,slnn,
.n«'<llicldc<; A 11:'1<'" Md ru1urcpro~pce1~. 811/l,t1nof6nwmalo/li<ttl Wt..sw'OOd.' lwwl nn thtco/l('('t(an in 11tt Bl'irislt ,\fu.sf!mn. \f,:Jt u
Rt"tMrtii. :".$, 1-,1. cllnp1ero11 mo/11/J p,,ru by /fJ. H<r1tSlm. Ph// D«. Briusb ~luseum.
2 u..sop, ~., .. 1993, Tr/limns mamuJJforr,.,ts.control pt>r,cnn,•I Vol 5 London
!111ttticldcs for ,~ue nnd 11}'/H1110$0mltlsls conuol using 11ttr~t1i1., ila/1 ..; a.,1.0-av. o.A.r .. rum-~ J Pt0,1.-,x, a., ,·o~ OCH..~e,.. f. ..\.&..&L\\'\:bsti.al:. c...
1«1tuiq11ti. Food ,ndA1;rh:uhu1c Orgonl,a1lon or th~ United ;>;~lion,, 19111. The t.\'J)ennumtal •1>rllcntion ~rfnsecucid<> rrom a hoh<0plt1< (Qr
-~om•. the contt0l of n,wln~ popuJ;1tlor1., of Olosslna ur.·/J/11otde. In \\ es1

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70 ,re. ,s n~-r: Aspects fntlut~ndng th~O('tuncnc~ of ln(Cctiou~ <H:...cases

\fr1c~ \111,Tlweff~bn: 1\,·o~pra~ '1PJ'llit~ufom:niO~h·5-o G?os~mtd~c:-1. Btt!it•tl111JJ t:111umofq:irnl n.~Si,"l)U/, :0, rs;-J~h.
.ondo,umm, and 0~1S· 1!:188 (dc.'t-1mt1hnn: oa (;. tr, ·ltim,ul,•;;.;;,,nJ J:'> m,xro, ...., , J9).i 11r"·.\atum1 HU'lOT)"'>/ l>t~tH'flr1.-,;. \h'molr ofrhc
non•t.s.1\:t!'t o:Jt,UU:Hns m tJJ>pcrVolt:\.. Trofllra! Ptr. \r:~nn;,(c•m,·n:.r..
l oncJ,,:1 , rnool 9! 1 rop,rod ~kJ1cml' nod I l~Wl..'Ol~, ~t\ 10. f ondon:.
:ttJ .. no.
N.,- l.::\111.
,S RA\.,'-'"" l!fl7, The, "t:0'.W' O( ,metl ;).Od S1jtht 10 CifoUIIUlJIC')rluttrlfllU,
:t.;- c \R\Ht H~1 t 1.• 1!.-l,'i~ Tl~t \'ln:"tof rintlcffk"'t aud lb rtl'hll.On 10 GfUJ..ilua
llulhm,, <1/ J;.,uom(Jio,:.iral Jr~sttud1. ;:U, ~;;Qi
llt(Jt~tk"t.1 \\ ~ 51\\. Rt1Ui'w1 ofbutunoftJi:.icu/ lk~i'rrrth, 2~. 3-.r.-142
(1 Buu-.. "·, '\Hf\\ \HI -.\ I t.-MI \'11\.•,n,. ,- ,~.a, lhl' t~·ding '-Ut'l'C'-... t>I ~ rA1-1,HnL\O .. J. sq:J8 EUnd("Jp1.•..1 in .\frlcan 8am1.>, /tmtnul oJ0.1mpamlllY'
ht.'I'<' flit....... (ilru.,i,m /'11111/tliJH'J D1pu.-rJ• Gio~lnid,tc tJO t>Xt.•n 1rc..i1t."t.l
/'atho{q;J'nt/11 nr1•MJlri,t,N. 51. :'.!&1-lfi,8
\u1h fl)7t.'lhroid pour-on.., :11 Gulnni1 R,mcl'. kc.n~·a. JJull,•titt CJ}
F ,,ltJmtJlngft·al R~rrlt. fl~. ~117-1 i2 2{t C.\JltrTr, K.).1., 1fto6 T -..~i,.l' U\· In Ar:1bfa. Htlti,h .W1•1ffr.:1/Jo11n:11I. l
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; ui..ua, t, M., ;439, ! hun.i.n tr)'lJ.1no.~t1mJ11,1..; Jn 'Uuth~:-n Rh.wdi..... t~,
Jfll 1-19:ffl. (r,nuat,·rlrm <1/tht lltn'fll VH·J,•:,•of1flJPlt11l .\t,,,t,,111rhlul 30 ciu.lirta. \ ... 196:,. \mdiornhon dt..-la method(' d~ ,rctcnnin:mon dt• r:1gl'
ll_i,glcn~...12. 729--7~2. µh}~I0.1olflqUt th•, ;tlch.~ 8t11ll'T11I d<' Ill SIH!lt.11,• tit l 1mlz,>l1>gf;o
E.taJu,111• 38. 250-259.
g nnn. w_1,., ~1 ,<.:Kt.'\:nr. P.\_ N 1111~0'\:. tt.1).• 1978 Tilr r.:fati\"l·
;1u:-..1cr1,-eness of donic.:~ ... <'llllll·.. ,ltt."trl) .tnJ go:tt~ to Clcwma motsitatu· JI t:n.w tn \. ,1,1 I \'11111f('JI I, F.L. 198~.A IWW marpho!ONIC".i! chil~C:ll!'( of
111or1tum.s \\'1.-..:t\,uod ~nd t;. p11/lr'dtJh'S >i.u.rtcm Dipc~ra: (jlo,.. miJnc•I Ill supcrio• 1.!.s\[M,.'r, :o dufcrent.l«tc ClO$$mn JH11lu/lJ)1'J .\u,um tmm
m1dJ:,..'i'tld .uea l'll Rhod~ta. l/utt.•1m <tfF.mmmJ/r>gimt H1~1t•m, ,,
;:i •.a. GltJ.r$[1:d Jm1gipalp1.l \\·u1ema11t1 ,oiptcru.: Glo,-~1nLd.ic !ttx\.t ':'o(ll'RCJ"'
·197-500. mul its ittJ,lil·,mrm tl.; !9-:~1~
9 RU\T. \\'.I•. ,uc~, XI.Ii. l•.t. .. Mbn~. h.h. 1. U:.\\1~ H, 19""'!.. ThL· 3:: c.1uWJJ1 '\.. ~ Ln'l.J>!>fi 10.. c • 19~ Un nuu\11.'".all )'IC-~ pour In C.'lpn.m;dt"
1mponancr ol th~do1\kt-\• fl.t]JIJ1>t1.1mu~· is~ nsowcco! fot>d ,md ,1 ~o'i!olnc;.,. C,fofJJt;u r>1p1t•nt. "l.tu,c,<Jat]l: fk)("ftptto,; c, ,, ...,u, ,ut It
reservoir of~rypanosome~ rm Cilu.-,-J.im, n:t:.r.\iuws\\c~t~\ lli:lH1,•Jft1 u;mun Cuin(rstfc.• fOlTic"t di! ta R«h<'rch S<"t1•mt/k11,a• rt 1 ,~·hmque
.Srft1ntt Vuu~. 6. 1~20 ,,•r.c· f.mom,,tng:l' m1..Vfta1l/#rt /Jarn.nrolng,, 11. ?.>t-262•
Oulri··,\/,•r,

10 IJO\'T, w.9•• ,:>,c,a;s-1.11;;. 1\~. ~ no,,. t • l!f-:"O, ,\n 3ueznp1 co dcmon~trnh' 33 n1. ,r>, \\ :\ to,. 1,1(),, \cn,d "J)~)1ngo£ 1~,~hc ni~ H;fontmi ,pp.) in
lht nuturat mutStni\i-iun ar ho\ine lr)J)aJ:lQSom:asu. by B$tl'fll.!'1- 01h,:r tthod~,i..l ,,ith uhr~, !t>\\' \t,lum'-'s of cndo..,ulfnn. Tnm.sae1iow·11/tht>
1hon G1os:!i1ui in llw S:lbi Vn1t1,r or Rhode-•· RJio<mw,, I, 11·m1t1')' llirO<lfNr, ~,·,mfic•.f..~orr<u,on. 5':', 12..:?t
Journal. 1, 7"-20. 3,i c:ttoRU,. I J... J<.,l•.1 !11(' h1onnms"'or GIOS$lnll 1,:or1iuuurn thcUrnnunl
11 tiMD\. , •• :973- C1fl.-ndi1m thom_gt""' m ._,.,,nu-d t'..\t"jtabllfty- thl'"orti::,n nf ll~ hL'it S.11,llht•tn Jthnd~1a1 l92:!-19ll, Rull,•1u1 r.ifl:11t()1nu!u;;i,"fll
ht't,:n1ou~ rhuh.uh in l~llj"(' tllc-5-tlld 01h,•r .inuna!..-. /11ur,111lr,f R,-utm. ,:. ?0, 17~ 301
l-,uamQ}(©' \1. SO. 79-95. 35 <.'ltOKI !-, J ~ 19:::~. 1:..,p1..~rrwnr!o.in gra<'\ fir1.•-.. .,~11r•"11 t;f,,,.~i,w mor.sltan.s
12 &tt.\O\, J•• 1s:-s. Huny_rr in dn,'" l't<.'l'4.'
ll}~ I he- nutnl!oni,J '°rtcl:.h. ,. <1t t.n ~,.uJi~rn Rluxh·.. ia, llutl•·mt o/ r.:,,u,muft,gtutl H,•u•,m:J: .:o. 1;-;-1:,0.
bdu1\1our Journal of l,1..~YI ('lty:.10/ngy. :! \, 1$\17 -lU"f, !6 c:uoru..1, Ji-.• 193~ ·\ .,,hon h?storical :iccount c,f l"'tl,,L· ln ~owhiem
l.l UK.\ln. ,. (I>· CiRl1 anH:., , ,.10~3 UJ•wind ll1tuu ,r: ..poni.c\UI h('I~ llh.""i RhotlL'Si.;i. / 11u«t'lli11s;,:.n,uf Tr,uu'nrlf111u rif llt, Hhmt~·,ir, Sr:,•1:tijfc
lCilt>Ssi1:«. l"IJ);>.) <O.pu:tA: c:h.s,lnlduo> tO .att10n~. octonu! .1~1d J.su,r1mto,,. JS. .J 1·SI.
phcuut-. in nature: J \ tdt..-..> .;md)· 8t1f11•tin e>,'F.u1r,r1wfogiwf H<>,f-t1f'(:/t. o;,,
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J':' n')C1t.1UU r,.1 • 1q6J f 1rclhn1nury in,·t"'tlrdtlon~ ,u, thr u~ oi ;uh,,m1C'Ql
3:~'--333. bn1tth k.Jller In ~ontrullin$: WH,fo\,1h on tJH.' ,~:i.,trm bent!,•, c1e.mnµ -
14 D:R..\D\°, /, .. Sltt:il.L\1. 1.\ ... J"J881 Umdlng fl~PIJJ'l"i~ o!th~ l!i.t!f'~ fly (jfo~stlUI Chlpinl}l Dhui(:;'-. RIU>t/fJi.-l ·lgrfrulwml /tJumnl. SB. 1 ;1-r 7":
111ur.:11,,u. ,,1of'.ti1tuu Wt.."!i.l\\f,od ~nu tht" 'lltJbh.· il) ~ttnnOJ:J'l(llklttmr, 38 ,0t~Kn111. ,. 1 t~Q:, Furtht.•r tn\·i.."'>Ug:)1ton~ 1nm ~-r mh• t,: -1 eh~,rur.,J
L l)\r,11•:.c Cfh»: ..1nJd,,t" ~ Mu..;.cidact to bt: c:~·.tn<l- .•.·hth,· rmucm, \ hru.,h 1o:ilU.·1 1n cc;ntrcillinj! ~°'vth 011 the '-'•ht~m boafer dcarintr,
1lllXt:1Unf")· 111,!~ . Bnll••rllt "'J fi11r,.mu,J,1~"'it11I R,-,r.nrri1 .. :'ft, 301-31 ! Ch.lr,in~ D1-.:nct Rh<x.fom .:igricultuwl foumnl. ~- 370-3:'3
I~ IJht(,Hr\\lU Jl. On.\~""11llD1 lt,0.,).;\0M."\' C. G-OlDflt, , T.nll\lll.,.\..e. 39 ~ut.:>.uu,. r. r 11,:q. 1ht'r elh.'<"I oruhto·low•\·utun11.· ;a•ms1olpphc-Jtit>n!t ot
,10,r~u. o. 198.~"c An~\,· lrnl", for G/b~·s(,w. 1,u!1Id/fJt1•.:. 'Frup,tul Jlt•.,t L"f\tfo:,i,u!fan up1ilh,:l n,::,.imn GIV$$.i'1cl t[)i111l't1l' Cilc.>~lnidac on
\.!tuu,1;v,m.·m 33. lSl-1 S9. popuJ;,rlr>ni- t>r nnn·\otrgtt cirganh,rns m snvanno. wuc,dlund ,n
1r, uuOt.t, o 189j, /'1t'llln11111rt• 1ttmorr 0111111.• B~uc.• F1J r>iJc..v,.~01 .\'uwmn. Zimhabwr,Rhude-,.1d BullN,n of f.,rr()Jtrt,l,)JJirof H,.,,""' 11. 6~. 6-15--6.15.
in ZuluJJJ,:d_ Durban: Rt•mwrt .111d D;i\ ii.. .;o COCJ,;BJll ..., •• , n"-1',mRr or Ii r-n~. rt.r., rn63. 11w co:uml ofl...et~·
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dt.·~·c.•Japmcnt of Tnpwm.\-.-111111 i;,m1l1:t·1ut- ln C,lo..v111a pa/1:'<tlls. Rhodt'.'&;, b>· rr:~J.I\..'of !n'I.L~1u;1de th'i,4. h11rged rrom ;s1rcn1n follo·,\\..-db)
Prtx,·«ll11g; of1h< II~'()/ '°'"'Ty ·81, 81,40;-o;t.;. :-etth'J11,·m ,,fmd gt'flf>U, µeqp!e.11111!,•rm a/Fnum1,,!,,g1mf Rt,x·w.:n. S4.
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Tr,mwc1:rmso/ 1/tt- Hoyal ~irry·, Lo,,da11 <B. ?41. 179-210. u (.0: VI'\. 1, • GlH-....;>".<-.. 1~~: ~fO~t-~ccitng beh.:a,1QUr ~J m.i.nog_cmcnt of
lS-elSf': ,4,umal ,r,.,,,t,•111 01 FmnmofogJ·, :r:·, 21 40.
1q ,mu.. , 11, *"-· 195,y. 1 ht' '.v.11tr h,,l.anc-c nr t:t-e1* l1f<·' tromr1atcm$ ,,j thr
Rr>)rtl F.nmuu,!ogit.·lll ~tkit'ty, l multm. l l l 205 ...23.5., it f.UIS.·\."n.:. o .. t!)Hq '" ,,,~etlJ!.~,,i!!, lJc"•l:tt. hllldi:.'~ \': S;:nt11l'W!o dt."
L'ln,tfrnlC" d ,·lt'\-ag,· et dl· ~h:dedni;: \'c1crlnafh."' Jl"'" PJ~' TrnpJC4U.'\
10 U\1Mu..t.L 1 •• 19()0 ":"hc~JT~ct uf humhfil) ,tr.tl 1cmp~1atur1..~ on th4.' t..ixtcnt
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0 1 abdr.imfna.1 p!grncnmdon In Cifos..,,11,11x1/lfdl1,vs.\u\;en. Bui/rm, oJ
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it au»-.rc.u.. '- 1960. The t•ffuct of tcmpl'ra.lUh! on the comumptlun of fat ,. ,1, RII u 1... t~8o L.1 hm,• ,·unm: Ci/o,L>·i1w 1Jt1lp<1(iJ J:llmbi,•rll/J
Uunng pupal dl'\~IO}H1u.-nt fn Clos.<hu,. Bullr:fn r,JF11:n11UJfo1!i(lll \ an-rlerphuik p,11 Un·hi:r.. dL" 1m\lt.-:,, lrr.uli(·,.. en Hautc•\i.Jlt.!·Etude-dt
H,wmch, 51.~.911.
p-..tn:am ....~r1; .. 011<:r.u1anncl:,.. Proen-dlns;:• oj rltr :>~ mpo.slll"1 an t10i01Jt" nrul
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Z? uuM~O.l... , .. >96ti . .-\.:,,ptCb of the llJglu mct..1bo1l!,n1 of L-..cUt• rn~~
lmc:rnauon11I \wm1t l~ttrgr Aj:.ency
c;fo.(;fimd. ComµartiJCJ·,, BltN.ll,•ml311'·"ntl J>hyslolo~·. 19. 809-818
4,: c:urtra, ,.t-.. l~fi1'. 'ltmt.• Obs.tt'\'iltion"on n:pmduc.:111,n ond th~effect,ol
,;s 111111'!1,. r.•• 19-,. S1111hc•is of prollnc h,• (~, bo<!y o: 1hc t<t:.se Ill ra\lU1tiun on Cilf,niua nusreut. TrutJ!lltrtrmu uftit,• Hui-.1/ \o.-wl) a/
c;t,us111aO"r.sfw11J.>: Mesabohr p,nh""Y!!-, lluaJ Bt«hcnllirr'}', 7. ·127-134. "Ttapf((f' \fttltdn,· tttuf ll)'11,U·•1ct, 62, 121
.::.,. nUR.'\tU.. .. 198; ThL· ttfoct ofwmd·bom,· odour~on t.hc din;~l1on of ,i5 r.1:1tn, c r., mu n,(, tQ'j'I Tln.'ot1..1ic..-nl ,1ud1n un dw 1-:,,. of
01ght m t(;t-ts.e lliet,., Glu.Jsllw 'J>J.l. Ph_\'>l1Jlo;p<ul E.1w.1mtJl()"/zy. 12 I ,9--1 i,b. mansloc.i1·0111 for the rnrurol of l~l~ Uu:') ;11,U mhn rll',,\'Uw ,·~um,
;1s tsun~n,.. r ,,. ,, 1 OR. JI'.. 19Uo. An eru.•t\;\ budget for G!,,....•um l>1ptl'rn: T11ror-,·:,'rnl P,ipuf11t1n,1 Bf<,{"1J._\', .! , il-90.

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Uhrtmar portuguth. ~ahre 11 murrolngirr da, arm.1dura gcmi1al J3 (; 6; •ut tHt, <•• l\~.:i. T(,iHl· ln thl• Trnn""\';'MI a!\d ~urrouur1in~ 1cni:orlt•... ,\n
pl.(1Jll1ilJ1t',l ,\us ten. l903 rt>lptcw), nolflm >l>t'lit,!n,t.-d;r {;JlUtltJ~. hbtoric-~ cc\ .r:\\ Fnum:olog) i\ft•moJr I, Uup~1tment of \l,.'TIN1tutt
.•.trif11111lnqm•, .?9, ~o. 125. ~-ction U. ~o. l. lJn!o, nr ,,.,u,h \Irita
\8 Q war..: 1 t ., 1980, Th, llis.:01)• ~,f1:(Pfi,i• f.'/y Ccmrol in 8ot~1ouur Ronc.•,wt.l M HIit Eh. r .-. ,1,.1'i.sQ~1 \M;... t~,:9, Ent(unnlogtcoll nnh~ un G/o;<Smn
Report to the- L>fr«:or o!\'ctttfnilt)' '4•nirl....., 8c-tS\\'W1.t
11alli1J11t,·~ U11Jt111 o(~Jttl: A[rlrn. Dt•ptuunem of \clrfrrl/tt111.• X'it•N.:t*
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74 >Lenn~ u,'" t\spec1s innucncing 1h~ occurrence ofinfoctious <Jisea,e•

17.J. xcnu_ cM. 1998. :\ nl"\,•!ook at ,~ct~e flid rind r~.ma.1n saulh •.\frlro. tfarmcth Rh.od.c,ia: tmd their rctc,,1:1nc.:r io .somr bt:lse c<1ntrol
P~per presenwd ~r the Sou1h ,\frlcan Vc-1crinttr\' Assoct:uJon Congress~ mea,.um,., SmirhNJfu. I. l-72..
jun• 1998. 1.9.: num1nsu.s. ,..c;.. ">-1.JJGE. E,a.• 196$. The utte of in~t1c1d!; :n ,U-re$ting31)
>73 001~,,o. \Ul. 19118. G/Q<.<fna 1mlll<li1><•s sin.I$: I!$ po,emiul foruS<: m :?.d1•3D(11 of Cfosslm1 ttwrsitttns Wt.-St\\'OOd In ,n~south·~nst h)\\'\"t'ld t>f
biolc>trical con1rol of1~et,,-;,, /ns.'t:t S<iwcemul fl,<tlpp/l(utlm1, 9, 399-103, Rhode~. Prot~~di111;1 and Trunscu.:1!0,,s oflite Rluxlt~,io St'it•ntifi~
1;,1 o,,."a,. ,u ,.. 1960. E1lic:icyoft.,uW.1rinc ,~.. .1 putcm at1ra,·ta.n1 ror '-Sttse WQC'/n/lOII, ;3, 1,-33.
Oi<>. IJu//,uu ofl\[ri(all ,,,..,., s..-i,,.,•.,, 10, ~-to. 195 noumtrw, •.\.G,; nu<..t J".1\,.XNl'f?'.l~GF.H, O.A... l>E.-SGU,.\, \.f...1~,;?. ·1 he
1(1. PAJl.1', P.O.. (jUftHlU, ,-~ .• At.tS<->1'1'. :\, t.- UL c..w .• !9';"2 .\ ia,gto•§Cllfc us" ufr~!dtwl Uh~cdcfc:k~ fn 1l1L• tt"Clta.motlon ot th~ RhodtiJ:t...
~hen,~ for the' c,:;idk:nion ar GIMJfnn morrit,wr ,>wnirmit\\'t!S-t\•,. in \fo,umblqu,• l'lorder rcoglilll h<:J\\'~cn 1hc SabJ.'Saw :,nd umpopo r1,.,.
1he Wes1<>m l'ro\'b\C<' of7.,mbill by ueri.tl L11re,lnw,\'olum,: appll(oiton from G!Ntuui m11r#m1u\\'1."'"'~14 Proc:,-wtJ,ijfJmtd Trtm!4c'li<m., of1hi:
of endo,ulfan. ll/ll/e1l11 of 1!11101110/fJfil,·nl P.11<'f1rtl1, 61.l':3-384, Rl:ml,;la Sclwtll/c ,l<S<>Clmim,. <;;, ;1.J..f.2.
176 P.»-,.;nn. Q. i. oltAnv, J.. 19\Yl. Flight bt'h.lviourof m:asdllc, 111 ,h,cl bush 196 uot,,f~')o~. r., lt()r,rn_.,, 11, 1,., wtLLlA'h, o, ,~u,.
Unlu:ri.ue ~nJJysh, of tJtiSe
{G/a,s!11n pa!/ldlpes lDlp1cra: Glossinld:ieiJ. 811/li!tln oftniomo!ogic:1// hllbho.t In 11'ccon1111on flybcl1 orsou1hemAtrkn u,ini;clfmatc and
RrsMJ"c:h. 81 Sl3-St6. remut.e:y se-ns.cd ,·egctation d~1a• .\kdic(II mul \:imrrinan' Ettu,molng;·
li7 Ptn'B TJ.1()~1.\$' ~:O.OCl,Hr.S (;Q'l;~;lJl.'rl :,C(. ~Ot\'ICU ZIMB\8Wfl h ~o Ml'.°'.IS1 tt,e 11 223-?3,1
.\GR1c.uuum U!'lirnm »:JNCJ')()M.1983,.A r~n~ ,sciw and
1.97 HOHT~:SO\, 1. St.OOCR>. P. ¥1 \\lJ,.LJ,\M'.'I, n... 199';". \lappJng Utt.Se bab1tat
1ryp.ino~omi~i~ e<>ntrol ~tud)t: Ma.lawi, ;\.Jt)z;imbique. l..arn1>1a A:\d
,ut1:~1ii:} In 1he common Oy beh QfSouiht<rn \(mu u,ing mulli,;111n1c
Zimbabwe. l'etcrThon1ns /\ssoc1u;c. Con~,ltlng ~•mce. Cl'l)'l LuL
:..nah'Slso! cllm~ueand rommel)' sen:std VtJtet:nio:, d:uo..'.!erlta,J tll'ld
10 B.:lkt!r A\'~.l1Ut!. Hatare. Z.1mbabWt',
V,•11•,.inary Fnum1nft>g.•. I t. 2:-tr,-z.t5.
1;8 PHf.1.P:,, 11,1, & lJUKHO'IVS, r.M .. 1969. l'upnrial dwa1ion ln (;/()$/inti
mfl"irtnu orienmUsundercondhton;. ofc-Qn:nanl tt!'mprri:1tur1.·.
19H no~, M. n.. l9;9. T~'™' poputauon dymmtlcs an41 dlssrfbuuo1\! A new
E,uom.alug;u EJ.1,ed11u,11u.,l1$ e1 A1111llcam, l2.. 33-;3, analrtlc•I appro~ch. Joi,trtnl fJ[tlnlmal ~IOKI'· 18. 8"~9.

179 PHru>~. 1<.i, &· uunROw,. r.:-._ 19G9. l'redlctlon of Ille pupal dum11on or 199 Kextr>K. <>.1. t9<11. S.ncllllc imagel}·, 1<01Se ~nd uyp0110,amlMls m Mric•
CIO$<lnn morr!:am orla111nlis Vt1n~\'l'pl11nk under Hrld condition•. Prr.:..-.!ntu;1.•\ ~wrinar, &,romolOf!J·, l l,-4!01-220.
Joumol ofApp!letl f'cnlogy, ff. 32.l-33,. st<>O n.ool..a~. 0.1, ~ R.\~CIOLPH s.t- 1978,. ~lerabolic s.un.tl?R,ieJ 01 rnnft a:nd
t8o PHELPS.. R.J. "'JtUB.lW\, S. 1.M ~. 1.969. Ltth:U temper.nure~ for pup~tf:t 01
1 r,male """" .UipJOra, C1osslnid1,cJ 1111h, fltld. 811/lttluof
G/(ISJ!tu; m.orWnust>rie11111Ji.f.. Eml)mo.Jogir. ii..'<JXrlr>it•malU el J\µplltY,ta. /:m(Jmo:osiCYII R,•,,arr/1, 68, 6'39--65•1.
12. 23-32. 201 fi("1ll<.~. o.,..- ft,.\~OOU'l1- s.1• • tYS.S. P() j"l13tfC):i tC'Ol0$,-•y o: C$f'iSt. ,\mmn.l
18) PH£l.Pt< R,J. • tr)VEMOlln. 0.1 .. 19~0. P,0, llO< CHI 39. Chi.<ipllc. l!orare. Rc:![ett• ofE11to1llhlogy, 30. l9i'-216.
z,mbabw~. Unpubli<hl~I cibscl\·01!11ns.
202 .tlOG~. n.1. & RA'-l>Ol~ -..f. •• 1886. Dls.-tributfon tmd ~bu"ldant"e of l.'>'Ct~
18i PU~. R.J. ~ ,1A.um 1:-., 1.• 1987. (;Jossma moMums\\'estwood at Ill'"' f(;/M.</11a"'pp... Jo11r1111/ of. \nlmn/ Ecotom·, 55, \007-lOS
1lck11mi111c lle;1eat<h ~••non, l.lmbsbwe. Tran,amons of1h~Y.lml,ah«"'
Scimrljk :ls.<a<'fo1lo11. u:1. 40-46. 2-03 1~00F"-""·n.J. • t1A~ncl1.Pn. ~.1;.. i9gi. '.\1ortalfty rnu.•s..and popufotmn
(hmslly of tsbtst! mes corrdatcd wuh ~ttUlte 1.aui,al.!l'\ L.•11.:u re> Vm,1rr-.
183 Pll£Ll'>, R.J." \'Alf, G \ .. 1978, Stud11..-. 0:1 p.:pulauon< of GIOM/111/
.Vawr,:, 35J 12ith June. 199 l.)
mQWU1r1.< m1mi1tms and G. 1,a/lldlpN (Olpteia: C!os.lnlduu) hi
llhodusla./01rrr111l of,lµplkrl F.<Olog;y, I~. ;~l-,60. ::1.14 ttom Jh n.f. ~ AA~tiou•u. SL, 199s. Distn'butfon of ~cc;.e ond tid,.., in

18: PlL\O~, ,u, .. uon-, ,,.1.. & ,um:;..-,t,1h 1•.tr..• 1g-78. The r.:lati\'e
11
Arrk.i: pa~,. pre"1nt anrl rumrc. P1,mSl1ol<>1.G' l'Olfny. 9, l6S-27l.
unn.ct{n:n~~ or c:mlc, !.het.>puntl go.sh 10 GlosMnn mo,s.i:ans mors/kmj 20$ Mv,01 . , U,<... 1960. 111l' O\"Ul;tdQtt C)'cll! tn GJ&•siJJtl mfJf'lflaJU
\~~,wood Md G. JXlllidipf; Au>1on IDip1era, Glossin:due, in 1hc Wc,twood 11)iptera; Museldnu1 and~ possible method m •&•
lambozl ,'<lll<!)·of Rhod,~fa. 1lu//Nl11 ofEmomol~irn/ Remm:/1, ll8. dctermlna1lon fo1 female 1.ci.c flt« by th• c;,..sminn1lonor their ovarlo'>.
489-lSS. fl'(J11,atttom <,f1/1~ Royal l!11romologlc,1/ So<·fvty. l;,ndon. 112. 221-233.
185 PIV>OX. n.o." 1.tC...<;.A1 r, 11,M.. 1962. A d1um:U .and stasoned study of the ior.,,,u"OruL<. .,.,,., 196.;. F.n~, of she an,l ,ampiing metl1od on the ,1,... ,,nd
~ting h!'h:ivfou1 nf c;1ouimt w1J11diJ11.'.fAu~t. Bul/t,ritt of F.monmlogictd CQmp053t-lon Of (\\lch\.°S afbe1j,(> fll\!.~ ((i/c,.CSfofl} Jlld r2bcU11di!I!'
R"'e:11'<//. 53.5Sl-562. !Oipternl, Btcllem, r,f f:n<Qmologi<al lleseor<II. ~,. 4Bl-4~7.
,..,1.• t90i', Bt:h:l\ mur~wdit..-s o!'Gl~mn
t86 1•1u;.o~.-. n.o,"' U1GC.i!\1·1.,
2o; S<:11:!\'TITlC t~\'tROX:O.it."\'"fM. \.10 ../l'tCltlNC CROUP. l9lt"' /mp(lrt ()} 1t4trl,,1
mor,.iums\\"cs~\\'., 1n the. fic:1d. Bullt!tiu(J/ElffOmot<r,611:111 Ratntch. 57,
spttl)-1hzgmK1 Ottot1r·IJt1it,,tf m~,, on 1r«1.t1..xl .:coS)11tcm1. tra·drut flu-
227-2S7.
BlogL'il~aph,c Univcrsuut d~>Saarlandcs. SanrbrOck<-n. FOl!eml
18'; PIIIE>., .,.. 1960, Umn nol'a ~pc!r:fcdo Gl<>,!intt do giupo r,wr,lrmu Republic or Gcrm.ini·-
Dipl~r., Mus<ldoe) (;/@i1111 bt'Jrge.,I sp, n•ll . /l()(eti11 So,.·/rdncl~d<'
&tut!o.,, ,\/Q::amb1q1ie29, ll:o. 125. ~ttdon D. :);o. 4 2A8 ~H-f..Jtr\iJ, w, 1990. Tst:1t!I! and Tryp.,no1tomin~i, t•mtcot Bronch.
DepartrocmtofV01urinary S<'rvicc$. l•.O. Box 8233, ~us•w~y. Harore,
188 vo!l.·1. ,.c•• 1&30. F.smllyClo!rro-sinlcfat!.. /m CR'JSS:t,,l::Y. fl,\\,, tcd.). Ct'ltll/(Jlftu:t Zimbabwe". Pcr;onal commtHU<.1ltion.
of1ht D/p1,taof1/zo,\jro1rop/r:al RegfM. l.nr.don: i>ml<h \fu>CUru o!
'\atural Hb1oi,. 2<>9 s,,ivso, 11.11 .. 19;3. 1no effec, of St•rilized malo. on o n•iuml t:'C:'l>t llf'
poptJl:ition. Bfamc,rrfa. U. 159-113.
189 1•r..rn-:. w.11 .. 1958. StctfH:a.tlc>n of l~t.~ mes (Glos.{ina) by 1i3mma
nuli:ulon.Amtal.s o/TroplcaJ,'Htdlrlnc wu' Ptlfasilo!~·. "52, .JS.S-J99. 2m sr~tH. J, .. 1,;UVJ•Elt, w.. 1969 b.xp\!!imems on olfnctory nnrncumb for
ue~ mes-. Cfo.<silm :;.pp. tDi1>11.!m:.Gloss.inidael in l,·ory CO"olSt. Twentieth
•90 ron~. w.t1 .• sgro. Sy.sccmadcs,.nd jdtt1liJIC'3uo:1 ofCtcm(na. In~
,1u1.tiG.\S, 11.w.. (ed...). 11:e A/rfc.,a11 TrypanlJ50mfasc.·s. tom.Ion: Ci1.'0T,RC!
~l""tln$ nJ' rn, ln!l!matioaal St'kimifit Council for r,ypano.som!ns:<
, .o\lJen a.ad u,m.1n Ltd Rt"'-•""-"""~"'' Co~11,nl \rnmh:.wi, KM}'n, 10-IJ April. 1989.

~9t PRO(,R,\,1,1t: \f.\l:O.!t<T ,\J."Rlf..\.~ TR\-VI\..... OW.M~qi, ir, T$tL~ cn1dfcatftm m ST£\'t..~.sos-,,t\., atro,. 1.. \9U:.A11fmt,l Uffl in..Afrita. l.Dn~on; \\llfiilm
on 7.,a,u.lOOr completed. R,:p()rt 011 ,1r., thirrl mr.ettngo/1/re progrnmmt fl1!l:,u:nrmn~
t:t1111miw.~. WHO 1/eaf/11111,rt.:rs. Cknu"'i. 2t2 STULK• "o!Kl MC:. tt,R.. 1969. Effective tcmpc:-au.11c ~~.in cc;t)logie.il factor In
192 JI,£;-.!',;; ~~• .&,D. tOM:.t.n-.N. \\'.11.R &- \\'ED.l,I,, CJ,. 1-g;.5-- T1:t- R~n, 11tc.~J of ~oulhi!m .-\frira ZOologfra Afrlt:1u1/J, ~. J.lf>-19;
Glo;,ln;1 pullldlJ)l"'<JI 11/1;/11. Report 01 the Eai.1 Aftlclm Trypano,omla,i. 2.t3 ~v.'\'x.,Tffl'O-.: c.r.,-, 1,st2.1. An i.:o'.'fflminonion or the htbe problNn tn notth
Rc:,e..r<h Organll'.a1io11. 195& ).to~1.1rbe. Porrugueiw l:n,-t Mrka.. Bu/11:Jin of£momotog!Ml Rt~1rd1
193 kOBE!ll'SO~, \,G .. 19~3. Th• fi:edfng hnblts of ~U:, :liKU\ llmbobw, 11, .l l ;;..395

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iu S-WY!\:\I tnt,,. c 1 ,,•• l!llS Some trU jW for tScls~mes.. 8ulf1.'"fin t,f a36 ,·,1.c. f, .., .• 1982 The :oll· 1:1fl.h~ omtcnntt in the ,waltablllt}· 01 N:L4\' ll1LI\
F.mo,110/1>1,;tcal ~'Jt:ttn:lt. ~4. sg...10J. I Ojp-:cr:i: Gto""'lnidJI.' tQ J>Qpul.1.1ion ,..:unpting u,chn.lquc.':'l rm,~.,t11ons
:as :i.w,, s0tto:,. c•-,,.. ,936. Thr C:>et>t'"U1t1·" of f.-ih~ . \fnc:o1. Tra,u,.:au111s1Jf ofthl• 7.imbabtn! ~rh·m1fil'.,l$S(l(:/..1lfa,J, 61 33-40.
th,• RQynl Entnmolog1Cal Sutit•ry. 1.ondon. 84, l J;g 237 ''AiJ.r c., , ,98:?. lhi: tr.np nn~rua,cd btha\iou, of t~~tst' m~
;!\6 1., 11.;1i,;f , . \\ • 19tJB. fa:nlo&,u.:al factorti, lintitmg dw d1•..i ribuuQn of G/Q,:mm
t<iio,..1rud.11..~l and 01her Oip,cra. Butfl:tiu nfEnumwfo,tlcaJ R,•i~'flt'ri>. ;2,
71-93.
11u1rf1u1111 morsirm1.~ ,md o( (;f1,s.tim• pt11//tlr1M •Ot1>1~r.a: Gtt, ..."nuJuc) m
Jnh,unh,mc l'rovim·c of Mozambique.. lnst't'I .'\f.~nM nnrl us-~pplla1.1im1, ~8 VM 1-. ~..1.. 1~. I h,• m,pm\t:tnlJnt o( 1ras:11• for bi:W- fli~ 1Dlpwrw
s.i3i-:M? Glo»in1d;u::,. 811/f,1 rt,1 t1/ 1:ttrmnnlog,rr,I R(l'M'<1f'Ch. ;~. !JS-lOb

21; 1tt \IJ. A.."''· JL, 1991 kccordi.. of the tsc1t;c tly (;/qs.,mm pnlluh'p.*f. m the !tJCl \'AU., r.. ,., 198.;, Tht" h~pon~ of (ila.ss/,u1 {Gtu~ltiuld.tt'l and oth\•t
nonh c.1...t.:.·m rt-gioa o!itmbah,..-e 'fr,mJ1rctror,,;Jjrl11· ~.iml.Jt1bui1 Dlp1orn 10 <>dour plum~ in th~ ftelrl, 811/Jt1/11 uJ £11wmv/11glc,1I R,·i<'<1rc//,
~ll·mfhr .Wonm,,m. 6~. 20-2J. N.113-152
~If\ 1 tll\~P"'I, Jt ., l::,i\: S'f:\V f«-Otd'lrOr lSC1"C fli~ in ~ol-'Jh..~rn /.irnb.ab\\\" uo \'AU. c...A. lY9J.. \'hu:tl Rl!~-;>onse~ or bi't,;:e OIL°" Dip1era: Glus.ssntdr.t co
Tw,w<tt11n.r of rhr lunbalilf'( :sn~m'Jk.-ls:son.atta11. bl>. 30-J.l odout·bAitcd la.Jtc~. RuH,•tin o[Emomolog/(<rl R,rJtmtll R1.
2i7-i89.
~1i\ t uo,ho~ ,1.f:... ,,s-:-.
llw ctft:et on t,G;.CJ\1! Ui'I..½ Glouim, "Pl, , o:
d~ltame1hnn npplkd 1ocattle ell.he, ;i, ., spr.i) 9r mcorptirall'd imu u1 't'At F. o,A. 1993- RC"PQ:1't~()I ,seo.c flu.~ (0fptcra (ilQS"S1ntdJi:l 10
eM·hlg.&,. 1'm/nt.'lll Pl."$r .\fmut,re1t11!11t. 33, 3.2:9-3~i \'C{:etauon in /.1mb:tbw,:.. i mptiC11.rioll3' f<>r popuhuion dtsr:ihLtion ::rid
ban >rtinJ. 81,11,'lln of F.mom(J/l)Jti<al n,-rcll. 88. Suppkmen1 I ,
.;UOTI-tO~~so,, 1,\\ .. MltLUIJ.L, ,1.. RU,,, tt.n.. ,t111u"· \\., :-(.J.lrn \flf Lb, Ut ,_ S7-SS9
"1L,<J~. • 19q, $111d1,~ nn •h•
cft\cucy (If delramtrhn11 lppllcd 10 fnnle
~,f.1. \'JU,. c., . 1q~ JQJm mu,sr,Ji11s1·n1ridesfor us~on cmrle
Hnn H<'pon-
fo: 1h1: c;untrol of 1.ieem fllt-s iGfq;{:S;,m ,pp,1 In 31Juthcm1 \lrk.i I ro11fr,1/
RL,g1nnnl T,ctS<' nnd J rypanQsomloMs Cunrrol Vrogramme. ~'.ofaw1,
:\nlnwl Jli'nlth uttd Producrfrm, 23. 2:?l-2.:?6.
\101,:.mbiqm: l~\mbht und limbabwe. li~m,~. %1ntb{l1;,w-:
221 1n1t11, 'i.f., 1Q88.. St!h.a.\·four c>( ht!tM• OW!>, (C!w.1Jw,1tn ho~t odour pJum~
~9 \+A.IL (j.,.\ 1,, t:UMMI""· It H.\.1 , &g:;6 Th,1 ulTcct~ Of '>elcc;U\'t't'hrrun;stfon 01
In the 1Md /1/J,1'Slo/l)/:l•'tll F.11/omvl"l{I·, 13, ;5;-1&;,
ho-.~ on., popul,.nlon of 1..et1J,t• 1Ut, ,c;t~'inu montttwt mors:tmu
~ tOJ'R. ~.I • 11}8~ Tht" hOSl•OritflUsll!d ~ha\'iOut f){tSl.!L"'-t' fli~ !li{UUU!tf \\~S\\\Olld ·V1Jm·~ Gfo...,mirll\J..' Bulleur: ofEmomrdt>gl(t1/ H"1rau1,.
and th~ tnt~rac:ion o(\'lsual;ind Q)f3(·tc>~ "'thnuti Pl:., ,,nlog,tal 611. 713-7;?!I.
f!.t1f(Jttlf1fO,'I)', l~. ;1~:-..310,
2.W \'AU t..••\ ,,.. tt.U.L. D tt. t48> The rolt, 011-octen·l•oJ. lltt!tone .int! c:ubon
~-.,. ro111n. ,., .• 11-1~0. Do~ rcs-pc.m~ oflSCbC: lH~ <?.0$1i,w to c:ubon dio,1d,• 1n 1hr ,mmc!itni of 1~,,i.s~ m..·~. t;tm1mn ~pp. Dlptura
dH;.\1dl.!• ..seer one ooJ oe:rt..'nol Jn the titfd Phtslofo;,t:ral J.in1omolo1;P•, I$ ('Josslni<l.at.!1tot>>. qdnur flullNfn oflimflmalf>g1fttl Rt---,r11rrl1. 75,
93-108. 209-217
:z::N toRn, ~., •. l{fe:.L .Rhl)<H1st~ of tsetse flit.~ trnpwr.11. Glo.sr.inld;u: ro A,J,t.. 1988~ Thti! olractor)' r~pon'IL,.,,_ of
~5 VAU. c9A., tl\!.l. D.R. &,<,.('1111,11.
w;ir1hu~ U1fztu<"'"°"''n1.( flt11hinplc11J l'.llla"''· Rul~·tiu of Fnt()mtJtlJRJ.:nl Olpwrn. (flo,,inld,u.,) to phcnoi.. a.n,I urin~ in
1.st:tM~ flft!' G!Cl.(Wlh >PP·
Rt~WJtd1, 84 411-119 the fidd. 811U,·1111 f)f huQmolog_ft"al ResL•bt1.:h. 78, .?93-300
us T01'.1",, 1.. 1tA1 t n.1t.,. S'1mt J.t., 1~\95, Rc~pon~or isctse;, !lie:,. Dip1er:c z...,GV.\U... <,..,,.. Jl.\~(,rtO\ l , J,\\ ,·oc MUU, (;,f, ~ -Pm:~ k,J .. 1986. Fu.:ld rnat~
GIOS>tn~; to n.i,umJ ;and sy:rnhc1ir nx odou,-... llull,•1m of ot b.in; to conlrot populum,n.... uf (ift1ssma morsittms mar.mm:-..,
l.nronmtoitral RA.,/,":trch, 88, I57-166. \Ve:\t\\"01><.l n.nc:I C.: µntlldith'1 At,ne11 iOipier:i; <1!U"-"trttdm,'1. Bult,1un of
F~11nmolog""'' 11..,,orrir. 76 I ~193
:Z.;!.6 TOM,~, HA.I l II.ft., Jlll!1 P), $t,f. & \! \U. (..,\,, 1997. \1tthod4 'or
d~p~>ng 1>dour aurncu1.m.; ~or tSC'l:SI! llie, Dipt.eru.: Gloc;..;!md:u-1. & t'OC l.."Jllll, c..r•• tq83. Odo:)r~b,iltl-d
u1 \'AU. f•~\ t.O\"E.\..lfllH r.> 1 .. H l:\"l, ,.
Bull,no ofFmomn/t;g1wl ReJ"'1tl'11.8,.m-311 t.u)t~·t;.. to cumrol L1,~h<' thct. G!0Js1110 ~PP. l)ip:c:r..1: G?o:.sinida~ In
ZJ.mbab\,t-. &,/l,•1/n of F.mQlno!ogio1I Resetm:lt. -:-a, :lt~~
2:li t01'N,, , •• tt()1 row.,,·, M.T P, 11 v,u.r. c..,.. 1992.. trr.p:0t.td per,"u:nct- ol
ln~kidc dupo~,b on ,ar};ct~ for conuolling G!t>~#nn µc11/fr11p,!;. ~,a, \U, c;•., .. ••1111,·~ M,f. 19:8. ,,llnpUnij prob!ett\\ wuh r,che •111t..
1n1;:uern: Glo~1mdu.c Bul/,•1/110/ Emomolt1gical Eli"•cw·,h. 8'.!'. 5i?S-63'J~ 1Dip1c-r.s: Gloouud>U..' . /t.mrnnl n; AJtplird l;tulogy. J&, ;1;;....7:?h.
~.Jg \'Al 1-.. c;..., •• \\1u:ox. 1. &. \ll!>O\:, J,. 199 ,. Pr0:tpccts (or usinR ndcaur~b.lltcd
22l 11>1Ut ,q '1 \S(;•,,1po. T.:'\,(:.. g. l &,\1 '· U,R., 199(t Rcspon'\C.. Qi (i!t)~lna
tr~ m ennui'>! hitt~t 111"'" B11l{c•ll1t ofHnfomologlt:ttl fk~"f.zrch, 8,l.
J)Olli//,p,·.• (1)1p1Cr•: (;Jo,,mldacJ 10 synrhc1ic nrp~ll~ms in rho 1101d
123-130,
Bu/16111 b/Enrom,,lv[/iu,J Rl",·t1rrh.8~. 609-61\i.
:,.so v.,~· M'-UU J,. 198'1 tn,·uHd t."lorim;u('tt of t me of po1,ut.\1mn incre.1sc iron1
Z,?910~M ...),, ,,.,,.l\t II , \..G. '° UH~U·flfol0\\l\,l,.t,.,. l98S:. t"hct~J)Qn,L.... uf ,;,,,,..,mm Oip11:z.a: (.lo,.;m1dnl' ~1~c clhrrlb,..tfom,. Bt1lltu11 of
f;lt1$Slt1t1 t>t1ll1d11,rs Au.c;1tn ;D1p1t-ra: Gto;;~m,dne - to (.)dour-b;med u.1ps
J:momolog:rol 11~1,orr/1, 78. 1S!H61
and t.srg<:t~ in :,om:\.lla 8ull,::Jn a/Enumrolog,u1l Rt'ti:orrlt, ;s 99 ..1ois.
~' ,, ,u. n 1~n 1...t1...- niJlin~ bt-hu\inur. lill«i-.uf ~ llnd hunger in
23c, nm~ r:11, n ~ .. t!)&i. :ht colon11.n1ion by 1h,· t.--;etY., CifrJSJina 11alltthJJt!S
Cilr.,#IJtn morr:rnttJ. lltQrsum,., and <i. (1Qllld11>t"S, PhyJtofo-_gtt11l
.\u\tt"n. of :a un1qu~ hDbaar-exqticcomferou.s pfa.niadon - with
Emomo/010· 13, ~';"S-;86..
~c:c:1nJ rdcrencc ;o 1he ~nbwc Vl1llt}'. fl:tn~'U luir,;c .,t(<:ttt<' n11d Jr.,
.¾•plt(!ll(l)lt, l . 24:J-?,18. :r~ '.\ \flSU '.'IJ t.. 1\l~:. .\c.:rha:lon of lht'\•r- ~~it.")O(l~elM: f (:/pJ.1tnn~J)J) an
rc:.pun(.c tn tio..1 derived ·•,-hnu.:h. ,\f1,t1kal and \'trtrinary J;1,rc,,m,!ug;\• h,
231 NR.._u1. D " ' "' ,~n,1. r.1 ... 1973, LuOOr;uo1yan~·,t"Sofn1,mn 1111\.t!tW 3,;9-:,s:.1,
mes D1p1. Cilo~..,md:1~). Htillrnn r,J E'mom,}lvg,trd R,.?ttll\'h, 6.2 _1 ;~-:\Si'.
~J \\'i\11,-:E..,.. ~,.L•• \',\:\' "'...: ll():o.M' .H* • ,,,. ( 1tlllt1lA, , .• ~H,Ut1,(,t. 0, nu111,llO;\ ,
'23:t ,. ,u. c.., , 1971 , AJtl.hd3l rufog,C:" fo1 lSCt~u 011:... t,lqsH1m ~pp.}. B:1/l,•1111 of r.1•, \HP.Rl '.\I \.\ ...,cu \Ul '\:G.\, ,·•• 19~9. E(fic;,cyo( ln"4!C'UCM.i: u ..·au•.J
Hmomu/og:t.Ytl /t,.,,,v,rth. 61, Jlt..350. ~ule •• J 'hamc1 10 rc-1n,·..,.lon u( 1wLie m,-.10 elea1<"1 ore:u 111
..133 , 1 ,u.. <;.,.\ , t<:t7~ I hun:spon~c.:.oft.scbe 01c'> {Dlptcr,,. Clo~)inid.'\.O: to north~cat.tem Ztmha.b\\'c, .\l,"rl!r1,t hntl l'Nc111u11,· £1:tomofog•, I :t
ms,blle nnd ·,Iation.iu; buh.... /h,/lr,/u off.Jtt<Jutl'Jf(lgh"W,tl R~~~o..nc/1, 64, l-;'-lA4
i>i!,-.588, ~ \\ATS<•~. 1-t.1.r. 198,_ Thcdom..111o1!c pJg.ns a rc~el'\'nir of T pm1bft•n~t.·
~J.l \'~.U. C,..\ l:i:", FittkJ rt:~pc.m~ of ~I Se JH~ llipr4,f;t; <ilo~ ..\rudk) lf) h1tt.·r11.t1rt011cJ ~i,,mi,(ir CJ:Jmmitlt.\• fur Trypan0$0mfruls Rrni01ch Sint/1
,ldow, of men. l,,crk ,,cld und ""1b~n dl1>x1de. 811/f,,tm of l'mmnolo,:i<al \!£.'e1ing, Cortitl:t:··
R,w,rd1, 6'l. 459-1"6. ,ss wm~ o.. t<lli:J. Th< focdini habit< or Gf,mma. 8111/e1it1 ofrl1C a'or!d
::is•·=· c..... 1980. Field Mucllc> c:>n the rt><~n= uft,..lw m,•, flralr/r Ofl1,nr.i;aM11. ia. 711 • 729,
GJo,,,nldaeJ:\lld othor Dlptcr.10 c:nrbon di~>cid~. occtono and ocher 256 \\lfT7. 6., 1s:--o. lf~hof <,los:dtJtr. hr ~ou1c..\.,, n.v;.. :ed.}. T;u•.ttlrt('nn
dtc=m1r'1.b. B:,ltt.tfn uj Eruorm;lngical RtstYm:h~..0. ;i\i.i-~;"0. f1)·1>a.110$0mia.~. l.ondon: ,,.,.orat: Allen and Umvm lid..

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76 ,rmc., o,c: ASpects inOucucing th<! occurrcnc.i uflnfeclious diseases

:?57 ,, ~ . t \., l9~ The imponn.nce of m(t('.h3rucal tr.immn,..s1on an th~ gencr.11 rp1dcmin!o!t)· uf C,amh1an ond RhOO~"S1u,n 'i!ct."f>ln~ s,c...i,~ m
epidcrruo1oj?v of nag.snri: A n.•,icw Tropz,nr .~1fmal 11,v1t1it mt,/ \frte1. T"iniJO.CIUPU IJ/ the Rnyal St>eit't}' of Tropimt .\ft.-lfrfn,•1m1I
P1t.Niu,·t!o11, 4. -:.i-aa. 11y11i,,,,•.59. r 1-1116.
2.;ff ,,11rru11An, 0.1.., 1985. UJse' of nntumJ product~ nnd 1hrir ;matogu~~ 1or ~ i. ,,,.·,Ut~1. R.
.!G $ WILLl-'~160~ D.l., U.\.. tl U, \.,<,;AH"-• t'J,ll., COOS, f.t.•• lt,\XU<l.
cornb.1111111 p,.-,.1>0! aJUICUllurol and public hNllh unr,or:uncc In Africo. P.\ 198l. lntt•tir.,tCun of ln~tt .\1crtlity .and lnserlidd~ for c:ontro1 ot
•&

/11: IILQI' r .. \. red.,. ltiort<gula1orsfa, Pf".J1 ComtQi, :i.merlc~ C:lit.'micttl Gl~fna mor,iuw~ 11wrslran, W~cw0od Oiptcra. GJ01;1.1nldA1.c iu
S()('ICt)' Sympo\r\lm ~riea- No, Zi6. fani'k!\l:t. \ lht• un~MCl ol S-(."t{UNUtul rt.iNiSl~(.l(~IerilttOO 1,l!tsC 0-il"'
2.5!1 \\1tt'rl'-"rn' , .s. .. 1q.s9. An cxp.:rimcn1 in c:ontro1 01 t"ICht' ,,·nh 1l111/l''11' orf.,11011111fog,w/ /11/St'<lt<II, 13,391-10-l
DDl-11ca1,>d ox,,n. 811ll,rl11 r,[Fnmm11l11$trt1/ /l,wr1r,·I,. 40. 123-13~.
211;; W?LSQS ,·.1 .. 197J. (_,_;1mo .:nd l:\t'l<C Hy in c:scwm /..amh111.. 0t't(J.J!cmttl
:t6o W1tot ..... JI. I. F1t.ESUI, \UI.• 194('.. Au OJ..ptrJml'.!nUi.lSIUd\" of Pi,,,-,,,, t,fth~ ,\'mumnl :,trl4"11tt11md ,\1tmur>u•ms ,,fRltt>df.i!n. :i, a.
Try'pallO-iOftUJ r1wdc'iicu4~ infotti.oa ln Zebu l"attie }Vllrtltll of 339-ll.f,I
c;,,mpn1111f1"t Pa1!111/og)• 55. 206-219.
266 t..\M&fZJ U\'f~fat..tr.. ASO u~ns l.n>.,:f.J\~B1A. l'-' ASSO<'t\TIO'.\: \~ttH IU)p
:161 ,, tll.l.\~1..~. a. 19'9J )fcde!s of nap S<-'(tking beh.1.\'lC>ur by s.stuc 11tcs nad .,J,..fl.\
t 1\·t-, · C)r., " ' " ' itJ .. u , • 110 NJ rm-Kt...\.-,D:'i. igq6.. Sm.d.l· of.lhi., ,,.•
'lnemon1.'0& 1<1111oiancs1s :u1d edge dot~tion. /our1:a/ of 11,,.,rttl<lll 1uahlrm ·nmmn111u,~,tg0lr,. Boi1uw,ur. \ "mniE)ifl und /.mnJ,tu
Biolug:.·. 163. 105-115. C.m,uh.u11, '"P"" 1111!1~ R,,:Jon•l T'l<!m, .,nd T~pann,omla,i, Conuol
lt:>2 n1W.\ '1~ n l)M~SFlll.D, f{, 81 OfUGlffi\~l t. A. 1991 The (Or.trot of hftSC Prognsmmc of \tal1J\\'I, "-loz.:in1bfqu.1,.'", l.cu(lbia .md ZtmtJab\,c. P.O. tt<n.
n1cs 111 relouion to tty mO\'Clll(mt ""d ,rnppmt= tllit,cnr~ Jnc,nwl nf \ 51i0 H.ratt, /.imbab\w.
,1pplie,t Emlt>g;•, 28. 163-H'9. :?..Gi ,1.u,\1u .... 1 ",,,.,u:-.t.tiM. u.1 .. 1,gy3,. \tetamorphoii~bcha\'!OUt.tnd
263 ,.,,LLErr. ~.c.. 1$fis. <;om,• nllscrv~tl<ln, <1111h~ z«cm tpldcml<>I~ or rcguJ,uion In Lo.ct~ fli'""' (ffo~um .ttpp.J CDlpwra: <il~1-i..nld.at1:., u."\it...\',
,tccplng <itkrn.-ss in 1hc N).in:e:, 1eglnn. l(c~)"· ond It, r.,faunn 10 the /llll/,•ri11 of 1,111umolO[llt'r1I fl{'l;,v,n·/1, IIJ, •l-17-~61.

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 3
Vectors: Muscidae

EM NEVlLL

Introduction reason. A second rca~on for southern African former.; ap-

parent apathy about these flies Is that ,hey are una\\'are that
In lhe Afrotropical Region (previously referred 10 as the so many species exist. each wilh its own requirements for
Ethiopian or sub-Saharan Region) the family :,.iuscidae em- sun1val and it,, own potential for disease transmission. For
braces 870 species in GO genera and six subfamilies, 1: How- example, at least 20 species of M11.<c11 are found on farms in
ever, only lhree subfamilies, the StomoX)inae. fanniinae, various parts oiSouth Africa, but are often confused 11ith the
and Musdnae comain genera of veterinary importance, house fly.31
these being Hae11uuobia. Haemmobosm. Stomoxys. M11srn. The purpose of this account is 10 prol'ide an insight imo
Hydrotaea, .Horellia. MusciJw and Fa1111ia. Species in these the life cvcles, disease tTansmission potentials and control of
genera are important not on!)' because they may cause irri· the muscid flies in the subcontinent. Identification of fami·
ta,ion co li\'eswck, resulting in production losses:• but also lie,. sub-families. geni?ra and certain species can be facili-
hecause of their potential as mechanical vectors of various tated by reference co publications by Walker.51 Greenbcrg 13
disease organisms. a face that llas l:wen generally neglected and Zumpt.55 These publications also provide information
in sou them •.\Inca. Greenberg 13 listed a multitude of organ- on biology and dbtribution. For a visual comparison of six
isms that have been isolated directly from \'3fious synan- common species of :-tuscidae. set> Figures 3.1 to 3.6
thropic fly species and which huve the potential to be
transmitted mechanically under the tight circumstances.
He reviewed the studies made lhroughout tbe world on Oy
Stomoxyinae
transmission of a wide varie1y of diseases. such as bmcello- In southem :\frica this subfamily is represented b) eight gen·
sis. anaplasmo$iS and summer mast iris. 1• era. the three main livestock,associaied genera being ilae-
The lilerawre is saturated with articles on 1he control of marobia. Haema1obosC11 and Sromo~ys.:;:; The adults of L11ese
the,e flies. In :-Jorth America much effort has gone mto the flies do not re,>d on nocrnr, but ;ire obligaiory blood-suckers.
chemical control of the horn Oy (/Taemarobia lrrirans irri- The}' possess a moslly slender, rigid and chlcinized proboscis
1a11s1 and the race fl)' t,\J11sca awumnalis). while In .\uscralia that enables them m puncture the skin of animals w obtain a
a biological control programme LO control the buffalo fly blood meal. '111e proboscb cannot be retracted. proiech tor-
<Haemmobia irri1a11s exigua) and tht' bush fly (,\fmm 11ums- ward and can often be seen from above (Figures 3.1 and 3.3).
rissima) by che imrod uction or e.~otic dung beetles was
started in the early I 970s. 3 However. the scable Oy ($romoxys
Haematobia and Haenzatoboscn spp.
ca/cirrn11s1. the house fl>• (/\'1usm domesrica). the lesser
house fly (Fannia canicularis) and the [alse stable fi>• (M11s- Common names: Hom Oies, horingvliec (At:rik.J
cilms1abula11s) continue to flourish in condirion,- created by
humans. especially In intensive fanning simatlons such as These are obligatory blood-sucking muscid Oies that seldom
dairies. feedloc,- and chickt:n lttrrn~.2' The 1:effuru; uml l11ic· leave tht, host t~t:olpt H> lay eggs in rel:tmtly deposito:d du11g.
nuil) of humans co produce more milk. meat and eggs do Female Haemarobin irrira11s irriums flies (Figure 3.3 have
not always Include efforts co avoid a greater Oy problem! been found to average 38.•l feeds in 2.1 hours and male; 24
Members of the Muscidae are not biological transmiuers feeds. 1r, This semi-permanenrly parasiuc way of life ensures
orinfcctious dise11Ses. Although they have been shown 10 be a food suppl)', and a larval habitat clo~e at hand. It also
capable of mechanically transmitting numerou~ disease- leaves them \'ery \'Ulncrable to many insecticides applied ui
producing organisms. they are seldom controlled for this animals for the comrol of ticks.
T,

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78 -=, o~,: ,\~pee,~ influencing 1h1· occurrence ofinfec1ious di sense,;

~
l
l
Figure 3.1 Figure 3.3

Figure 3..2 figure 3.4

Figures 3.1 to 3.4 Common synanihrop;c flies. namely S,omoxvs ca/cit1sns(figure 3.1 ), Far.n,;; caniculens figure 3.2). Heemacoo,e irr,tc?n$\Figure
3.31 and '1t11scina scabulimsffigure 3.41 Tne,r s,ze. :yce of rr,nuthl)arts and shape o' the fourtn w,ng \e,n M1~2 are impona~: dragl\Qs,·: c!·aracte·s
(lllus11a:iot1S !ly f Gre1)o: ,n Gl8enberg, aema,o Fliss end D.sease. Volume l CopynQh, 8 '971 Pl.iP fl!pnnted by perm,ss10n o' ?rm:e:c~ um, ers.;·,
Press)

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 \'l.,r1<1r,,, :>-111,cidae ,9

1 l
figure 3.5 Figure 3.6

Figures 3.5 10 3.6 -.~ s;,eC11!$ of llie genus Mus:a name,, W dumesc,cetfigure 3.St and M S(Jtbe-.s !Figure 3.61, Note the similar shape o' tts
founh wir.g •.e,r 1,:1-2. which ,s d1agn0St1c for th': genu5. Also com;,a1e this vein w1lh those of t,e iou; other genera of synan1h1oplc flies dep·ctsd 1n
Figures 3 1 ,~ 3 ~ 1lllusuat1on$ by F Gregor 1n Gieenber~, Bama:d. F.'.es and Disease. Volume i , Coovr,g~, 8 1271 PUP Repnntsd by oerm,ss1on of
Pnnce:1m Ur ve!'Sltv Pr~ss,

Three species of 1/aem(l(obia h,\\'e been recorded from African buffalo In !>oulll Africa. Since buffalo are curiers oi
~ourhetn Africa, namely H. rhirow:i po1n11s, fl. ml'ridin11a the foo1-and-mou1h disea~e ,irus, the possibility exists that
and H. spinigem, all of which are primari!}· associated wilh this horn fly could transmit the virus mechanically from buf-
game, especially African buffaloes CSynterus mffer).3 ' In falo to nearby cattle. Two attempts ac the Onder.tepoon
1937/38 H. rllirouxi pou111s1,·as repor1ed to be a problem on Vetcrina1y Institute to 1ransmi1 the ,1ru~ by thi~ means from
cattle along the coas1 from the Eastern Capr Pro\'ince 10 infected t!) o;usceptihle caule (af1er m·o hou~l \,ere
southern KwaZulu-NataL9• ..~ The problem disappeared unsucces.~ful.48
after ~yntheric organo-chlorine lnsecticidt!s were used al the ;\ second genus of horn fly. llaemmobosc<1. "'as listed by
end of 1l1e Second \\"orld \.\'ar.18Today. horn flies have again mosl earlr authors under 1/aemarobia or 8del/ol«ry•11x.s5
become a pest on farms. eicher where non-insecticidal tick- Pour species of llt1em(l(obo.<c-a have been rt'Corded from
detaching agents are tised for tick con1rol. or where no rick southern :\frii:a. namely fl. 1mMeriara. H. larifro11s. H. m1gu.<-
comrol is prac1 lsed. tifrons and H. :.u/11e11$is. 1~ Only che first rwo specics are
In somhem Africa no 1/aemaro/Jia spp. have so far been com mon.55 Neremacolmsca uniseriara was firsl recorded in
pro,·en to be \·ectors of a11y organism lhal cau~~~ dbea~e in tile then Transvaal Pro\i.nce 100\\' compnsing Limpopo.
game or livestod,. In many other pans o:· 1he world Haema- Gauumg. '.\;Orth \\'e<,t and JllpumaJa nga provincit:!s) in 1933.
10bia spp. are biological vecrors of Srephanoji/(lrta spp. t \\'hen :,iieschulz and nu Toit3R iound il reeding in relanvely
The presence of stcphnnofilari(lsis in ca11lc hns not yet bc.:c·n !urge numbor;; on 1hc belly nnd leg:, of hor~cs near 1hc
definitely confirmed in ~outhern Afri~c1. but in the 1980s and Onderstepoon \'etcrinary !11sri1u1e. Zu mpt 55 caught speci-
early 1990s stephanolllariasis-like lesions \1cre seen in cattle mens on a l'reshly killed \\'Srthog (Plmcoclwerus aetliiopi-
in lhe KwaZulu-!\'.atal Lowveld and \'enda in the Limpopo cus). This fly was also collecred 011' cau le on farms in the
Province 0fS0uU1 Africa. and in Swaziland,31 where I.hey oc- Kumman arra in the eastern pan of 1he :,.:onhern Cape
curred in association wich large numbers of II. meridinna.l7 Province during an unusually wet yt•ar (19i4J. In subse-
l·lai:mmobia rhiro11xi 1101nm is found in large numbers on quent drier years il was present in much lower number~.:?!!

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80 ,,~'t''" '" Aspects influcntmg the ouurrenc~ ofinfoctiou, dh,ca-~

Stomo.ty s spp. f:trm hygiene and treatment with residual insecticides ot the
resting places of the adult flie:, and the animal~ themseh·e.s.
Common names: Stable mes. ~talvliee C:\frlk.
/1 variation on some of these approaches to control \ltilizcs
Eight species of Sromoxys have been recorded in sonthern 1he preference of adult stable flies 10 rest on t·er:atn surfaces..
J\frica~ 2 with 'i. cnlcirmm; (Figur~ 3.I J being the major pest It was found that translucent white fibreglass panels. trade-
~pecies. In South ,\frlca Swmoxys 11£ger (= S. nlgmJ i, seldom named · Us}nl1e·, c<iuld be used 10 construct an .mractlvc
collected. although it can, on occasion . occur in pest pro- stid,-y samplin~ trap for thcso flics. 5·1 If treated with a pyre·
1>ortions in the wetter and ho11er parts such as the Lmweld throicl thh de,icl.! could also be used to control 1hem.2 ·1 but
of .\·!pumt1langa and north-eastern KwaZulu-:-.;a1aJ.3 1 By the 1-alue of this approach in t11eir control ha~ not yet been
contrast. S. nigerwas rated in Mauritius to be a serious pest in\'estigated in sourhern Africa. Traps baited with symhctic
of cattle and deer while S. calci1ranswas con~idered co be of odour attractant!,. de,·eloped to lure und catch tset~e flies.
negligible hnportnnce.27 also auract stnbll' flies (and flies belonging to the Taban-
Stable fiie5 are compu.lsory blood;feeders. but !.lie} differ idae). These traps are predominantly bright bh;e (ptha.logen
from horn flies in that they only spend shun periods feeding blue\ and may be made of clorh, e.g. 11:IJ:,; and H traps.2° or
on their hosts. digesting their meals \,·hile resting on nearby of hardboard painted with a highly sticky substance ,uc:h as
surfaces. They feed on uverage t'A/4ce a day. 16 usually on polybmene or ·Temoocid'.~1
whichever host happens to be a\·ailable at the time. If inter·
rupted they 1,~11 feed more often. This bcha\'iour makes
them ideal mechanical transmitters of any disea~e organ-
Fanniinae
isms present in the blood or their host. The main nuisance fly in this subfamil> is Fn1111ia C(ll1/c11lnris
\ \'orld1,idc there are numerous examples where swmoxys (Fi1,'llre 3.21. the.sti•called lesser how.c Oy. It is often confused
spp. (particularly S. calcitra/ls) ha,·e be1m shown to be eithe, with the house fly (Musca domesrim) but is slightly smaller
invoh-ed in. or suspected of, mechanically 1ran~mitring dis- and more ~ll'nder in build. It is a cosmopolitan fly associated
ease organisms such as Brw:elltl ,pp. Borre/in spp. and primarily with lntenSi\'e poultry production. where it breeds
equine infectious anaemia.55 In 6.itain it hai, been pro1·ccl ex- in the litter. Its sheer numbers make it a nuisance a.nd it often
perimenwlly that S. cnlcitrr111s can act as an efficient infe:;ts neighbouring properties. These flies are commonly
mechanicul vector of capripox and African swine fever seen in daylimt• $wanns cirding in midair between cage aisles
\'iruses. 25 ll Is also believed that this fl~ may be capable or on poultl} ranches. in work alleys in milking parlours. and
mechanically trar1.~mitting foot-and-mouth dbea~e virus. under trees or other sheltered places. Unl!ke house flies. Fa11-
These findings are particularly significant ior southern Africa nit1 me less lmponant as household pests or disease vec-
where lumpy skin disease, •\frican swine fel'er and fuot-and- tors.23 In high-density poultry houses in South Africa rhe
moulh disease are endemic. In South Africa it ha~ been main conO'OI approach for it. as well as for the hou,l! lly. i~ the
shown experimentally that S. mlcirrans can mechanically ;iddition of a cyromazine-bas!.'d additive 10 the feed. C~To-
transmit.4naplasma margi11a/ewithin a group ofranle. 13 and ma,dne inhihil\> the 1om1ation of d1itin ~o that den·lopmem
that it can, in the field, .1c1 ~ a mcrhanical ,·ectorof Besnoitin 10 the foU01,ing larl'al stage is prcvented.3
besnoili, rhccauseofbovinc bcsnoitiosisorclepham ski11 dis·
ease.~ These flies are therefore a Vl'ry real means of dlsease
t ran~mlssion bcmoeen animal, in a herd or fePdlot
Nluscinae
Members of the St0moxyinaedifferwiclel~ in rheirbreed· \!though this subfamil~ contains ta genera.u on!,· species
ing requiwments. Hr,ematobia and Hnemmo/Jo.ica spp. of the genus Mu.<ca are commonly found on livestock in
breed in freshly dropped dung from animal$ grazing in the southern Africa. However. low numbers of other gtmt!ra
veld. whereas$. calcfrr{U1S prefers fermeming organic mate· such as Muscint1 and \forellitt may sometime$ he collected
rial. such as rotting heaps of discarded animal feed or bed- olf livestock.
ding. that may or may not comain dung. In 1997, at the
Johannesburg Zoo in South Africa. ,he sourCt' of a severe S.
Muscina stab11lans
ralcirra.m problem was found to be rorting klkuyu (Pl!1111is-
ewm cln11cles1inru11) lawn clipping~ lying in windrows on the Common name: False s1able fly
lawns in 1.he public area and adjacent Zoo Lake Parle For
1hesc reason~ S. calcitr<m.• is u~uo.llr as~ocimecl \\ith intcn ·1ne false stable fly (Figure 3 ,4) b also often mistaken for thC'
sive fanning and suburban situations, "hile 1/nemaro/Jia hou"e fly be<:ause of Its simllorlty In 1:olour, size. and food
and Hmmwrobosc:a spp. are. more commonly found where preference. and its habit of emcrlr1g homes. ln appeanince it
cattle are free-ranging. fall~ in berwecn Sromo.q•s and \Jusc11 in that its \,ing vena-
\\'hile conrrol of horn mes is relatively easy where facili- tion is similar to S1omoxys but mouth parts resemble those of
ries exist for trea ting livesroC'k with insecticides. S101110:rys M. domes1ict1. It ieeds on and deposits its egg;; in a \dde va-
,omrol is more difficult and must rely on a i;ombination of riety of anfaial excrement (poultry, horse, cow. domestic

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 Vectors· )lu,<:tdae 81

pets) and deca)injl;,·e~etable maner.23 !ls larYae have been which bri:eds in dung and dt.-ca)ing organic maner and reiu~c.
known 10 prey on those of other flies. 1 all the other .\ llc;m ~pp. mentioned above breed in fresh dung
in the wdd and their lan'llc all have to feed and sunive in this
medium. Over thousands olycars. evolution has resulted in the
!vforellia spp.
development or a ,-aiie1y of $pt!Cies. each with ii~ own pro:fer-
Co11unon nullle. Sweat fiil!S (!11u,s fo1 fuoc.l, Lt:1npcn11ure. u1oi~turt: w1u uthc1 1.:quircmem~.
Twenty-five ,1/usrn spp. have i<o far been recorded in
Thelie Ilic, have been collec1ed off caule whel'e th!ly ,crotch southern Africa.~ ·•· They belong to the subgenera £11-
the skin surface and lap up exuding blood. In Sr,u1h Africo 1111,sm. Byomya ..\lm·ra. Philaemmomyia. \li1,iparomma1
they have not been found in high numbers. They arc- also of and l,issosterna. :2 lhe 12 ,\fusca spp. listed in Table 3. I he·
interest be('l!U$C like M. sralmlll/1$, their lacvae prey on long to 1h<:- first four ~ubgenera Only five species of the last
other larvae in the dwlg in which the) de,·etop. two subgenera have bc>en recorded in southern Africa~·· a.ncl
these have ~eldom, if e,·er. been collected in South -\frica.31
The observations on Musca ,;pp. below are therefore re-
Hydrotaea spp.
stricted 10 13 South African species.
Common names: Sheep head flies; plantation Oi(',
(HJ(lrotaea irrirans) Subgenus 131111111.SCll
Common names: African face mes. Afrikagesigsvlrcc (Airik.l
Although species of I lydroraea hc1ve not yet been recorded
as pests In southern Africa. certain sped~~ elsewhere in the Musca .\·nmhomi'las is ofwn the commonest -\frican face fl)·
world ha,-e warranted serious auemion. In Europe. espe- associated "ith can le in the ,"eld in South \frica. It is found
cial!)' the '\etherlands. Denmark. German} i.lelgium and tliroughout the yt>ar m the Limpopo Pro,ince Bu~h\·eld but
the UK, Hydrouu:a irriums is regarded as the mechanical is less common in the South African High,·eld. It lays an a,·
vector of summer mastitis.~ 5 In Czechos1ovakia, H. armfpe; erage oifourbatches oi33 eggs each and may sur\-h·e for up
ii; suspected of being a mechanical vector of lnillct iou~ bO· 10 35 days at 2.; 0 C.33 \111sm xa111/1amela,f has only once been
vine keratoconjunctiviris.8 collecceu ~outh of latitude 29•.!8
M11SC11 luI01in is the largest of theAfrican foce flies. !tis com-
mon throughout the year in the Umpopo Provin,e 811slweld.
Musca spp.
but, as is the c;;sc ,,i1l1 M• .xamhome/tl$. is also !es, common on
Common names: House flies. hui.svlice (.\frik, ,: face tlies. the Highveld. The fomale 1s ovoviviparous. depositing one late
gesig~\·liei! IAfrlk,J first stage Ian a a1 a time on fresh dung, and Is abll' 10 produce
up to 27 lan":le in her life-timc ofup to 75 days.~··
E.~cept for the cosmopolitan M. <lomesrim (Figure 3.5). the Musca 11e11illi dosl'ly resembles M. xa111hom~t,1s in ap-
vererinary Importance of 01her species of the genus .\fus.:a has pearance but produces icwer eggs.22 :i3 It diffe~ great!).
been largely overlooked in southern Africa and there has been however. in hs seasonal prevalence. being common only
a tendency 10 muibu te problems caused by them co,\/, domes- from late Januarv to Apdl.
rica. In the 19;Qs and l981)s, however. various studie;. in South M11se<1 (l(:1hio11; Is rarely C'ollected off can le. but ha~ been
Africa 011 the transmission of the filarial \\'Om1. Parafilaria IJO· reared rrom the dung of both cattle and Al'rican builalo.
vice/a, in the Umpopo Pro\i.n~e Buslwe!d;l<l n Oy prohlem in Musrn m1mmi was never collected in rhe sun·ey~ sum-
the "'orthern Cape Pro\ince,34 the tran~mission of bo\'lne ma- marized in Table 3.1 . On occasion it is quite common on
ligna.m catarrhal fever in the l.lmpopo Province Bushvclcl,3.; cattle farms In parts of KwaZulu-~ai:aland the £aqern Cape
and in :--:amibia on the epidemiology of anaplasmosh in provinces or ~ulh ,._lrica.
caule.3 z have led 10 an increased intere;.1 in the .\/t1$Ca spp.
as.~ocimetl \,ich canle and, co a lesser extent. wild·li\ing game Subgenus Byomyll
animals (Table 3. l). Six Musca spp, rrable 3.J, were common M11Kn co11fisc111a [= ,\J, fascimal is a very versatile species in
10 most of the fanning situations examined. naml'I) .\/. .mmh- that it may be found in reasonable numbers in both the
omeltu, .\1. lw,·oria, .\I. domestica. M. ronfitcata '.= .\/. fa.~ir1w). Bush\'eld and Higlweld areas of South Africa. It brct'cL~ in
M. 11e1,il/i and .\J, scrlxms (Figure 3.6). The presence o( certain both caule and wildebeest 1Co111wchaetes spp., dunjt. ;md
other,specie.; wa~ dependent on the geographical arna. tht> 5ea- foedb on caule and presumably ah,o on game. llb abilif) to
son and climatological factors. It was onl~· near hotnl!Stt>ads u tilize ,-nrious ~ituation, would therefore w<~ll qu:llify thi, flv
that the ")'llanthropic M. domestica and \/, sorbf!/1$ \,ere im- to act as a mechanical transmitter of disease organi,ms.
ponam. In t11e \'eld. awa) from the homesteads. there is an ."111sca sor/M11, (Figure 3.6). llke .W. nmfi,rota, ma)· be
imerac1ion between various symbovine \Jusca ,;pp., and the fou nd 1htQughout the year in the Limpopo Prmince Bu~h-
dominant ~pecie$ change according to both local and the veld but. Hke .W. domesr/ca, it is associated with human
broader climatic factors.. \\'Ith the exception of M. do111es1ica habitation~ and dot', not brl'ed in Animal dung in the veld.

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82 S«:TU>>. ox.: Aspec1s influcncln<t ,he occurrence o(infcc1ious dbeascs

Two forms of .\1. sorbe11s have been recognized i11 sourhern Feeding habits of Musca spp.
\frica.41 Bolh are anractcd to fresh human faeces. which ,\II Mmc<l spp. require water. sug(lrs and protein ill order 10
appears to be cheir preferred breeding medium.' 1 survil'e and reproduce. Flies tha1 live in the veld obtain 1hese
,Wusca 1empes1at11m is only encountered in the 1~cinityof basic requirements from fresh dung. eye and nasal secre-
1,ildebeest. in whose dung it prefer~ lo breed. tions, wound~ and sore11 on animal~, carrion. plant and
,vtuscafrcedmaniis a lesscr-knom\ fly whose preferences flower ~ecrerions, and !.he cxcrcrions of sup-sucking in·
hn1·e n01 as yet bee11 observed. sects:'3 • •1~ Protein h essemial for the development of the
Musca lasiophthalma is occasio11nll}' collected in 1he ovaric~ of the Ole$. Protein hunger in female face flies (.\f.
Bush1·eld in wimer. but has been collected in large munbers <mt1111111a/is) ha~ been clirec1ly related 10 the Sl3ge or oogen·
on the H igh1·eld in both winter and eaily spring, e~b and b significantly diminished sometime before the end
Musca co11d11ce11s has been reared from canle dung of each gonoirophic cycle. 50 All M11.~ca spp. po'-•ess a
samples collected in the Bush veld but ha) seldom been col- sponge-like lapping mouth-pnn or labellum maclu up of
lected off cattle. II is possible that the llyiclemified as M. COIi· numerous medially dircc1ed channels or p~eudo1racheue
d11ce11s in southern Africa is different from the nominate which radiate out from a cemral oral opening. Fluid ls
species. which is found in A~ia.40 sucked up along these pseudotracheae into a central cavity
and then , ia 1he labial-guuer-epipharyngeaJ mbe into the
Subgenus Musca digestil'e system. 10 l:.xccss lluict Is stored m a large sac-Uke
Mu.sea domesrica is divided imo three ,ubspecies in the crop from which it is either passed 10 the mid-gut for diges·
Afrotropical Region.42 Two of these subspecies arc indi· tion or rcgurglratcd by I he fly if more auracci,·e food is
genous 10 Africa. Including southern Africa. These are .\1. d, found. Tht. fluid Is also used 10 dissolve or dilute food that Is
calleva (an oucdoor fly) and M. d. curoiforci!ps (more domes· too coarse or viscous to 1ake up through the pseudotra-
tic in habits). Hybrids ma~· also be found.39 During the writ- cheae.53 The II) mixes the regurgitated fluid \\ith the food.
er's studies In the Bushn'ld. ,\f. domestlc;a was never ofu~n using its prcstomal teeth for the purpose.
identified 10 subspecie~ level. It was common near home· All .\!u~ca spp. have one c,r more circles of prcstomal
stead kraals in the first ha.Ir of i.ummer bu1 was seldom teeth. They ate ~eldom visible a~ they are <inl~ ex1ruded
reared from cattle dung samples collected In the l'eld. during feeding. and are 01herwise hidden by tht> enveloping
l,1bellar lobes of the mouth-parts. The first row of these
Subgenus Philaematom.y ia pres1omai 1ee1h ranges in size and l'onn according to the
Alusca crttssirostris is a 'veld-lly', apparent!\ common only species. For examph?, •hort. relatively blum 1celh are present
in dry years. in the house lly 1,\1. dm11t?slica) and an ,\frican fan, fly

Table 3.1 The relat1\'8 abundance of various Musce spp collecied of! cattle and reared frcm ine ~uno of cattle and blue wildebeest IComochoe1es
raurinus iautim,s) 1n S1lUtl1ern Africa
COlLECTED OFF CATilE (%1 DUNG-FEARED I •,J
MUSCA SPECIES RSA' IIAM1BIA RSA· HIGHVELD RSA• BUSHVELO RSA· SUSHI/ELD
N CAPE
KURUMAN WINDHOEK PRETORIA LYOENBURG VARIOUS TI-IABAZIMBI l HA$AZIMB1 O!STIUCT
DISTRICT OISTAICT DISTRICT DISTRICT DISTRICTS DISTRICT CA.me WILDEBEEST
1973-1977 197s-·gso 1974 1974 1973-1976 1987 1987 1987
M. lusa1ie 11.4 25.3 6.6 4.1 !.i0,7 12.6 j~.4 i ..6
M. x;m/flamelas 28.2 52,8 0.7 2,1 9,3 27,8 4.1,9 o,.:
M. nevi/11 3,3 1,l 0.2 ·o.3 15,9 4.9 ..a
M. domes/lea 18, 1 S.8 23A 26.2 18.8 32.5 G.6 a:
M. conftsca1a - 6,7. 12,0 7,5 36.~ 7.3 7.l 12.0 ~l.9
M.smoens 2,3 0.1 1,6 14A 7. 3.1
M. crass,rosws 20.4 0.7 ·.o 0.2 9.2
M. re,.,,pes:a1um 0,4 0.6 5· .S
M. freedman, 8,8 0,5 0 '. 0.2 0,3 0, l
N'. 1as1opo1t1a1ma 1,2 13 60,0 16,9 0,3
M. ae1hiops 0,1 1.6
M. t:Qr.ducens 0.2 0.2 ·1.6 2.5
Fly ;c;a/s •• 4()
3262 8000
1 889 19S 20~5 2380 2631

• Re,:uohc ot So,nh Ai11ca

• • Accoromg to Pcot'2 M. ronf,scaui 1s the re;:iac~ment name for M lasc1ate

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 Vi,cr<1r,.: M.:scida., 83

(,\I..wmlzomela.,): 1 longer sharper teerh in the face fly M. au- plasma spp.. Chlamydophi/a sp. or rickeusiae (see Chapter
tumnnli;. which occurs in Europe and :-:orth America and 140: Morruella spp. infections). ~o work has yet been done
two African face rues (.\1. nevi/Ii and M. lusorial; and large in southern Africa on the transmission of any of chcsc organ-
coarse teeth in .\1. c:rassirostris10 (Figures 3.7 to 3. 12,. isms, but detailed studies have been conducted in the USA
The feeding habhs of flies differacc-ording 10 the ~1ie of on the tranhm'.!>l>ion of.\l. lio11is by 1he face fl}'·,\/. n1111111111a·
prescomal teeth present. Musca donw.~rfca and .W. .wmrllo- lls.' 5 The :-.onh American studies proved conch1sively 1ha1
melas !Figu res 3 .7 and 3.8). feed mainly on the lachrymal the damage done 10 the eye surface by the prescomal recrh or
secre1lon~ and wounds alrnady present on an animal: M. a11r111111wlis "hile feeding predisposes the eye, of can le
M autum1wlis Figure 3 .9) feeds direct!) on conjuncti\'al to infection.~·1 tn addirion, experimental smdic;: have ~hown
tissue"· 44 and .\1. /11soria (the southern African counterpart that ,w. /Jol'i$ may be present in material regurgitated from
of M. t111111mnalis) (Figure 3.10) 10 ma) b<' <'xpencd 10 feed che crop qifface nle:,, and that infection of the eyes of caule
in the same manner. Ir has also been shown thrtt .\I. m·1•illi may follow exposure to infected Oies.'· 12 Furthem1ore. a
(Figure.3.1 1) has prestomaJ teeth almost identical to those htghcr pre,,alence of 'p111keye· in the field has been related
1
of M. mirwmwlis (Figure 3.9). 6 Muscc1 c~assirnstris ~eldom 10 face fly numbel's. The,\frican face Oies fall into the same
visits the eyes or wounds o-n ca11le, but crawls under the subgenus as ,'vi'. a1111111111nlis. and have similar feeding habits
hair of animals to rasp away the skin surface with ics large, and prestomal teeth (especially M. lusoria and ,\/, ne11illl).
coarse teeth until blood is drawn, on which it will engorge ·mey may therefore 1ransml1 JW. bo11is in the same manner.
(Figure 3 .12). .\s an obligatory blood-feeder,,\/. crassiros· In South Africa invcs1igations have been conducted 11110
rris is therefore the Musca equivalent or Stomoxys. Musca the possible role of .W11sca spp. in che field transmission or
co1ulucen$ has a similar feeding mechanism. although the anaplasmosis, foot-and·n1outh disease and malignant ca-
mouth-pans are not as heavily developed as arc those in tarrbal fever. Unfor1unatt:'ly none of tl1ese studies has bcc,n
.\1. crassirostris. sufficiently imensh c 10 prove or disprove the vectorial role
<>f these flies.
Disease transmission potential of the Muscinae
The control of flies and disease
Very little is kno1,,1 about 1he transmission of infectious
agents by /\Jusct1 spp. in southern Africa; that whkh has Under intensive condit ions A mu lti·faceted approach is
been recorded ls based largely on circuns1antial e1idcnce. necessaT} for the control of so-called filth flies. the most
.'1,fmc11 co11fisca1a (= M. /(1sck1ta} has been ,uspec1cd of me· common of which Is the house fly (M. dome$1ica). This
chanically cransmitcing lumpy skin disease ,inis.~2 ,\Jso. should include good managemcm, Lhe judicious use of
during on outbreak of enzootic abortion in ~hcep in the pesticides, and 1he encouragement of parasites and pre·
:'liorthem Cape Pro1incc in 1975, a C:lllamydop/1i/a <p. was dators.
isolated from Musca spp. collected around sheep. 19 Good managemem practices are essential lO reduce and
In the search for the vectors of Purajifarin ho1•icola.30 eliminate conditions Lhat favour fly development..13 These
20 000 specimens of 11 ,Wusca spp. were collened in the should include:
Bushveld ofSouth Africa and identified (Table 3.J ). Oithcsc. • a system of manure management focusing on irs collec-
M. Ir1,<oria • .\/. :mnr/1om1:lt1s and M. llt'vil/i, all belonging 10 tion. processing, storage and 11tili;ia!ion as a fenili1.er or
the subgenus E11m11sc,1. were shown 10 be vectors. These compost.
tlies accounted for 65,3 per cent of all the specimens col· • proper water management. including supply. drainage.
lec1ed. As African face !lies breed in fresh dung deposited in and dbpo,;al sysu:m,. taking into account above-normal
the veld, ii is only in these sl1uations that they occ:ur in large rainfall fn the area concerned. and
enough numbers 10 act as vectors of the la.n·ae of chis wonn. • general farm sanuation involving che routine disposal of
Face flies are unable to de1·elop In the dung of caule which other animal wa:.tt.'~. spilt feed. organic debrb. and e1 en
have been fed a supplementary grain diet. as this dung fer- household garbage. 2•
ments: iL is. however. highly Sltitable for this purpose for
house flies. This previously unsuspected predominance of Although certain r1.>gistered insecticides may be applied di·
African face flies in the veld has stimulated thinking abouc rectl)' to livesiod.. insecticides are best used as residual
the possible involvement of these and other Musca spp. in sprays on resting surfaces and incorporated in attractive
the u:an.mis$ion of huectious agent$, not only bc.lt\\'een baits in feedlo,& and dairies. In this way only resting and
c:artle but also between carrier game animals and canle In feeding adult flies \\ill be kHled and 1101 the wasp para~ites
souchern .\frica. searching the breeding medium for fly pupae 10 parasitize
One of the diseases most likely to be transm.ined me· nor other anhropod predators in search of egg. and larvae
chanically by !lies in the vcl<l is infectious kennoconjunc- on which 10 reed. Certain poultry feed additil•es sl)ch a,
civitis ('pinkeye'). which may be caused by a number cyromazine. which inhibits the moulting of fly larvae.
of different organism~. indudilig ,\Joraxella botl/'s, M.1·,·0- are harmless 10 non-dipteran parasites and predators.

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8.J ,,c,10" ll'<I: 'lspecis fnllu~ncilig the occurrence of infectious diseases

Figure 3.7

. j

Figure 3.8 Figure 3.11

.;_ ::/
Figure 3.9 Figure 3.12
Figures 3.7 to 3.12 Scanning elewon m1crograoM c· the orestomal teein of six lvluscz si:;: name, Al dcme.<rica !Figure 3.7 I,, x,r.:ro ;,11 iiS, Figure
3.81. M euru:rna, ,s(Figure 3.9 1'1.1 tusor,a.Figure 3.10 , .-., nevi/!i(Figure 3.11 •anct M. C'E~Nost,,s Figure 3.12t. (Bycour;es•, 1l · D·s;. 8 B~:e ana
!I J Ez,ni;a, ~ar.sas State Umve1Stt,' US.41

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 \'eetQr,.. :vlu$cidae 85

C}TOm2;zine is also available in a formulation that can be ap· by 1'1:evill et a/., ls who drasucally redu ced rhe prevalence of
plied direnly 10 fly-breeding material. 31 P. ho11irola infestation In caufe by weekly to fortnightly
dipping with a pyre1hroid insecticide. An easier method of
Under extensive conditions Field studies indicate !hat face II\' control is to auach pyrethroid-impregnated tags 10
most fl} specie~ 1, hich bref'd in the veld seek out the nearc,t both e:irs of an animal. In 1his way. in one British study. a
host animals and do not fly far. unless the animals are re- reduction of up 10 90 per cent In face fly numbers was
moved from the area. Preliminaiy proof of this has t:ome achieved during a rwo- to three- month period. and infec-
from insecticidal trials against African face flies in thf.' Lim· tiou~ kera toconjuncthitis wa~ almost complete!)" elimi-
popo Pro\~nce Buslw<'ld of Sou th Africa.- A marked fly re- nated.'10 Th is principle of fly rcducrion should be applied in
duction could be maintained in an insecticide-treated herd situa1ions where losses due to fly-transmi tted d iseases wa r·
of cattle while the control herd, only 3 km away, had a high rant the effon and expenditure. In South Africa no impreg-
Incidence of Illes. indicating thnt these mes hnd not come nated car-tag5 are registered for fly control in livestock. This
into comact with the created animals. is because treatment is aimed primarily ,It ricks, for which
Reduction of fly numbers on cattle in the \eld should ear-tag~ are unsuitable. Ho\\'C\'l?r, pour-on p)Tethroidlt.
therefore lead 10 a reduction in the transmission of certain which are often effective for fly control, have become
infective organisms. This has been clearly demonstrated popular.31

References
,llt!..,o~1.1.. ,,1Ut..lrT, ILL,, it,A.1110, f.1, ~ 111o;:m. ~->-. t~.: Tr.i.nsmb':>tan n1 p)Qkcyc an cuul,...•.\ re\,e\, ~nd ~ymh1.~i."toflhv rde\'ant hlt'ratJrc.
\/Qrn.tollfl hwls Imm bl~oll og,r cuhun,,; io Hcroinrd C4tll~ b)' f1Jl:f! Ill<$ /011r1m/ of \/<'(//1:(1/ F111011u1/og;r. 2l. 36 l ·JtiS.
OiJllffl: ,\fU-iCIUue1. /f>umal of l;'txJuomir Lmomolos,•. ;- Js,;-.3~8. 16 ~UlU.lf, 1A l'I 11tAZAll. l.D .. 1971, f-lOm fl[~$ a.nd >t.ibl~ Oi~!\;
ll~lOU'i • .K.1
2 DIG.'\l.Kt JU> , IQf,8, ~tW C01l('C(lts on thu tpidfm:OIOfPt"'Jl h:aru:c..'lo of h:,·dlng nc-tt\'h), ,;,,,wt. of tlu• linrnmu/og!Cal Sodt'tvo/ ·}mufec., G7.
bovinr bl"SnOitlO.)lS 05, dcumnin1..'tl by laborrm>r}'A..lld field 69)-1Jg,I
ln\'e.5fi&iltlon.s. Ontft,r.<11.,poort ftmrnal of\ #twrmor:,· RPJ,4.,i,rth. 35.J-i 38. 11 u1nuu, c.P,, J'ittt• U~\·cJn1,.mcm nr 51ttp/u,1mfi/Jttl11 srifr~'i'in tht' horn tl)
J JU1JtXl-. '""'""'· r,.F.. 19,;,:t. t'hl" Al"Jrnlhm Onng Rt-.,Jlr Rr'4c,;vrh t 1nit 1n Juim:ttl uf l'11m#111loro•. ~2. a91H198
Pteton~ ~utlr Afric,m Journal a/St1tnc,•, -~. 257-260. 18 uowl!ll. ,.. ,.. 1r..s. \'t'tctlnar'!, t•mon'mtow,1 en suuthcm Atriro..
l IR(l<;J-. ,.n. l!l$, i;,lns.JS ~UII~ Unt,~rSlly ll!, \. rcnon.1! limomut,,~· .·.~emair. Xo• .;.1. DL·l>.trtm1·nt nr AJ:ric:uhlm1l T1.-chnir.al
commun.tca1f<m ~rvic~~ R1,1puhltc nt '\nurh Mri,·a

:; lk()CJ;, ,.,, • " " " · ' ILi., 19n ,. Comparh<>n of prc,.mn,,u cttth 1n tho 19 1mwr-;11 c. J. 1r,;;. Ondt1'Wpo<m \'t•tt•nnur,1 lnsurut('. ~uth :\trk:1
tac" ily ,\[11$.Cn tJWum,w!ti, :snd 1hu houi:c fiy .,tu.l<'rl domct.•Mla,) Uupuhh,h,-d dn14.
D1pt••~ Mu,cidoo: Jo1u11nl of .\1<-111,11/ F111omol()f() • I 82-115 :o ~\Pr:\trllJI: . h :qoo, \ Ot."\'.1) dt."\'CIQpt·d odour·b.dt<"d ·H cnap· (Qr the
6 8Roct •.\.II &. n.:tlt..:(~\ l'l.J .• IQS:8. J(a.ns.:l:i. S1+ite Un.l\.CM~ USA. fH.•1.:on:11 llv'\" colti;:cnonof (i/qi$inn lirei 1ipal11u and (i. mr,.{t,·m fOlpter.a:
cnmmunl~tiun. c;le;,,.. intd,,c 1:l So\nh.\tric~. o,ide1$fl.1XXlfl /111,rm,I nf\'ert1111u,ry
Ht'S,'ttr.fh, 67. 15-2$
; t ..,rro~. o.G ... 11)8.J. C~·hnlnthiin. .-. n•Jw s~1uhedc P)TNhrold- cfficacr
""r.1b1bhcd ;a,gnin,t Arri.can face: f11ts "1bgi•nu'"Fmu:&al in South :u KAl•PMtUIH'lR!t~, ._.,, n ' TtM, (.;,J,-> \.HUI l,.\l,. !U)01. The\.r\'illuatlon o:

\frka.. Pux~ft.li,,gs a{,;,e 13Jh Wotld Ci.mgr~ss t>n D~..1..~.s ofQml~. ~tkJ..}' uup'II ..1nd 1hrfr m1J.1fU\'t•zm'nt .o .s 111t>1htcufRg toul fur GWJr.a
Dwban.198;,,1~2-i.;i. mASttm llnd Ci b1",1rx1lpts Oiptcra; Clo,.,;lnidael inSou,hA!nc..,.
Unp11ol«hod d>1•
ft. OVSSt\llllr., s,i.. .!'>OU>.'UPU\'A,-' • GRCC.01\, •• ,:. ):JU(Cf , 1982. Tht> rolr o!
22. i..uvsuA~~. , . •.;i.. 1fttl7. \Jui.ta ,,.•,,J//i')p. nc>v. rUJptcra: \tu011.ddac , n
llydrarnra nrmrp~ F"-11, Diptt'Q: ~hl~.cidn.<' in 1he t.nnimlssmn of
inrectious bcn1nc kNa1oconJunc1M,Js. l'o//11 PoriJSltolo/1,l('O. 1Pmhal. 29. dun11-br.·1:dlng 11) Crum ~Omli .\frl,.1 0111frrrttf)O<,rt Joomn/ of

:9-83. Vf"tt"riru,,,., ft.-...mr~·h, ~4. l !&-l Hf,

~ U')()~Jt'-. r .t:. 198:1 Commu,1 E.xtemnl Jlt1rnslti.•$1>nd l'estsuf Ut..wrocl.: um/
s on Ton "" 19l8. The horn ny tl..)·1,..,o.<11111111111w. In South \ir(et1. Joum«I
of ,h~ St1uh": ~v,1,,wV111ennnry MC'ditYtl~\ssa('UUif'Jn. 9 136-143. 1'011/rr)·lll CJJlifornw Un!l'cNl) ofCallfornl:t. !>•vis, o~,,._
Cl!Ufornlo.
2.1 \11:.IH.Rt, D.\\ ,. ~.\ntll!,O'.\ A.,.,,, \\'111T1WI O, I .. I.A•IIRlCQIJ[., Q.t:. ~
10 U.7Jsc.,. R..J, « erRor,il~ , ff.. 1936. l.')l.>eJJa.r modiflc,umn~ af muscomorpha
wt mH.\.U, tu· , 19;tt. Unique ~1Unt'.1am•1nmc.,m ~~;stem to c:on:rohtable
Ui(-S tDJptcral.AmmU 1,>/1hr EnromofQg,cal $oic'1try• f1f.o\mffirll, 79,
1 ~()...209. tl)· populdtJ01ts. /fmrn11l ojErounmtt.• Emomo/010·, "i I. 290-29?.
2.5 , 1u.L<>n. p~ m• ut:-.:1, 1t1•.• \\ILKS~~,. 1•.1.. 198;. 1\l«bauic-31
4
U t;l!J\HAJU;)T, ft.R,, ,\I.LL', I,\\., ORl:I SI'. \\'Jf. 4o ,,tmi P,C., 19:8~ The role-or
,r.ant.1m:,..~um of Cl'lpnpox ,,rus. and Afnran ,..,,.int rc,•e, \1rJ\ bi'
face tli<-, an an epssodc Qf miecnou,. ht.,;.·im: ltm1tocooJ\lnnhiH... Jom,~ot
5rom1JXY$ ,ntnrrm,s. Rt1~1rtlr iu \ i•:,-·rtnary SdNU~. ,j,3. l~l 12.
of lhf' .,\ru,m,·,m \ f.•11·mml)' Mt·rlrr,,I Ass<k"I.Dliou 180. I ~6-£S9
26 ,uHU>,;., :ttx.H 111c ~/J 1 r..ip- .t .. irnp?,· cnvirunmc.·nH.r..cndly doth
l:t ,r,t..~. n ...... .r.. C'.ll\st.,11u,, k-ll ml:k. Tr:an,ml~1onof ,\ltir,,A,•/ln lm1•Uby
trap <n, IIUing OIL-.. th,, d<>t', no:"" lnsecucid,., .\,-;i!Lwl~ from: URL:
rcgurgl1otlo~ rrorn 1he <rop of1he foe~ Oy (Olpt=· ~1u.cldatl./011m,,/
lmp:tlinfomuuia.,npl .c,4! n:l.lhul,•.T.l1tm (Hus.tcd mthe lmematlon'1J
o/Economi, Emnmolog;. -;-;- 399-101.
Centre of Ins«! Phy,11,loJ;)' ond c<ology, Nairobi. !-enra
13 C.JU.l.~1tor.. h ltil, F1il">nnd OisNu•• Vol, 1. Ea>!tJ~~. Clt1.~.n1trmton. tmd
~ MO~"n r .. 1972 ·\ rt.'..1C\\ ofrhr.--,;abk-lly p1ob~um in \f.turitiu,. lt1R1'1:o:Jt
8/orlr /i.J.sorlmfrmf- Prinre:1""· ~t•\, Jt:rst.·~-: Pnnce1on Uni\er.;11, Pre,.> 1l[ll(tt,lt• .-: Sutrf,...,, tf,• I'll..- :.Jt1uru 1•. S l 13-29.
,~ Giu:L\BEwr., u., ,m. i-11,~ 11,rtf ms,,1.1,.., Vol. II. ftlo/Of,)· tmd Vl."'1.0f 28 :,.r, llJ.. L ,, .. J!f8o. StudJ~ on Por,1/Uuriu J)O:•k'Olt1 Tub,3ngul Jn SOuth
Tran.smis:stoi:. Prlnceiun. New Jc™')' Princeton Untvc.-11:y Pr~,. Artie.a \,'1th panicuhtr tt."forcncl.! 10 tht" soft, pl:ayt.•d ~ fn11,1.,c1.. iTI :h,
15 H.W. !lO. 19&.1 Rclatlon<hip of tho r~co flv (Olptffll· ~1u....-,dae LO tran:.snl.s..~1,m ind dbrnhuunn. D.SC". th..-Ji!\, Un.lter,Jty "' Prl'lorf.i.

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86 ''", ""•N: A~pccts inOurnC'ing th~ ncrurrcncc oi inftctious disease,

09 ,n,u. t-~1 •• t98:!. Ale,; o! ,'C<crinnrv lmpononcc on c:ittlc. Prc1<w1//11F;t of rltt Dip'"" oft/11• ,-Vt1>trovtmf H,>;,~M London: Bnn1h ~lu«,um ,au,ral
Su,:po,mrr. (Jt1 tcW/JtlttlSiti!S. ofrt.lltli.•. 15-16 ~I.arch. 19~. Ptttoria pp. IU«.10~ ••
86-90. 43 1'Q1<.o1ft1 1' t .T.. ..ur111 •L\-.;t>, e." 'Ph.G~. H.c•• 1981. \uc1r.p;, tn UoUl>rnil
30 '-t \•11,, t--'1 .. 1985, Tht• cph!cmm1o~ o( /'11r,1/JfarJt1 l1tu·:rola 1n 1he .~11,,pln.m1a rr:arginnlt• \,ich f/fpfJ"I~,, ntfip.:~ .1nd .,mm..,~-, ,ak,rruns.
1 r.m,\.1;d 8u1,h\'eld orSclutl1 \!rico o,ulrttt•'lJOQrt Jour,uit oj\ Nrnnlln CJrrdrf3.if'JX)Qrt /owunlofltNtrumry HcrHnrrh. 48, t !~l~
Rt1••r1n.•lt. S:!. Z01 ·20i ..q ~IUu.\nt. r., .. C..A.\JJl8.UJ.. J.U.... uuuso~. D..lL, IUSB'-,C."il •. \VIOU lt.G.£
31 "1 v1u. t ,, :Jno1, Ond~htC)?fl()rt V(.'h:rlnnt1 lrhdtutr~ ~onth Afrlnl. <;L\ \:101". o.(:.. 19:"9- ·\bituy of t1,c face tl)" to <"amt.• d::unngt> co t..')t""' 01
u11publl<J1cd d,1ta. c;,n1tl. /mu11nl OJ F.cmtamic F.nwwola.t()~ :":?, 6l3-63S.
32 ,t\'11..L l.\1. mcc.s.,u.c &L\.."<GC.,"HO\'E..~#1.w 1980 011dcn1~pnon .is SOi.. ,.. 1~ Cuntro1 nw1hml-. m ~ummtr- mA.Stiu,-: Jlw Jmp«,HJnt~ uf Hy
Vi"tcrlnru,,· ln~tli\.ltl" So\lth \frkn. unpubl~l"J dat.a ,,j
t'(»mror Procrrdin~ 1lw 13th World (",<,r,grc•$s rm D1sea1t!so/C.t111le,
l.l11rban. 198-l, 23ti-!~2.
33 ,r,·1u t ,,. ,. '\1H>n-f,u .,,P. 0... 19.87. Th<" cv!o:u.auon..a.nd lift,ryclcs.or
,\/tLita tusoria. \fWL'tl xanthomclos .md .\lusrn Jlt"t'llli, \'l"CI01'f.Of 16 Mnt.'h,, c.n., lit:b, fftt."i:h ur 1.~h:rnaJ and lmem.:il ~\nhro11enf p,,,.J.,.i-u:"
Pnmfilnrit, bc>t i<'o!n m Son th Afnc.t. o,ulMh.•poort Jmmml of\ 'tterJnnl)' on dnmr ..tk 11,.t-"tod. µ:oduaion. lnmtt,/ Rt'i'lt.·~·o/l.'l'f!<>nw!~\:? !
Rt....kVlf(h, 54. f,0;'...6,11.
! 5:t-1';&,

l4 "'t:1.10.. t . ,,. '-~,,.>\UT, ,., .• 1917. ()ndcrs.tepoort. \'t."tt'nnory lrumute. soutJ, ruut,. 1M, .. 1~18;. Cnttl..l'- rl~· comroJ UiinJ! .;ont.mtlc:d,reh:·4."1.' Ubtt"t<Clde?S.
.::i
\ i!lc,.'nlUII}' ] 11/fdJttfJ!t>g>,. 18. ~'?J.;I •
..\Inca, unpubU\hed dat:a. •
;8 11tn,?-S.C\, , ••. lJQUD.l D-\S &flA,\t~. l,.LO., ,.us.u,.;~J ,, u.. ... IIL\:GI' ax...
35, ~Fvtu.. f ,,. ~. u !\,·En. r..1 .• 199lt Ondl•hit:pomt \'t:tvrin;sn· tn~Ututc.
1988 ruiur,,..af 1/atmauH..Jiu rhfn,u:ri 111)ratu fBe1.:u1 to t:-atlbmit
S0111h Mrlc•. unpubli:mLod dRta
(c>t>t•,mU•mou1h d, ....-.i..c ,·iru-, Jnl'Ch:micnHr bt1wcen \1t,,u•rnk .md
36 sf1\"l11 1:...~1 •• WH.Ji:1s .s. C"-1\, It ::1.At..:K.1~"\IO~, r..• J')H;, th~ comrul of Parnfilnr,a
Ml-"'-CJltlb1~1.'"~ttle. Otuln-,t,·~,t: /nunwl of V~1~irt11? P.t.,,•un.·h .·,:;,
bo111eo!o uansuu1>s1oa in Suutb Afntu.. Omf1•rs1,•p0Qrt /<mrnal 11/
l~l-122
Ver.~rmary Rl:SNVdr, fi.t, ~H7-550.
49 \'AS (lf;f\1, t. &.· Rkot 1 , .tt . 14)85. Sigi1lnt:\nce of C,lltl<' di.srhMg<'s :md
a; ,.um"' ... , .. ,·t:ru.nR. A.1.,1. • J,\r..oai. r.r. •J9fJ:t !so\'inc ~fn lc1,fon~
st'\\
~L"Crt11ion, n, pr,,win ',(Jlllt't·" for O\':trlnn dc\'tolopmc.-nr In 1h,: f.H't.' tJ~
or pos,iblu lilarinl origin U$>0ela1ed with ltea11· hum fi) infe,1.,11,,.,. •Oipt1:r£ ~t~jd:,t, Emfromnemnl Emom"ltJtJ:>·. 14, 60--64
( Hu~11u11r,l1tt1 m,•t1llitmtJ). Omkn1..1roar1 burnnt (): ~ 'rtrrllltu;,· H,·H"tirrh,
5,, VA'.\ r,11 ,1 T ,., 01ton,.,,9,, 19SU. Flue10;.,tfon ... 1n the prokln .and
50, 73-73.
tiUbolwd~te tQntrn1 c,f thl· crop related lo periodlcitl~ in O\'Tln;an
Jll ~1r~-1tJ1..t. o." ou 1'nn. IL, l9l3, Thr 1'ltturr,ml'e r,t Bdttl!ultu/1" d('\,•Jurnient ul Ille (cn,;ile lft<'C ii\ \l.llpten,.; Mu"'1d:w ..~nunli nf tit,•
11nts..,11mu '.\.lnllO<"h, • blood-,-urklng fly. m 1h,· Tr•n" oi.ll, /utmuil ~Itit, F.momd,>.:,rwal :itX·1t·t): o{Amrrla,. n). 1-6.
South Ajrir,m Vftl'ntta,y V/ed{eol.k.:.qx-fmum. i , 2?1-:tu.
,.;1\\'Al.);f•. , •• 191)~. ·rh~ .snh:opoch, oJ human~ .1nd <lQnie~ltL Jnim.,L, - .1
39 r.,n.nso~. u.l •• 1~1. Population gont-lir .®di~,. in .1rea, of O\'Crtap or l,'lllde •• J>rrlm11n,\l') ldrnlitirntlon. l.ond,m: Chapman l. !l.,ll.
t\\r),uh,pect~ of .\/Ug:;J d.t>mt-stlm L In: Dw1, n.11 ....... :l"Ci l. Fcotogk11J 5:J wu:,..,,, .._i;.. 19:tttC Ulmp~ ,ktn d1sca.s~· ,in.h l 1n,/r.,gy Mr,rt1J_:/4[Jh1. 3
Strttlfts /11 Sow/:m, ,l{rien••l/01101,,rn1ml111• Hlofug;·. 11w Hagu~: \\'. Junk. 111-13:.
,VI PA'' ' u...,w, H r 1gft1j: llnl\"("r;.:1ty 01 Q11t•,•nJond .\u,,r,1h:-i ;.l(H"\t'IO.;\f 53 \\U'1. 1......... 1951 l'ltr Jlo1ut•/l>· lth11co1, Sew York: Con,:,1oi:k l'ubll:dun};
oommunu;atitm. Comp,ul) Inc .
•u P\TUt...:o~. If.I;;. & ...;nft.RIS. K.k,, 1970. rhe Mu:Jeh u,rlNIIS C'ClrllJ'lt"x: The 5.; S:irl..y uop~ ror !):Offlpling popt.1!a11on,;, of Stmn,,.~.,.
\\11.l , ... ,:,.. o.F •• lt,73
rrla1wc- itJ;tu~ or tht! \u:).traltan and t\\'u African r,npulaih1n.s. .~.\Uull(m ,nlc-itrun>.. /cwrnu; rJ/ t·,mu,mu; F.nWm(JfCJR}', 66. 1z;ll··l?80.
/1J1m1<dn/7.(}<J/1,g.. l8.~l-24'i :» 1.U\tflf. i-... 19':'J, n,., !!,to1m,,:,.i,tr.• Biting Fll1.'1$ o/1l1t h'r1tt(( ",tuHjt,.Ut:
-1:2 JoO'.'i t. ,.c 1.c.ao. ~umil> ~h1$ddne,. /u: cnl)s...-.,a.:n. 11.,,., ,"1 1. Ca1,1/r,,z.11rof liu"'lJ\' f 1schrt \'N!,1~

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 4
Vectors: Tabanidae

EM NEVILL, R J PHELPS AND BR STUCKE;\'BERG

Introduction ln soulhern t\l"rica. Chainey and Oldroyd 10 summarized the

larval biologyofthe~e flies vervsuccinclly as follows:
The reason for the inclusion of a chapter on mes of the fam-
11) T.tbanidae in this book is their great potential as me- Tabanicl larvae live in wet places but need 10 reach the
chanical rransminers of infectious disease agencs. atmosphere periodically as they breathe air 1hrough a
Tabanidae are commonl)' known as horse flies. deer mes. posterior siphon. lhe usual habitat is the wet mud at the
clegs. and, in South Africa, as blinde\•llei! (Afrik. (blind mes). margins of ponds. lakes and $treams. but t11ose of some
The large size and \'lU'iety of genera and species of Taban- species li\'e in sand on the seashore. Some Tabmws lar-
idae in southern Africa should have made thl'm an obvious ntc inhabit pockets of damp so'il in generally dry areas,
choice for stud) b)' biologists, but studies on them have and llaemmop<Jtn ian11e occur mainly in areas of wet
been moslly restricted 10 Lhcir taxonomy while those on soil. Tree-hole hrl"eding occurs In a few Tabanidae: lar-
their biology. role in disease transmi!.s!on and control are vae of Tflrwmnsrocern for example living in the accumu-
limited. lated debris in rot-holes of palms and forest trees ...
larvae of flnemacopow and Tnbwms feed on other
Biosystematics insects. crusrncea. worms or their own kind. but many
iabanid larvae. such as those of Cluysops. feed on
The taxonomy of the Tabanidae of the Afrotropical Region \'egetable debris.
embrace, three subfamilies (Pangoniinae. Chl)•sopsinae
and labanlnae) containing 31 genera and 727 species.q· io. Adult tabanid Oles are generali)' lnrge. usually ,,ith \\'CU·
39
l,- Four hundred and ten of these species occur in south· developed oycs. Their momhpans are adapted for both
ern ..\frica, and ofihem about 80 percent belong 10 only live sucking and lapping, with both sexes feeding on nectar. The
genera. namely IJaema1opoU1. 1'al)a1111s. Philo/it:lre. Rhigio• females of most species also feed 011 blood. which is needed
glossa C= ;\lesrmz,vim, and Cluysops. for the de\'elepment of the ovaries. The mandibles and max-
Flies belonging to the Pangoniinae are usually of little illa are serrated and are used to cut into the skin and lacerate
known economic lmporiance.30 Variou~ spc('ies of Philoli- the dermal tis~ues, and thus induce the formation of a pool
ch,:, of Llie subfamily Pangooiinao are. howe\·er. persistent of blood which is then lapped up. A small amoum of blood is
pursuers of blood meals from cattle and horse;, in localized trapped within 1he component pans of the mouthpart~
situauons.46 Of the Tabnnidac recorded from sou1hern /\f· when the)' are withdra\\'n. Mechanical uansmission of dis-
rica, three genera of the Chrysopsinae, namely Rhigioglossa ease organisms is thus facililated by 1his action. The coarse
(39 species). Chry.<ops (23 species) and rabanocel/a (10 spe- structure of 1he rnouthparcs and the mode or feeding result~
cies): and five ge11era of the subfamily Tabaninae. namely in painful biles cnusing cattle to 1ry 10 dislodge or kill the
Haemcuopora (13i species). T(l/}(mus (68 specie,;), Hybomi· °
flies by striking them \1ith their 10ngucs.~· 3 Funher details
tra (pine ,pccies), Ary/oms (sLx specie,;) and A11C'nffl (tour on lhe life C}ci~ :ind blonomic:s of Tnbanidae can be ob-
species] appear in a list of tabanid-transmitted disease tained from a chapter in the IJook by KeHle ..io
agems.?.? The densiry of tabanid popu.lations is closely as~ociatcd
The truwnomy of tabanids of ,\ngola, 1' 16 Mo1.am- with breeding site~. Popularions around dam~. \'lei,. and
hique, 1~ Zimbahwe2• and South Afric.i~" has been reviewed. narural wecland" may be !ugh and composed of only n f(:w
In general, the biology of tho Tabanidae ha~ been poorly species or perhaps e,·en a single dominant one. which in
studied throughout the ll'Orld, but thi~ is e\·en more the case South Africa is often a ~pecie~ l:lf Uttema1opotn.
87

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88 ,>.CTio, ,,,,: A<pccis 1nnuenrins the occurrence ofinfectfou~ diseases

In ~0111hern A[rica. 1he breeding site:.of many 1abanids i11 (e.g. l-le1t!111ncopoca ,pp.J may achieve high de113i1ie3 wl)ere
:,.1ala11i'n 35 and the larval feeding habit; of Tabm114S 1hcrc arc ~uitable breeding sites:"• Wooded areas. or
bigwrruus \Vied. nnd ,1ma11ell1' emergens Oldroyd in South patcht", o: indigenous torest. favour a greater diversh:y of ca-
\frica8 have been recorded, but published information on banid specie, :ind may harbour den~e populauons of these
the biology of adult$ Clf specie,. of the .\fro1ropical Region b me~ :\ common a~mblagc nf fli<!S ma) consist for ex-
rather sparse and deal:-. mainl) 1,ith tligltt seasons. abun- ample. ol speLles of the genera Philolidie. Jfaemaropora.
dance and nocturnal ac1ivi1y. 111• ~8 It b considered that cer- Tab,1110..-e/l(I and pcrhap, 11,rumellu, and the component
tain phy,;ic:al adapmtion~ aimed at di5rracting the am:mions species "111 l'ary from locality 10 localit) . Members of ~uch a
of 1isually orientated hiring mes ham evolved in some ,\fri- iuild of species aggregate around hosts that are potential
can "11dlife ~pecle~. •· ii has been demonstrated in the sources or blood m.:als; 5uch hosis ma) range in si.7.e from
north-eastern KwaZulu-Natal Province orsoutb Africa 1ha1 small antelope 10 cattlt!, and the flies wiU folio\\ them with a
1he StTiping of zebra mnkes 1hem I lnuatly undetectable co marked clegre<> of pers1scence. Movement of the host omside
1se1se me,. ((;lossin11 spp.) and 1ab~nids.aa Field observa- the fores! or woodland resulls in selecche 11~thdrawal of
tions also indicate 1hai ant<'lope species 11ith a coumer- tho~e tabanid $pec!es 1ha1 arc unwilling to remain in open
shaded haircoa1 panern. charactl·riied particularly by a pale areas in full sunligl11.
belly that ellminatcs chc roundcd appearance of the abdo- \!any 1abanid~ hal'e preferred feeding sites on animals;
men which is enhanced by shallowing in sunligh 1. are evi- lht! ,mailer 1abanid species tend 10 <ettle on tl1e ltigs,
dently unanracdvc to rabanids.4;; In contrast mammals whereas the large species usually seek blood meals on
with unifonn colouring. such as elephant. rhinoceros and 1he sidl"> of the bodr or on 1he flanks. This beha\·iour ha~
hippopotamus, which present a strongly rounded appear- been interpreted a" a fonn of resource panitionlng through
ance in sunliglu. are highl)' auracti\'e to large cabanid spe- which interspecific competition for feedini ~i1es may be
cies such as Tabmws 1ae11iolo P. de B. and Philolic/re reduced. This aspect hm, 1101 heen studied extenshely in
at>tlliopica Thunberg. experimental e\idence has shown Africa. apart from obsen·ations made in Uganda.:!! and
rhat a dark. single colour. hori?.ontal cylinder is a most at· 7.imbabwe.4 l
tract!l'e shape for tabanids. and indeed the Harris Fir Trap :-..1os1 1abanid species arc active as ad,llt flies onh during
represents essentially such a form.w rhe rainy '\Cason \lass emergence of the mes may occur
Tabanidae are unequally distribured in Somh Africa.'~ In with 1hc advent of 1he firs1 rains. or in meteorological condi·
general. many .pec[e::. or the rich ,avannah and tropical lions in whkh ,uch raln Is pol.'Sible. Consequently. in 1he
woodland labanid fauna of 1\Irica occur in the easrern winier-rainfall area of South Africa 1abamd~ occur com-
coastal lowlands as far south as the Eastern Cape Provinc.-e, monly from hue winter 10 early summer. wherea,. in sum-
and also range 10 a much more limited e.xten1 into the sub- mer-rainfaU region~ their occurrence may be highly
arid Limpopo \'alky and onto Lhc H lghveld. In the southern correlated wi1h 1lw min) ~eason of ~ovember 10 ~larch.
and south-1\·e~tern pans of che V,estern Cape Province a Some species nHI) per"ist in low numben; throughout che
1n.xonomically very dis1lnc11abanid fauna is characteristic of year in lowland area~ \,1thou1 frost, and a ie11 fly only in the
the macchia fynbos) and 1he fold mouncalns. as ,,·ell as lhc df) ,;eason. In hot climaws. some species can be seen drink-
adjacent Karroid areas. lhcse 1wo major fauna! units ha1·e ing ;11 water sources such as the edges of natural pools; ic has
few species in common. 18 been observed in the Kruger National Park in South Africa
Regions wflh a high rainfall have the highes1 number of that cememed water troughs provided for game are con-
speciel'. Topographically diversified areas also have a more siamly 1·isitcd by specu?s or 1 abanidae In drought years
diVerse tabanid fauna. Consequently 1he plau,au slopes, 1hc \,hen namral watl'r ~1,urct:S arc scarce. Rearing record,
.Mpumalanga-Limpopo I.owveld. the coas1al plain of Kwa- sho,, that males and females occur in equal numbers. bul
Zulu-::-:acal, the Eastern Cape Province. and the southern the male, may seldom be seen a, they do not take blood
and south -western J)llrl nf1he Western Cape Pro\'ince have a rneab and '\Oare not a11rac1cd to mammals and only rarely
much richer 1abanid fauna than do the interior plateau 10 iraps. 11 ' ' io
areas and the arid western and northern re1,oions ofSoULh Af- Large 1ahanid!> mainly au.ack large mam:nals (ca ttle,
rica. iH horse:., large antelope. elephant) and their presence ma} af-
A large proportion of species occurring in rhe northern fecl or modify behavioural patterns of animals_ Th<! blood-
and eastern lowlands of South Afnca is shared with iMozam- sucking ac-1i\•ily of three common largespedes oflabanidae
bique. Zimbabwe. 7.ambi;1 and Angola. (Tnlx11111s l>iguru1111$, T. 1mmiola and ,.\111:0/1, africa11a Cra)'
Certain species dominate in part:cular areas in souihi:-rn (Figure 4.1' have an Important influence on rhe summer
Africa. In Souch1\frican sand-dune areas along the southern diurnal ncthity 01 hippop111ami in nonh·eas1ern Kwalulu-
and 1\'estem coast of 1hc Western C.ape Pro,·ince. a dis1inc- 1\aial.1 ' Because or the pre~ence of large numbers oi these
1ive 1abahid fauna ls e1'ident. wilh Rhigioglo;sa edemula flies in conjunction wirh high solar mdiaLion, hippopotami
Wied. being particularly evident}'1 The grassland plains of are obliged to remain more or less ~ubmerged in tht> water
the Higlweld support a limited fauna. though some species during daylight hours.

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 YCCl(}l'S: Tabanida~ 89

,..

Figure 4.1 Ancela afr,cana

Disease transmission susceptible animal. ' 7-:1.2, 211 • 53-SG Transmission is facilitated
by the fact that 1101 onl) are they generally large insecb and
1bc Infectious diseases or agents ,,;th which horseflies their presence wonying to animals, but also that their bites
have been implicated as pla)ing a role in their crnnsmission are painful. Feeding is Lherefore often imerrupced and dis-
are ~ummarized In Table ~-I. The method of transmission persal 10 alternate hoMs frequent. The factor~ which innu-
may be either biological, which is uncommon. or mech,mical. ence mechanical transmission are: 1'
Only two disease- producing micro-organisms arc • the survival time of the pathogen ou1sidc the host:
known to be transmi11ed biologically bi· rabanid mes (Table • the proximit) of infected and susceptible hos1s:
4. 1). Sali\'ary gland fnfcaions of Haemopro1eus me1clmiko1ri • vec1or mobilil!,,~
are known to occur in 1he deerfly (Cluysops callidus 0.S.) • persistence of reeding the less persis1em feeders are
aftt'r fol'dingon u1r11£..,_ i; The other is Trypanosoma 1h<'ileri probnbly mo~t important in mechanic3l transmission):
which causes a benign infection in caule. The resuhs of • vecror size (the larger tabanids would appear 10 oe be11er
carefull~ conducied trials in Germany suggest that trans- transmiuers of palliogensl;
mission in the ficld takes place l>y contaminarion of the oral • the total tabanid population (this appears to be the mo$t
mucosa of cattle \\ith metacyclid stages from gut conte11ts or consistent phenomenon associa1ed \,ith peaks of lnfec-
faeces of tabanids. when the canle defend themsel"es and 1ion);
kill feeding tabanids with their tongues.. • painful bites lhh. can be directly correlated with disease
Ofinterest b that although both sallvarian and stercorar- 1ransmission):
ian ll)'Panosomes undergo development in cenain insects. • the host rllSponse to the bite of tabanids may be dimin-
infocrion rate.s of :.alivarian trypanosornel> In the \·ector are ished during the acute stage of some infectious disease,;
often low, whereasstercorarian trypanosome infection rates and cherefore the early febrile periods of chronic and
mny approach 100 per ccnt.:io acut<.· infeaions may be the critical pcriod5 formmsmis·
ln addit ion to infectious agents, various illarial 1,orms sion as pathogen titres in the blood of the host arc often
de~elop in. and are Lrausmin ed by. ~pecie~ of tabanids highest then;
throughout the \\'Orld (Table 4 . 1). • die quanticy of lnf~tious material mmspor.ed b)' indl·
'fabanids are the insect group most frequently associated ,,idual vectors between hosts;
with the mechanical transmission of pathogens-a result of • the amount of Infectious material depo:.ite<l at a \\'ound
a fly feeding successively on an infecti\'C host and then on a ponal of enuy;

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90 ru:no, o•;i: Aspeels intluentin~ the occurr~nc~ of lnfocuou,, cllseas~

Table 4.1 O,s:Jase agems associated w,th taba~:;1s'' ?.c cem parasi1a0mia of ,\naplasma marginafe for 50 da)'S
AGENT ASSOCIATION ma) be equi\-alcm. as a source of infection. 10 an 11nimal
with a parasirncmia of 50 per cent for one day):
VIRUSES
Bov:~e leukaemia ET • the densi1y and age of the host and
Cahfomia ence::,hahtis • prevalence oi in!ec1ion and degree or immunity in a ho~1
Jameslc,·,r. Can·,on sarotype I population.
taCrcsse sl!ror;pe I
Equine inlectious anaemia ET. Nf
Hog ch1>lera I, :i Ni Tabanids possibly played a role in the mechanical trammi, ·
lnt1uema EE sio,1 of na·gana {ttypnnosomosisl du ting the major epidem-
R,noerpes, cT :,,:r ics in nonh·t·as1em KwaZulu-Natal. 3· ;;u Spor.:1dic oulhreaks
TiC<·borne encephal1t1s I
Ves,tuiar s:oir.a1.;,s i:i of the disease still occur in this part of KwaZulu-,atal.0 and
Wes,em eq~1re encephalitis I biting flies such as tabanids and horn flies (HaemC1Wbia
spp.J may well play a role in it~ lransmission..ffi
BACm\lA ANO RICKETTSIAS
Anaplasma m?.rgmele H.Ni in southern Africa the only recorded cxperimemal work
Becillus anmrac,s I. E'T, NT on the mechanical transmission by tabanids ofa pathogen is
81ucelle abotcus ET that done on 8esnoi1ia besnoiti.' The tlics that successfully
Bruce/1/J mill,;ens,s ET
Broce/la suis ET transmitted thl' parasite b} bite from cauie to rabbit and
CIO.Sllidium r~auvoei ET from cattle to cattle ,,ere Alylows nigromaculams Rrcardo.
Closuilfium PErftingens EE Tabanocella d,mticomis Wied. and l-/1umu11opora albihirm
Cox1e//a burnet,, I
Er;s1palo1hrix rilusiopa//1iae Karsch. the only tahanid species which could be capmred In
Ei
francise/1;; tu/arensis I, ET. f'..'T sufficient numbers m the On<lcrstcpoo11 Veterinary lnsti·
Fusobacterium necrophorum EE tute. In three instances only a single Oy was nece.,;sary to
lis:eria ma.1aq1ogeni!s ti Lransmit 8. be-$110i1i. Flie~ remained infecrive for up LO
&rtelia blirgdotfe,7 I
!'asuwrelle mul1oc1da :,; 24 hours:'
chrhcf'Ja ,1';;.•cii .. Cenain large species, notably T<1ba111L.< i,umiola and
Plli/oliclze aetlriopica. are uninhibited by iarge expanses of
PROTOZOA
BabEsia ovar? EE open terrain and are capable of sustained, powerful flight.
&.nc,tra /Jesrv1111 ET.Ni Their ofren noisy lliglu. persistem attendance In large num-
Haemup:.1/eJJS metchmkov, 0.1, co, B bers and painful biles may induce re~Llessness in dome,ric
Tr;panoscma rftgilen 0.1. Ei. i'.;i, S
Trypanosoma evens, ET.N- stock.'"
,rvpancsoma erjUl/1i!lfium e- In ~outhern Africa no wotk has been done on 1hee\'alua·
i se:se·oor~e ll)'Oanosomes tion of economic losses due 10 tabanid bites. This would be
Tr1panosoma v!vax EE difiicuh as lhe appearance of a large number of these flies is
Tr1(1imoscma congotense ET
i()panosoma simtae EE ofwn localized. sporadic and can occur simuhaneou~I)' ,,ith
T1vp2.,csoma tpucei bruce, H othar fly genera. in 1965 the bites of horse flie, led to cattle
ir;pancsomif b:uce, gamb:ense losses in the US,\ amounung to 30 million do'.lars from re-
lmodesiem;e sirainl :1 duced weight gain and JO million dollars from reduced milk
HE! MINTHS production. ;i Cows treated with synergized p)Tethrins
loalo., I, O.ET NT. 8 showed a significant increase in butter fat produc1ion. and
D1rofilaria re;iens 0 (e~p.1 B
Oiroiliaria 1aemsti 1,0.Ei, B similarly lrcated bel'f cattle gained 9 co Ill kg more than un-
E!aeopt,ore scMe,ce11 1,0, Ei 8 treated cattle. over a 38-day period.'

8 "' b,o!l)gica! transmission Control

0 = ae,aelopment of organism observed ,n tatian,d
EE : epidemiological evidence Mechods or comrol generaUy encompass either the use or
ET = e~rimemal transnuss,on in unna:urat manne· baited or unbaited traps or insccticlde-,rcatcd anlmnls.
NT " natura machan,cat trans:nrssion However. no concerted effort is made to apply the,e meth·
l~•P I ~ in eioenmentallv infected fhei ods in southern Africa and there is also no published work
t = ,soiat,c.~ !rom wild tallanicls on the comrol of mhanids in the subcominem
Srudie.~ in Zimbabwe on the react ions of tabanid~ to mo-
bile and stationary baits, with and withom human and ox
• the titre of the pnthogen in the blood of the inrecred host; odour, ~how('d that, although od(lur d id entice more Oil!~ to
• seasonal changes and the persistence of infectious station~ balls. the main a1traclion was \~sua1.•t 11 has been
agents in lhe circl!lation of the mammalian host {e.g. shoM1 in Zimbabwe 1h01 l-octcn-3,ol und .J-mechylphcnot.
under constam vector pressure an animal with a I per which are kno\\'n anractants for some Clossi11a spp. ,~. ;;o are

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 V~tors: Tahnnidat' 91

s1rongl~ amactl\ e 10 Tabamis p11//11lus Austen and T. rope- more hos1 species. In OklahQma, USA. it has been shO\\'ll
mani Au~ten. and possibly a1tracth·e 10 several other spe- that Tabanri$ a/Jacco, Philip is capable of flying a1 le,ist
cies. ' 0 The indications are lhat ll may be po:-sible to develop 2.4 km and that the e,1imated number or host-seeking fe-
odour-baited traps to control some tabanids. males per hec1ar,:, can exceed 2 000 mes per day. 11• 11 I low-
Control ofadulttahanids in other parts of rho world is not ever. a noticeable decrease in horse Oy numbers. mostly
commonly practised. Various approaches have, however. 'fil/.la1111s 11igro11iuaws Macq .. resul ted when 300 10 ,30 box
been tested. Because of their visual attraction to balls. use traps were placed around chc perimeters of Cape Cod salt
has been made of Manning or box 1raps. :-'lanitoba or marshes in the USA.~2
canopy traps, and sticJ..-y traps.'· 1~ 3z 1,i1h on\ithou1 the ad- Since both male and female tabanid, 1,Jsi1 no\\'ers for
dition of carbon dioxide. In Georgia. USA, the effectiveness nectar, they will pick up pollen and so may be pollinators for
of the canopy trap in capturing female l abanidae increased some plants.. \ potential side-effect of the widespread use of
significantly when l -oc1en -3-ol was used a;; an aurac1ant.i~ odour-bal1cd. lnsecricidc-treated targets and traps to con-
The success of these uaps often dep~nds on their being trol rscrse flies might be to reduce tabanid popul::nions. as it
placed in unobstrncted flyways such as footpaths or the has bec11 shown that a1 leas1 some species or tabanid~ are
edge of a forest.'12 strongly artrac1ed 10 such targets and traps. 10 This tapic was
Another approach ro tabanid control is the use of re- lnvcstigared at Rekomitjie Research Sration in Zimbahwe.31
sidual insec1idde, on animals. Permethrin is lethal to horse By compr.uin~ pollen tak!:11 from tabar,ids 1\ith 1ha1 from
flies and persls1s on Lreared animals for about lWO weeks. z. 27 plamo available at rhe same lime, 1t \\'a$ dcmon~trated that.
11 has been found 1hat 10 ppm permethrin I$ needed 10 kill at this localicy. tnbanids were not specializing on any parti·
1abanids compared with only 2 ppm for stable flies. z; cular plant specie~. In addition, direcl obsen•ation re1 enlcd
Use of 1rapping or insecticides fonhe control of tabanid~ t hat plant.~ \iS1tcd by tabanids attracted many other insect
may be wa.rrnnted under fairly intensh·e farming situations species not known to be auracted 10 anti-tsetse de\'ices. At
but. under ex1ensi\'e conditions. their control may not be Rekomitjie, an} reduction in tabanid numbers would 1101
succ~ful due 10 large and mobile population~ of cine or have an impact on pollination.

References
B,\Utll a. 4, WI IIU. u .. tW7,1. \"crsudw mh fll1!gen(.ilit.'1'l 7\IJ i:i ct><>,t.'*'• l.;\1, 1,, \\ntc;HT, M.f
t9S7. Might nmg1."'.: i.md d!,pf.'Th-aJacti\1tyof
Tobanldonbt-klimpfung aur dcr Wold,·. T/ernr=1/ir/r, l/msdw11. :!9. the ho!i~·'-'"et..lng h,1r~· Ry. Tuhmms ,wvrrro,•H)1ptnn~ I ab.tntd,1i:J. in
20&-210. nonhet:ntr~\l Okfal1<11n.i. Em•1mmnc11tal fntomnftJgy, !ti. 2tl-217.
2 n.n. o.t ... 1t0-,.,rr>. s.,.. ,. u,mu°'", n.L.19;li. E\Jltun:on of .t wnlhl"tk 13 (X)(•~~n. L,, ,,. \\'Mu·.u:. tu , 1~9 ropulnuon t."'.\t1m.11lfl11 nf tht· hom 0\',
p)><ihrotd for 1,\banld control on hQrM'\ und u1tlo. Tht Sor11/111.,·r,•r11 Tn/la1111111b"""' (Olp1,.,., Tab•nldae,, ln nonhcon:in! OIJJhom.s
lit111J11wlo1;i.11. I, 198--203. /ourmdo[.\J.'lllea/ Entomt>lu~·. 26. lli7-Jf2
J 1u.:0110Ro. r..-\...11,. :g;:: Repon on rhc uan:,nt1s,i<Q11 of'\;'a)!.trua in ih~ ti! otA.o;;.1.A TM.\\ '-'-.!-11!>~,'\'.10~. t960. :"\o,i:t cantt1buicnouu 1$1\Jdu do,
Xt,1ban3.n:i ;and M.hL.1tuzc Seu.Jcrnenl~. 1..uluI,11:d. J>..\n I. Eft•t'i.•t11h ..~ taban!dt":09 ·rnptera: rub.\n!d.\e) d(! \flgoto. IJJ.,blf.:ncoe~ tullutt'ti>
n,.. ~lftit Rc1x,.,u ,,f11Je l>Jrt1cmrtJ/\i.::,.1ri,u1f}' Etlutmion 01111 IIMMrclt. 1-125
(.'umpanliitttfi• r>ftWUJllli-1. tf1,•.A11S,Oiil. ;J,
275-300. 15 m.,, 1..,. n~,\ -"'"'"0:,. ,.,,TQ). 19tW. Tabn.n!dcos (l)ipten:,,.. l ,1tlnnidJ,•, d,·
.; BtG.\I >ff. R.n., 19.u8 :'\ch' r.Qncert~ on 1hc cpidcm,o!n&ical reatun::' of M'><":trnbiq\11.•. Con1cib1.m:,11;1 JMrd o 't'l' conhc..-cin1eoto. ~'\1 l:?tS:Sf'J•.
bovmc b~•no,110,.1, a_" dut<"rntiu.:-t1 by labo1":\to:'y a.uJ fu:Jt! l..ourenco ~brqu1:1o - M.\pt.m.).
1n\'t:\tfg:,,t10n).. b,:tlctstt·JX,0,1 /ourm,I ofV.:u:rumry Jl?J(·,ud1 , l.S. 3-138, 16 or.,s. f,.,. rn,w ....,~--.,-\,su--..,&. :i.I.NM'-O. r.~ur.. l96:-.Al,tun:,, nu\oidaJo~
3 h6,1., k HtH l>HOl-T ).;.T.. OlBJUCll. :\., HlkCUUI, c•. a. llO.,lfYf'.R, I.• t96j. ~bn• o~ 1.1b-;1nldt~ t>lp1l"r.t· f;ib:u1:d.,e} d~ Anr:.ola. R,,1iJJU &uu/p,
Trnh'!>ml~~ifm tll T')))(J/1Q$1Jntfl ,11~.111,,,; to ciut~ by t .ihamd.il." ~'t·rnh UnJ,,,.,,.ttuio, dr _\/tx·f1mbiqu,.,_ •I, .Wl-S.:.10.
Puras//of<1g1· Rownr.·/1, ;a, 121-424. II £-()ll."' n., 193S, Tab~1n,d, il'- Vt'ClOTS()f d(S,(93se-~gem<- PnttlilllJ/rJ~
6 &~\\A.~. ,.i• .. 1S1)0. Trypanosam1a:s1-. 7 n.mgr,l,•1JSi.1i m SOulhAtru:·.i, Todny. s. 36·%.
O!kv d,, t, ·111un1111fcmnl dt~ EpfZQQti.~ Di~1ut 1,1.{orm,:rfmt, 3, ;s-ao, 19 fQlt, t.P AU,\,\t'•· \\ \., l''-H CI• '- J'l..hO R. 1~81, T~ba.nld Oiph•tJ
; tnmn. \\· ..,.,. OH J.Fit t,,c., l9:iL Tnb:anid cnn1rul on d.a11) omd lwi•f cattle pupuhufr,n,u...... o,:i.ttt'd \~lth ,m t1quin.c infccdou1.1 Jnc.•mUluutbr~ u, an
..,·1th ~~·nctg}lL~ p~ n:thrin, /oumal of f;c-0,:u11t1, !,,,o,m1Jui::·. 44. in;1ppartntl,• ln(L.-atrd hvrd ut hot"'~~ /ounwl of.\1edfml f:'mm,wt~·. :!1.
15-1-159, ZS-30,
8 c.,11..,s. 1.~1<'.(;... 1980 L3M:l fcedlng.habhs n! T11IX11tus lJif$1t!mus :ind tY HJII. Lo .•.,o\..,,s. "'-~ .. ,H ...u .,u:i., J.:'t.N 1~1-1 r;.1 .. 1937. s1oodm1:.Jl
:\mo,:..•lln,•mNSt>JJ,"1n S0u1h Africa tDlplcr.l.: TiJb:tnitf!JcJ. Rt.•i•ut·rlr n?':fdu,.. on 11001hpMh of1illxw11sf11sti(o#11ws (Dlprom, TabJnld~l'
Z<>IJ/agh· ..Jftiro/11, 94. 791-7!M. nnd rh" potcnll:il for 1nt!'chnnknl mmsmission or pa.thogctb./ut,r,w/ u,f
~ c1,1111-.;1·,-. r.r .• 1sa-;- \frouoplcQJ Tnb.inldo~ Diptcr,u lb\.... ,tcnu-. .\l,•dft'flf Ftu~11wln,.:.,•.1.1. Rll-6Hi
Hlu'gtoglo1.>n \\"h.-t:fonwnn 1823 (Including .\fw,myf'1 M.1eq;.rnn ltlSQ• .M 2() J.011 ,.o.• "Rf\CH. {).n .. uon Jl.{i., h5I:l., c..1 .. UPXJN,~l. o., )KM,V•V~.
2 ,ubgenus,. ,\n,111/,ofthe Nara/ M1•~1tm, 28, 137, 159 Ttan-..mts'lon ofboVi.ne leukem,a v,ru~ by
r,:,,1. ~ ~tc.H•. ~.,_ 19ss
10 cn,J.,LV, J.1 ._ 01.DRO\"'D. u .. ,sao. F:unily Tab:muiac.• /11: CJ<u~liiX.\-. n..·A'., Tub,mus /11.s;u:,n.rm,u ,lmrnrtut /om,ml ofVcwrlnnry Rh·eilfch. JO.
ed.' (".acnlogut10/IJ1n D1p11..•ronf1he .~01rop1~1/ R•18Ulfl. t.nndon· 8m1(h 1;;1.1;-;3.
\tu~um 1X~uur:tl Hts101')·} 2, TOIi, &.n .. ,, ......., t I ... '\It,\,\\~. ,, ,._ ~ l~t'.I. c.1 .• 1q$3, ~h."'<h~lul
11 , •.,.... J.E.. 1!'68, St-a~n.11 abundunuofTab;IJ1!d;10 al ,;..,,bul.a, rraru-fllh.,,iunol equtnC" inrc.•i..'l&ou" ant-i'lt.lu \1JU.~ b)' de~r flit...., c1t,,~µ1
/...amhl;, P1CNc't"lli11gJ ofthl l<oyal liJuomo/t>g!t:tJI SV<i\>1)', /.qmlmt '·"'· -13.
9
ftm:td1,,, a.nd suiblt Ult!"- -)tomcuy,, t,,lritrn,z.~i. ·1nh·rfcan Joumnl uf
108-l:?I \'ttc>nmrry RQ,"irrrli, -14. l:",~)56.

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92 •rn1,·, <N- .'lspeccs inOuencing the t>ccummce of inrenious diseas1•,

il l-OIL LI>.. $fGUt (' L. fRL:--.:cu. 0,1),. 1s..,l1 .. C,J.. ,1t. \IA ,u~. J.M., OUftftli.A(.. 10 ,isu.il •nd odour ottr>ttun« In Zimbabwe. Jo,m:t,I oJ.,frtmn Zoolo,::,•,
c.t , st., •,1~[\' R.t \988. Mt.">ChJnico1 rrn.n!l.tni'i~ion of ho\int' ltukcmla 106. 3;1-380.
,·,ru, b, hondhes ·olptor.: 'J'obJrtldae)./oum,11 of\T"llc,,/ ..n PU~U'", ~1 ., 111;J111Joww. '1.1 "·· 1'!!>U- Altg.htlng ,ht"' otrerna~ Tabtlmdi:ic>
F.mot1tolt1~·. 25. 3i.J-3;"'G. tl>1ptcra: ~t fft•koml1J1c-. l.imb;ibwc. ~1<.-,U«1t cmd Vc.1,•r/m:ry f'ntamnlo;,o·,
23 Htt..,LH, F.t ._ "'USh o.L, 19\S9~ l•oc-c-en·l·lll. :in cifi:ch\i\! a1irae1nnt for •l 3,l!f-356
T.JlMnidJI! 01ph,•r,1> Jounwl tJ{Mtttlical 1:utom'7fnJ,', .26. ~59-4b ! 42 1tODF.1trs..1u, ... 1978. Hnf\t" flh:,. J.ntl det-c fltc,.
FumU~· tolb.1.ntd.!l,J Ju·
2a1 1;n()r>trH. ff.. 19U.;. \ list urSouth~rn lthode,itm honefiic.•" IOiptt.'rn: ax.,~,. ILL
t'tl, ~rmlt!llln11r,: a1:,IC(J/locticm tJfr.r11110/J()'dsof \',•tt•t'1fuc)'
Tabonlct..,·1, f.m<1mulo~-ut's Mom/J/y Wt},'<t:lm• 102. 9'l- 03, Jmpurumu, USl1A .'\h-rricuJtun:,l llnndbook No. S18.

f<t:pon on the bh>ncuulc:,,. uf thi' bi'IY.• fl\' Gl<r.;5lua

2,.; u..uuu"-- H.u.t,P.. 1q~('). -13 noutn1~u ..... \.C• , ;gS;r. C)b,!..\•r.utlOrl). on the- ~:twnnl lndt!enre and
p11Wd1pt(' ..\m,t.) VrmindAI Adrnlnr. . u.u.ion of ',;t,t;il, PiN1.·m,olci1Yht111i. -abundmlCc.· of h;,,C'mau;~pha~uc: h1ghC"r Dip1c:rn <>Lhttr 1h;ut u-.~1,c 0JpU:ra:
Glo~sinldoc m the :,...,b, \'olff:y of /.imb:,b,,.·c. and romnan,Qn.. with
~. t< \JUU.,, 11..K.t.1·. t9.SO, Rl·pun on 1hi! 1r.1ppl.tt~of lstI~ lltt'- wi1h
ll1ussrntiuns·. P'ictl'rm;'ltilzbu~: The Nnrru ,,~nnc~. lad
1a.bnnid t-lt1:t:1:,- from dk!wheir 1n 7Jmbubwr tm,,mcmmsoj ,h,.
/.,mb.,lt,,, •.'\ct,mt1fi< ·WOl1nflo11, 61 -i l-S5~
~: 11-.'\AAI') R,I.. ttl'Olllu-Jt. D.O •• 1976. Centro! of i:ahwi,tl~ on hon-e:s. n,r ,a.; )11-Jut,-.. co, t~- EffN't't o! L"~ti:rrtal and lnu•m.i.l 3nhrarmd p;-lras11ct
${1wlm'('ltN11 Eirromolo;gm. t t9J-l!r:. 011 dom,:,.tir h\·,~~t(K i.: prQ<fuc11on •.-l.m1ua/ Rt"t•fPt4 ,,ff.'uu,,,rofott,•. 21.
:-:ft ,,..,fl. c J. "ro1L. LD.• 1q.$.: ..:mdlei on equlr.p ln(tctlou~ ;uwmia \1N' 155-1;8
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..W~durum, l84,29it-29i N11c111111 S 1.J.5-1 ·i~I
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R"\\l•ruorl ~:ulo11:1l l'oArl., Uyt\nd~ Pl'1,cc..'ilwtp oj tlw ,.,,,, &u, A{flenn Picu:t111t'rh1ba1 K-- !-<.10111 A(rJ\.tl. Unpul,lbla:.J J.u""'
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..1; 11,u\ '- l 1i.,11: ~unte ol,\.1--'l'\'a.tlo1'1$ on cc-mdn t~h.J..nld !Jic,s In notth
10 ~LE. o.~•• 1984. •\1edfl:"I ,md \ .tterinQJ)• f.',,ramott,10·. t ondon und ,•.1.,iem Lulul.,nd ,mptN'3 Tab-Jnid~). J'roc«1/IngJof11t,• /lt,)111
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Jl KAAMUt, "-.. l!t!lt, Ui.: &vfl!lllllllf 11(1n rnlm11frltJJ 11/s !10/linn11?1)m mul "'" ~ V"'tHJC. ,. J.. l9;l. ,\ rc.'\"IL''I\' Of lht.• ~uth Arnc:m hor!'t'fl} fa:.anJ 0tfH4.'T~
poW1tl<l/r (;efi,!trdu11s tf11rclt Tms.,J/1eg,,nl.,.k,m1fung.vrt::onm In I nb:lnrdu• • ,·\n11at1 oj thr N(IUJ/ \lu~J-um. Zl. ~5~-.307•
•\mrbabn'f!, D1plomnrbcn, ln~1hut fOTB:ogcoK7ar,h1e.•. Un1w~1hu 49 \Alt c. \ ,. il.\tL o «.. 1a1ts., Thr ,ateo1 l·«tenol•:1:·Ql. ucclone and
SIUlrlnndo,. S.fllrbn,ckcn. CUIOOII dto\1dr in •he amJ<tion or i<N~e ilios. Clossittn iJ>P Ulprr,,,:
J2_ ltJU~~l,.")' W.L I~. \11fma} Jf~e,1,t--agfnh lr.ln>tmfnt..'d by hor-c Olt:, :.tnd Clo,,inl,IM' 10 .,, udour 81,1/;r/11 ofE1110111oln1,.,rnl R,•,.,rrh ,$,
d,oer m~,,Ot111cr,.; Tabnnidacl, Jo11n111t of ',/,./Ira/ f11111mnl111{)', 13, 209-21';'
ll.i-275. :,0 \'AU c;,.\., lUO u.Jt, .. (;OUCH. ,\.,J.L.1988, Thoolfactnr:• nspa:~~(1(
33 xu\·t. , .., .. l~lO. Report on a j<HafflQ)· to chl.' Ltun~,·.1. \ :iUI!). '"''"' Ille< Citomtta ,;,p, 1Dip1ora: Glo,sinidae to phenols ..1d urine in
;,(9nh-Fa,1~m Rhnd,,fa fro,n Jul, u,,._,p,L-mht-r, 1910. 11111/<'lmQf the f1~ld. R11/1'.•1,n oj l:"lilomol(>S;ittll R,-wx,trh. i8, 293-300.
r.mamol11tknl Rowm.:/J, J. JOl-317. .5,1 \·-411 Lo.A, to •'tt1·1 v.. k.1 .. 1~,4. ~01~ ... un lhu lluM-OmJiug l11:h.i,.u1u; ul
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conuol "'1uno1>111:f~i.'llhead Oil!,, m,pt"1ll: Tnbanidno, on t:ape C'1d .
ihr1r lsft- h1s.tOTI('S, 8111/etln ofF.nromQ/og1,nl R1.•Jo1','1rd1. 5. ?87-320. \l,U.~:J.t:hu,it'!H,.. F:m•utJ,mt.4.'t1/lll Entomolt>,J', 8. 3:' J-375

J4 ,~1-\lu. r ~, ?001. Onderdsmpoort ,·uierin. J) lnMHUH". Ondcnr.c.poon.

5..l \\1J.<t~ulrnzfl', r. 1~;5, Rt°'"'-1.'.:,.rch inw tJlC reJ:uh"C lmpun:iner or
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Ill 10 ~auth ~fnCll. Unpubll<hcd o!l«ll\·atlon<
lm"'II~•"''" of rhc bd,a,·lour and fe;·ding h3blt, of hbanlda,, In
:r. ot.ono,-u. u .• 19;p. I. un~mau,po:n i1nd ll1111KK,•u:t,m1. ntL' H"r~·Jltt.., rclo11on :o <'Ulllr /or,mal 1,f \11111r<1/ 1/istor,•. g, 38S-:19t
tt)Iprt,a T,1l1'mid1wl of,11, £1/u'o1Jiarr /k"'f!.ft>n London~ Briii'\h \lu~um
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t011m:1n: Tn1>nm,t1l8Jnftltf' F.1ii;()pirm Ht~i,m. Lt:>11don: 6rill\h '-lv5-tum :\mmul Jltwltll mlfl J•rcd11t!lm1 1 ; , 14-22.
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3U 01 UM<tVU, 11 1957 Ill. S1,.1b(am11i~ Chf)'\Op:n.,e. '!Cttp,.1din11~ .,nd lah;anatu~ rthptcra tn rr,..:rh:mlcnl dlwa.S<! trJns,mti.~ion. I\.:\
f'1in_gonHnac .Uld ~ ro\ised C.h.l\.'\Uic--.alion. '11«• IIOrtt··fli,•s 10,,urw· C'onrribuuon to ,h ... L'pid1.•m1otogr of bo\1nc trypanorornillM~ int\frH:-.,
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Hl1i1ocy s, """'' ·, ,~ ;~. \ll'lllc.11 ,nd <ctorin:m lmponunce of hor.•-lhr<.. Sl>ut/1
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 5
Vectors: Culicoides spp.
Synon}111s: Biting midges: brandassies (Afrik.,

R NIEIS\VIXKEL, G) VENTER At\D E :VI :-:EVlLL

Introduction than one million Culicoides can be captured in a single

blacklight trap set at a horse stable on a wann ~ummer night.
Culicoides biting midges arc mosquito-like in their behav- Remarkably. such numbers probably represent only I per
iour with the females of nearly all ~pecies. being obligate cent of 1he Culicoides population actively seeking blood.
blood-suckers ffigures 5.1 and 5.2) and being active only at and lllu~trates the nighLJy levels of irritaiion. and possible
nigh1. !1$peciaUy when environmental conditions are calm risk of infection. thaL livestock may seasonally have 10 en·
and wann. The male5 feed on plant sap. Culicoidl!s are l to dure.i 19
3 mm in ~i:r.e and are 1hus tonsidetably smaller than mos- Certain species have a proven Involvement in !he rrans-
quiwes but are more abunda111 0nd. on occasion. are found mission of at least two of the 15 OIE list '1\" diseases. namel~
in enormou~ number, in climates ranging from tropical to bluewngue cBn and \frican horse sickncss(AHS,. They also
temperate. .\lost species pOS.SOlSS grey- and whlte-panrrned transmit epizootic haemorrhagic disease (EHD) of deer. and
"~ngs; these patterns being a very useful aid in their identi- equine encephalos1s (l'F.). 110• J2ll Thus. in sub-Saharan J\f.
fication. To date l 2 l O species of Culi,YJides have been de- rica. Culicoirles are of great economic and veterinaf)' signifi-
scribed world"ide~11 but it is certain that many mor.: await cance. Indeed. It hns been sugges1ed thar Culic"oides midges
di~covef)~ for example. one third of the 112 species kno\\'n were Involved In two of the ten biblical plagues of ancient
10 occur in South :\frica arc 11uw to St"ienre.- 11; Egyp1.a10 This is not Improbable:. a~ thesupprl?!>~ant effect of
C:ulicoitles are well represented in the fo,sil reeord (:i9 C11liaoi<lt>.<·transmitted (lisease mus1 have mnnlfested soon
species; with thl' lineage of the famil} Cerotopogonidae after the domestication of livestock some len mille!mia ago.
going back at least 130 million years.38 .\Hdges dearly recog- In support of thb contention is that epidemic~ 01 dC\-astat·
nizable as Ct1lti'oides have been described from upper Cre- ing cxient have occurred in the modern era, the most not-
taceous amber (88 to 93.5 million years agol. and it is able being when .\HS swept through 1he Near and :Vliddle
therr>fnN' r>rnhahlc> rha1 01/irnitfp;; nnrr frci nn riino,aur, ,\ 1:;t~1 in 1959/60 causingane~1ilmued 300 00(}death$. 11" An-
modern equi\•alent is the exclush·e association between other example is the AHS epidemic that decimnted iO ooo
some Culicoicles spp. and the African elepham (Loxodo11ta equids in the then Cape of Good Hope Colony 111 southern
(if,1cana}. 223 .\t present 38 subgenera comprise the genus Africa during the 185~ /55 ·sea,on'.19 The unprecedented
Clllicoides but many more await delineation. This taxo- scale or the latter outbreak. which claimed olO per cent oi the
nomic di,·ersity is testimony to a long :Jnd complex evolu- colony's total horse population In a mere eight momhs. at-
tionary history reflected in the fact that Culicoides today tests not only to the eflkiency of CrtlicoidlfSas disseminators
feed 011 a broad spectrum of hosts including reptiles, mam- of infection but also 10 the modern-day necessity of prowc-
mal.s. birds and humans. It has been shown that in south· tive annual vaccination. In eastern and somhem Africa
ea,1 Asia i.ome C11/icoides spp. will e\'en feed on blood- s1>ccilicall~ 1his prophyl.ictic regimen cannot be abandoned
engorged mosquitoes. Their blood-thirst\' auacks on as ii ha~ long been known that rherc is a dear link between
humens in some parts of the world are legion. and have above nveragc rainfall and outbreaks of Culicoiafs·borne
earned them scientific names such as <lan111osus. irriums. orbiviral tlise~~.
1,e,xm1s and tlial,olicus. These auad."S can be so intense as 10 ·n1e first studies on sub-Saharon C11/icoitlesdate back to
retard de,·elopment and production in industries such as 1908when two specie, \\"ere described from :-=amibia 1:!:! but,
forestry and tourism. 15"· 201 In Africa animals appear 10 be whil~t of 0 high standard. lhe sub~equent continemal re-
the hosts mos1 fmensively bincn. Under exceptional cir- search effort has been mostly short-term and fmgmemary.
cumstances. such as prevail during overly wet pt>rlods, more The la~t decade. howe\'er. has brought sib•nificant ad\·ances
93

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 94 -<c:nm, o,c Aspects influencing the oc<:urtcncc of infeeriou~ di,t-:1scs

••,• f
fi
,;
'\
\

• J•

_.. .. .._,,-, ........ -··

..___ ..., _., ..-···

•'
r
..... . , '
,
-
/''
I,'
I
~

..,
[,
figure !i.1 Culicc1desiu1ue/1Sls fellll!le

Figure 5.2 wlicoiaes spp.. lateral view

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 Wero,,;: C11lit11itf,,t ,pp. 95

especially in our basic underslanding or Culicoides sys1ema1- monicoring 1001 used for the capture c,f Crilicoides is the light
ics, which include$ the discove1y that the most important Old trap. However. other trapping methods of some varieri have
World vector of BT and Al IS, C i111iC'oln, lsbu1 one member of been de,·eloped over many years for the c.ipture ofaduli Cu-
a complex 1hat comprises al least IO ~pecics. :,,;01 only does licoides. Those mos1 commonly used in A1rica Include \·ari-
each species ha\'e a unique biology. bll! laboratory infection ous models oi white· or blacklight traps. emergence traps
studies have sho\,n chm a recently described <pecles \\1thln (for larval habitm ~tlldie:,). asplra1or~ (for the coller.tlrm of
the complex. C bo/i1illos, may in certain instances be a more live midges off human and an imal hosts), and truck rraps
efficieni field vector of bluetongue virus fBT\ i than C. imi- (which Involve thl.' use of a large net mounted on top of a
r:<JTt1. FU11hcr insights gained from field outbreak.~ strongly slow-moving vehicle). The trapping melhod employed de-
support the laboratol) findings around C J.;0Ti1i11os. and thus pends upon the purpose or the resoarchas each mer.hod en-
the ·single-vector s1aru, quo' that has ,tood ,incc the rime of joys some advantage,. not shared hy the oth..-rs. Their use,.
du Toil's ,;eminal studies11" on C. imlrola more 1han 50 year< and the merits and disadvan1ases of each, arc briefly
ago, is gradually being eroded, ln addition, 1.he long- reviewed below.
neglected study of the interaction between Q1/irnides and
African game animals (which nacurally cvcle the above- BlackUght traps
mentioned viruses) has finally received some ,memion. and l'hese traps are preferred o\'er those with whitelight as d11.•y
yielded much that ls of interest. This growth in basic know- appareml) attract larger numbers and varieties of insect~.
ledge was not only multi-disciplinary in irs embrace but also This, and their flexibili1} in application. make the blackligh1
formed part of an attempt 10 'modernize· the study of Cu/l- trap the lOOI of choice when bi1lng midge abundance le,·els
coides-in Africa. Another consequence was the production or are co be determined during ou1breaks of disease. and when
three satellitc-based predictive risk maps for C. illliwlti, the large-scale faunistlc surveys arc being conducred The Culi-
first for any specie:, of C11Jic:oidcs in the world, Also harnessed coide.~ are captured in o bowl or liquid (suspended belo\\ a
for the first time were molecular techniques ,uch as the ran· suction fan. which in rnrn is below 1he light ~qurce) in \,hich
dom ampllficd f)<)lymorphic DNA-polymerase chain reaction they die. In a manerofhours these ·wet" capture, make an1il-
(MPD-PCill and nmochondrial DNA {mtDSA) ex1rac1ion. able large numbers of midges ~uhable for a number of pur-
Not only ha\"e chese srndic, resulted in a partial phylogeny or poses that include \'im, isolation. blood meal identification.
the lmicola complex but. imf)<)rtantly. have also reaffinned age-grading. and ta.xonomicstudics. When the light trap is set
the stability 01 ·otd-fashioned' morphological identifica1ions, near lh·e$tock(usually at a height varying between land 2 ml
and so reas..,ure us that the s1eady advances being made on a few small drops of detergent are added to the bowl of water
the unique life cycle of each species do indeed reflec1 ·reaJi- to break the surface tension. This is necess31)· because
til>,,' in na1ure. These advam:es are re\iewed here. and be- midges are too small and light to drown and sink to the bot-
cause the research has focused principally on \'CCtOt'$ of tom, \\~th 1he result thot in the morning. when 1he trar> i~
orbiviruses Lhnc affect livescock, this chapter concentrates on s\,1tched off. a number of !hem may recover and fly off. \\'ith
a.s5embllng much of the knowledge accumulated thus iar on minor modifica1ion the same trap can be used to catch large
1he cwo proven ,·ectors C imicola and C. boll1i,u,s. Included is numbers of live midge,- for \·ecmr t'ompetence studies in the
an introductory preamble of !he standardized prutocols used labormory. In this instance a gauzed cage replaces 1he bowl of
10 collec1, ~ubsrunple and age-grad,;> Culicoide,, which were liqu id. AhematiYel}. the bowl m.iy be retained bu1 is partially
developed principal!~ in South ·\frica onir the la,1 30 y<>ar... filled with loosely <·rumpled tis.~111' pllper so 1hn1 the li\·e. <'UP·
and are now used in ,·arious parts 1>f Africa and beyond. It is 1ured midges can creep imo rece!'Scs away from 1he incessant
essential 1hat daca collected over a wide geographic area and wind created by tl1e ~uction fan. ,\nother advantage or the
during a number oi sea~ons and years be comparable so that blackllght trap is that It i~ easy to u~e as ir can be rigg,'CI in a
accurare dls1riburion databases can be created for 1he suhsc- shon space or time and. furthermore, does 1101 have m be
<Juent development of disease risk-maps. At various junc- moniwrcd u 11cil morning, when it can be dismamled e<1ually
tures in this review an indication is given of the rc5carcll '\trll quickly. \.\"hat is time-consuming, however, {<; the journey to
needed LO refine our und<'rsranding of,\frican Cu/icoides and disease ou1break poims and choices 10 be made as 10 which
the diseases they rransmi1. sites will produce optimal catches. In addition, the journey
has 10 be made twice a., the rrap$ nrns1 be retrieved the fol-
l0\,1ng morning. Ught rraps cannot be left in position for
Collecting adult Culicoides
more than one night, nnd should not bl' allowed 10 hang m
The majorit) of 1m,es1iga1lons conducted on Culicoides sunlight as the captured midges, cspcciaJI)' 1ho&e In a w<1nk
worldwide deal primarily with the mm1icoring of disease soapy de1ergem solution. starl 10 decompose very quickly in
\'ectors as 1heir acknowledged role in epidemics of tropical or subtropical dll)'time tl'lnperarure,,. Another disad-
disease impacts directly on management practices. such as vantage is lha1 most light traps require electricit,, which
vaccinaiion, and quarantine and expor: of live animals co greatly res1ric!S 1heir use in niral si1uations. Whilst it is pos-
imemational destinations. The primary. almost sole, sible co run chese trap> using a generator, the tTansport and

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96 , i<'I'"' u,t ,\spec1s in!lucncing tl1e occurrcnct-af infectious dlsi,a~e,

movement of surh heavy equipment mean, that very few Aspirators and sweep nets
geographic points can be $8mpled per night. Ligh1 traps can 1\spirntors (or 'pomers'J and hand-operated swe:eep ne1s are
be adapted 10 nin on 12-volt hattcries bu1 here. mo, the usod in specialized host prefcrenqc swdies when lil'e Culi-
transport and maintenance of heavy batteries can be bur- coide.~ must be captured off tethe,cd animal,, 01 humans. \
densome and impractical. In this comexi It mll!-t also be hand-held poo1cr can be used but this require;, that each
borne In rnintl that the:, various n\uc.llficatiun, made tu trap~ lmlh·Jt!ual mic.lgc bc !ucalt:tl, u;,i11i; a mt! turchlight. prior 10
to capture Crr/icqides in ou1 of lheway places must not re.5ult caprnre, whit;h is time-consuming. More commonly tl1e
in their 101,·ered performance as this will prol'idc dam oflim- 'sweeping· of marked areas of the host with a ~mall hand·
ited comparative value. A funhcr disadvamage is lha1 1he held domestic vacuum cleaner is the prefened method. In
traps are equipped with a strong suction fan, which ,:an th!$ way more Culicoides may be captured more rapidly.
damage Culltoides. TI1is can be particularlr vexing when which is necessary ifit has 10 he es1ablished (>xactly which
rare or new species have been collcc1ed which must be in a areas of the host are being aitacked within a specified time
suitable state for spec;iali1.ed taxonorpk smdies. l!cingpow· slot. In life cycle ,tudics in which the caplure ot gravid fe.
erful. these traps also capture large numbe~ of other insec1$ males for O\•lposition determinations is required. the
including moths and, under certain condition,;. this can re- midges are besr obtained b~· pootcring those atuactcd to a
sult in collections that are extremely diny and tha1 take a source of light,
lc>ng time to clean. Larger insects can also damage the Traps baited with carbon dioxide have al;a b<>en used 10
smaller midges and is another reason why blacklight traps collect Culicoid<',~- The particular advamage of these traps is
should be covered \\ilh a fine gauze mesh to exclude the that the~· et\able the collection of diurnal ~pecie,. l.imita·
capture of all insects larger than 2 mm. As mentioned. ex- tlons of carbon dioxide traps arc 1ha1 the~ require dry ice
ceptionally large numbers of Culicnldes can be captured in and char the unregulated release of carbon dioxide gas may
blacklight trap~. To be able 10 deterllline thb total. and che pro,·ide concemrati<>n~ that are attractive to ,ome ~pecies
exact number of each species c;ollcrted, It ls es~emial co be and repellent to others.
able 10 subsample catches ea~ilv and rapidly as hundreds 01
collection,; mny be made during a large sur, ey. A method of Emergence traps
~uhsampllng C11/icoideshas been de1·eloped32:, and i~ widely These are madf of fine netting, arL' conical in ~hapt, and
employed. Because of their srrong attractant power. black- have a collection bottle at the opex that b lined \\ith a
light traps are considered biru,ed ,amplers and therefore in sticky substance or contains a liquid.2 ' 6 An em.,..rgence trap
more specialized studies. such as those concerned with host (built to co,·er a specific unil areal Is placed o\'er a ~us-
preference. other more ·objective· collecting techniques pccted C:ilicoides larval habitat and remains i11 siw wh1;1re
must be employed. it can be monitored hourly. daily. or week!~ In this way
emergence rate, and species-association profiles can be
Vehicle-mow,ted traps ob1ained. The chemical composition and structure of the
For ob1-ious reasons the blackllght trap is useles-; forthestudy soil. and the amount of water i1 holds can. in addition, also
of Cuiicoide.s that may be active by day. For this. \'ehicle- be determined for a particular emergence sice. Further-
moumed traps (commonly rcfen·cd to as ·,ruck traps') can be: more. during emergence. if larvae or pupae must be
used to capture n~~ng Culicoides throughout the die!. and in re1rie\'ed samples of the substrate can be extracted and
this way houri> acrivity rates may be detem1ined: the results taken co the laboratory \\'here a sat mated sugar solution Is
can also be rela1ed to pre"ailing meteorological conditions added to them co airer the specific i,rra\·ity in order to
such as temperature. re!alivc humidity and wln<lines,,. Ve- induce the larvae and pupae 10 Ooat to the surface. The
hicle-mounted 11aps are particularly successful for the collec· pupae are rccrie,·cd \\"ith a spmul(I rtnd placed in individual
tion of large numbers of biung nudges during the hours of l'ials for ecl05ion. The resultant adult and ns as~ociatE'd
dusk and da\\11. In addirion. male swarms. which may not b(! pupal pelt can then be moun1ed on a single g!as~ slide
attracted to light traps. can becapturw in this war, a1 times in for iden1ifica1ion. The rearing of individual laf\·ae so ob-
large numbers. !f collection periods are limited to 15 minutes, wined is more complicated bu1 can be achie\'ed on agarose
the midges captured will be in pristine condition and there- media to which cultured micro-organism:. are added. This.
fore suitable for 1he preparation of material on glas~ slides for or a similar. rearing strategy is essential tO\\'ards under-
taxonomic study. Because Culicoules are host-oriemated, standing more precisely the e,ology of ind1\ldual Culi-
tntck trapping great!~· enhance,; the po»sibili1y of capnaring cr,ides ~P?· n~ it h the availabililr of th~ larval habi1at that
Cttlicoides near large groups of wildlife that are either danger· determines primarily the prevalence uf a species in any
ous and/or scaner when approached on foot. 11 is. in addi- given area.
tion. ,;ery difficult 10 set light traps near ,,ildlifc: this being It should be noted that much more research ls still
compounded by the fact that herd5 are constantly on the required towards more clearly defining the lan·al habitats
mo,·e. Disadvamagcs of cnick trapping are the cost of fuel for and food requirements of the immature s1agcs of the ,·ec1or
the vehicle and the 1in1e required fordri\ing and trapping. species of Culicoides. including C. imicola and C. bolici,wJ.

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 Vectors: Culicoides spp 97

Table 5.l Micro-organisms associated with lhe genus

Cullco,rfes worldwide (cominued)
CUL/CO/DESS? MICRO-ORGANISM REFERENCE CUUCOIDES SP. MICRO-ORGANISM REF£RENC1:
C. amamiensis Macacanema 27 28 Haemoprc1eus hamfai 240
formOSona Eufifa,,e/la 11anle11ae 11
C. 8/Ull<JC Onchoccrc;i sp. 3i 347 f rte//ce;a 11
C. erakewae !etJCOCViOZOOfl (Aki/Ja} 1, 2. 123. 170, 1S3. E kalifei 237
caullery, 24~250. 284. 317 Oncizoce,ca ceMcalis 133. 134. 157 22g.230
lemdanmae 313 0 g1b$0fli 31, 60,124. 201
Onchoce1ca gu1wrosa 98. 113. 201 318 0. gu11urosa 99. 113, 201,318
C. 8//JOflCOla Haemopro1eus 7 0 re1icu/a1a 244. 245
meleagr,dis
01pe,a!onema ozzard/ 205.232.257.3'2.322
C. adersi Hepa1ocvs1is ·J7-13S. id6. iA7
(Plasmodium/ kochi C obsoletus Onchocerca ce/'V/aJl1s 75. 151, 157,180,201,
C us;eni D1peralor1em!3 16}. 162. 270 231. 301
perSlans C. odlbi11s Leucocyto1oon (Ak1/Ja/ 146
C CI/Cumscnprus 1~5 cau/feryi
LeutOC>/lOZOOfl {AkJ/JaJ
caullery, C OX)'SIOma Onchocerca gibsoni 31. 60, 124, 20i
'C C/BPUSCtJlaris Haemopro1ous i4S C or,e111alis Onr:hocerCJ g1bsoni 31. 60. 124. 201
danilews4yi C pa(loli One11ocercacer.1calls 76. 151 157 180. 201.
H fnngil/ae ·45 231,301
Cnandletella quiscafl 163 C. p/1/ebo1omus D1petalooema omrrJ, 205,231.257.3•2.322
C dovmasi Haemoproteus "3J. i4S, 2S£ C. puiiC:JflS franc,sella 1Ularens1s 176.273
ne1rionis C.pungens Oncnocerca gibsoni 31, 60. 124,201
C. edem Haemoproreus 1 3. ·o C. riethi D1pe1aronerna oll/Jrd, 205,232,257 312,322
meleagridis C. schultzei Leucocyrozoon (Ak1ba/ 146
Vavraiasp. 9 caullel'VI
C. fblvtrhora)( Hepa1ocys1is '37-13S 146, 1~7 C short11 Onchocerca gi!Jsom 31, 60. 12.i 2.01
IPfasmodiumJ kochi C sph;ignumens,s Trypa111;sama avwm 25
Onc/wcerca gutturosa 98. ii3. 201 3;5 Haemoproreus 146
C. lbfeflS D1pe1a/onema 204 c.r.achires
ma,moserae H danilsws,f(yl 146
D. oztarrfi 68.59 132,205 H mansam 7
C. graham11 Diperafonema i 1,. '56, 270, 271, 292. rl /Pa,ahaamo· I 146
Del$1ans 293 proi:eus/ velans
D. srrep1ocercum 68.69. 156 C. srifobeo1ol!iES 1rvpanosoma avwm 25
C haema;opo:us Haemoproreus 7 Haemop:o(eus 146
meleagridis fr,nglllae
C hmmem Haemoprareus 7 H. (Paratiaemo· 146
meleagridls proreus/ velans
C. nolle11s,s D,peralonema gracile 113 Chand/ere/la 17
D. caud1spma 113 chltv.'OOdae
D. 1/ewellyni 35l C 11avi'si Chamfleretla 17
D m;;rmosetae 204 ch,1waGdae
C imm,Jtip.annis Difi'll/;;/um1mrJ 114, i56. 270. 271 292, C. mfasc,el/us OnchO!;efCIJ gu11uros11 98. 1lJ. 201, J18
perstans 293 C. varilpennis Trypanasome sp. 161
C ir.sinuerus D1pera/l)f1ema onardi 205,232, 257,312,322 Hepa1ocys11s bray, 2J9
C. k,ngi Onchocerca gutw,osa 98. 113. 201 318 Onchoce,ca cefl!icatis 133,134, 157 229. 230
C knOWl(Ofll Haemoproteus 7 D1pe1e1onema 011ard1 205.232 257 3"2, 322
meleagridis Culicaictes sop ll?JShrnaniasp. 70
C. krame11 Oncr.ocerca gutturosa 98, 113. 201 318 Plasmcdium matarlae 91
C. ,r,IJ[ksi Or1chocerca sp. 31. 348 129
leucocyuuoon neavei
0. gibsoni 31. 60, 124,201 L schou1edeni 129
C rr;ultider.aws Spfemfidoli/aria 341 L lll.kfba/sp 12. 126. 127, 187, 29'
califom,ensis
Haemoprateus 147
C. nubeculos/JS Trypanosoma baker, 238 canschtres
T chabaudi 67 H sp. 26, 79, 125-118 ·30,
T davirfmolyneuxi 67 144 147
T ev/ilem 57 Ormt/iofilaria 5
T. gentillnii 67 fallisensis
Hepa1ocys1is bray/ 233 Dnchocerca gibsoni 31, 60, 124, 2C1
H levme, 40. 1!15 0 volvulus 141,326

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98 "'CT'"' "'": ASpectS influencing the occurrence o( infec1ious diseasus

Livestock and wildlife diseases/infections subgenus AlJ(/1'/tia alone. l 5 rrom C. brevltarsiJ in Australia.
associated ,vith Culicoides and oight from C. imico/(,( in Africa. with BT\/ being common
10 both continents. :\)though these figures may be biased in
Summer seasonal recurrent dermatitis that Lhey reflect both the specific direcrions taken in Culi-
Summer seasonal recurrent dermatilis ls mos1 commonly coldes research and the panicu la r drive of I he research team
referre.d to as sweet itch, bu1 is al$o known as Queensland involved, they do show that the tropics and subtropics are
icch, "dhobie ilch·, kasen disease. allergic derma1iris and particularly rich in both bunya- and orbiviruses.
sommerek.zem. It is a chronic. seasonally superficial allergic
dermatitis of horses that usually affects 1heskin of1he mane, 81111y1111iridae The Simbu group viruses (see Chapter 94:
tail and \,ithers resuhlng from the bi1es of Culicoides Oiseases caused by Akabane and related Simbu-group
midges. Various species are i1wolved. This condilion was viruses) that are found in southern Africa 90 include Akabane
firs1 recognized in France in 1840. but has since been inves- virus. the cause oi abortion and congenital deformities in
tigated in manycountries.55 · 5"- 78· MS. l88. zs&. 323. 3• ' In lsrncl domestic ruminants. Akabane ,~rus is widespread in south·
C. imicola was implicated as the pnnc1paJ species involved ern Africa occurring in a1 leas1 25 species of wildlife.3 and the
as 70 percent ofl95specimens caught on a horse showed a prevalence of antibodies 10 il in African buffalo (S)'ncerus
clear preference for feeding on the dorsal ridge, which over- ca/far), blue \\ildebeest (Com1ocl1aeres ta11rim1s) (as high as
laps the Itch zone.1 2 tn Lhe tJK a significant and rapid reduc· 69 per cent in '.';amiblal and African elephant ll.bxodo11w af
tion in the prevalence of swee1 itch was demonstrated once ricanal srrongly implica1es them in it> n1almena11ce in na-
affected horses were more regularli• stabled. 23 ' \•1 rtualty no ture. Sabo vims.<><>· 321 closely related to Akabane ,irus. also
studies ha,·e been conducted on the condi1ion in Africa but occurs u1cattleand C11/icoidesmidges in South Africa. whils1
it is suspected to occur in South Africa.324 Shamonda viru~ has been isolated from C. imicol(I coilected
near canle at the Onderstepoon Veterinary lnstimte in
Diseases associated with Nematoda: Filarioidea Somh ,\frka.211 !n additio.n. in S0u1h Africa a strain oiShuni
Some 20 species of filartaJ nematodes are transmitted bi• virus has been recovered from the brain or an adult mare
Culicoides midges and include five species of Oncliocerca in 1ha1 died after showing nervous signs. Subsequent serologi-
caule. horses and water buffalo (Buba/is b11balus). and two <.'al surveys revealed positive antibody tines in horses to
species or Dipemlonema in monkeys (Table 5.l}. 201 Akabane, Shuni. Sharnonda and Sabo ,iruses, with the latter
two predominant. Their presence Illa)' explain some of the
Diseases associated w ith Protozoa febrile reactions and other vague clinical signs observed in
Wotld\\~de, at least 33 species of protozoa In the orders Eu- horses seasooally.i r,s Although they have not yet been tested
coccida and Klnetoplastida are known to be transmirted for pathogenicity in mminants. South African strains of
only by Cullcoides {Table 5. l J.201 Except for a species or Sabo. Shamonda and Shuni virnscs induce marked 1erato·
Hepatocysris found in several specifll> of monkeys in Africa genie effecrs in chicken e mbryos.~0 Alno vims. which is
and another in the Malaysian sciurid squirrels of the genera closely related nntigenlcally 10 Shun! virus. has, in Japan and
Callosci11rns and SundasciumJ, the other IO pro1ozoa occur Australia been implicated as the cause of congenital
in birds. One of these. J.e11cocyro~oo11 caulleryi, is an eco- anomalies in caule and sheep. 110• Jrn
nomically importan t disease of poultry in sourh-ea~t Asia.
Reoviri-d"e Within this family, the orbiviruses of BT. AHS.
Diseases associated with viruses and EE are commonly associated with causes of livestock
Of all the pathogens transm iued b~· Culi,oides, Viruses are or diseases in somhern Africa. Other members of this family in-
the greatest veterinary importance, and. more especially. clude EHD. :-.:yabira. Gwem a nd Kaslr.i (: Chuzan) viruses.
those that cause BT, AHS. EE and Akabane disease. Other,~. Both Nyabira and Gweru are liule understood Palyam-sero-
ruses closely related ro Akabane \'irus have also boen iso- group viruses (sec Chapter 107: Palyam serogroup orbhti-
lated from Culicoides (Table 5.2). Although many of 1.he rus infections) that ha,·e been found only in southern Africa
viruses that are Cu/Jcoides-associaied ha\"e been isolated but. like Akabane ,irus. are linked to abonions and teratol-
from. bur octur more commonly in, other anhropods such ogy in canle, goars and sheep.l 14 • 3•1·1 Viruses of the Palyam
as mosquitoes. argasid and ixodid ticks. and ph.lebotomine&, serogroup have a particu lar association \\'ith cattle. and the
there rema1n 45 percem which have been isolated only from many isolarions made from Culicoides sugges1 that they are
C:ulicnides midges. competent \'ectors for Pal)•am viruses irl southern Africa.3 ' 5
World\,ide, somo 64 named and l 1 unnamed arbOvi· Nyabira ,irus, for instance, replicates well in C imicola and
ruses are C11/icoides-associated. Of these. 25 are from Aus- C zuluensis, bu1 transmission trials ha\'e proved inconclu-
tralia and 23 from.Africa. Of the i5 viruses listed in Table 5.2, sive.s.• There is serological e,~dence for the widespread dis·
46 have been isolated from pools of identified C11/icoides 1ribution of Palyam-serogroup viruses throughout southern
spp. rather than from pools comaining mixed species. ·\frica except for the cooler winter rainfall region of the
Twenty-three o( these viruses are from the C11/icoides Wes1em Cape Pro\i.nCe of South Africa. Since 1hey may be

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 Vec1ors: Culicoldes~pp. 99

Table 5.2 Viruses i$olated from I/le genus Cu/lcoidesworldw1de

CUUCOIDESSP SUBGENUS VIRUS ABBREVIATION REFERENCE

C. actoni Avaritla Warre{lo WAR 149

C agarhensis Unassigned Alphavirus sp. 53
C a/gecirensis Mo11oculicoides B0111ne ephemeral rever BEF 296
C a.'Srropelpalis Similisgrp. K\JTlijnurra KNA 149
Wongatel 225
C bedfordi Unidentified 266
C. bolirinos A~riria Af•ican horse sl~ess AHS 226
!31uetongue 16
·tetsitele 266
C brewrarsis Avat1ri2 Amo AINO 111
A~abane AKA 87 99. 108. 111,308
B!~etongue BT 302. 307
Bovme ephemeral lever BEF 87
Buny1p Creek BC 87.88
CSIRO Village CVG 298
O'Agu~ar DAG 110,111
Douglas DOU 304
Ep,zoot1c haemorrhagic disease EHO 277,306
of deer
Kimberley KIM 351
Ngaingan NGA 110
Peaton PEA 310
T1brogar9an TIB 89
Tinaioo TIN 86,304
Walla! WAL 175
Unidentified 111,305
C. brev1rarsis - C sdlultzei .. Mixed Un1denufred 305
C. bund)1!ns1s Unassigned Belmont BEL 149
C. ccarcretlft Unassigned Bovme epl\emeral fever Bff 34
C. cockerel/ii Sylwcola Blue tongue BT 190
C. cornurus Monoculico1des Epizootic haemorrl1ag1c disease EHO 16
oi dee1
C.dyce; Unassigned Wallal WAL 110,111
Warrego WAR 110
C. e,specieror Similisgrp Blue tongue BT 266
C. filanfe,• Unassigne(! Blue tongue BT 242
C. fi1arifer - C. pusillus Mixed Blue,ongue BT 149
C. fulvus Avarir,a Bluetongue BT 298
C. fuiws + C. oriemalis Mixsa Bluetongue BT 291
C. guloenkiem Avamia le,s,tete 266
C. hisuio Mei/erehelea Thimin TIM 298
C. imicola Avaricia Akabane AKA 4.34
African horse sickness AHS 34,266
Blue-tongue BT 16,34,SJ, 143,235 266.339
Bovine ephemeral fever BEF 3<l
Equine encephalosis c-
.t 266
· Lemtele 266
·Nyab1ra 34
Sabo SABO 66. 175, 197
Shamonda SHA 175, 197
Simbu 266
lJmdentified 97
C. imicola - C. sct,u111e1·· Mixed Unraentified 97
C. msigms Holfmam11 • 81,:ens Arm 140
Bluewngue BT n. 145.242
Sweetwater Branch 225
C kir,g1 flemml;; Epiroottc 1laemorrhag1c disease EHD 235
of deer
Umdemif1ed 235
C.megnus Culicoides 0

LetS<tele 266
C. marksi Unassigned Belmonl BEL 298

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100 s.r.cno~ 0~1. Aspects influencing the occurrcnc:e of infect[ous dil;eases

Table 52 rcommuedJ

CUUCOIDESSP SUBGENUS VIRUS ABBREVIATION REfEllENCE

Barman BF 2!l8
Eubenangee EUB 109.2S8
·HumptY Doo 298
"Leanyer 298
·Mudjinbarry 112
·?arker's farm 298
Walfal W/l.l. 110. 298
Warrego WAR 110. 175
Unidentified 300
C m,1ne1 Hoffma~u; Akatiane t,,l(A 34
Bluetongue BT 339
C new1/i Remm,a Epizoouc haemorrhagic disease ErlO 16
of deer
C newS1e2d1 Cultcoides Alphav,rus sp 53
C. nivC!PJS Meijerehelea Th1miri TIM 225
C obw/e1u5 Avatitu, 81UoU)llQU8 BT 236
C ozysrcmo Remm,a Al:abane AKA 192. 193
Sun\'IP Creek BC 225
Kaslla KAS 194
C. pallidothotak Unass1gne:i Wongorr WGR 298
C p.raensis UooS1iigned Ananindeua ANU 175
Oropouche ORO 196
C. pereyrmus Hoflmamil ·eeamceHill 298
C. peregrinus -C schultzei· • Mixerl Un,dentif1ed 88,305
C. punc!arus Culicoides 8aru1 BM 174
C p!ISilfus Avarma illuetongue BT 2~2
C. pycnos1,ct11s Meijerefli!fee Blueiongue BT 266
C. schult.te1 • • Remm,a Bovine ephemeral fever BEF 97
Bun\'IP Creek BC 298
D'Agu1la1 OAG 111
Ecizoollc haemorrhagic disease Et!D 196. 198
of deer
Kaslla i<.AS 19~. 197. i98.2!1
Keterah JCTR 259
Mar1aka1 MAR 298
C schultze1·· - C 1m1cola Mixed Kasba KAS 97
C scnu1ue1 •• .. C pereg:i/lus Mixed Marrakai MAR 88
C schu/a,;, g~ Remmia Unidentified 256
C s1etMer Unass,gn~d Vesicular stomatitis New Jersev VSNJ i91l. 340
C /CfOIOeflSiS Awmt,a Nairobi sheep disease NSD 97.339
Bluetongue BT 340
C :>anipennis Mcnoculicoides Bluetongue BT 135. 136. 190,203.285.346
Bunyavirus sp. 154
Bunonwillow BUT 153. 154. 175. 190.258. 287
Epizoouc haemorrhagic disease :HD 145, i 72, 35A
of deer
lokern LOK 85. 190.258
Main Dra,n MD 85,121.268
Vesicular stomatltis New Jsrse'{ VSNJ 175. 190,340
C !·,aaa, Averic,a Akabane AKA 311
C luluensis Hoffmi1111<1 Letsne!e 266
Umder.t1fied 97
C. 1u/uensis - C im,cota Mixed Unicenufied 97
·w1ico1aes spp Se1fla Lo~ern LOK 190
Main Drain MD 190
Vesicul~r stomatnis New JarSlly VSNJ \90, 340
Culico1des SO{) Mixed Afncan horse sickness AHS 97. 175. 233. 266
Aino AINO 175
lll<abane AKA 175,321

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 \'c•ciors: Culicoidn spp. 101

Table 52 {continued/

CU!ICOIDESSP SUBGENUS VIRUS ABBAEVlATION REFERENCE

Blue!ongce BT 32. 44 lS7 198,243.206. 283.
309
Bov1oe ephemeral fever BEF 95. 97
·surimana 150
Buttonwllfow BUT 175. 190
Ccnao-Crimean haemormag,c CCHF 197
fever
CSlRO Village CVG 88
O'Agu1lar DAG 300
Dou;;las DOU 198
Ougbe DUG 197
fas1ern equine EEE 177
entel)halomyelms
Ep,zo01ic haemorrhagic disease EHD 197,198,266
of deer
Equine encephalosis EE 265
0
Gwe11, 345
Israel turkey Ii 53. 175
meningoencephahtis
·11acalunas 149
Kesba KAS 197 198
Ko1cnkan KOT 179. 197
0
Lets11eie 266
Marraka1 MAR 298
Mitchell River MA 110
Nairobi sheep disea.sil NSO 197
Ngaingan NGA 110. 178
'Nyabira 345
Palyam PAL 266
Rift Valley fevet RVF 92.94.96
Sabo SABO 65,175.197
Sango SAN 66. 197
Salhcpen SAT 66,175.197
Shamcn!!a SHA 175, 197
Shuni SHU 66.197
S1mbu 266
Tahyna TAH 150
Uunga un 186
Walla! WAL 110
Warreyo WAR 11 l
·weldona 63
Unidentified 197. 256. 300. 32l

Arbov,ruses not registered in the lnuirniltional Catalogue' 78

Most ii 001 all. of these virus isolations should be anrlbuted to the Schultze1 group l~ S1Jbgenus Remmia, as the species C. schultre. Se/1SU stnc!o
d~es not occur outside Africa

abonigenic in cactle. there is clearly a need for patbogenicity from a mixed pool of two Remmia spp. (including C kingil
trials in Uvesrock. 345 A third Palyam-serogroup vims (Kasba !hat a n EHD virus was isolated in 1he Sudan.235
,·Jrus) has been associated with congenital abnormalities in The first isolation of equine encephalosis vims IBEV) was
calves in Japan and has been isolated from C. 0).ysroma.'1A 1 made from a mixed pool of Culicoides consisting mainly of
This Culicoides is a member of the subgenus Remmia and is C:: imicola collected ut the Onderslepoon Vc1erinury
widely and abundantly found In sourh-east Asia and lhe lnst!tute.321
eascern Palaearctic. Similarly, in Africa. 1he Mediterranean
1egion and the !\ear and .\1iddle East seven or eight species Rluibd-0uiridae In this iamily is induded bo\ine ephem-
of Remmia can also be found in some abundance, panicu- eral fever virus (BEFV). The seasonal occurrence of BF.F (see
larly in honer. drier regions: these therefore deserveconsld- Chapter I 00: Bovine ephem eral fever) suggested insect
eration as potential vectors of various reoviruses. I! ,,·as transmission to early investigators i.n Africa29• 30 although

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I 02 ~11CT10i, o,r. Aspects influencing 1he O{:curr~nce or infec1ious diseases

firm evidence of insect involvemem was not pro\1ded until Vector competence
the causative ~rus was isolated from a pool of several spe- Vecror competence is a measure of the number of midges
cies of Culiccides io Kenya in 1971.9~· 97 l.t has been recov- that actually become infective after feeding on a viraemic
ered from Culicoides only twice in southern Africa, namely hos1. Ths comperence is dependem upon the genetic
from C imicola and C. coarcta!ll.$in Zimbabwe.34 l!s low iso- makeup of the vector midge and external environmental ln-
latlo11 rate is possibly due to it being panlcularJy diJlkuJt to Ouences.31&· a•~, :}.IS
isolate u slngscandard cell culture techniques, and is further The vrrus vector tompecency of a Cullcoide; sp. can be
complicated by the fact that there are many other rhabdovi- experimentally assessed by first allowing a number of
nises closely related antigenically 10 BEPV.303 In Ausrralia midges to feed on a viraemic animal or. in the laboratory. on
BEFV has been isolated fTOm C. brevitnrsis.m the sister spe- a blood-virus suspension through a membrane. The en-
cies co the African C. bo/itinos. Both species breed in cattle gorged midges are then kept alive fo.r the e>."Uinsic incuba-
dung. with C. bolitinos also being found in the dung of the cion period, i.e. the period between feeding on infected
African buffalo and blue " ~ldebeesr.213 .
Two other rhabdo~ruses, Bivens Arm ai1d Sweetwater
blood and rhe appearance of virus in the salh'll of the midge
and is in the order of one to two weeks. Following ingestion
Branch, have been isolated from C. (Hojfmanfn) i11sig11fscol - by a susceptible arthropod. most arbo\~Ses infect and rep·
lected near a water buffalo imported into North America lie ace in cells·of the mesenteron before penerrating the basal
from Tri nidad 1'10 but have nor been linked co naturally OC· lamina to be released into the haemolymph 10 set up more
curring or experimentally induced disease. These viruses are cycles of infection and replication. Further barriers co infec-
related 10 Tibrogargan vi rus. which is associated with water rion appear to exist at the levels of organs such as the ovaries
buffalo. cattle ai1d ho rses in Australia and has also nor, as and salivary glands. A ~rus must infect and replica te in the
yet, been shown co be pathogenic. 90 The Culicoides mil11ei salivary glands before the extrinsic incubation period of the
complex, which, in Africa, is represenced by at least 16 \~rus is successfully concluded and the arthropod is able ro
species. is very closely related to species of the subgenus transmit the ~rus by bite. Tt is possible for arthropods to ob-
Hojfmania. They can occur in large n um bers and must tain ai1 infection ,~th great efficiency b)' ingestion of a ,i-
therefore be considered as potential vectors of rhabdo- raemic blood meal. yet they may still fail co transmit virus
vinises. because 1he infecrion has not spread to the salivary glands.
Hence It ls lmportanc In vector studies in the laboratory ro
disiingu ish between arthropod infection rates a.nd trans-
Viral maintenance and transmission
mission ;ates.313 The ability of these midges 10 transmit
Vector capacity virus iS then assessed by a!Jowing them to feed on suscep-
Vector capacity is defined as the average number ofinfective tible animal hosts or on suirable s11bstitutes.
bites that will be delivered by a Cu/iccides midge feedJng on After a ~eries of failed experiments involving mosqui·
a single hose animal in one day, and isa combination ofa!J of roes,r-2 Du Toit in 1943 conducted the first successful Culi-
the following: coides vecmr competence studies at the Onderstepoon
• midge density in relalion to the host 8Jlimal, Veterinary Institute. t 15 He fed field-collected Culicoides on
• host preference, BTV-infecced sheep. and after an extrinsic incubarion pe-
• midge biting frequency. riod of 10 days. was able to transmi r the disease co suscep·
• vector competence (re~ewed below), and tible sheep, but he did not specifically identify the species of
• life span of the infected midge and duration of infectivicy. Culicoides used in his studies. rt is most likely ro ha.v e been
C. imico/a as it I~ by far the most abundanc species collected
The successful transmjssion of an insect-borne virus from around Onderstepoon. He is reported to have similarly in -
an infected to a susceptible host is dependent upon the fected a horse \,rjth African horse sickness virus (AHS\/l by
complex relationship that exists between the virus, its insect CuUcoides bite.334
vector 8Jld the vertebrate host. with each being influenced Th~e seminal findings l:Jy Ou Toit that Culicoides are In -
by particular en,ironmental conditions. 152 Four criteria de- volved in the trMsmfssion of orbrdruses were later con-
termine the competence of a particular vector species:6 firmed elsewhere in rhe world. i.e. in North America.
• the ~rus must be recoverable from field-collecred ar- Australia. anc;I rhe UK, and then again in South Africa.329 The
thropods whose abdomens are free of fresh blood, laner study involved C. imico/a. !n the laboratory,329 C. imi-
~ the ability of the arthropod to become systemically in· colawas fed, through latex and chicken skin membranes, on
fected by feeding on a ~raemic host or an anificial sub - blood comaining nn1 serorypes 3 and 6 and AHSV sero1ype
stitute muse be demonstrated, l. After an incu bation period of 10 days at 25 10 27 ' C, the
• similarly. its ability to transmit the infection biologically infection prevalences for C. imicola for Bn' serorypes 3 and
by bite muse be dernonstraccd. and 6 were established at 31 and 24 per cent respectively, bur no
• there should be field e\~dence confirming the associa- AHSV could be recovered.
tion of infected arthropods with diseased vertebrates. In 1998, in th e laboratory. a second species of the lmicoln

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 Wctor.s: Cul/coides ,pp. I 03

comple:<. C. bo//ci11os, was incrlmlna1ed as a vector of most experienced person. Funhennore, 10 per cent of African
BTV.335 SignificantI~·. for Lhe Lhree serorypes ofB'f\' tested. Culicoides spp .. e.g. C r<11111s, lack a wing panem. and for reli-
higher infection pre\'alences, and higher vjrus tilres per able identification such species (indeed for all spt>cies in all
midge. were found in C. bolflinos than in C. imicoln. In complexes) must be dissected and mounted on glass slides
addition. ii was shown that these higher infection preva- and ex-amined by light mkroscopy at 100 to 400 x rnagnifica-
lences and mean virus conceturations per midge in C:. l)o- tlon. In slide-mounted speomens identification of the remale
lirinos were not influenced by the incuba1ion period (2 10· Culicoides to -spocies level Is based on the precise shape and
20 days) . nor by incubation temperature 00 to 30 •C) at number of thespermathccae (Figure 5,4), and the shape of the
which the midges were held.281 These findings suggest that third palpal segment and 1he distribution of u1e ser1smae on it
C. bolirinos may be a m·ore competent \'ector of Bl'\' than C. \Figure 5 .5). the conformation of the space between the eyes.
imicola. In a subsequent stud/32· 33·' three more species of and \\'her.her the chitinous areas between the ocetli are, or are
C111icoides were shown to replicate BTV, and so broadens not. adorned \\1th hairs (Figure 5 .6). Perhaps the most useful
our understanding of the epidemiolog)' of BT. In the same taxonomic aid for the identification of females to species level
trial the workers were unable to demonstrate virus infec- is the precise number and arrangement of each of 1he seven
tion. after incubation, in 14 of the 22 Culic:oides spp. exam- types oi sensillae on u,e amenna (Figure 5 .7), but unfonu-
ined. naiely. in the majority of iaxonomicsmdies published on Culi-
Artificial feeding of 17 s pecie~ of field-collected Culi- coides world1,'ide, these arrangements are not complete!)'
coide,< on blood containing three serotrpes of AHSV suc- enumerated or described. This arrangement and number of
ceeded in inCecling only C. imicola and C. bo/itinos.321 The sensillae on cbe amenna are also very useful for identification
findings suggest species-spec;ific serorype dependence. It of the males :o a species level but they are also a neglected
appear~ cogent that 1,0 AHSV replication could be demon- source or taxonomic infonnation (Figure 5.8). The shapes of
strated in any of the remai ning 15 species assayed, but for the various pans of the male genitalia are. in addition. highly
most of them only low numbers of midges were available. It species-specific and are always used in identification (Figures
was sho1,11 also that of seven species 1.but nol including C. 5,9 and 5.10). Unfortunately the preparation of such mate-
bofitinos) fed artificially on blood infected with the Bryan- rial on glass slides for taxonomic study requires exten$ive
ston serotyp e ofEEV. onlyC. imicola (22,3 per cent) became dissecrion. which is both laborious and time-consuming
infected.aili but is essential if chc small differences that exist between
Available data clearly indicate thal considerable varia- species are to be discovered and used in descriptions.
tion, in terms of susceptibilil)' and prevalence of infeccion Whilst the more modem approaches, such as mtDNA ex-
with different orbi\~ru~ species and serO!)'J)CS, appears to traction and RAPD ·PCR. ha\'e recently been employed to
exi~t amongst the various Culicoidesspp. Species of the tmi- separate species of the Jmicola complex.290 ir is 1101 fore-
cola complex appear to be the C1tlicoide.( most susceptible seen that they will ultimately replace the C'heaperand more
10 infection In the laboratory, and are therefore the likely rapid morphological method ofidemification. Rather. they
vectors of these orbiviruses in the field. Other data, such as will be harnessed to solve the more elusive issues sur·
high levels of abundance. seasonal and geogrophic pre\'a· rounding the identification of particular species within
lence, and fieid isolations of virus, support this contention specific vector complexes.
(see below).
Taxonomy
Historical perspective and problems Nearly LOO years
Biosystematics of Cu.licoides
have passed since the first Culicoides was described from Af.
Most Culiroides haYe a ,,ing pattern that is composed of grey rica and still no regional monograph has appeared that
and white spots that ls unique to each species. Although this treats the Afrotropical fauna as a whole. There are. hO\\'evcr,
panern may vary only subtly between species. and so compli- a number of taxonomic publications from South Africa,-•
cates the delineation of intraspecific varia1ion. ii remains an ex- 131 Angola, 62 Kenya 1~2• 181 and Nigeria.37 There are also

tremely useful aid to the identification of the midge. The landmark studies that deal with signlficam pans or the
patterns can be fairly easily observed under a dissecting micro- fauna such as those by Carter, Ingram and Macfie,h5 Clast-
scope; those of the 14 species of Cuficoides mos1 conunonly rier.71· 73 Clastrier and Winh.74 Cornet. 80 Comet and Cha-
found assodated wiu1 livesrock in Africa appear in Figure 5.3. teau,82 Cornel and Brunhes,81 Cornet and :S:evill.83 Comet,
As is ~videm in the ligltre, Lhespecit!:> ar~ 4uile easily separ.tuh, Nevill ancl Walker."" de :l<tcillon, 100• io; houa and Comet, 171
on wing pattern but this is because they are represemarh-e orsix Meiswinkel and Dyce, 22·1 :-1eiswinkel,?13-?IS. 21 • and Ne,ill
subgenera. It is \\1thin subgenera tJ1at Lhe patterns become and Dyce. 269
more broadly similar. but what hampers rapid. easy identifica- The taxonomy of South African Culicoides for the larval.
tion is that subgenera can often comprise li\•e or more species pupal, and adult 1nale and female stages is shown In 1he dia-
complexes. II is within complexes that identifications based on gram in Figure 5.1 1. Only where the rele\·ant areas are col-
wing panems alone are unreliable. even fn the hands of the oured in Lhe clio.gram has the particular life stage been

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JO~ "'1£1lOs (l\~ A~pecis Influencing tho occurrence or lnfectlou.< dis<ia«1s

r .;.,r· · I . ..
,,. <. ~u..,
. ~
. ..
>, -
~ __ "')':-
....

:~~
a n

~ .... .... . . . ~ -
..;:;::'-::~--
...,__ _,<,
CT ..
~
... ~~-~~
- -

•
~ - ...:::~-)
.
~ :--
.. ..::.~
" ~..__,>, ~-~~!~~~ ·. '
-- ~ ·-....·- "'~"?.:;.~~~~
. "'~ -. . ~~
..... ·,--~~
. -~~ ~~;;1
-~
-~.
,~..
,~ ·
... ~~-:
' ·,-•". ' """'
~~~~~-
;
~":-~;. ~~,~
'' ,~.
. ,.:
·,-;.,_., .
·.,~·
.........
..
~.:
·~~

--~
,.,-
,,

...,""-- liii
! ~ ~ -'- --~
. ~-
- ---:,. ' ~
.,, ~"!;~~
~-
d
k

~- ~
.,.,
-~--~~ ~,-,. ~

~ .. ..

figure 5.3 A wmg o! each of e

the 14 species of Culic11rd8S
rr~t commonly round
assoc1aied wrth li\'e}tou rn
soLMem Africa
a= C imicola!limb. SJ
b~C. bolhitl0$ (Afr OP32)
c= C. zu/uensis(Ol'6SI
d = C. magnus !limb. 11 I
e = C. gufbe/lk1e.n/fA!r 0?31) m
I = C. fufvitbora:ttl imb. SJ
g--C s;mifisjAfr. OP34) ,-_,. .
h •C. 1rop/cs/is(S.Alr.8I ~ . i f - - - ~
,~ C. neQvei(OP1 7l . · "-., -...::.
1• C. st;/lu/lle.ifAfr 0?27)
k = C. engul;and~i !Zimb 14)
l = C.bedfordi(S.Afr. 2'1)
m =C pycno$/iC111S(Afr. 9)
n =C /eucostiCIVS (SAfr. 10) g n

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 Vectors: Culfce1itfe3 spp. l05

,-.·" fi,)

,, '• .,,....
. "
...... \
\I
.......__,,. ~
onderstepoonensis

I (_;- -· -
' ·- '.::,. i ftr ' .,.....

walkeri
/1

.,.. . Figure 5.4 The spermalhecaa of 1he

femares of three South Afrn:an species
mgenaa
of Culicoides

described and published. Yet undescribed species are num- parate collecting b~· early French, German and English
bered in the diagram: 1hey comprise a third of the 112 spe- workers is 1hat the holot)'Pes of many of their species are
cies collected over 1he las1 20 years in South Africa. The today either lost or are scattered amongst a number of Euro·
diagram shotvs too that the immature stages of a1 leas1 pean museums or research institutions; or the holotypes
iO per cem of the species still await discovery and descrip· that are extant most are in a poor state of preservation. Ex·
tion. Annually, two or lhree species are added 10 this list.216 acerbating maners for practising sys1ematis1s 1oday is 1ha1
and if this Is representatlve of the situation In the rest ofAf. mos1 early descriptions were 100 brief. were based on a
rica, it illustrates the amoum of taxonomic research still to single specime,n (usuallr female). and were often no1 ac-
be done on A.ftocropical C11/icoide.t companied byillus1ra1ions. This means that many character
The first Afrotropical species described were C. sclwltzei traits. crucial 10 identifica1ion and assignation of a species 10
(Figure 5.3j) and C. herero from i':amibia in 1908: 122 by 1925 the correct subgenus or species complex. are mostly lacking
nearly a thlrd of the 156 Afrotropical species recognized from these early descriptions.
today had been described. An unavoidable result of 1he dis- One of the mos1 prolific sruden1s of African Culiccides was

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!06 <en10~ o,~: .~~pects lnfiuen(lng the occurrence of infectious diseases

-
0

b
Figure 5.5 The eve-space (al and five•segmeoted palp (bl of a female Figure 5.6 The eye-space (al and five-segmented palp (bl of a female
Cul,co,des imicola The third segment nears a sensory pit Cuf;co,des tavtJs. The third segment bears a sensory pit

the French abbe. J.J. Kieffer. who described 20 species be- holotypC'S were disco,·ered in the Paris Museum 60 years
tween 191 J and 1921 from ctiverse localities that included later. 191
Lhe Se),chelles. Kenya. the Cameroon (most species), Sudan Contemporaneously (1919 t0 1925. 19-17). 21 species of
and South Africa. To a great extem the research ofKieffer 182- Culicoides were described by Caner.&; C.aner. Ingram and
18~ has remained esoteric. For example. 14 of the 20 Culi- ~lacfie,65 Ingram and Macfie,167- 169 and Macfie.Z07• 208 ~lost
coides names made available by him are 001 in use today of their mat.e rlal emanated from Ghana bm also Included new
even though the species be desc, ibed are probably com- species from the Sudan and Malawi. The taxonomic studies
mon. Hil> ·aoating' Species are a/bosparsus. bisigrwm.t. of Carter. Ingram and :vtacfie were of a high quality and, im-
de111a111s, g11i11ee11sis, kribiensis, nilogenus, nilophil11s. ponantly. included the first descriptions of Culicoides larval
octosigT1arus. quadrisignatus, remows. sig11a1us, siluesrrii, and pupal stages, and their habitats. Influenced by Ingram.
rrisig11ams aud xamhogaster. Indeed. the names of a funher De Mcillon (J936 and 1937) 100• 101 commenced the first bio-
three species, rropicalis, imicola and alticola, had suffered a logical studies on South African Culicoidl!S that culminated
similar fate. bUl were resurrected when their respective in the description of ne1v species and their pupae. and the

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 \'cctors: C11/icoidesspp. lOi

Sharp-upped trichsoea

'~!/JJ IJ J Basiconka
Long, b!unHlpoed [rlchodea
I
\~-- l,:ij}.,' , Chaet,ca with alveolus
\ 1~ \If¥_ ./ Short. blunt upped

~\r-~ ~:· ] . / ~ mchodea

I
Coeto~omca

\¼;
1

\\"··ze( I\\ d/ / -
.
~t#l \ ·~-'
~~;/t/Jl'::~ I /
"'"-l''.u '~ ·./;,' / ~
~~
t1- \ .--1-./ I ~ 0

~,\t/Jj1) \ ,=) . J
"""' " 1/ ~
~J~~ l .• ;
1i,/ v'./
\ ~\'~0/ \\~ ={~~/// /
,. -~ ($ /
'~;\~~ /
\'J>~
' ·;.
"\.:,,r-----

Figure 5.7 The varioi.s ~ns1llae that occur 011 ne d,s,al 13 flagellar Figure 5.8 The vanous sens,lla;; !hat occur on the disial 13 ilagellar
!egments rantennomeres'I of a female Culicoitfes segments of a mate Cul1Co1des

first observations on breeding sites. Unlike the situation example is C. pal/idipennls, which was subsequently shown
with Kieffer, many of the Carter. Ingram and Macfie species 10 be a synoni'l11 of C. imicola Kieffer, 1913.
names, such as similfs, accraensis, bmcei, pycnosricms. Besides tl1e possibility of funher synonyms the most
prmctirliorax, nigcriac and bedfordi. arc in uso today. How- vcidng taxonomic problem to remain is thut most oflhc spe-
ever, it must be noted that, like some of those of Kieffer. a cies memioned above represent species complexes. the
number of their species are also borh rarely recorded and most notable being the Accmensis. Grahamii and Nigrlpcn-
poorly Lmders1ood. These include adersi. ausceni. n.is complexes. Each complex is represented by l Oor more
DC'hrochom:i;, eriodendroni. inornarip1mnis, 11igripem1is, species that are rarely collected and are very difficult to
citronem. clarkei. (011f11s11s. mtilus, arenarius, corso11i and identify. One consequence is Lhat precise delineation of the
lambomi. One explanation is that rhe couJ1tries from which nominate species of each complex eludes us to th.is day.
most species were originally described have since only been which means that the description of new members of these
rare!~· sampled for Culicoides. More co6ent is thar these au- complexes are either held in abeyance or are being
thors collected mostly in the high-forested tropics. an area published asC. sp. It;\, B. ctc?7 or are given numbers. 142• 223
harbouring only a panicular subset of 1be Afrotropical Cull- Ultimately these systematic problems require resolution as
cofd~ fauna, and one that by all accounts appears robe rela· each species within a complex fills a niche chat should be de-
tively depauperate. It is pertinent that Kieffer and Carter. fined as a first step towards estcibUshing whether it includes
and Ingram and Macfie described mosr ofcheir speeies from the transmission of pathogens harmful to livestock. The
the equa1orial coasral rowns ofKribl (Cameroons} and Accra range of niches is almost infinite and includes differences in
(Ghana) respecti\'ely. It is thus likely that these authors. who geographic distribution. seasonal prevalence and abun-
seldom referred to each other's work, have to some ex'tent dance. hoSt preferences and vector capacity. Evolutionary
described 1he same species. An obvious and significant radiation u1to a particular subset of ecological conditions is

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108 >JSC\'IQ,' o:-r: t\spcets influencing 1he qccurrencc of lnfocrious diseases

--
N
E
E
c:i
'
~'-
'
l ',
•
;

I
••\
'. l
i
'/
lE
~..,E
. c

'

a ~"'r"~
_,.I
a
I
,,, {;>!-ergum IX,/~

;
Paiamere tI ·.1··'
I
• • • 1I
''
\
./ ""·-_,. /

I
•
r··
'

I
---
,,

'
•
b b

Figure 5.9 The emire mate genitalia (al and tergum IX, aedeagus and Figure 5. 10 The entue male genitalia (al and 1ergum IX (bl Qf
paramere (bl of Culico1des onders1epoonens1s Culico1des irmcola

a hallmark of ¢peciei. rr,mprising rhe recently discovered their pn•sible involvement in 1he 1ransn1i!,sion of /\lcahanl'
Imicola complex, and because chis complex contains the and BEF. Blue1ongue virus bas also been isolaced from
most important transmiuers of orbidruses known today in C. g11/be11kia11i, another species of Avariria, which, like C.
Africa and beyond. it is discussed below in greater de1ail boli1i11os. breeds in the dung ofcatde and is therefore closely
with special emphasis on the two vector species C. imico/a associated with livestock. However. it probably plays only a
and C. bolitinos. minor role as it seldom becomes abundant and is restricted
in hs geographic d istribution.
Vector species In addi1ion to the three livesmck-associa1ed species
Of the more than 1 210 species of Culicoides tha1 have been mentioned above, there are seven species of Avariria par-
described. abouc 70 belong in 1he SL1bgenus,li•11ririt1.At least ticularly closely 3S$Ociated with larger African herbivores
30 species of this subgenus occur in Africa; of these nearly such as the elephanr, ho1h rhinoceros species (Ceraiorhe-
half remain undescribed and so would bring 1he world total rium simum and Diceros bicomis), buffalo. blue "1ldebeest
to 85. Worldwide a mere IO species of Culicoides are in- and the plains zebra (Equus b11rchelli11. Not only do these
\·olved in the transmission of orbiviral diseases that affect Auariria spp. reed on these hosts but also use rheir dung as a
livestock; fuUy half oflhese belong in.4varirla. The mo51 im· larval habita,.•1. 118· 21 :i. 213 · 223 · 261. 207 ln the tase or C. boliri-
porta'nt vectors in Africa are Av(lrilia spp .. namely C. imic:o/t1 nos, it ha~ broadened its resource range 10 now use not only
and C. bolitinos. Both transmit BT and AHS {Table 5.2). 1i; buffalo but also cactle dung as a breeding medium. This has
and are the suspected vectors of EE, but less is known about enabled it to maintain a widespread prevalence in Africa.

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 Vcc1ors: C11Ucoitlesspp. 109

which is in contrast to the elephant-associated C. loxodon- man.36 who noted only one difference between Mediterra-
cis, which remains restrlc1ed in its distribution presumably nean and South African populations. This difference was
due to its inability to invade other dung types. With a bur- due ro an error that appeared in a publication of
geoning game farming indt1SlJ: in sou thern Africa. !he Meisv.1nkel. 213 and so confirms that C. imico/a semu siricro
translocation of wild herbivores can be accompa11ied by the is morphologically homogeneous across its large geographic
establishmem <>f pathogens to livestock in hitherto ·clean· range; Lhis homogtmicl11 being suµpuned subi,equcntly.220
farming areas. For example. zebra exported from Namibia The indcpcndemspecies status of the ,·arious lmicola com-
were indub imbly lin ked rq the ou1breaks of AHS that plex ta.,;.a sensu Meiswinkel217 was recently validated by
occurred in Spain becween 1987 and 1991, in which several RAPD-PCRanalyscs:28:l DNA (RAPD) patterns for seven spe-
hundred horses died or were destroyed as a result.288 In cies of the complex are shown in figure 5.l2. ln Figure 5.13
this instance C. imicola played the principal vectorial an unrooted neighbour-joining consensus tree shc,ws the
role.zi~ Sttbsequemly. it was confinned in South Africa relationship and discrete clustering of 19 popularions of
that cycles of AHSVoccur in zebra and tha1 this can pro<!eed these seven species collected in South Africa. Botswana.
in the driest of en\"ironmems wh.e n Culicoides populations Madagascar and the lvory'Coast. Until 19i2 studies on the
are depressed. 13· 217 There is also evidence thac where taxonomy and biology of C. imic.olt1 sensu seticto appeared
new populations of wildlife are establlshed on game farms. under a variety of names that include C. pallidipennis, C.
the associated Cu/icoides spp. will also become established iraqensis. C. mitwlltS and C. pseudomrgidus.
in the long term, and in th.is \\'a}' new vector foci are
created.216 Vector compe tence Incriminated in 194-1 115 as a \'tCtor for
It may not be circumspect 10 single out species of the BTandAHS. C. imicola is still regarded as the most important
subgenusAi:aritia as the sole vectors of orbiviruses in Africa Old World vector of these livestock diseases. It is also consid-
as potential candidates from other subgenera have been dis- ered a pc,cemial vector for BE. 328 Depending on the envi.ron·
cussed and listed in the lherature.330 Ni noted above there mental temperature, the extrinsic incubation period of BTV
have been sporadic isolations of ,1ruses from a nwnber of in C. im ico/a is 7 to I 2 days.50 • 115 More detailed information
otherspeci.es (Table 5.2). ln addition, in laborntory inrection on serotype-,pecific infection rates and virogenesis is given
studies. 335 evidence has emerged that more than two spe- abo,·e in the section on 'Vector competence'.
cies of Culicoides may be capable of replicating STV; chis is
the first indication that there may be other vectors of greater Host prefe rences Culicoides imicola has been found to
or lesser imporcance in the field. One of the strongest candi- feed on cattle, ~heep. horses, pigs, goatS and poultt,•.42 • 45• si.
dates is C magn11s. which is a member of the subgenus C11- 3 7• 16:1, ZGS, 26~. l.'16, 3 19 In Israel It is inclined 10 bite cattle and
/icoides and which can at times become quite abundant in horses along the bockline; in cattle it prefers feeding in the
the cooler cemral and southern regions of South Africa. It darker coloured areas of the animal. 13 There are indicarions
has been recorded \\idely from t.he continent. and is closely that C. lmicola is somewhat exophilic in its feeding habits
related to the European Pulicarls complex. which is also sus- showing some reluctance 10 enter buildings221 but these re-
pected 10 play a role in the transmission of orbhiruses. 233 quire ,-alidation in situations where ve11· large populations
Below is summarized what is known about C. imicola abound. As regards wildlife. eight females have been col-
.ind C. bolirinos as regards their taxonomy. geographic dis- lected off a darted African buffalo.213
tribtuion and sea~ouality. vectorial competence, host pref-
erences, gonotrophic cycle, dispersal, and associaLion with Gonotrophic cycle It has been esthnated3 :lll that. \\ith a
outbreaks of disease. In Table 5.3 the broadscale Ian-al habi- gono1rophic cycle of four days. C. imicola females might
tat differences between these two vector species are com- take five blood meals during their lifetimes: it was similarly
pared. and the resultant impact on adult disrribution, calculated subsequemly-~0 that the mean period between
prevalence and abundance indicated. blood meals was 3.3 to ~.6 days. In lsrael 49 C. imicolt1 was
found to complete 11 generations per annum \\ilh each gen-
C11/icoides imicola l(jeffer, 1913 (figure 5.3a) eration being approximately 23 days In duration. II ls gener-
Ta-xonomy As a taxon. C. imicola is most closely related to ally accepted that under warmer environmental conditions
C. 1111dipalpis Delfinado. 1961. the latter known onlr from the life cycle lsshonened. and thus there is a greater number
south-east Asian islands east of the Wallace line. Cullcoides of generations and adults produced per annum.
imicola is a member of the lrnicola complex. which com-
prises 10 species. all restricied to the Old World; eight of Larval habitat Using emergence t.raps, hundreds oi C.
these species occur in Africa. Detailed smdies have done imicola were-first reared at the Onderstepoon Vetennary ln-
much towards claril)1ng their taxonomy. biology and distri- Slitute in 1950 by du Toit and Fiedler260 in a swampy kikuyu
bution.21 ~21s, 220 The complex was re1~ewed in toco In grass (Pe11nise111111 cla11diste1111111)-covered area next to a
1995.217 The ta.xonomy of C. imicola in the Mediterranean leaking cemem dam. Nevi11 260 was the first to show chat the
region was comprehensively reviewed in 1991 by Boor· pupae are susceptible 10 drowning as they are unable 10

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110 SEcnox o'<c: Aspects influencing the occurrence of infectious diseases

. 'i
D c1
D Pupa
D Larva

Figure 5.11 A diagram oi the iaxonomy of Sou1h African Culicoides for 1he larval. pcpal, and adult male and female stages

.~
C:
C:
Q)
.9,
,2
"' ., ... ==~
.;
.!!>
-~
0
0
.s
~
0
.Q
C:
-
%
.g
0 .. ..
0
M
0
~ ~
b
:,
.,
-
.!;? ci
Q)
.§ 0
.Q C: ~ ci. Q.
ti ti ti 0
II)

<.i
"'
0 0
M M

... """
-- 1 000 t:p

--w --- - - S00bp

Figure 5.12 Representatwe individual random amplified polymorphic DNA (RAPDI pattern~ for the seve~ morphological species oi the Qiticoides imicola
complex. using me Prlll1t!r D17: C. im,cola ss from Ondersiepoori. C. bolitmos irom Rn1noce•osfontein. C. m,ombo from Maun. C loxodon11slrom
Phataborwa, C. sp. 0 30 irom Pon Edward, C sp. #107 from Mabula lodge. and C pseudopel/1d1pennis from Abokouamekro. Ivor,, C0o--s1

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 Vectors: Culiroides,pp. l lI

Table 5.3 Broad-scale la,vai habitat differences between Cullroides imicola and C bolit,nos. and iesullant im;>att on adult distribution. prevalence and
aoundance216

CUL/CO/DES IMICOlA CUUCOIDES BOUT/NOS

IMMATURE STAGES IMMATURE STAGES
- Inhabit moist soil that are organically enriched but not wateilogged; - lnhab,t exclusively the dung oi herbivores tncl~'<ling canle, Aftican
predom,runc in moisture retentive clovoy soil~ wh,ch moy be buffolo ~nd the blue wildsbeil$t
unvege1e1ed or covared ,n short grass as in pastures - Following depc.smon oi eggs by adults 10 fresh oung Ille feivae requ,re 7
- Absent from qu,Ck-drain,~g. micronument-l](!Cr. sandy coastal soils to 10 !la\'S ,o reach maturity
- Upper layer of soils must remain moist for a minimum of seven davs to - Will be found h any terrain in which the above herbivores may occur.
sustain larvae through four moults: this resrrrcts the species. at any irrespect,ve of degrees of landscape slope
alt,ruae. to flat. low-lying. slow-draining areas - Achieve great abundance in areas whP.Je dung 1s l!epos1ted on sandy
- Numoers drop d1amat,cally in sloping terrain likely due to water run-cfl so,Js
and lsaching of soils - The la.-vae. beil\g coproµhihc. are independent of soil mo,s,ure and
- Ou11ng episodic psnoos of above-average preti;:i,atfon C 1m1col;1will Qualny, can 1h~s occur 1n txlth flat and slop:ng 1erra1n
disoerse and increase 1n numbers as moist so Is oee-0me more widely - Average annual prec1p,tat1on does not apµea, to affect its geographic
available distnbot100
- Restricted to 300 lo 700 mm rainfall isohve?s as soils become leathed
of nutrren,s at higher prectoitauon levels ADULTS
- Almost as widespread; on ave<age one Older of 11'.agnn~cte less
ADULTS abundant ,han C. imicafa and thus very rarely found in cc-dominance
- Mosl widespread and abundant Cu/1coidessp3Cies in South Africa. - like C. rm1cofaa1e catholic ,n their choice of hosts
ieeds on av.rlde variety of domestrc~ted livestoc;1' - Adult abundance vartable but orooonional to the amount of dung
- Show large variauon in abundance; 200-fold inaease in years of avadatte in t1.1·n dictated by ammal b1omaSS/un11 area
above-average :ainfall - Largest single •ight ltap collectton recorded' > 3 000
- Largest single tight uap collection recorded > 1 000 000 - Currently ..nde-staod to be reSlricred to sub-Saharan Afr;ca. recorded
- Widespread m sub-Saharan Africa. and on offshore islands. also occors from only eight continental c®ntnes {South Africa. Lesotho. Zimbabwe.
1n North Africa, ana thence up to the 43ro parallel in European countnes Malawi. Kenya, Nigeria. Ivory Coast and Gambia). an:i from the islands
bordering the Med1tei·anean Sea: occurs also in the Near Middle and of Mau11tius and Madaga51:a1
Fa; East up to southern China and V1elJ1am

float on the surface of watec as do many Culicoides spp. bers peaking in late summer and amunm precisely when the
pupae. The requirement for a semi-moist larval habitat by C. numbers of adult C imitola are at their highest. and when
/micola appears 10 be a characteristic shared by all species of diseases such as BT and :\HS are most prevalent. In another
the subgenus Ava.dria and. for C. imicola, was partly con- s1udy2&7 1ownumbers of C. imico/a were reared in dry as well
firmed in lsrae1• 7 where it was reared in rich mixrures of as in wet ki.kt1\'ll paddocks especially where the son \,as clay-
organic mnuer and water-saturated soils. but despite this it like and medium moist. ~lidges were also reared in continu·
was also the species adapted to the driest breeding condi- ousl)' moist area.~ where Lhesubstrate consisted of sandy soil
tions. The breeding habitat of C. imicola in Israel has been covered by vegetation and a layer of organic matter. and
summarized as being 'in and around animal pens in water where the moist but silly soil of irrigated pasrures had very
irough overflow. at the margins of animal sewage and drain- little surface organic maner except for some rotting plant
age canals and puddles created br leakage from water panicles and,or diluled manure. It is now fairly \\1ell estab·
pipes. '33 In Kenya:i.10 high concentrations of larvae were lished that C. imicolt1 is mosr abundantly encountertid where
found in mud and dung mix.lures su.rroundi.ng caule pens lhe soils are clayey. such as the black cotton soils at the
and in associated effluent ditches. In Cyprus236 C. imicola Ondersrepoon Veterlnary Institute and those of the Laikipia
was found breeding where there were leaks from irrigation plateauand the Masai Mara in Kenya. bm it is vi nu ally absent
pipes. These were small seepages with little free surface where the soils are overly sandy. 222 It is postulated that sandy
water. They were generally drier than the type of habitat pre- soils, in draining 100 quickly of moisture and being nutrient -
ferred by other species of Culicoides and were usually covered poor (especially those found along the coast). are simply un -
by a growth of fresh grass. There was aunost no contamina- able to sustain Lhe larval stages of C. imicola for 1he required
tion ~y animal droppings. but organic matter was present in 7- to JO-day developmental period.218• 219 The proposed rela -
the soil in the fonn of decaying vegetation. In Laos163 it was tionships between soil type. prevalence and abundance of
reared in mud mixed with dung. In an unpublished srudy C. lmicola. and the prevalence of AHS during the 1996 Soulh
conducted at the Onderstepoon Veterinary Institute up to African outbreak of the rusease is illustrated graphi~ly in
382 C. imicolaf mlwere reared in islands of short kikuyu grass Figure 5.14. It is probable that the rearing record~; 4 . 161 ofC.
in the middle of effluem-enriched drainage ca nals.215 They imicola, documented under the name C. pallidipcmnis. from
were. in addition. reared throughot1t the yearwilh their num- rotting banana and pla111aln stems do not refer to C. imicola.

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112 Sfl."TIO)< o,-r~ Aspects influencing the occurrence of infectious diS<>ases

, - - - - - -R
! h\nocerosfontein
0.05 59
r--"-'-1
~ - - -- Pen Efizabelll

I ,--- - - -Pon Edward

C be/it/nos

l . . . _ _ _Tamatave
(Med~scor)
, - - - - - -Allokouamekro
C. pseudopofJ/dipenms
- - - - - Maun (Botswana) c .miombo

___ ___,, - - - - --Kimberley

94
'------Port Edward
I c. S? 1130

, - - - - - -Mebula IJxl_ge
3
'---"'--I
C. sp. II 107
~ - - - -Etf,sras

- - - - -Skukuz:a
S7 OnderS!epoort I
53 ' - - - -- - Tshipese
C. 1m,cola ss
' - - - - - -Ph.ilaborwa

'--- -- -Mtunzml

, - - - - - - - - - -Mas,nga
97 (MadagaSC3t)
, - - - - - - N'Oyala Elllsras

- - - - -Phalaborwa C. lo><odorrtis

'------Nsha¼v

Figure 5.13 Unrooted neighbour-joining (NJI consensus tree ol 19 populationsof the seve1 morphological species of the Culico/des imicolatomplex.
based on tile d,ss1mi!a1,ry value~infened from i96 RAPO fragmen,s. amplified using :ouraibiUary primers. Onlvbootstrapvaluesg1eate1than 50 per
cent obtained from i 00 bootstrap resamplings of the original dataset are shown

lmmaturestages Contrary 10 a published sta1emen11·12 and of C. imicola been smdied and mapped on a national
the questioning of their possible identlty, 38 the descrip- scaJe.20• 23• 392· 333 On a smaller, but more intensi,,e scale are
tions260· 262 of the larva and pupa of C. pallidipe,mis are in- the area-based studies in South Africa. of the Western Cape
deed chose of C. imicola and not of C boll1inos. A study of 1he Proiince,267 eastern Free State Pro11nce,330 the Kruger Na-
pupae of five 1axa of the lmicola complex ha~ revealed that tional Park and adjoining areas, 21 • the greater Onderste-
1hey differ markedly in morphology from one another. 268 poon area in Gau1eng Provi nce331 and the Pon E.lizabech
area in the Eastern Cape Proirjnce.218 and in Lcso1ho.3Jo In
Geographic distribution Cu/icoides imicola was originally addition, its abunda11ce and prevalence were derailed dur-
desc:ribed from a single female specimen collected at Th,~ ing an outbreak of AHS in the cenrraJ and east em regions of
on the southern coast of Kenya. Today, C. imicola is kno\vn South Africa.219 These studies all took into account the exist-
10 be one of the most widely distributed C11/icoide$ in the ence or sibling species and in this way do not conflate data
world and is found in at least 35 comuties throughout Africa. from various lmicola complex 1axa. They also demonstrate
in most coumries surrounding the Mediterranean Sea. and the extreme variability in the prevalence and abundance of
in the Near. Middle and Far F.ai.t a~ far as ~0111h11rn r.hina. C. imirnln in the vicinhy of Jiveswck ranging from it being
Laos and Vietnam. Only in South Africa, Morocco and the totally absent 21 8 to \\,idespread and superabundant.219 In
Iberian Peninsula has its distribution been mapped in some each s tudy an auempt was made to e~plain this variation in
detail.214. m. 33z terms of climatic or edaphic factors. Ir is emerging from
these studies that other than extreme cold and aridity, the
Seasonal abundance and prevalence Only in Somb Af- degree of topographic slope (inducing water run-ofl) , soil
rica. Morocco and Spain has the abundance and prevalence type (whether drainage is slow or rapid) and soil fertility

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 \'cctors: Ct1/l(n/dt•J spJl. 113

revealed b~ weekly collections. made over a full year at -10

sites spread throughout Somh Africa. are discussed in
greater detail and the various implications of the findings
assessed. At the periphe'1 ofirs range in the :-.liddle and the
6
Far East the disLribution and abundance of C. imicola are
mo:,tl) unknown.
5
Dispersal Under normal conditions Lhe flight range of C.
imico/a probablyvarie~ in the range of a few hundred metres
10 a few kilometres. Clearly this is dependent upon the local
availability of hosts for feeding and larval ~uhstratcs for
0
s breeding. It is gcnerall1· accep1ed that Culicoirle.s are rduc-
s tam to fly if the wind is above a certab1 sp!.'ed, which is when
numbers caught in light traps diminish. but it is apparent
m s too that light traps become increasingly lncfficicm as ,,ind-
ss u
2 s spceds increase and do not then accutately reflect biting
s
gg midge activirv rates.1! 1• " ' There is rnmpclling evidence
that C. imico/11 can be transported upon prevailingwmd!I for
hundreds of kilometre~. This is considered the most likely
way Jn which outbr(luks of UT are being precipitated in
tc
0'---- - - - --------9- - ccCCCc coururiesaround the Mediterranean Sea.i9o brn It is difficult
localities to prove a~ the cland,,srine movement oi infected animals
Soil also merits consideration. Similarly. can the occurrence of
tvpe
C imiro/<1 on islands off the African east and wes1 coasts
CLAYEY SOILS SANOYSOltS (Madagascar, Reunion, :-.lauritius and the Cape \'<.<rde) be
moisture-•etent\e. rich qu,c~-draining, ooor explained by wind transport or were they moved toged1er
1n m,cro-organisms n m1Cfo·organ1sm~
with livestock? Nevenheless. the evidence to u11equ1vocally
figure S.14 A schema1,c rep1esentation or me proposed relationships suppon wind dispersal remains largely unsub,rantiated as
!letween soil tyjle. numb.era of Cvhcoides ,m,cola and !he number of cases the capture of large numbers of C11/icoid11s at \larious alri·
o' AHS during the 1995 ou1break ,n South Atnca based on 52 msect tudes O\'er sea or land has yet to be made. 1\nod1er, as yet
collett~ns at 47 s,tes Sttes we,e allocated :o ;en reg1on-gr011ps ,\ll1ch, unsubstantiated. mode of dispersal forC. imicola, is the pos-
mov,ng trom reft 10 righ: ;hov, adeoease ,n AHS cases and an ,,crease
tulation that C11/icoid/!$ may be moved over long distances
mso,1 sand content Reg,on-groups are: Kaalpla.s farm. Onderstepoon
in enclosed horse 1ranspon (loats. 219
\etennar, lnsutute 1kt Preiona (p); Johannesburg 11); sou1h-cemra1
MPllmalanga {m); eastern Mpumatanga 10\weld (IJ. KwaZulu·Naral [nl.
Free Sta,e and nonhem Eastern Cape Is): Graatt-Reinet area oi Eastern Association with outbreaks of disease Since Du Toil's In·
wpe tgl. Unenhage. 30 km inland from the soutrarr coast (ul: and crimination of C. imlcoln as a vector of both lJT and AHS in
routn-eas1ern Cape ccast {cl 1944. it i~ remarkable how few attempts han• been made to
isolate one or the other virus from this species during field
Olllbreaks of 1hese diseases. The \larious isola1ions made
(availability of micro-organisms) are additional imponam from C. imico/(1 (Table 5.2) were invariabl} the result of ran-
factor~ affecting che dlstribution and abundance of this spc· dom sun·eys for Culicoides-associated orbi,iruses and were
cies. ;\s to its detailed seasonal preval1?nce a1 various sites often nor linked to an ou1break of a disease. ;\ number of
throughout South Africa reference should be made to two these isolations cannot be linked to a specific species bm
srudiesm· 333 whose data were used In conjunction "~th emanated rather from mi.~ed Culicoide:; spp. pools. As far ns
various climate variables and satellite imagery, to model the can be established. 1hc firs1 isolations of AHSV from identi·
prevalence and abundance of C. imicota.21 v\11ilst rhe model lied pools of C. imirola nor containing freshly blood-
e:-.1,lained 67 per cent or the obscn·ed \'Orianccs it also sho\',S engorged females wen: made during the I 987 to 1991 out-
some anomalies which are probably the result of too heavy break or the disease in Spain.233 Four serorypes ofBlV were
teliance on 1he effectsofrab1 and tempermure. This h'3S fur- isolawd from C. imicola in lsrae1+1 before these dates. In Ar-
ther demonstrated in an unpublished preliminary African tica II is equally remn.r kable to note that only in 19!!6 was the
model in which C imi.cola was predicied to be most abun· lirs1 im·escigation made 10 establi~h which specic6 of Culi-
dantly encountered in equatorial Africa, i.e. precisely wht1re roides [Table 5.4) were mo~t abundant at stables in which
it is known 1hat C. imicola is \~rtuaUy absenL 171 · 211 In a ~p- losses during an exten~i\•e outbreak of Al IS we.re occur-
arate section (see be!owl the more precise seasonal and geo· ring.219 11 was clear that C. imico/a wns the most abundam
graphic di~tribmion of C. imicola and of C. boliri11os. as and pre\-alem i.pecics and 1hus. for the nrs1 time in Africa, it

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11 ·1 st.(."lm~ oxt:: Aspecis influencing 1.he occurrence or i11foc1ious diseases

Table 5.4 P~alence and ranked allt.!ndan?:e

oi tne 13 Cuhc01aes species mos1 commc~lv
ABUNDANCE CULICOIDESSP. C..OFAtl PREVALENCE ABUNDANCE
encounteted in So ligh, 11ap collections made
RANK I~ CUUCOtDES JNS6 LIGHT RANKm
at 47 sites dunno ire 1996 outbrea• oi TRAPS
African horsesickr.ess in South Africa l' 9 1 C. imico!o ~~.2 60
The last column 1s the abundance ran<1ng 2 C. wlue.'lSiS 3.0 4G 5
or 1.t-.ese spei.1es accm<{w~ tu an tlotller sw:Jv"1 3 C. magr.us 0.7 25 10
a C /J!Jfrtinos 0.6 34 15
5 C lellcosrictus 0,4 33 7
6 C. p,1:r.os1ic111s C.1 28 6
7 C. nivosus <01 15 8
B C mtlne, < 0,1 13 17
9 C. nea,lli <01 11 16
10 C ent!e1iein1 < 0,1 10 2
1i C subscnult,e <0,; 9 2
.·~
12
~ C onaemepoonensis
C gulfleni.,an,
<0.1
<0,l
9
9
20
1i

Table 5.5 Summary or blood meal ,aer.1,1,ca11cns tor C\JlicoiC'~scecies m South Alnca and Zimbabwe
CUUCOIDES CAffiE SHEEP PIGS HORSES CATILE/ 80Vl0S HORSES/ HORSES! HORSES/ MAMMALS AVIANS/ TOTA!.
SP Ref ~, Ref 5 ilE· ;• Ref ' SHtEl' Raf !: v\ntE SHEEP SHEEP• Rel.;· CHICKENS
z."! .:is~ mm 5.• :~ ~
' ~ Re~ Rel 5' Ret CAffit Ret ' "
,a,a Rei 11 21 : .!1i: ."Ji
~1

C. im1cota 247 459 27() 371 l; 1 11 1 16 126 9 1 621

C milnei 4 7 120 2 16 18 273 ~~l
C sp.near 16 139 58 n 8 38 336
g121Jr,penms
C. fulvlrhorat 167 s 5 105
C wtuensis 11 15 74 7 40 181
C leucosr,ctus 5) 5·
C. oodfordi 40 40
C. mag11us 17 14 6 1 39
C bruce, l 2 20 4 3 31
C pycnosricrus 1 1 2 2 i6 22
C gulbenkiam 1O 5 2 2 2 21
C bolitinos 10 5 5 21
Sehultze1 group 16 2 ;g
C. coarctarus l 2 4
C IB'/t/S 2 2
C niVOSIJS 3
C engubande, 2
C nelJ',ei 2
C s1milis 1
C enderleini 1
To;al 332 687 828 i 16 42 2 16 199 ~02 3053

was posid,·ely linked geographicallr and temporally to an phase El :,;ino weather e\·ents.2? These usuall) bring above-
outbreak of AHS. El'en in this inwsligation 1\0 attempt was average precipication and iL is this factor that strongly points
made to isolate virus from identified pools as the age-~rad· to C. imicola ha,ing been principally involved in past
ingand selection of non.engorged remales from the million$ epidemics of the disease. As shown in Table 5.4 C. lmlcoin
tbat were captured were just roo overwhelming. '<lore re- was the only spl-<:ies of C11/icoides found to increase e:-.-plo-
cently, a single isolate of AHSV serotyi,e 7 was made from sivefy in numbers during, and follo"ing. the heavy rain, of
pools of C. imicoln collected during the 1999 outbreak of the 1996. This pauern of' oucbreaks and the occurrence of
disease in the Western Cape Province.111'! A recent anatrsis of Al IS as a plague following excessive rains was noted as
the 14 larges! epidemics of AHS 10 affect southern :\frica long ago as 1863:?'11 and was subsequently corroborated in
since t 7L9 were shown in 13 instances to be linked to \\'arm· 1921.JW

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 \'ec1or,;. C11/icoidwsp1,. 115

Culicoides boli ti11os MeiS\vinkel, 1989 (Figure 5.3b) l mmacure stages 1'he pupa has been relrieved from cat1le
Taxonomy (:ulicoides l10/itinos was ontr rccogni1.ed as a dung. and morphological smdies have confirmed tJ1at it dif-
species separate from C, imicoln in !989 when the type fers Crom chat of C. imicola.2oa
st1ries was reared from African buffalo dung in the Kruger
'.\a1ional Park. South Africa. It is most closely relmed taxo- Geographic dlslri bution Culicoides /10/irinos occurs in \(.
r.omically to the Orientru/Australasian C brei,itarsls Kleifor. rkan tropical and sub,ropical regions south of the Sahara
191 i, which is regarded as the most imponam vector of Desen. It has been recorded 1rorn l I countries, namely
on· in Australia. In the past C. bolir/110-, has been confused South -\frica, Sotswana, Zimbabwe. Lesotho, Malawi.
with both C. imicola and C. bre11irarsis: these misidentifica- Kenya. '\igeria. 1he Ivory Coast. Gambia. \ladagascar. and
uons have been corrected. 213 Before 1989 C. boliti11os was Mauritius (misidentified as C. breviwrsisJ. It is likely that it
recorded in the $outh African li1era1ure as C. sp. # 49. Culi- will be found to o,cur throughout Africa where c,1ule are
coides /;()/iti11os is a member oi Lh<l lmicola complex but. kept. In South Africa it is almost as widespread as C. imicola.
allhough it is morphologicall}' \'Cl: similar to C. imicola. the antl has been found in all of the area-imensive Cu/icoides
two species differ markedly in vario~ aspect, of their surveyl! referred to above under C. imicola, l'unher details of
tespcctiveecologies. These differences are listed In Table 5.3. Its distribution, as determined between 1996 and 1997, are
given below (see Detailed dlstribution and seasonality in
Vector competence Cu/icoides bolitillQ$ wa~ fi~t im:rimi- South Africa . II~ occurn,nci.: 011 ull'~hure •\fricau i,hmds
nated as a vector of :\HS in an outbreak or the disease In bcgcs the questio1t a, to why C boliti11os has not also pen-
South Africa in 1998.22" Almosrsimulcaneously it was shown etrated into rhe Mediterranean region where caule are
in the laboratory to be a competent replicator of BT\' and is found In abundance. or has it been overlooked being raxo-
therefore also a likely \"ec1or of the disease in the neld. 33" In nomically \·cry clo$ely related to C imico/a?
addition, it is considered robe the likely principal vector of
EE\' in pans of the Eastern Cape Province. South Africa. Seusonal abundance und prevalence Only in Sou 1h Africa
where C. imicola Is ab~ent.280 has the seasonal abundance and prevalence of C. IXJliri110~
been studied.2:1.? Before 1989 most studies on Culic-0id1Js in
Host preferences Blood meal analyses show that C. boli1i- South Africa failed to recognize the existence of C /)0/irinos
11os reeds on canle. horses and sheep (Table 5.SJ. It appears \,ith the resuit that the data on these two species are con-
LO be more endophilic than C. imico/a ss ir can be captured tla1ed. This is also panlall~•tme for another snidy conducted
in large number~ in horse stablc.'S/ 21 thus dispuring 1he 1hrough much of Sou tit Africa in 1992.3 nl-!owever, the area-
long-held belief that C11/koides do not enter building,. intensive studies mentioned above under C imic:o/a, and
published between 1988 and 1998, arc reliable as regards the
Gonotrophk cycle This is unknown but is probabl) very informaiion on C. boli1i11os.
similar ro that of C. lmicola.
Dispersal It ls presumed that the dispersal of C. holitino.~ Is
Larval habitat Culfroides boliti11os was first reared from similar to that of C. imicola. Of illlerest is the record of 1hou-
cattle dung in l 968 but at the time was mistakenly idencified sand1> of females or C bolirinoscapmrcd at dusk in the pres-
as C. pallidipe1111is (=: C. imico/d).2~ The larval habitat of C. ence or a slow-moving herd of60 grazing African bu!Talo.217
l10/iri11os was described as dry canle dung pats with a very The specimens consisted of nulliparous. parou~ and gr,1vid
hard cmst, a dry sponge-like centre, and a moist lower layer females. which implies 1hat the first two categories were ac-
about 25 mm thick in direct comae, with moist soil under- companying the buffalo in search ofa blood meal fonhe de-
neath. This describes a pat that has aged to some eMent. It i~ velopment of their eggs and that the gra,id females were
probable thai C. !10/ir/110.~ O\'iposits on!) ln \'Cry fresh dung seeking fresh dung for 01iposition. In this way C. l10liri11os
still emirting the necessary volatile amaccams. but this sup- may disperse a number of kilometres in a short s;>ace of
position remain~ to be substantiated. It has been speculated 1ime. The results of an investigation concerning the pro1ec-
1hat In Australia the closely related C breuilfJrsis oviposits 1ive s1abllng of horses against the attacks of C. boliti,ios im-
in ageing cattle dung over a number of days. Culicoides plied that. in the absenceoi cattle dung in a 500-m radiusoi
bolitlnos has also been reared from the d1111g of African buf- the stable, C. boli1i11os dispersed nightly from neighbouring
falo and blue wildebeest. 118· 213 On occasion more than fanns over distances greater than 500 m to attack the ~tabled
l 000 of these midges can be reared from a single buffalo horses.:!2 1 The question remains as ro how C. b0Uti110.r ar-
dung'pat.216 The duration of the larval and pupal srnges in rived on the islands of Madagascar and Mauritiu~. 1\ mo·
dung has been estimated Lo \'ary berween 8 and 30 da) $.?13 lecular 1,iudy comparing a Madagascan population of C.
In South Africa, buffalo dung can produce C. /1olici11os for boli1111os witb three populations from mainlandAfrica failed
up 10 90 days especially during 1he cooler months when it to reveal an)•degree of genetic dilforentlation, 269 and there·
does not desiccate a~ rapidly as during the hot summer fore suggests the invasion of l'...ladagascar by C. !lolitinos to
momhs.216 be relatively recent. ~~11ether the introduction or 1hc midge

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occurred at 1he time caule were introduced 10 1he island or and George). This indication that C. bolitinos is a
whether ii predated this. cannot be answered. wee- and cool-adapted species must be 1reated with
camion as cattle. not prese111 at all sites ~ampled. deter-
Association with ou tbreaks of disease ·n1c recem out- mine primarily the prevalence of C. bolitinos. :'\ever-
break of AHS in rhe eastern Free State Province of South theles~. it is clear that significant popula£ions of C. boliti-
Africa Involved C bolltf1ws. and Is the first substantive 110,< are encoumered In areas somewhat depaupera1c In
proof of its invol\·emcm in field ornbreaks of disease.2:t6 .:u7 C. i111icolt1. a facl that increases the tO!al area at risk to
In this ou1bre11k live isolations of AHSV serotype 6 were orbi,iru~ transmission In South Africa. 1\s revealed in
made from identified pools of C. bolil/11os as were four iso- f igure 5.16 ('Thabazimbil ii is clear !hat C. liolitino, can
lations ofa11 unide111ified sermype ofBTV. Furthermore, C. also occur abundantly in warmer. drier areas.
/Joli1i11os wa, the domi nant Culicoides captured and W\b • The lowest annual abundance, of both C. imico/(1 and
d1e only ~pccies to be found at a.I affected stables. In the C. bo/ilinos occur in \,imer and spring (or weeks 22 to ·16
same ourbreak C. imicola was rare in numbers. and had a or the year) <Figures 5.15 and 5.16). Their abundances
lowered J>revalencc rate of71 percent. Large outbreaks of increase gradually in the early summer 10 peak eventu-
AHS have in the p:ist been reported from Lesotho, which ally during late summer and autumn (weeks 10 to 20).
immediately adjoins tho eastern Free Sta te, such as the one During this period the number ot older. parous midges is
in 1953 when an es1ima1ed 5 000 horses died. II ha~ been also at its greate,1. and so increases further the risk of
shown 1ha1 C. imic.ola is rare in Lesotho. and it can only be ,irus 1ransmis~ion. Thi~ perfectly mirrors the late
assumed rhat C. boliri11os could haYe been the species in- seasonal appearance of BT and :\HS. which corroborates
volved. C11licoides bolitinos b al~o to be the likely vector of 1he maxim 1ha1 1hc geographic:al and seasonal occur-
EE\/ in the Eastern Cap(.' Provln=e.280 specilkally in 1he rence of an in$ect vector mus1 correlate ,,ith that of the
Pore Elizabe1h area. where it comprises up to 90 per cent of disease.
au C11lii:oides capmred.218 • It i~ generally belie\'ed that area.~ el(J)eriencing che
coldest "1nter,,, including snow. are also those with th!!
Detailed d istribution and seasonality of Culicoides smallest p()pula1ionsof C i111ir:o/t1. This holds true forthe
imicola and C. boliti11os in Sou th Africa222 ,Itel. of Giant\ ea~ite in the Drakensberg. and Ellio1 but.
From September l!l9610 tune 1999 the distribution and sea- ('l'Cn here. C. imico/a \,~II appear briefly in lilte summer
sonaliry ol' these two vectors was determined b} weekly col- (weeks 5 to 10]. Unexpectedly. ii was revealed (Figu re
leciions made a1 •10 lives1ock holdings spread almos1 5.1 5) that C. imh·o/a is rarer at. or absem from, three sites
equidisrnnily across South Africa and the results are lisied which have a far milder. frost-free climate (Port Eliza-
\'ertically in Figu res 5.15 and 5.16 The daiashow chat: b<.'th. 51ruisbaai and Alexander Bay). These sites, sirua1ed
• Culicoides imlcoln b most abundani in 1he warmer on the ,andy ~ouclwm and \\'es1em coastline, demon-
northern and ea$Wrn parts of South Afric;a when> it can >1ra1e that wi1hrn certain limits. cli!i1ate is secondary in
be captured 1hroughou1 the vear, including winter. Si1es de1em1ining the abundance and distribution of C. lmi-
include Tshlpese. l)ofokwane lformerly Pie1ersburg), wla, and that the availability ofa suitable breeding habi·
Phalaborwa, l\omatipoon. Lydenburg, Middclburg and 1a1 Is therefore parnmounl. '\\'here known, i1 has been
Onders1epoort Veterina.!)' lnsifrute. These a.re.is have found chat C11/iroidrs larvae inhabit the upper layers im-
long been ron~idered endemic :rnd so may an 11• sourri.> mPcii~trly hel('l\\ 1hc• ,nrrac·l' of thP soil.~5 \\hl'l'P rhr~ ri>-
areas for d1e southward spread of Culicoides-bome or- quirc adequate te,·els or moisture and nutrients to
biviral disease~. However, the data also show clearly that complete their, · 10 14-day developmental period. How-
C. imicoln can be found in some abundance. and also for e,•er. if these upper laye~ of the soil drain too rapidly of
prolonged periods, in 1he souihem half of the country moisture. as in sandy coastal soils. or in sloping topogra-
(e.g. Kimberley. Joubertina, Lain~sburg and Porterville): phy, the larvae desiccate and die. This means 1hat O\'erly
In the past 1hcsc areas have also experienced de11151aring dry soils mos1 E'ffectively comrol the numbers of C. imi-
outbreaks of discuse. 1hose of Al IS being especially well cola. Similnrly, h is assumed that soils too poor in micro-
documented. 19· 120 ·fhc figures also reveal that C imicola organism~ will nlso impede larval development, Such
can he found in most, but not all, parts of South Africa. poor soils arc commonly found ln coastal regions as well
which means that there is probably consiant cycling of as in areas where nutrients are leached by higher rainfall
, BT\/, Al IS\· and rmv ln many areas, and that when Such barriers do not apply 10 the replication of(:. boliri-
circumstances become particularly fa\'Ourable for 11os. The fact 1ha1 i1 breeds In cattle dung ensures ade-
C. imit:ola. outbreaks will occur. quate le\·els of moiswre and an abundance oi micro-
• As regard~ C bolirinos the data in figure 5.16 re\·eal it to organisms. Thus sol! type and quality play no role in
be an order of magnilude less abundant tllan C. imicola. de1em1ining its pre\·alcnce. which. in rum, means that
and with the most numbers captured in cooler. higher C. l,olirinos can invade areas free of C. imlcola. buc only ii
rainfall areas (e.g. Middelburg, Ficksburg. Port Elizabeth cattle are prcse111.

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• lt has been argued !hac winds (such as prevail along the require the inclusion of other parameters such ns the pres-
coastline) induce high mortality rates in C. imkola:"!O chis ence or absence of cattle.
may be anoth(.'T reason t0 explain its ab$ence along pans
or the South African coastline. Howe\11r, chi;: persistem Rost preferences
pre,•alenceofC bolfiinosinanareasuch as Pon Elizabeth. Understandably. ne11rly uJJ ~cudies on African Culicoides are
\\'h1ch Is known 10 be the second ,1indies1 point ln Sou1h strongly biased towards the welfare of exotic breeds of
Africa and where some of I he largest capru res of C. boliti· horses. cattle and sheep. As several of 1he orbiviral diseases
nos have been made,2i 8 reveals Lhat windiness rnay onl) affecting domes1ica1ed 1i,·es1ock have their origin in African
affect midge activity on particular nights, and does not im- wildlife. it would seem appropriate to study the relationship
pact upon overall foeding and breeding success. Presum- between African C11/icoides and game. As has been re-
ably the same could be said for i1s sibling species. C viewed.= nine studies in Africa have purported to deal with
imicolll: this therefore points co factors 01her than ,,ind!· this relationship but nearly all were conducted in unnatural
ness inhibiting its establishment in !he Pon Elizabeth S)'Stems in \\·hich wildlife had been removed from their
area. habitat$ and maintained in subverted conditions. The rela-
• Of significance is that these data clearly show 1hat both tionship between some species ofCulicoides and 1heAfrican
vectors of AHSV occur in some abundance In the AHS clepham. zebra and buffalo are brieflr re,iewed below.
control zone of the Western Cape Province of South Af. These herbivores serve as hosts for three specie~ of the lmi-
rica. This means that 1he zone is alway,; at risk to incur- cola complex, and for other speciei; of,\t'(lritin whose role as
sions by the ,irus. as was amply demonstrated by the vectors remains unk.J10\\11.
death of some 30 horse~ from the disease in the Stellen-
bosch area in 1999.2 ~ The prospec1 that this can recur at C11/icoides and U1e African elephant ·1here is oni)' one
irreg11lar and unpredicu1ble intervals is real. Vector-free study in which an attempt to clarify the life cycle of Cu/i-
points (Figures 5. 15 and 5.16) occur along the southern coides found associated v.ith :\frican \\ildlife wa~ made.2 ~J
and western coastline ofS0U1h .\frica. ·n1is dealt "ith five species of the subgenus .-t1rariria, which
were discovered feeding in considerable numbers behind
As noted. an earlier data set"32 similar 10 the one above was the ears of the African elephant (Figure 5.18) and to breed in
used to model the distribution and abund,mcc of C. imico/a Its dung. The midges were collected off culled elephants in
in ~outh Africa.21 \\11llst the model satisfactorily explained broad daylight and included specimens that were fully and
67 per c:cm of the ob~erved variance. slg11ificanc anomalies fresh!~ engorged: this clear!~ shows that certain species of
appeared as regards the geographic distribution of C. imi- Culicoides feed throughom the die!, and no1 only ar night as
cola and its predicted abundances. This may be due to the is commonly belic,·ed. It was learned too t.hat all the Culi-
data being representative of climatically ·average· years. but coides ~pp. collected on the ears of elephant were also those
which fnll to model for the superabundance of C. imicola tha1 breed jn elephant dung. indicating their co-e,·olvc-
follo,\ing excessive rains.~1~ This would seem essential mem. a iact thar is further substantiated by the observation
when one consider$ that fl~ superabundance at such times that 1hese most!~· new ~pecies are exceptionally rarely col-
correlates perfectly \\ilh the statement by l11eiler320 "thnt lec1ed outside the range of elephants. It was observed tha1
horse sickness appears as a plague following heavy rains'. gra\'id iemalcs of these species oviposn in fresh dwlg in
Interestingly this observation appears tO be nuuradlc1ed b~ broad daylight, and that Cuiicoidet are among the first in-
1he central clwnping of abundance data in Figure 5.16 sects 10 arrive ;Jl freshly e\'acuated dung. Gravid C11licoid11s
where it is clear that the largest numbers of C. imicola ap· females mus1 therefore track elephants in small swarms.
pear in those areas experiendng a more moderate annual and when dung is evacuated the}' immediately descend
8\'Crage rainfall varying between 300 and 700 mm per upon it to Jay their eggs. O\iposition Is rapid and is a likely
annum. It remai.ns co be demonstrated whether this appar- part of an evolutionary adaptation enabling the Culicoides
ent contradiction is e:q>licable in terms of soil fertiliry. i.e. in 10 relocate the elephant. which wiU have proceeded little
areas experiencing a pel'$i~tent annual average: rainfall of more than 50 m from the original point of dung deposition.
above 800 mm, do the soils become coo leached of nutrients By once again auaching themselves behind rhe ear to obtain
to sustain 1he larvae of C. imicola? Ifso thb may explain the a fresh blood meal the;.e elephant-.i1ssocia11.'d Culicoides re-
,inual absence of C. imicola in the equatorial regions of peat the feeding cycle needed for the de\'elopmem of their
Africa where suitably high c,wironmental temperatures pre· nc,x1 batch of ew.
\'ail. but where rainfall is excessive and ,oil fenilily is notOri· ft appears that, af{er blood -engorgement. 1hc Crrlicoides
ously poor. Another factor di fficuh to Incorporate into any do not stay on the elephant, bu1 rather abandon the host to
model is the extent to which humans aid the establishment reside on vegetation during the two or three days required
of large populations of C. imicola b~ maimaining livestock for egg development. Here they \,ill await che return of
in confined spaces and by irrigating pastures. Whilst a siml- elephant 10 once again trnck them but only if they are
Jar model has yet to be developed for C. l.lolirinos It would ft11ly gra,,d. It is well known that elephant herds. led by

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118 '><1KJ< o": \sp<:C\$ inDuencing the occurrence of inftc1lo~ diseases

G=4Cur:o
Mtun?irw
Po<1 Edwatd
lio!lsgrue
Georg&
J• )f J> "f ,, ~ ~

-
M tl ... ,ff . . H ,.., II Q ' I

•••
J I I I Y f I •; U :

.. • ,~.• -• ••
h tt, M I• •I 1 1 .'. 1' - lJ :1 rt .. fl' ;'t 1' JI; ) 1 .If ll ),I ~

•
Vryho,d
El1to1
Flel<ll>u!!j
v,ode
Veceen9ng
Port Blza.batn
M<laell>"rg
l)'(!o<\bul$i
~omabpoon
O<>do ""'''"'°" I
•
Wam>b.d
Pje:e,sburg
F'nalabo•wa
TN!baz,mbl
E111$<,\0
Ku'""""'
Svu1~1
Pono,v\P.t
JOlll!ernna

_ -
Fon Stauforl
UCbltnbllrg

•-
Weli<Qffi

.
T~
Klmboday
$1
I
lall!Qsl>Utg
Veo:<><olid
C~'vu·ila
O&Aat
Up,ngton
Sp11nol!Oi<
81:aulort \v~
AA>~andOr Bay

figure 5.15 Culicoides 1m1cot1r log (r.-11 rumbers ca~wec ·.\eek!y a, 39 livts:oc~ nolc ,ngs spread across Scum Africa between Sep1ember 1995 twea,
371 and September i 99'7 1week 36}. s,tes a:e :.i~ed from h,ghe~r ,o fewest :ainfall Ii 200 mrn to 250 mm an_numl. The wee~iy C.:ticoides imico/a to:a:s
are colour-coded -.·.hi1e 1no collec:ion maoel, g,een (0 C 1mico1a). yellow (1 to 91, orange 110 10 9SI. red [ 100 :o 999i, p!nkl1 000 to!! 999) and dari( :,Jue
(10000toS3ll99!

Glal\l s Castle
Mlunzlni
P~rs!:dward
H<)qsgate
Ge1>,se
Vr,-hald
ElhOt

-
FIIC~tburg
Vr~
Voreon,glng
Po~ Ellzabe!h
•
•
Mlddolbvrg
L)'d~l>\J'll
Kom.'ll1poon
(),v';·-
.....nnbad
POlO<SIMlj
PP!a!.l?>o<w3
Tha!l4tmb,
Elli$,us
Kuru!'Mn
•
•
S!rulsbaal
Po~o!Vllle
Jouberllna

• •
Fon 8e•ul0<1
L,<l\tent>u,g
'Metkom
T•rupes•
Ktmb~rioy
?t,osl<a
la•t19sl>u!ll
Van:{tf&tad
" • - •-----~
~rvin·.a
.O&Aar
Up11191on
Si>r,I\Ol>OI<
Bo.a~fort Wast
Alo,ande, Bay

Figure 5.16 Ciit,co,:tes oolittnos reg (n-1 l r,umbe1s caotured wee~ly at 39 lrvestoc< noldirgs spread across Souih Africa he1we~n Seo1emJer I 9S6 !wee~
37' and Septembef 1997 lweek 36); sites ..·e ranked from high;;s! to ,owest ra,nia! 1 200 mm to 250 Mnvar,n~mJ TM wee~r; c~1,co1:Jes 1/otiwwstoralf
are colour-cooed· wh,:e fno colle.t,on rrsaei. green (0 C 1mico1a;. ye1fow 11 tc 91. ~,~rae I iO to 991. rec I1O>J to 999). pink Ii OOii ;o 9 SS91 and dar:: blue
llO 000 to 99 ~3

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 Ve<:tors. Crdicaides •PP· 119

(
7~ I ll - •' \

'-~~~
I \ ! e

~- J)
'\./,:,,?)
l .;, . .

I
'
J .
\ -~
!( /""":" "
~
- L ~ - :¢;)'--/
'::..:;1:1

Figure 5.17 Di39ramma1,c repiesenunion o: tna imeractlon bet,..,een b= dung ;:a1s from large game or lr.;;s1ocll
'::ulicoides spp. their hos!s and preferred habitat ':l,>pes. fach Cu/lcaid/:$ c =uee-MJes. plant and rock cavi1,es
sp. requires a spec1f1c sutst,a,e fo1 the development ol ns larvae; :oese d = rottrna 01ant s1erns. fruit and ful1ll1
arva. habitats may be d,~,oed rnw four mam :vpes Cuhcoides SpP lllqu,re uie blOOd of sJ)ecific
a= sunace water and s:oll ,merfata (organically enncned or 0011 host anima1s !e) in order to deve!op the11 eggs

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120 .,,·no, o, ,\Speers influencing the occurrenre o[ infectious dls~!lse,

Figure 5.18 The eteonant possesses no s.~ea: glands aoa so coo,s 1is hlooci bv p.;mp:~p large amounts l5 :c ·2 ( /min.I thrauyh ~'1e ears The sr.,n a! 1ne
b2tl( of these hea·:11·; reined ·1adie101s' tS S110l)!h, and ,s /,r.ere huridr6.;!S :,f Culiccides'P.£C Th,s Cllotogra~h c' e,ght Culico des~ me ea· o' an
e'ephant v.as l?~n a, 15:00 and ,ncJuceH fres.~ly blooo·!ed !ema:-e

matriarchs, move In well-defined home ranges. and will arc easiiy identifiable and 1hey occur exceplionally rare!)
daily traven;e the same well-worn paths for feeding and wa- else"here in S0u1h ,\Irita. This indicate, that their moss
tering. This would facilitate rhe del'Cilopmem of sizeable mo,·emems. if any, over large distances, either do not occur
pnpul:uinn< of l"leph1rn1-:is~nria1Prl r111/rniri1~ in rh!' !mmP- or hn\'t'}'1'1 to br ohsPr\'ed. 21 •• m Furthermore, an ime11,ive
dinte vicinity of these routes. An elephant bull ,!ail}' con- t\\'o•year survey or 1hc Culicoide.~ that occur in and around
sumes 300 kg or vegetation and e,·acuatcs I 00 to 150 kg of the Kr\lger National Park which ~ielded O\'er fi\'C million
dung.m More rhan 3 500 Cuiicoidl!~ have beeri found 10 midges, faill'd 1<1 reveal the presence of significant 11tunber~
hatch from 5,75 kg ot elephant dung.21 " Thus the dailr ac- of elepham-associated C11licoides on livest0ck farms Imme·
crual in numbers of elephant-associated Culcoirles amongst diatel~ adjoining the Park. 217
1he 7 500 elephants of the 'Kruger :\arional Park. in .:.omh Af.
rica, could total roughly 500 million Individuals. C11licoi1les and the plains zebra Recently. another nt.>w
One of lhe specie5 associated ,,irh elephant is C lo.t·· species of the lmicoln complex, C. sp. " 107. was di5co,··
orlo111is, a member or the lmicola complex.211 ae~-ause i1 is ered.ii- almost solely restricted to 1be \\'armer arid drier
closely allied to both C. imico/a and C. l10/i1i110,-1he ques1ion cemraJ.northern. northern and eastern pans of South :\J·
arises as to whether C. /o:wdontis is not olso invoh·ed in the rkn. In the 1(,-ugcr Nntinnal f>ark h enjoy, n clo~u~ochttion
iransmissiQn of orbivirus.es. It was suggested that the el- with the plain~ zebra (Figure 5.19) which is a hO$t of
ephan t ma) be a namral host for J\I 1sv'JJ but subsequent Al!$\. u 5ignifica,ulv C. sp. 11107 breeds in thl' dung of zebra
smdies have failed to bear chis out. u It would \oeem thar ln and. unurnally, is rnohl prevalent during the cooler ~easons
Sou ch Africa at least the elepham play~ no pan in 1hc epide· but 01lly in fro,1-free areas. Whilst C. sp. " lOi never be·
miolom• of Al lS\' as the disease appears in large areas from comes abundant. no doubt due co rhe limited m·ail3bilit,· of
which elephants are absent. Elephnm-associated Culicoides Its breeding medium, ib occurrence duringwinterrai::,ei. the

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 Vectors: C:11/icoid,,:,; spp. 121

Figure 5.19 The plains 2ebra ,Equus burche/11): it 11.1s been shown :hat all nine known ssrowpes of AHSV c11cula;e '" zebra. Eve11 under ·.e.ry dry
condrti0ns tn the Kruger Nauonal Park consicsrabie numbers of Culicoi(lestC 1mico/a) can still t:e cap1ured

que5tion as to whether or not C. sp. * 107 ls capable ofmms- cent B1V-lnfection rate has been found in some buffalo
mining Al-IS at a time of the year when the disease is not herds in die Kruger ;s;ational Park.2"' but this does not ex-
considered a problem bllt has baen shown stlll to be cir- clude the im·olvemem of other Culicoidesspp. in their infec-
c11 lacing in i.cbra. t 3 tion. Of the hundreds of fight trap <:ollections made in the
Jne quesl ion is potentially signitican t a~ C. sp. /1 l 07 can Kruger ~alional Park the largest numbers of C /Jolitinos.
b.- found in some abundance around horse stables where it numbering in their thousMd.~, were capmred in the imme-
in all probability breeds in the dung of horses.217 It also diate \icinity !lfgrazing buffalo herds. In these colleccions C.
breeds abundantly in the dung of che white (square-lipped) bollri11os was Llw dominant species. comprising more than
rhinoceros (C. simumi.217 80 per cent of the captures: C. imicola comprised less than
3 per cenr. 2 t 1 The only other species of Auarilfa that ls likely
C11/icoides and the African buffalo Both C imico/aand C. to be associated with the buffalo is C g11lbe11kia11i but 1his
/Jolicinos have been collected live off a darted African buffalo has yet to be demonstrated.
(Figure 5.20).zt, There ls little doubt that C. l1<>lili110$ espe-
cially I:, closely associated with rhis host: in 1be Kruger Xa· Culicoi.tfos and domesticated livestock Because of 1heir
1ional Park more than l 000 specimen,; of C. bolitinos have minute siu> and generally noeturn:il habits it is difficult to
been reared from a single buffalo dung par.116 Ofvecerinary collect or identify C11/icpidQSOn any particular host. Virtually
importance is that C. bolirinosalso uses che dung of domes- no bait-trapping has been done in Africa. and therefore the
tic cattle for breeding. This ensures that this seemingly effi- presence of large numbers of a particular species near spe-
cient vector of BTV occurs almost throughout Sou ch Africa. cific livestock species is generally assumed to indicate that
and probably plays a major role in tbe Infection of cattle the Cuficoides feed on lhcse hosts.= The \,~ng patterns of
1,'irh the virus. In support of chis is the reporr that a 90 per che L4 species of Culicoirles most commonly encountered

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122 ,u,10, oxc Aspects Influencing the occurrence of infectious dlseases

Figure 5.20 The African ou'!alo (Synceros c;,ffe/1· in the Kruger Nat1or.al Park thousar.ds of Cultcoides 1>80 pe-, cent C. bofllma-s· have been captuie.i
around such ~os at dus~. More than 100) adults of C. bo//rmos. car. oe rearec ircm a sir.gle buttalo dung oat

near llvestock in Somh Africa are shown in Figure 5.3. much merit, reliance for Its verification ls sllll placed on
These. ranked in order of abundance, are C. imicola, blood meal identification.
C. $tibsc/111l1zei. C. mag1111~. C. zulmmsis, C bolitinos. C. in southern Africa Culicoitles blood meal identification
pyt:nO$liCrtl$. C. leut:o.ttie11is (= C. disrinc1ipem1is In earlier using various type~ of precipitin tests and an enzyme-linked
South African smdiesJ. C. 11ivorns, C. sc/111/nei. and immunosorbem assay !ELISA) has been limited to srudies
C. 011derlei11i. These species made up 90 per cent of rhe 86 conducted in 7.lmbabwe and Sourh ,\frica (Table 5.5)."7• 265·
Culicoides spp. collected coumrywide. 2 n Although close 16. Blooc meal identification can sometimes give a biased

on ·I 000 collections were made in rhe particular survey impression of the potential vector importance oi cenain Cu-
only 75 per cent of the Culicoides known lO occur in South licofdes spp., especially If these species are collected in large
Africa were captured. indicating that many species either numbers at isolated sires. F'or example, 215 Ctt!icoides spp.
have highly specialized feeding preferences or larval near glabri1n1111ls te.~ted positive for the blood of four hose
habitats. A similar panern of species pre\'alence and animals at one site, yet this species was rare in catches made
abundance was recently reported l'rom Zimbabwe and at the other 23 sires in the Onderstepoon Veterinary lnsti-
2'S4 2s.;
Botswana. rure area (Table 5.SJ.
• Another approach used to determine the host p,cfer Culicoides spp. which usually feed on mammals will OC·
e.nces of C11licoides spp. is to couni the number of sensilla caslonalJ~, feed on birds iflarge numbers of them are present
coeloconica ('pi1s'J found on 13 of the 15 amennal flagellar in an area a.nd their primary source of blood is scarce. !'or
segments ('anteunomCi'.es'J or the female. 18 It b considered example, in Zimbabwe. a small proportion of the large nu m·
that a preference for mammals is indicated by six or fewer bers of C. imicola. C. ;:ul111msis and C. 111i/11ei collected on a
pic-bearing·amennomercs·, while rhosc with 12 or 13 show poultry farm of 75 000 birds had fed on the poulcry !Table
a preference for birds. Although this approach may have 5.5). which indicates that under specific circumstances rhe)'

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 \'cc101 ~; C:11/1t0Ult>S spp. 123

may become opponunL~tic Feeders.;;o Conversely. kn<J\\11 cent hone\·,i60 Fure.her details on che life cycle or five species
bird-feeders. such as C. le11cosrie111s l= C. tlislit1cripe11ni:1 in or ·ll'llritit1 found in association with the African elephant
early studies on South :\frican Cu/it:oides) anti C. pyt·110Jtit"· ha\'e been reported. ~23
ws. mus1 occasionally feed on mammals (Table 5.SJ. as In :mempts to explain 1he possible eifect of \'arious cli-
suggested by the fart that In South Africa two BTV sero- matic factors on the ~ur.ivo.1 and seasonal prevalence or Cr,Ji.
rypei: have been l~olated from c. ,,.1-c110.<rlctu$. (Table 5.21. coldes ~pp .. espectally c tmi(o/a, Lhe effect or desrccatton.
Epidemiologically chis could be significanc in the irans- submersion in wa1er and exposure IO low temperatures on
:nission or virus from birds to mammals and /'ice 1•ersa. different life ;.rage;. have been investigated.2611 Eg_~s showed
Culicoides pycnostictus was recemlr also infected in the some re~lsmnce tQ desiccation. for e.xample. l per cem oi
iaboratory wi1h an· 282 However. the darn remain conti.ts· egg~ allowed to dI)' at 8;3 per cent rela1i\'e humicliry did not
ing as in a more recent study2 z1 it wa$ found that at leaM collapsl.' and laler hatched on immersion in water. Low tem-
five bird-associated species (C. leurosticrm. C. vyeno- perature, ad1 erscly affected egg \tjabili1~ . If the eggs were
sricrus. C. 11ea1•ei. C. bedfordi and the Ondersrnpoonensis kept ai 6.5 •c. viabilit) started 10 decrease after seven days
complex) would no, enter horse stables whereas se,·en while none hatched nf1er Ji days. The immcn;ion in water of
mammaJophilic species did so readlly. Wheiher this thousands of eggs. mos1ly or C. imico/a, had no ad\'er;;e effect
reflects a strong exophilic tendency, or whether mammals c>n hatching - rhe lar.11e of seven species. including
are uuly unattractive to C. P}'Cllostic:111.< and thu~ very C lmicola. survived immcr;.ion in water at 22 •c for Gto 13
seldom a!lacked. remains to be clarified. days. Fourth instar larvae of C. 11/11os11s. C. µyr11os1ict11iand C.
zuluensis present in breeding medium sun~ved refrigeration
Life cycle at 6.5 •c for 10 to 14 day~. end completed their de1,elopmer11
The life cycle of Cu/icoides spp. worldwide was revie\, ed In normally 011 return ro room temper,uure. On Immersion. lhe
1984 by Kettle. 180 In Soulh Africa an early attempt to darify pup:1c of all ,pccies. except C. imirolu. wriggled free of the
the ilfe cycle of some specie, was made when C. imic:olt1, C. breeding medium and Jloated to tl1e surface. C11/lro1c/es imi-
pyr:11ouicms. c. 11i110s11s, C. leuccs1ic:w.< ,as C. distim;ripe1111is), roin pupae. however, Illy on the subsmue below the water
C. e11derlei11i (as C. sch1dn:e11. C. ;;u/11e11sis. C. ued/ordi and C. surface and drowned within two clays at room rnmperaiurc.
mngnu.s were maimaincd in the htboratol') under controlled .\t 6.5 •c they sur.'ived for at least six days under water. The
c-onditions. 260 After a blood meal had been taken. it wa~ clfcctoflow rcmpcratureson adult survival and longevity\,':!$
found that the maturation ol eggs in C. imic:ola females took also tesied.261 This showed tha1 at temperatures as low as
two days at 27°C and th rel' to four da) sat 22 •c. ·n,e ·sausage- - 1.5 C. 15 per cent of C. imicoi<1 adult> lived beyond 15 day..
shaped' brown eggs were 400 µm loni; b)' 50 µm wide and whereas the last C. pyrnDstic111scliecl on the 5-llh day.
were laid in a staggered double-rowed pauern somewhat re- The re~uJ;.,, or the,e laboratory studies Indicate that low
sembling footprints. .-\t 2 l to 24 •c the eggs of five SoUlh ,\lri- wintt'r temper<1tures do not klll any stage ot Culicoides but
can C11/icc,ides spp. cook three days to hatch. The number of merely slO\\' do\,'11 de\'elopment or induce diapausc. It can
eggs per batch varied from a mean of69 for C. imicol<1 to 162 also be deduced that soaking rams have no ad~·crse effect on
for C. 11it'Osus, but the number of batche~ wa., not dl.'tc:'r· the egg~. larvae and pupae or moM ~p~ci1h, but that they do
mined. In C. i'l1rilpe1111is. a 1\onh Amerkanspecies. up to four dro\\'n C. 1111/coir1 pupae; the larl'ac.> of C. imicof(t wm not pu-
batches maybe produced ina life-time.?51 A blood meal muht pate unul conditions are sufficiently dry.:\ seven-rear srndy
he taken h~ th«" fl'mall' in h«"IWPPn ('arh barc-h /\f egg,. of rh!' sra,onal abundanc-e tii r imiro/11 at the Ondlc'r,11,-
lar'\'ae of Culicoides go through four stages. arc eel-like in poort \'eterinary Institute ,;howed a drop in aduh r.umbers
their movements and burrow in and out of their breeding eluting ~u,taincd rainy periods followed by a sharp '.ncrease
medium. As is shown by the variations in thestmcture oflhe in population~ during The drier periods that followed.2''3The
epipha.rynges of different species.is:: the food preferences of greater tolera11ce of C. )J_l-c11osti1:tus adults to low tempera.
lan·ae vary: many are carnivorous and Feed on protozoa, ro· tures ma}' panially c:1.-plain whr it wa, the most common
tifer$ and nematode$.:mo The fourth stage larvae oi C. zu/11- species found in two smdit!s in the Free State Province 0 1
ensis and C. 11ivos1is ha\'C been reported to be cannibalistic South Africa, and why C. imit'o/a comprised less than l per
on second stage larvae. 260 In the laboratory at 21 to 24 'C, cent of the tolill (.'11/icoines collcctcd. 17
larvae of C. ni11os11s. C. /e11cos1icm~ (:: C. disr/11c1ipe1111is in
earlier South African studies), C. p,1•rnosrictus and C. 1,/!d. tanral habitats Knowledge abnut the habitatsofC11/iro1de.s
fordi. dt:veloped readily in a mi.~ture of moi~r soil and cow biting midge~ has increased greatly since the early 1960s a. a
dung and reached the pupal stage in 10 m 20 dayi;. The fol- result of studies. invol\ing the collection of live pupae from
lowing developmenral times for six South African Culicoide.s moist situations in the field and rearing them through w the
spp. were: larval period 7 to 25 days. pupal pi,riocl four days. adult st:1ge.Jt,i. 2118 Rreeding material suspected to contain
oviposition 10 fir~t adult 20 days.;\ generation (egg to ~>gg) immature stages has also been Cl)liected and plac1.1d in special
requires a minimum of25 days. Adult Cu/icoides suntjve for emergence boxesorcnges, 11a 21 ?· 211• 261 or allowed roerncrge
benveen 14 and 21 days if fed on water containing 10 per in .~iw in specially constnictcd tent-type emergence trap~

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124 sn:n<>, """ ,\SJJCC!s influencing the occurrence of infec1ious dis<?ases

placed over breeding grounds or tnaterials. 2; 6 Tiiere is a com- lageri. C. eriorlendroni, C. p1111crit//ora;1: and C 11igripen-
mon tendency to believe that Culiwide.~ breed only in moist 11is) are known or suspected to breed in these habitats.2&.l
low-lying areas but, although this is true for some species, but owing to the restricted size and availabili ty of such
others have more specialized lan·al habitats. ·111e basic re- rain-dependent habitats these species never become
quirements for C11licoides larvae are moisture and a medium abundant and are rarely coUected. Birds are thought to be
containing organic maner. Thus. Culicoi(Jes may breed in their p1imaty soutce of blood. From rhe wet boles or
situations varying from those whic:h are almost aquatic in na- crotches o f 11 genera of trees in Zimbabwe more than 500
ture (such as pond margins) 10 those where no fre<: water is $pC~ir1ens of Culicoides representing the );lgdpcnnis. Ac·
present but the humidity is dose ro !00 per cent (such as the craensis and Eriodendrortl species groups were reared. 11
interior of dung pats). The various larval habitats <:an be .\s nored above species from these complexes are \'ery dif-
grouped imo four main types (Figure 5.17). ficult to identify.
• Surface water and soil imerface situations (Figure • Roning fruits and plants (Figure 5.17d). These larral
5.17a). About half the mown Cu/icoides spp. in southern habltatS have still to be investigated t11oroughly. Jn South
Africa make use of various combinruions of soil and water Africa a ne1, A11arltia sp., closelyTclaced co C. pseudopt1/-
as a medimn in which to lay their eggs. Soil may lidipe111Jis from \\ est Africa, has been reared from the
vary from coan;e sand to the finest clay. and lhe basic rotting fallen fruu:s of tl1e Sau.sage tree (Kigelia africa,1a)
medium may be enriched 10 a greater or lesser extent and tl1c maruJa tree (Sc/erocaryt1 birrea). 117 In \Vest Af·
with decomposing plam mauer. val')1ng from intact ma- rica the larvaeof C. grt1hamii (also ofchcsubgenusAflari-
terial to humus, or with fresh to well-decomposed dung. 1ia) have been found in the rotting stems of rhe banana
such as Is often found on irrigated pastures. The water plant.z09 The cultivation of bananas by humans may thus
may range from fresh flowing streams ro polluted stag- lead to the creation oflarval habitats for 1his vector of the
nant pools with varying degtees of acidity. alkalinity or human filarial worm. Mansone//a rnrsrans. A concerted
salinity. The degree of light and t11e presence or absence effort to rear C11/icoides from rotting fruits, flower bracts.
of plant cover. which m ay be c:ther tall or kept shori by plant srems, and large fungal fru iting bodies shou ld lead
grazing animals. are additional imporcam factor:. that to the discovery of the breeding habits of several known
govern the larval habitat of certain C11/icoic/es spp. [n and. perhaps. even new species. Some of t11ese habitats.
southern Africa mosl ofrhe major stock-associated species and many 01hers. were sampled in Zimbabwe but wlch-
{C imico/a, C. ;:11/11e11sis, C. magn11s. C sc/111/czei group. C. our success.41
pycnosricms. C leucoslicrus, and C. 11foos11s) use one or an-
other of the above combinations as their larvpJ habi1a1.r.~
Control of Culicoides
Similarly. in Zimbabwe, :1 tl1ou$ancls of specimens of 16
species of Culicoideswere roared from mud samples taken Around livestock. Culicoides can occur in stupendous num-
at the edge of water bodies. Shon notes on the larval habi- bers especia lly on wanner nights and during periods of
tats of six South African ·groundwater· species have been above average rainfall. At such times more than a million C.
published.269 imico/a can be captured in a single light 1.rap in one night
• Dung pats of large animals (Figure 5.17b). In Africa these (Figure 5.21), 21 !1 and. if the estimate that this may represent
larval habitats were only recognized in 19G8, and rheir im- a mere 1 per cent of the number of midges active o n a parti-
portance appreciated since 1985. 118-~ 12• 213• 261 At least 10 cular night in the area concerned is correct. it graphically il-
Culicoides spp .. all in the subgenus Auarilia. require the lustrates the intensity of attacks that must on occasion be
fresh d ung of certain animals to complete their life cycle..~. endured nightly h) exposed animals. It b impossible co
For example. the dung of t11e ,\frican buffalo, caule and eradicate $Uch numbers of Cuiicoitte..~ and in :\frica $peci-
the blue \\ildebeest is used by C bolitlnos, which appar- fically. the first line of defence against Culicoides-borne or•
ently also feeds on these hoMs.3 13 Orher species. most of biviruses must remain vaccination. However, even in the
them still to be described, breed in the dung of the case of a 'lOLifiable disease such as the highly fatal AHS. and
elephant. the black rhinoceros (Diceros bicomis) and despite the social and commercial value of horses. i Lappears
white rbi.nocero!> (Cem101/1erium simwn). and the plains that less than half the horse population in southern Africa is
zebra.212 being vaccinated annually. Recourse to additional methods
• Tree-holes, plant an<l rock cal'ities (Figure 5.17c). These of disease control. such as Lhe eliminatio 11 of the larval habi-
• larval habitat& Val) from deep. dark, water-filled holes to tat of C. imicola. has received no a,tention in Africa perhaps
shallow, exposed but moist hollows \,·h.lch may comain because little succes:s has been achleved \,ith 01her species
various amounts of water. decomposing leaf lirtet and of Culicoides elsewhere in the world. Under restricted situa-
sed iment. Tree-holes are norrestricted to dense forests. as tion~ it rr.ay be lea~ible co reduce C11/icoideJ adult numbers
even the more sparsely treed savannahs are rich in uee- b}' rre~unent of!heir breeding sites with compounds such as
hole-assocfate-0 Cullcoides. About 15 per cent of African Temcphos. 158 Howe,·er. C imicola's potential for. an explo-
C11/icoides spp. (including C. accraemis. C. c/arkei. C. oI.vs- sive increase in numbers and its rapid radiation over large

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 \ ecior<· C.'11/l(oitl<1s ,pp. 125

l<AAL1>LAAs
IS. 3. l<}~b
1,054, bZO
CJ.le aiJe~

Figure 5.21 The comulat~e total in these :wo :xrn'es amoun:s to>· 000 000 Cullccide;'92.~ per cent C ,mrt:c al ca~tu'ed in cne n gh; n a single
t'"'\:l1gh1 trap set r ear he·ses on tlie Kaalolaas 'arm oi the Onae·s:epoo:: Vetennaiy trs:,:.ta !March 19961

ar~ru, as $Oils become sui1ably moiSt under comlnuou~ for emfreseasons.fll ( lcl\\ever. the modem era has ~een a re-
rains, would make the widespread application ofTemephos ducrion ln fam1 5izc and a concomitant increase in stocking
not only expensive but impracticable. ror the same reasons raws. Sustained irrigation further incrnascs the numbers of
the biological coiurol of C. imkoln lan:ae would also not be C11Jicoide, bret.>ding sites and , in the ca~.- of the moist soiJ.
feasible even if such a measure were arnilable. Similarly. th.: lo\fog C. imicoln and the cattle-dung bre('ding C.: l,o/iti11m,
daily treatmenr of animals with pyrcchroids to reduce adult it is obvious that few areas will escape being colonized by
biting rmes is labour intensive and expen$1ve. Despit~ Jim. one. or both, of the;e vector ~pecies.
Iced successes reported worldwide. 25i and re\iewed In some Beside~ vaccination only ~tabling appears to ha\·e the po·
detail below, the) merit further study. particularly in Africa tential of more reliably and pem1anently protecting animals
In Israel certain extracts of plants from the familie, ~leli· from the attacks of C11/icoides. More than I 00 years ago
aci:ae and the Lnbiatae (including oib derived from ~age. horse ownen; in South Africa noted that stabl1:d hon;cs
rosemal) and oregano) were found to ham a weak to rwgli- rorelysuccumbed 10 AH~. but despite these earlrand persis-
giblc repelle.111 effect against C. imic-oln. "' Recourse to mon• tent obscr\"ation~ onl~ in two studies in Africa. t 1. 22 1 nnd not
traditional or esowric remedie, •urh as the smoking of many more elsewhere ill the world, liave the merit:. of pro-
~tables and the feeding of garlic would appear equally in,:-f. tcc1iv,:- housing been explored. 111c results of 1he Afril.·an
fectlve. but such control strategies have receiv<?d little scrl· slUdie,; revealed that both C. imicot,: and c:. /}()/iri11os readil~
ous attention from researchers. enter stables, and indicated that stables were only or value if
In the early years of animal husbandry, fanners soon well sealed. The cx1e111 of sealing required \1'35 ,issessed:221
noted that distancing their animab irom low-lying moist an 80 percent shad~cloth Induced a 14-fold rcdurtlon in Lhe
areas reduced the pre\·alence of disea~c. and it became cus- numbers of midges enrering stablt!s baired \\ith horses.
tomar,· to move stock to higher-lying terrain at night or even Since Culiroides midges of most species are active mainly

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126 ,t.rnr,~ n,~: Asp<?Ct~ inOuenting th(' occurrence of infccuous d.iS<!as~s

during twilight hours and at night, ,·aluable or young ani· and relative abundance. Therefore, the remol'al of dung
mals, whose Immunity is not established. should be stabled should lower the numbers of C. bolitinos but this is only
at night. Because it is now evident that C11/itoides \\ill enter practical!}' feasible where caule do not range exlensil'ely.
stables, it ls ~'1\ential that stables be \\'ell built and a,- many Funhermort-, such hygienic practices would hal'e 10 be
of the small openings as possible closed. Thousands of implemented over an entire farming community us C iloUli-
C. bolitinos (and C imicola bUL in lessernumbetsl ha\'c been 110.<would be able ro disperse from neighbouring farms that
captured at night in an open stable in tht: Clarens district of migh1 harbour C:. /Jolitino.( if some owners are less careful
the eastern Free State Province of South ,\Irica. The exclu- abour the rcmcwul of dung. In the case of C. lmkolt1, itS ca-
sion of C11/icoides can be more successfully achieved by pacity 10 breed explosively under rainy conditions make" it
screening stables with insect gauze or ~hadecloth,2:!1 and an even more formidable insect 10 control.
will be made more effeclive if their surfaces arc m,au?d with :--o publication~ exist concerning the efficacy of repel-
a p~Tcthroid inseaicide. For exceptionally valuable animals lents against Culi,·oidR.< 111 the African situarion. Th<' chemi-
a stable can be made virtually insect-free by the installation cal control or @dult biring midges by direct treatment of
of extractor fans which mainraln a negative pressure so that livcst011k with pe51icide~ is not usually prac:cical under e.\·
odour.; from the animals d1ac remain inside go largely unde- tensive farming conditions. but it may be justifiable for valu-
tected by the Culicoides outside. h does not :;eem that light able animals $uch as racehorse~. Many pyrethroid
traps operated outside a stable a1 night will collect C11/i- inscctici<!es are elfeCli\'e against the Diptern. .'.Ind, depend·
coides in large enough numbers ro si(:n ificantl}' reduce the ingon the fonnulation used. have a reasonably long residual
auack rate on animals inside. effect. Sprays can he used weekly and. in cattle, in~ecricide·
Another strategy for disease comrol moowd 40 years ago impregnated eartags may be effective for four 10 six weeks. 111
is 10 maimain deCO)' hosts, after it \,·a~ suggested that t.he or even a, long as 10 weeks against Cu/icoides. 159 lnAusrra-
presence of canle protected nearbr sheep from contracting lia ii ha~ been found lha1. after Hereford rearling caule had
BT.I 16 ~i;., 11 must be pointed Olll. however. th:11 because C been given a ~ingle ~ubcutaneou~ injection Ofl\'ermetcin at
boliri11os is dependent upon ca tile dung its numhcrs are likely a dose of200 mcg/ kg. the mean mortalil)' ol engorged Cuti·
10 increa!re \\'here,·er canle occur. and l11erefore the r>r11sence cofdes females 48 hour~ after feeding was 99 per cent for 10
of cattle may actually increase risk. lnd.c cd th<.? implication of day, afrer trcatmem.'\00 An unfortunate adverse effect is the
cattle in the epidemiology of AHSonly ret'cntly became obvi- lethal effecc of ivermeccin on the dung beetle founo which
ous after an outbreak or the di:,ease in the Clarens area of the arc ~o impomult for sanitarion.
eastern Free State Pro,~nce in 1998. when it was revealed that ln a recen t and chorough smdl 52 very limited success
1he maintenance of unvaccinaced horses in the presence of wa:. reported against C. so11ore11sis in :--orth America ,dth
caule is risky. Decoy animals are also an added e:\-pense and formulations of.; per cent pennc1hrin or 27 per cent pirimi·
considera1ion must be given to the fact chat llii) i11creasc in phos-mechyl applied to the dorsum of calves. It was found
animal biomass on a farm 11111 result In a concomitant in· that engorged Culicoides exhibiting subletllal 111101dcation
crease in local Cullcoides number, and in viral transmission. recovered and subsequentI~· matured average-sized batches
Since many Cu/icoltfes spp. breed in moist, organically of eggs. However, a belly spray of 0.2 per cen c pem1ethrin
enriched soil, large concenuatlons of them can be expected s11bs1amlally reduced numbers of engorged fomalei.. and
in wet low-l}~ng areas. Such areas should. if possible, be lowered engorgemem levels on the third and seventh da}
avoided h)' animal;; between dusk and dawn. i.e. during the po~Hreaunem. but by the tenth da,· liule effect was noted.
hours of midge ac1ivity, and especlally in mid· to late sum- In the c:lse of C. so11ore11sis it was shown that most feeding
mer. However. it must be borne in mind that C11/icoid,1s in was done on the belly of the calf. but in Israel II C. imicolu
search of a blood meal are quite cap,-ible of dispersing over was found to feed preferentially on the dorsurn (70 per cent)
considerable di.i;1ances. ln addition. for the 1racking ofho~ts of a horse, with less !28 per L't'nt) feeding on tl\e belly.
they are equipped \,ith acute sensory apparatu~e1> ,on the Th11se studies on the chemical c:omrol of Culicoide,·
antennae) and rherefore will locate animals wherever they show 1ni~ed promise and so meri1 addhion3l research.
mar be kept on a farm. In the case o[ C. l>oliti11os the pres· and par11cularly in :\frica where attack races appear co be
ence and amoum ot' cattle dung will de1ermine its presence high.

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1 "'1""· ~ .. i9&). Studi1.1, on 1hc U.uu~ )'lfJ:;otJII fotmd in ch-: chlc.U•n In ,wull,•n., b\' C:11/i(Ouf,., amkn1t"-'tt. fapt11h•&.• /ou11ml 4f\l,~:"rum1)· ·*lti:r,·.
Japon. II. On 1hc tr.i.n:;ml,<;foo of L r1wll.-ryl by Culh:oidts nmJ..1w.v.1e. 11. l2,-,2a.
/np,1114•'\,• /<Wl'fUll DfVt'ltri,:nry· $rli•tta 12. :<09-J t~
3 ,t~Jlu~u m·, , .. llAMDU'.\", c t.. • u 1 oa. W1il'•• sl)M:--. ,ouua.lv.ln); .1nUbodl<"~
~ .u:,ft.\, >,;., i.:ct1\t>,.:.\ ~ . P,JU.L\.1,. 195~1. Murli1."' on th1• lJ.!11ttx.; <a:A<J11 O> AS.aNm• 1.inH• in (C1.'t"•ll\ini; ,1,,;ld anlm::i.1< in Afrk.1 Tt~11>lt'11I A.1u,nal
found In tho chlckrn ln l•P•n Ill Tmn,m,<,lnn ()(u•11n,c,v111:oon H rnllh 1,tld /'ru,lutflQt: 19. I Y7-202,

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 Vcct()l'S'. C11/icoid1>s~pp. 127

~ 1.m<Amv. !S.~. & \trU.OR.. r,..,s_ 1991. lial~umn and idtncifit.o.uon of

,1.0
d~<• and satcnltc ,m~i•" ,1rrl11~s of\ ·,ro/"11)' ll>uppl•mon: Hl.
arhU\1nb~ from the Sultdntuc ur (_)nt.1n. E,pidtmiolo'-'·uml lnt,:,:110,,. 13;. l~l.
106.,;03-113. 2..; acu rt., ... 19~,s·. 0\1tbteJ.1J, oJ .-\fncan ho:~C' s,d,t..-'"lcr.$ in the C.:JJW Province
~ ,,nr nso:-:. ILL. 19.so. Th~ life C'yd~ ;md ,("m.oru1J lrQ.n!i'mb.~,mn uf ol ~Qu<h Afnc,. 1hr i,·1,•n,inry• Rt"tOrd. 144, 48:l.
Onmlt0Ji/nriafn/liH•1,.ss1Andcr.son .•-. p..-r.1,,w or dom~cir ;ind \\ild 2.5 ut.,:\tT. ~.r 13;0 l~vcfopmtnl nf U)'JUU11>MU1't·\ oftht' 1. mirm,
du<k:<, c:a11lldin11 /ouruul 11//.0<>/ogy. 3•1. •165-3?3. (otnplL'X in th~ lmcru:b-r.sw ho!!:, Cmw.dttm J1mrm1/ nf loo!tJSJ)·. fH
fi •'-'"'r.,~ .• 1967. Arl,01·tnt'>i~ mrd lfumnn DLc..•as,.• \·."odd l·h.•ahh 9-~s. qi;;
9

O~mit.itlon. Technical Report.S.:rk,"" Y,• :tli ,L1:,sn r..., . ..,.toc","-'- ,u~, tll:-;. Ornithor,hlhc.· \"ee10:-,;.,bf twfan
; ,1x1»0~. c. T.. 19cw. Uhta(IMJC1ur1..· or thL• ool.inttes of JfntmtJµro,~u, hcmntozoa 1n insulor ~l"',,•fourid.land. C111011Jn11 }fJurn.ol ofZ.oobJJ:).', il.
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'.Ptato7.o-.... I laemr."lpurh,,,, in Florfd,\. pc,tcntinl \'ec.tors .am.I prL"\:tlence .\laa,(--CZ q'C'lop/J. IJ,ssu,uw"" ..U;srmrts. 21k 2921 ..2922
in tliilurally irtfoc:Lt..'t.1 <:ulltoldl'$ lDlpttta: (\,r..ttnpogonidat JnwntJ) 11j :a nr-w., .R, ~-f -!'o:: r.,r.nm, ~M. 1. .\., 1~. The m,uial p;1r-4'io1:c- .',lnwrnnfma
.'11,'tll~al 1!11w,r,01Dtc-· 25. 3~4. formo,mm,, from the 'l'n1"·an cnonkl'Y .in.d fJ.S de\·,,:io-pn!ent in ,·nriou,
9 A1i,;1s-so:,,-, c-~T., 1990.. Fine :!ltructurt and ..,pmo~1riic dt!\i1..·lopm~n o! n. .:anhropo<!s.. fqrmo.wm !\c-t~m·,,. 22 l....fi8.
\ <ll'l<!insp. ~llcm~porid1:c Plcis1op1,11ndu,•: in 1hc blunt 111id)le 2g kl\'"'· Lt..\.,_. 1901. 1'rcltmlnary \\Ork •>n the >U·Callud ,ufl'nc~· cu
Q1llr.'uu:ln t-dc11l c01µ:era: Ccr:,1101,c.,gonld"3t-,. humal of lm•..•rtcb1utl' ,,!
'lhlt.'C·da)'-,tjOOW~~· <'4U1c Jn Rhod~l.a. /tJU,11(1/ QJComP<lft'Jllt'4•
Pmhology. SS, 105-111. PmiztJ,'&~-i· ",.,.J
n,rmpt',11"1. 20. 1o.;....113
1() ,n:1:\$:0S, c:r .. 1991 . Vector-,. 1Jpi1,oc;t0oiogy. ;ind pathogcmc1ty oi u\·1:in JO ts8\'.\..' _L1.,, .• 191!.!, Ephcmtr...tl ft-t~t or dut•(• d.t)"). Jlelmt·~ of c,mli..•.
5,pec1e-s of H<lcmopru1,•us (Hnttmospu,-im:r 1tat."IJ1aprntc.i<k~('}. G1ttrulln11 \'t'tt'r.uu,".t· Jotmml. (-.8 . ..&!",8...461.
/oumnl o/ZIJO/Oftl, 69. 1880-1883.
31 fll\'f.1UOGE ,,.JW'1MfR(t\\,1 I wun,s:os.1•.1ir:COl'J,M\.~,l').S .• 1ft81,An
u IJAtS, o.. 1.980. 1\-.·o fdarine bf 1he gcnu~ Ea/U11ru1 m n,rr/u,. 11wrulw 1m't.."StlJt3tiOn c>f t:)1Hng nud-goin rt-latlai: m their pol.t!ntinl it:,. \"t"(;lOr.i. ot
d("\'c!opmcnt in C,,brouJ,,s nul1"cult1·ws. Anna,,·:itlt• Para~1wf,,gi,t l>0\·1r.e onchOCt:l"(.~i,, 1n oonh QuEN:fW.and. f(luruat oftl!~ ;m1rul!1111
1/umn.i.ni!et Compnrr.-,. 35. 583,-390. buamf11°'-rfl."ttl Sott,,tJ', 20. 19-15.
u tt~R. J.SI•• 1910. l.t·urocy«r-0011 in F.umJH.-·- Joumat ofPmNuollJg)·, 56 :A 11.u.,n,<-.,,.., r.. rn.-.5,\t>, to, ~'Rl\""->T,\\"\, J\.x .. 199,1 L"ulirord"'
(Supp!emcm). 16. (C:eralOp()&Onnfll(') ias \'t.·r1i>r11 of bhu•iun~u<' ,iru:,. ,im:c>Js uf 8/df'IKJ,
Q lL\_R.,..I\J.U>. b,J,11+, 11}~.J-Circufadon of Afr1c:m horsL•si'ckuc..s vhLh ju :n·b1a (I 11d/11a11nJ. ID. 1711-l~U.
1F.Jjuusburdwltn In tho l(rugor 'l:ulonol !'ark. !>outh Afrkn, as m~,~ur<'d lJ 111Hlf">'. '1 H tlR-'\\I U\t\,, Y. 4' llU~U. \,.... 198,1. l0\\'3(d$3.~t..~· n{ chc
b} tho prc,':1l~ntr of l)p<· ,podfic >ntiboJ1.,.. 011,t;,r<1q1,0011 /011mal ~r ,ur.h~I nm: :md pcri0Jk•11y 11i "SOmt Culit.~,ut~ 'f"-"<"ie-.,. in hr.td,
\'t•ttrlm1ry R,-A.wrrlt. 60. l I 1-117 rn,1m11ml•n1UI F.rlUJmol"lt(. 13.-1?4-l!.1.J.
u "-.tt~.\.fU), 11.1.11 ... 1~ir.- Some i.:tcco~ gt't\'t•m1n, ltte ~ntry oi C,41/r-<,f,f,._~ ~pp. ~H bL\C~Uk..'-,' KI ?<ol'Alu.t l • t'ULU•l!io H,1.. ,,n.,, \"itu:,C'-' bolo.tC'd fttm,
! Diptct-'I: Ccrnto1>os;onttl:s(" tnto stnhll"t\, Otrt.lt.,Jt1..'J)()Ort Joumul nf Cultr:(l(Jr.s 11)Jp1cra.: C"..tra1npOJ.;(ln\dc11..•J cnur:}\1 ~t tht• ,·c1crfnn[)· re,ir.s.rt:h
\'tr,·ri11n,,, Rt!.W'tlrth. f;.• :r2;-233_ farnt. \ta.znw~. i'Jmh.nh\,·t·~Jnumul of tit,, /l111t>mtJloglcaJ ,W'iNJ'n/
15 11.\'R;'I.Atw, H,hll .... UL:-.:r.1, R•• j,;UT. O •uu;..:ut. t •• 199,:, £!.p1demiu!ot0·<>f .,~mii,·m -ljnm. lij, 3.11-.1:iu,
.\fr1can hurs.C'ltknL'$$: a.mihodJcs In ft~·h\1n~ clepha.n,~ fl.o.'ft>dmua 35 nt~'-<-"',. •u~ ,\. \()O~<.. ,ur.. ,. ,to1lDur. w .• 1qq4, The rnir:oh:.1>11~1 of
11jri,111w, and 1hcir n....pon."r to \l);JUrrim\•ntaJ lotectron Ond,•r,<t,•P()()rt <:ulrtoJdttSfmr:nmctot.uJ ·uJpt...,ll:. C:c-rn101">Qt.1,on1dtll•l lan'ol! in Scor,~nd.
Jnur,ml ofV.:t~rilu1r;' ~.'t·<1rc/:, 62. ?71 ...z;s. /lt,lt,•tin t,JEttu11m,lo,:k11l R1.•.<1."11rtl1. 8J l9S-~(lll
16 n.\R~ \lU), n.J,H,, c;t JU1l:S, (,.U. Se: \1fl~W1Xn.L. n., 19,s.. Somc;s 3-6 1UJ0R.,1.\...._._ J.. 1391 A rt.•\·1c:w of Cullroi,ft's '-Ul:>Renu.s..:\1wn1Uupec1L"6
cp,dcmlologi<al :i,pec" of n bluctougw-hk~ d1sea,e in :;ouch ,\fno;u- 'lrht"l.'.t.s, Otprern. C:t!riuopngonidat"l vcct(lf"S of ,im)t"S of-sht'<'J>, qmlc:
l!:'$"3196 and 1997 o,,.,f,.,m·tXH"t Jmmml <1/Vi•.n·,i,wry• R~W'orch. b3, and h1'r'-~~ ,,1th pnrw.:u.t&r rc·fc:onco ,o Cuhroit!t'>-$ imfcoco 1n Eurcm~
1~5-IS1 3nd thl" ~fedtrcrraDt"an ~1un. A rep<tn dm duped for 1hc O\ct.,ea~
t7 rt-,ktu.n. C....\1.4' .\'<Ufft-.o,. ft.<:., 1980. l)t..~X>li,pment of Cluintlltrr«•lla Dt·wlopmi•nt.·\dnuni~tr.1uon, pp.. >L
chiru·mxla<• JFila.no1d""' Q"chQC('rcldocl in C:uli,ofd,•,srllo/,,..,:i<,1dc, 37 UOOfC..\t\'\. ,. t DtPTOLU, (), o. 19z9, A lA,\0:lOnl1C ')IUd)" of aduJ< ~i~crian
and CIII/C'Cnde,,; tr<T1"1i fHp1t!-ra· Cl'r~to11or.omd~, ("nna,llm1Jm1mnl n/ C.ullct,n'1lit1 J,Juteillc (f'lipt(lro..: Cur.nopo~uo1-dac-s;pideli>I E1111>mol11g,ral
.i'..ar,/og>•, 53. 1002-IOOI.\. ,"io<,,,ry of \'igt:rin t 1n:md11nal PubllMtiou. ?.2. pp. I Z1
11 Ii.nu. c.,F., 1989. B1uctonguc. P1"(1(:t1..,ding$ OJ rite. .s.,~mnd Jm.~1111uio11al 3~ D<>tt•f, \., 19~~ 1li1lni; midge, from UPI'"' I :t,w,ctou~""" Jetw)
Congress/orSltt'V/> \ ~·11..rrlnnrirm&. ~l'\\' 74•.ata1\C \'ett-rinar'} A,-,oci;,tton. -\sube, H:~racopngoi\!d~c,: Oipter.i .r\m,•fi(,m .\1us..•1m1 Vnrirnui. 3 IS9.
~,.,..,)' Unlvc...lt) P,llmrm~n '1/otth, 12-1;; ~,·bMt')' 1!189, l~9-3:o7 1-:!9.
'" MVUY, T.B.. 18:;6. '1/otr<oa tho hoN•-,lck,:es-,J< lhoC'npl'ol'Goo~ :i'il bOf.tKI., t ..\, 41 \\'lnn1, W,\\. I~!\:" World Spttic,. r1r blt-inginldg~ ll>iptt"r.t
Horv. ln 1$1-'55. Cllpc lown: Saul :i<ilumon i.. C:o .. ~icam Printing Ccnuopogonld,"'!.t'-), Httlttti,z oftit, .•\J,urlen11 Alttti•unt of \'a1u1al lilslcJ~·.
Ofiir.c. l,;n11m,lfkct Su-""t. 2.n.1-2,\7.
20 8.·WU'i. XI-. DOU•~\"(")U:\'t, II .• TOUTI, J. Ac El. u.,:-,..- '\001. u.. 1998. Us~ of !O llOUtAAb. ,·,. L\~DAU. t. \llllt...,. f,. ft\C<'.,'1, U. t,ttU\l?lfJU'.\'•S,11111, 1••
tHmutfc doto. .1nJ ""'tt1Hte imngery 10 mudl•I U~ :1bundnnu.! of liJS;i. lJl1Tas1roC"turnl obw.r,:utmn~ on 1hc- ~J)Oh>J,:oh) In HrJNUOC),,,-tiJ
Culit1Jft11-s lmlt"ofa. th~,t-.ccor of A(ric-J.n hork,fCkJ\~ viru,. In l,::•lttri. h~mopl'Otddl1n pat~'!-'.ite or Ao.,.tru.llau Chiropu~rJ. In Cnlfoold1:s
~toro«o~ \lt·tt·rbuu,• mu/ .\fi'dicol EllfontO/l)J,.'l'. 1..2 l55--26K mllh'<uf()..<lll.. Prt>tittvl<1sk11.. :! l 93-105.
21 u,,., 1s, ,, .• MCJS\\'lS~H.. It. & \'E.'\"JVt. ,~. , •• l9$S9••, prethnln:iry ram?mpr 10 ,11 nn.,\·lR.\t..\N. Y. 197<1, Ch:1rn,1~ristiC> ur C:u/i('()/tlt6 !Oip1cra~
u,~dhmuc l(:.l\.a umJ ~utUitt' lmll):;-<l't)' U.J 111(.,t!d lh.: ul1u11UJlll.:l' 1Ult.. <:t·1,11op1.1l,,"U11ic.t,n.•) br1.,-dln).: ~1fm.t:'> lhr•u So?hhut). RI Jil..,_,.,,,'1-. a-t,r,,,;,wf
,hsu1..bt:don or 1.ullt:'tnd,t$ fmlt'()ltt (Olptcrru C~opogtinid:u,.. l in /'Jrtomolag;, 3, 163-1,0.
'.\Uuthl!:n Aftiro.. Jourt:ol o{tlr~ South Afrfrtm \ i.'"t.lJrilUlf)'1'w·,xwtifJ1t, ;'0, 1:t ott,\\'111\l\S,, , 1988. l'rdmcd lar.dfng ~le:,, o1 Cctli<cntlt"$ ""flC'Cle,
80-89 lV,ptern.. (,.era\41pogorud.a~1 on n horw in lsr;.1c-l ..snd 1b relt\1lnt,: tu
n n.,yu~. ,1 •• :.rn1Qr,i , •."'.',..~ 'lflS\'t'T:\~1, R.. 1~:..~. Hoti1e~ick11~, nnd t:~l:>O sum ml.'; i-ea~onal rc-c-u.rn.•nt d"'nno1llt~ i,...,wc1 ,tchJ. Et111m1· \ (°tt"fltUU)'
ln S0u1h t\fric.,, .\'Ufm'1.39i'. 514. Journal. 20, 4~&-4Z9,
~I 11,1,,.._~ "·" """1.1,0, P.• 1998. Modelling th,· distnb11don ,Uld 43 I\M.\\"1:P'tt.\~. ·.-... ;~,s~. Uu,1 dN1.,""tlan, hourly .,uidl) ;md p.rC:'ftirrrtl htrlns;
ab1111dance of (.'r,li,ol,t,.., lm/ooln in Mo10.:wand lberl• u~111g cllm,ulc ~it\'"~1:1! Cttli<eldes imlcoln ID,ptc-nr: Ura,opugumcJac) on a caJnn S.,-rn~•I.

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128 ,~c,1," (,"· \spcct~ influencing 1ho occurrence of infec11u11~ thsca.es

ltr. WAJ..l'Ox, r .r. , O:.,,BUft.X. s.1. 1cus. a!uttont,.'\IC. Afrir;m horsl? '-fdntss. oowr.~. P.A. tssin t~1'1llOn ofTt..'te -.cmgroup bunyi\iru:it..~ tmm
and telatdtf ocbh.iru~. Pme.,~fitl.g!Oftilt' settuul iu1tn1tui.011ut ur:1ropoG<on1du~ col!~a1cd in Colnr.uln n,e. imme,111 Jmir11al of
s)•t>1/J()$(11111, Boe~ Ro1nn. Flor:dn. U~~A: CRG r>ress. Tr11pir,1l Mi'1/l,·111r1111tf /f.VJ!ltllt. 5.~. .l H-3 f 8.
,U llRAVl:R('.t.\N", Y.. R\FV'.lf.\1, T.. rRJ"II, "· k11UKlS<\, '\1., H)~S Blt:ctongue\lnr!!< ~~ (".\I\Tl"lt, u.1-.1919. ~cw\\'~1 A!nc-.,Ut Ccraropus;omda~• ..\1111nl.t<1/1rc>,mw1
1)0hHions (nun pool~of <:ulicordn\pp. in s.~roel dunn& 1961 m 1963 Mtd/<1~~11111/ Parn,w,!11;:J· ID. 1Jl-13lS.
Pmg1-.•Ji.-q11 C111m~·1I Rfofogu:al H,"Stvuch ':M t 91-1 ~1.3,
65 c.\l11HL 1f.T l:W,ft•\'\1 • .\,' \t.\011 J,\U1,1 l~JO. ()b..('l'\~01'5-0n the
;:; Kfe,\\'f:A\ t.\.'=, \,.. il,Qftf.H'\M, 11 f _t.. & G.U.U'.\ M l~':'1 The nris:in 01 C1."raiopogomn.-. midg\.~ oftht C:<>!d C:~i\">I "i:h dc-scrtption~ of lh'W
bloodm~, o: lcm•lt• C11/itnld,•, p.11/flll;winis 1ruppcd in ,1,h..pfold ,n ,peci"' l'an II ·\1111al, ofTrO/JM>l .\1,•tflrln"ind l'nra.lto"'I!:)·. 14,
!\f:u•I. Ju11mt1/ of .\fr,//rnl F,110111r,/og;·. 8. J,CJ-:td l. 2! 1-?:'·I. Pb 7 8 .
.:6 URA\'l'NM...\.~.r. ,. CUIZO\ •mNY,J.UJ~- ~'. J!l-97- Rt,)tllc1tq of~~ nt.bt'tlt attd 66 C'AUSU.', O.K,, ),.h\U>. (..,1 G\US{Y c. ,.,. I U, \",11.. 19:-.:. l..ol,Hion nf
pla.nt-dor:\•t.id p1upanllon(; for CuHC'ol<k, imiro/n ,\kt/Ind ,md Sfmbu-pnup Vll\L\C:<i ln n,~d~. Nl~~rt."1 Ulfi.l-19().~. indu,Hng the Ill"\\'
\:tHt•nunt)· Emomol<>J:.'. 11. ;n>-360. !)pc., :..ngo, Shu111ondo. "4bn ond Shuni .\nn11u nfTcopl(n/ .\/,•didn,
J';" HRA\'l:Jt)t,l.... , Y,, G.\J.U\, x.,,;., IJ\, ,r .. 197.s, Rrtedin,r~t:e- of 'C>Ini! m:d 1'rrmsfto/1;gy. 66, 35T-3iiZ
Cullcoules.iped....., ·1>1pwra: Crn1topo1,,"0nH.i...1t!l In l~tUll!'l. .\!O!lJui:o .\'l•u·,. o; ('.J:L\."'.ilfSIH<, , .• U~U,\U, t. C<1 8.\C'U'-1. 0 .• 1~~ Sur h,*\ l.J'!,'pli-llO~m~
:K303-308. d"oi~i:a·Jx c..lritd1Jttt:. It, Et\JJt~ bwl~h1u1.·,-. .·l11nnl.~ dr Par,1:Jito71>8l"
Th(! rdallon-.hip tx·n~t.oen the
-18 IIR..\Yt.8).t,.\X. ,-. ., 111,.n.LEY, P,1 ,, 19;-,:1 ll11malnr "' c:01111,nr.,~. t,'3. ?43,..252.
numl'4,!t".S of some antcnn.,J nnd pa.lpill ,('flq- org~nS, nnd host prcforcnr,• 68 GH.\Rft0\11 ,1 ~·f'Ul, I •I 1949, ·nn.'"Ot4'Urr«.:nc1.•C)(filana ln tbt• n.•glt;n or
m soml'.' 011iroldt, fUip1cm= Cc(a..mpt1g\'>n1d:){l) fmm i,c,11tham :\lr1cn. <:ottu.tlL11,,1t~ nnd tht• 1rnn:sml~.s.ron of n1p,,.·111loth~m111trepWrt1rer, h>
Journal o/..\fediM.1 F11tonwtu~·. 15..; l 9-12:4 C:u/i(0/1le,">gt111'<lmiJ. ,-\nr.alr~S<,cfrk' lkl,:,, d,t .\ftYllrtnr Tra11Tmt.· :?':,I,
.:9 DR.\\?»L\..,., v. 1- usu.,·, J.n.. 1&88. Parlt>· .1nd \.nhin1,m of ~,..,cra1 9!>-119
Culico/des 'PP· 1Oiptero; Ce1u1opogonldac I in !;rod, ""deitrmtncd bv 69 C:IHMlK't,u. ,1 4- fllif·L, ., 19;1. Re-warch m1 thcocturreonct 01 filari.-:. in thl"
1wo t:rnppmg me:Jiod~. Journot of .\Je,ltml F.Juomolog,-, 25. 121-J 20. rtogjon pf r..oq,.01l,u,1lft.· c.nd thC' tr.sn"imi-....,ion nr l.>ll)f'talartm11
SO bl\,\\'fltM,A~- , ... USLEl.J.U.. , MARt:u,, A..~,,,..... 19il!i, S'-',lSOf\,al lttw11"'·~·,rv1 b~· C11lk1Jld,•s gmluimil AUSWlt M1111wim /mtlru: R<JJ·at
~urvi\'tt.l tuid ~Pl'Cl,ttlnn or infccttvc Uit"01 Cuh,m'fic,.I :,pp. Cu!nu(cl 81•/J;lf..4>eaio,J ,,~ Scli.•11~s ,,·mur.-1{~ rt ,\/tldlwlr.t.. 19. :t-83,
(lliputra: Q,ra11111u~nldaeJ In 1..oel, wl1h lmphcatfon~ for
70 CHJu~TCIP11lHS,:fi" It .. .... nnn. H.E. & fl~HKAUI>. r. ,, 1.92..~ C:ulfroldrs1utd
bluetong,uevinn rran,.mfi.:sfon ;utd .:icump:ui~n r,fth~purous rate ii, C. i..:a.Ia•..\.'.nr In A,,,,,;1.n1. ltilflnn Joumat of.\.f,,1funJ Fr(·~t!tm'lt, .3. l 7S--.l ifi.
imiroln from l<rwd iltld limbah\\'e. /01,mal oJ \lrtli"'' f.lltQmolog> . .'??.
4i6-IM. ;, cu,,""'"· ,.. ,9si, 'l:ote,;,~r le- CcrotopoMonfdl!> I\. C:t'm1op0g1>nldt"-
d'Mnqut ocddl't1tJI,• franc.!i><, . ..lr<li/i,-, de /'/lt'1Jt11t P#lrl'llrd'l!s•r:e.
;;1 ul\A\IJ'A"'-A'· , .• ""r1trlll',. n.J, 1~1 ~p.l.'ct~ct,rnposition ~uJ b!ood•rneal
36. l!U 2.58
ld<nlific.it!on in sample, or <:11/i<Oid,•, Olpt,:ro: <:ora1opogon1dJl~)
collcciod near ~•li,bUI)'. Zlmbab'-'<'. in lG,6-77 Jo11mal oj1/1< ;-: CJ..A')tnu1M, J., ii;S'l, ~oh:·~,~· h':!C. CtCr.uopo~Qnfd("\ \1. C1!at<>f)Qt:OnidC,
Fnttmwtosn·"I ~,d,•1)10/Sourl,rm Afri,,•. . ;.s. 315-323. d'Afriquc O-:.'cld'-tttl,1h.· (t"Jnahe (Jl, '\r,·11it1•JC.dr l'ITl'lihlUl Pn.mm,
d:•tJgltN. ~i7, lf'ir-197
~ nfll\\'liR'lA'-. ,,. A Huiu x-,. ,, . 19;ti. Lighl rrnpping of blUn8, insect< in
pouluyhou~ ut hrnc-l. lt1Ylf'l/our,~11! o,'/AIJ!Q~·. J. 95-101. :J CL-'~,xn..1t. J.. 1960. ~otc.·c. ,ur I'->< arotopo!_.:onu:!(' ... lx. (Cffitoputonld ....,

Sl ax.,,1:.~1.\.,, ,·. •um!\.,,.,,, tri, ,usu,.:.. 1.9,Kt, Pnchattt11h ufvet.Uinar,· c.lto la Rt'pubUqut• du I" :ont,.'O•.-lrrllwt·) th• l'Jm.11,·m 11,uuur t{~llgrrh.. 38.
lmpononce i<ol.11<-d from mosquuoe, and bi tin~ mid~~ In lsmtl. 1,,,,-r1 ;9..!03.
S('/MttApp/1<11UOt>, ~- 1';7-161. :..1 t:l.,b'J'11lDt. •• .&. \\'ttfUI, \\•. \\ L96t. ~OH,!( .. ur ll.... C:t•r.JIOJ)O(tOl1idC~ XI\'.

34 u~wwc.,A;,,., y, "-.•., \!\ti•m 1.. 11.- t98"1. tnfccuon nml 1rW15mission mDh C4!ra1opogcmldl!< ,le Li IIA.'glon ethlopienne 121.Atrl,/J\•·,It• l'/1:stilllt
PflJM/1 tl'Ali;M,, 3U. :102-33;'
,,ith Nyub1r3 ,1ru.., m ,ll'dc,s flCJ!J7Jfl iDlp·i.:ra;. r.uUeidne) nnd two specu.:-S-
or Cuh,Y>id.cs <ll)fn~.m: CcrntupogonidaeJ Zimlk,tm¥ VN,,.ri11no·Jo11mt1/. w co~to ,:r.ri. 19.a6. ·\J~uri.. Oilkoitf('.{ do ·rr.,n~\·nn.L A111inl1 t/1> lru11tutu
12. 13-17. JJt.\J,'f1i£1111J Tropm>I. 3 21:-266.
~ 81'1\\'1:R;\1,A,, ,. fJ'.'\G,\f'•W,\RO~. K .. ffU~H. ).,. ADUn. n., D,\Sltl ;. ,. • 8\J,.l.R. ;ti torus~. 1tc. l. Jo:-.;r.s. It.tr.• 19;8. L:tbor-110"'' trdn~rn!s.1ifon of O;,c:/Jott""rro
)..r. ,- Qor:,~. P.f. 1'8..~ EpiJ('nuo?og,~"41 ind ,mmunnlDi,.rtCal ,:tudfC"\ of rt'fr/tJ,iif wilh CJ;lfa1itlt'~ t>tJrlf/h'llllla. Am,.riam /tJurnr'II a{'fwrtftai
SWCl'I itch in ht.>~ in l\ml"I. n1"• \'e/lltttll,Y R«u,,J. 11?. 5~ 1-32,1 _Ht•,tld1:e oml Hygfrm.'. 27 n;... 50,
;:& kRctuRh\t\t, u. At tAA~"-· A •• 198-:.Al.!~:~l" d~nt'UtttU .. f \:\\~C ft.Chio! • , cou.1""1S, ti,\, "P,\ft.Pfl(, t 1.• 1965. lxol.ctlQn ~d ldcnti.ficauon of
l<clandlc hor- In S\\c'\l~n: All epldcmloloWrnl "udy. l:q,1/11~ bhicto~c \'lfll~~ a ~erot•,p~ nl~W to lhc IJ.~. Ptt,;:.r~ ill Clittie<,I
l't'lninllfy}uumill. 19. 229-2.36. R~""tcY1rth. 178. 319-327.
57 kilt)\\.!\, M..A., ~01mua-: I t u, IL)• .\.I .. Ui\.\!IIIJ\'. l.., \'l"~t l 1\, C'... J•. -l.-\JQ'JU)U'f... ;JI CO'-!\,~. R..•,:. i u.oro. s .• 1988. Sta,;,on.'\l ,'111ttQ;u: drm1.1utk 1n ,h~p. 1'ut
,,.. , 2001 • .Ho,t pidcwnc~ "Jtud!6 nf C1,liroitlt"i bl ting mfdg(!j.. Tfo· Ror11I Vtt,•rir:tll)' f(t't'Orll. 123. 3..~5-337
EJt;omnlogiJ:tJI ,'iot"rttfl' 111:rr,uuional S\1r.1,osmmo11 lmrY'lSU1UI l"Jiu,1.w,
1()..12 Scpt<mll<,: 2001 i!'1 COOK". n.,. 1!171, llaimOJ)f()l1,."'tJ,S i(.""'lJSC 1800, /n! Da\'tt.. l.\\'.. {cd.J.
1>,f,'C!iQUSt111d Porautir Di1<·,u,s11flrl/d 8:rtL• lowu; low• St~t<
S8 nucxu--v, 1.,...... 193.3- A notl! on I.hr d~·a-loptru:nt nf 1-·u,irlt1 o:.:artll In
Unl\~N11,· l'rcss pp. 300-308.
CuUcotdes furt-•m Pot•)·.Journal ofHelmmtlzalogy, 11. 2.57-~58..
so ~>Rst-r. "·• 1909. L,>s C111/ro11t~ U>ipitr<> Coro1opo~omdaoJ d• l'Oue,;1
.59 svoa.hY, 1.1.c.., l93-:, On 1ht• dc-wlup.1n1.\flf m Culltui(ltsfurt'n1P01:v ot
3.friah: t li!re note· Ct,t: :\111 ,!,t J'Off!a, ,trJo Ht"'('ltL·tt"11, ~ltnrlfimw t•t
l:Uaria (i.\lmuoucl/J1 m::.uuli .Mnnson. 189t Jownul 11/lltlmi11d1c,ff>ID··
T1.vimim1r011trp,,..\frr S<·ric 1-:nwmolo,at,, m,'dtMI<' t'I ParaliJt,log~. 7
12. 99-118.
3-:1-36-l
(>(I On O,h.,.tndcs as a \'t!CCor c,( OudttltV'rtugfbsonJ
IUJttr.an. 1.1.c•• 193&
81 roesf~. >1. & onu~uc<. J i,i94. RC\·lsion de> ~-Jk'CC> de Cull(o;tl,•s
•,C:khmd and tohn:oi(1n. 1910). /ourm,lajllrlmuuholus.>', 10, 1:?1-J:"l(t.
•17pan-,,1tl,,s a C S(l111/= ,EndorMn, l~llSt don, lo regiQn afr(llmpicale
~t ftUflOU!LL w 1.. :8,.: Trtn,•/1 it> tho lntmor0J'S011rh ·lfrira. \lot I, 1 Ojp1cm, Ceruto1,osz0n1dao>, Butl~·ri,, <le 111 So<itrlt En:o,,mtog:qrh'd\•
l.<lndon; Longm,1n Hu,,a, Orme. Brown. and (:rcon. Fra11a. 99, Nll-11>1
6:!' CAUHO. ,._,,.r. .
t9GL Concrlbui~ao p:ua ocSwdo das Ct!r,,p~ctMAngolanM
82 toRSliT. ,1 'lo'c.:11\TUU, ~. ,~;o. Les C111/1:0"4<"5:dr l'Oues1 ofncain !t.•
do genotll l'.u/lro1d,.., ~lleilh•. 180911)1prora: (:¢m1opnsonida•)./r111111 a
no1i!'1. E.ip«c~ ill)pJrtn1~e.s C. simill$Catwr. tngrom ri M:iclie. 19:?0
lft· lm~'!jrlw1f"c'$ Ci1,mlfi111. do Ultmmar, F~w,lu.<. fimn!us. t' D«unumtm, IDiptem. r_.muopc,!(or.id>c,. Cai1ter,,w /'OJ!it, dr 1,, Ri'c/11:J't'l>r
8S, 1-:159. 58 pl,. St'ientifiqut it Tt'Ahnf(Ju~Out~·,\l.:1, s.•.ir.t 1ititt>mo!ogk mtdimk.-1
Pnms,toloyft-, 8. J.: J.. 1,J.

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 \'enoN: Culi,·oide,, spp. 129

83 a,n..,n. ,1. & NE\1u.. E.~t.• tgS-0._ C11liroid~s 11tnci..nros1Ji n. ~p .. une Re<.OI<h11nd ,pi:c,es from ~ou:h Afn••· Publlcmto11.1 ofrlu, Sou/11 ,1frka11
nouvelle osp~ee d',\lriqu~ du Sud IDlpter.a. C'.e,atopognnidnc). a<«<: lrutttu1t• for .'l!tdfcal Rt!St~nrcit. :. 3?3-385.
unc nOt<" ~ur la <"1Xonomft <"!~s 6l~t>S ~thfopfonnH ft 3llt-S ~ni tttht"!i
t02 Dt MrlUO~. a., 19~2. ~Xl"\\' ~cmatoct.'f'3 from the f.thlop1;m Rt.1;fon
C.ahier.s d'1 l'Olftutlc la R«h~rclJIJ Srl11mifiq11t tt T«lmiqu,.,ourr~.W,•r.
/011rtU1/ ofsilr E,,romulogirol Stxfc1yo/Sout/lern .Vrlta, 5. 87-98.
S8rit Emomolog,c rn<'dlr,t/u1 P1misltuloi,sc 18 383-389.
103 r>t ~u JLLOS. & .• 19.s2.. C:t"ratopogontd:u; (IJ1pt. ~i'mamn•ral from
8,i COR~tr• .)I.. ,,;-1;.v1u.. &.-.\t.&•WAI..Khf'c, A,.." ,19;4. ~CltC.S\Jrlt'S Ct41ft.uldt."S
southt.•m Rhodeotio.. TrUttllil'llon.r of1he R./1()(/i'$ltv1 &£t•nt(f.t \.uoetndon.
COip<cr.1, Cc,"u,pc,guuit.lae) Uu ):JOUJh'-d~ C. milnc-i AU~(,.m, 1909. \."11
..'1.friquf Oric;<iltale ci 11usuttlt.1• C(I/Jit'rsdt l"Ojfit-t,fc lo R,'Chcrr:h,• 39. 113-119.
Sr~ntiffqur '-'' T1."tl:nit1ui.r Om,c..J\lr:r. .<eril' E,:umwtr)gl'r m.'dimlr-11 10.: OE ,1ou.os. a., tq,-11. Simullidae 3.nd Cenuopogonlda-, Oipr
Paraii,olo1fi<' 12.231-243. Nc-matocem} fmm the 1.-otnJ\}' n( \tfocnmh£q\u.•. £n11Mo.-Vui•mt1larit11 tit'
85 ~~'(. Q.)L. {LUU.., M..(.. }';ft.ANO". n.u. t. (;..\U~ln:8, c.11 .• l983. :\tho\'iN.4;eS
l.tJ111r1tto Marqws, 26 pp.
!tom wesiern Utah. 1967-1976./oumtl/ a/ lfrt~eal £11101110/cgy. 20. 10,; DB ,11:.1WJ;\. a.,1943, Nt-rw record~ and nc\\1 Spt-Cft.~ c,f Ntotnttla<'\n
l9.;-300. , Dlp:ern f1om the fJhfnptan Ri,gjon /o:m,n/ ofthe Emomotogi,•a/
86 ~111ss;;1, 0.11.19S4. Ooug4ts and Tin:truo ,1ruso: two Simbu g,oup <.«lrl)•of S,,ur/,rrr, ,\frlra. G, 90-113.
arb<),irus..'S infccting C111/coidi'S bflltlilau,s and Uwsiock in Au.•tralia. 106 DE ,,uuos. I\, 1947, Nu,v r«ordt,, and spcciriv of biting, in,ects from ,ht:
.-irwrolinn /ourr.al o/Blolo1Jic<1/S<:lt11e,,. 3,. 91-97 V.1hlopfQn Reglnn. II. /01,mrtl of rlir E,,romologleal Soor1>·0/Scu1hm1
a; CYf:11~.sKt. o,H. & MULLER. M,J•• 1990, Js.otanon of-crboviruscs from c:tltde- ,ifrlca. 10, 110-12•1.
Q.lld insects-at two~uncl sifcs in QucJtmsland.. Australia l9'is..8.5. 10; •>• "111.LCI~. k .. 1952.A nt,v ~pcci<'>OI Orllcordr; CDlp,era·
Australian /oumal ofZOOloifY, 38, 25-32. Ct,.to;,oi:onlciac) from Uganchi. Procr,dlng, ofthe Roiwl Ensomolog,ca/
88 C\"'Bi;i.;m, o.n. • Pn.·iimmury chltmc1(.•1iso11on of
ST. GroKGE. T,O., 19&2, Sc'1ifyQf/,011dar1 (fl). Z), 173.
D'Aguil~r ,irus a.nd three PaJ;,un group ,irusc; n~w to Au.<1ralf:t 1o8 00Hl!1tT\", t4.L.C.kJ,UJt\', ,.c; .. HUl1Pf0f, ~ .. tv..Rk(,)W, G,J,. wn.so.~. A.&
A11strailo.11Joumolo/Brolog:erJ/ SciMCI!, 35. 343-351
••ows. , .. 19;,. l;pltfcmlologu;al ;rvdle,,of 3rboV1ruses. Rcpor, 0/1/10
Clg CYD!~il"l; D.11., ST. GEOR6f.,. T,P,, STA/'.QPAST H.,\ r,, MC:GfU..GOJI, /\,, 1980. Q11wuk111c//11sti1111,o/Mediral Rrm11clr, 30. 6-13,
lso!ation orT1brqg.ugan ,nus. a HL"\\' Au11ruhan rh~bdov1l'U$. from
109 OOTITRTV, Jl.l., CAJU n·, J,t... Al.1PP10l1 C .. GM\A1T, 0,1, IJ.AJ\ROW, G.J.~
Culfcoid,,, bn:t•i/al'.is. Veterinary Mirn!/Jio!ogy 5, 301~;108.
.,.,.,.., n.. 19;6. Epldcm,ological sn1d1cs of ~,bo,iru>es. Re1>0rs i,f1/1<•
90 o., COSTA MFNUF.5, \',M,. ig&.; The isol3tfon and ,mJ)Qnantc of Sirnbu Qw,emfa,uJ /11stttmeof.\Je,Jical R,-s.areh. 31. 7-J.l.
group ,·lru$tS in Sout.hAfnca. ~1.~led. \'et. (\'ir,) Tursi:;. Unive,s.ity nf
HO OOJIEirO', R.I .., CNU.EV, },C.,STA...-..DV'M"l, JI.A,. o,-ca:. A,.1.., A;W. 8.11, •
.Prerooa.
,so,,'1)():,:, ,,.u\... 197~ Virus.stf\\in1oi isolntW from :mhropodsdun.ng ITTt
91 D.\S GUPT1;, R, f< PAL. N.. 19,S, :i,,tnlarj~ QOC)¾'i u, C:1tlfc-0Mc•s .:i,p. In ~pi:,.001ic o(bnvirit t!J>l1cmcml fe\'cr in QuecnsJ:tnd. ..\usrmlltm
Oftr1eel!ng. Trw1St1C/i~os ofrh~ RQ)'(Jl s«:ral}'of Troprcal M<'<llti11, nnd \l<turlllhry /ow..aL 48, 81-86.
Hygiene. 10. 89-90.
111 oonurn. Jt t... CARu.iv. J.r..• ,.,..,-'"tw.,~'T. JC •.\.. ov«.A.t... AA,·. t.u. &
91 0,\\"16$, i.~. & HlGlfT'U~. R,B,. l9tl<.t. f'O'Mtbl~vcc~o~ o( t<tft Valley f-evtr In SN0,,'00~. w ..Ao 19':3, i~llulon.\ of ;&rboviruses from mOMtultoes biting
i.:onya. Tronsacr/ons of11,, Raya/ S~er;· o/Tropicol .\/edlcine1111d mfdgM, "'1ndfll« and ,..,ri•brntcscoll<C11!d In Quo•n.land, Hl\\9 ond
H)'g/"1rf, ;4, Sls-816. 1970. Tron.mC1/011s of:he Roynl$llrieryoj'r,opkn/ .Wedlt111, 1tnd Hyt:fonc,
93 OA,1£5, F,C. & ones(), s .• rs;;. Eleph•ms Md ,:,bra,; .is Pllh>lbk tCSOt\'Olr 67. S36-543.
hons ro,
African horn-sidwc>., vlru.. 11:c \·,u-rirmry Rer:ord, 100.
lt2 l)OHt:A:n·. R.L. STA.~UJ,MT. H.A .. ova,••\,L, CA~. f.fl .• f.OR}~ •. ,, ..
:?91-292.
MUPPICH c kNW, n..1r. 1978. Mudj1nbany ,·lrw, .an orbhiNS rc!ated co
94 DA\1.L~ f,(i,•• $01, 1t.-J;.,. an~EPAL. \'A. t9SJ.. .\mean ho~ ~rckrn:!J) ..;~~ W:ulul ,iru, i;olnted from midge< from 1ht K<inhom territory of
isoloied In Ken;• 71:t Vrwfrwry Re<'Ord. 132. ~.o. Austroba .~ustmlinn /011mnl o/B:o/ogtc11/Scie111:,-. 31, 97-103
g; 19;• The isolation of cphcmcrul fe\'er-iru>
DAVU\S, F.c. 6 W<\UCl!ll• .w,,. u3 nousu.. , .. au~s. , .• \\'NU c . ..-~nutn:·>:n·, ».. 1990. Di.suibutkm of
from ~aulcnnd C,,1/roid,s m1dR!!S In Kenra Tit( Vrtcrlnmy Rrnml. 95, microftl:niac of 01,cJ10t:t'fco lt,~Nqfis and Onciwnucn guau,oso in the
GJ..6.;. skin Ol c.ittle &n Genm.my :ind thc-1rdtvutop1nc,nt in S:muft1m1 ornowm
96 o»srs, r.c. • WAIJtl'JLA.IL. 19;, Th~dl>1ribu1lon In Krn1·• ofbluNong,.:e and CU/it:oitlts 11ul,e,:,1!Q1tJ.t fottc;w.inp- a.ni.rkial infes1ation. i«t1i,uuy
\in.u. and ant1budr. uncf ,he (:ulknldf'..~ \'4><'t1'r. /m,mn/ of Hy,:i7tf-'. Paras:zofog:,·. 36. 325--332.
Cnrnbrldg/', n. 21i.;..2;2. 1111 pu>,:-E. e.o.L.. t9Sli. The intill:e o! th~ microiUari.u: ot .'tc.l!IJ/li0tlr~to11vma
9C DA\"IIS, F.(;.. WAUa:R, c\.lL, OClllt.'-C, r. • '"'"' T.. 19;9. Arbo•iru~r, i,trsums b)' C,li<oill<$ 1,'rolramir and thefr sub~uent d(,,.·clopmcm
fsola100 from Cirllr:old,i midges fn Kcn)'ll. Joum,d of<:ompnrar::, Annals, a/Tropical ,\lcdid,u ,rnd Parn.1i10logy. 48 . .; 1~20.
Pa1Jro!ogy, 89. S8i-59S. 115 oun>rr.1t..:.t. 19.u.. rhe tnm.~ml,iiion orblut1ong.ue and horsesic"kl,w
g$ 0.A\1£S, J,lL. TM~,A.J, MCC\1.L P.,J ... B0¢J.A.RIL. N,J,, Tk();\,U"SO~. >1,C... b)' C11/rc»ldC1 Ondi,,supr,orr Jo11rn1d u[V~scrinnry Rl"Scflrrlt. 19 7-lfi.
MCIO:IJ..AM, ,u. •
'"'T!Vkl,A.$., 19119. On th~ pos<ibllit) ofbo,ino 116 1>u TOIT, lt." .. 196:z The role pl•yl'd by bo\1ne, in the 1:.1nsml.<sion of
Onc/:occrm speoh.·~ In(t."fflns,Slmulium dnmnottmt unsu Imo In th,: bhmomruo to '1>.eep.Jo11mnl of 1/1tSc111/1 ,l[rico11 l't.<'ri,:aQ· .Hrdicnl
fore;r ,.one ofSlom, Leone. ii. Bl ting dtnslti!K ~nd lilari\11 inroctlon• tn .AJ.!i0t!.orto11, 33. .;83-l90.
StmiUum spp. and Cr,/fa,fd,sspp.Anmlls o/Ttopltal Mr1lldn~and
Pttl'll$/tology. 83. 60:Hil4 117 01c,:. A.L, 1988. <18 Quo;,nsRd .ASquilh, N.S.W. 2017 Unpubllshod datn.
n8 ow:1; •.,.&... ;., ""4K!\t1.,u.. B. o.• 198,s• .An e.arty record or Culicoitlf., .spe<:ies
99 OEIJ.A·f10RTA, "·'·· '.\IU1U't.W. M,O.• IIAR'rl.r,·. \\',, .. n,\'.\'Df:A$T. fl.At. S?
GtORC:l. r.o., 19T:,. Ai.abane ,;ru,. Rrf)()rtoft/r~CSJ/1() DM#t>n oj,v,imal iD1ptor.i, Cet1tnpogo111doo1 de,·elopmg in 1h• dung oi game •=Is in
Hcp1tl:, l9-2•t. wu1hom Africa. 011dmupoor1 Journal oj \'11,nr,ory /1.:1,nrth, S6. 85-26.
119 E&l'JUCARD, M,L &-1,owR,r. R.C., 1.979. Dcvetopmt>nt of l)ipqrakmc·»a
100 Dl stULt.0,·. ti .. 1936. Entomological itudl••· ~1udfes on in=, of
medkal lmp,,nanc~ In South \frlca -Pan Ill. SouthAfrkun gmrllt and D, cmrd,splrtn to ,he infective ,,age,, In C11lt<o1d<n iroilus,sss.
Ccra19pogonldn1?. Pan ii Some new and unre«irded $!)<!Clh. /011111<11 ofPruQsirology. GS. 89-95.
Pub/katio1110/1h•Sourl1 .1/rlcoJ> /ns1i11trt/or .\f,Jfa,/ (l,..,:nr,lc ;. 1:lr;) 201soru:-.. , .• 1895,. Ca))(! He>rsc-Sitk.nC$s. 1hr \ ~terlnarlan, 68, SCJs-GOi
141-207
121 tlJlEL k.f,, CR""- <:.t." CAIJ.l-HfR, c.H .. 1977. Arbo,in1s twlollon rrorn
101 ot >fDLL!):<, .u.. 193;. Enwmologlcal ~1udfes.S1ud1es on insect> of sou1h,,-e.tern Ut~h oncl nunh""''tm Arl1.onu fn,ccn l9n-J 975.
medical !mponance In South Africa and ndjaccm 11:mtorfe. (l'a,11\1. 2 1/otqtr/ro S,ws, 3?. 401-.505.

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130 sh:. ,o~ ,,,i: ·\sp<>c1s influe.ncin!( 1hc occurrence ofinfoctlou,; diseases

t ~ f;.':Ol'lU.EtS1 c;.. 1908. 2. C"..ermopogon:dar ~e11t Ccnitopog,vnuum m1, tn~lpm In fllll 1cht ond rr1.itt'd tn 1n,rogurgan v1ru, of Au,tr.aliu
Sudai,ikn /11: 7.oolui,,..che und on1htup<>low.clw Et(lebm"• c>1w1 \ "t'H'IWOT}" ,\Jt('((Jl)tulo;r.•. 19, 1-t1-lSO
For$chun~reis(' 1m \\'f..:dichc-n undr 1,.c:nrotcm ~Hdafrik.J au.~~t·iOhn 1n
1.u v1tho'\, 1 , . ., \:-.( ou, \\" , .. 1952. The rdauun'ih1p a( (t,l,,:'>ltleJ tU1pr,'r.\:
dcm Jahren t90J- 1903. lits1er Band Sys1~ma1ik und 1,eri:~aphir.
llel(!tdoeJ 101ho Lmnsmission of Owl,o,w<n 1>11!•11/11.< Juumnl of
t.\Voiw LiC'ft!run~ ln~ch.· F- DfJucr:t f>1mksrl1rtf/lm di!r
PttmMMft>~·. 38, 3 f >-'.t?O.
.\frtli:i11i!-th•.\O:uru•1:,:.,r4..•nstltf1Jfllrh~" Ci.'"S,.·ll,,tlwft :.11 h 1m1, 13. 4S9-161.
~ PAq1:11., ;,,:. ,. tm .,n T,"'. 1,.-;-s, C11llm:df'$ ·1ml·nw11,·(.\,.,..,., rn lOJ: a 1,u t.iU<.:lt • 1. 1990. C111fri,f,tn ll:untt m1dg<!.,, I OiJlCrt,l. (.~r:a:10;,ugo1,tda~~ uf

pos>ib!e, «tor of Akd;qctmll~ry-; in penin~ular ~h:shi}':.sa ,prchmlna~ 1,,;enyil. tomtml of.\frdlr,d ElrrfJ11u.>logy, 2-; e · -! 95.
rcpon1 ,C:oji.tw \t·un»alhl(,1w11sapor,·). -. '.:l2-Jb. 1:.1 t.Qtn,\ln 1 ,; MIU"u.,1, , , tgfl~. Rluc.·tong,1~ 1n hmwl. /I:· •.,Mnu. r.1.,
1u HHR~l',N. 1.. \ ..\,..:\,. J.: "''"":uvu-1tr,,n·. u., 1~8-fl. ln~cMit)nof miuutilru1M· 1oc1n:,l :-.i.,, .• tt:d'>.l. Btm:mu1f111~<1ml R('lmti-1 Orbi,~iruR"'J'. ~cw rotk: .\l;m
horn differem ~our(t~b)' CrtlittJith'I m,lw11/o!'tlrJ tDlplcr~: K. I.;~:,,.
CCr.)lOP<)J:;onid:w) thruugh,.m ;anlflcl.il mcmtm,nc. \·if111n,rory• IH c•1\l l '-t'JI. 1·~,. 1N?5- r,.::v.sicnc~· alld 1x1ccmint ~1.'Cl<tJ' of lfu,•111upn,11.•us m
P<1ru,i10/QJ;;· 28. 3!5-120. Wbro;~ mourn1n1: dO\\'> Jormu,/ nf Wilt/I/ft /IIR'f1'1'$. I I I '.>O-IS6
u.a liU.U.S...\..\1.. Jg;8. Bitlng.mi(lgt.,~ t<:t>r.ttopo)tonlJac) aslntt.tnnediatl' 1.:5 01Ul'.':l a. 1;.t-. MRittM, f L... Pl;.\Jtq}_,,, r. t _ li.iU.\1T.R. \\ .L .. tilHU,., i,P.r .. 19a:,.
hosts. for Umm:oprori'UJ Qf duck:,._ Pt«l·~•1m!JJ of1/:t' frmh Jn.l&1nw1ioanl Orbi,ir.istl'3, 1mm Culicofdt•s u, Flttnda. Prag,t't$ :.o C:/lltl(<1I IJ(QliJFi"'I
C,,r:gr,.u fl{ F.ntpr,io/r,g,· I l!J.,61, l, !la!. H,•JMri:h 178. l!J.'i•:!00
l26 f..\W.S•.\.M., 196.:. On chcdcwtopmi..nt:1,d tranfflli~,fon or
l.1:ttmc:y:o:oon ;,;nd Hr.eme,vrottu~ Prot:t«lmgs oftlJ&..• FJr$1 h1t1•t1lmfonof
ttO <..Htt 1~r111. ii.< 1c BL.._M ..... Go.I i9-:- C:.ullroitf<.'1 .,, \ ,•cm11i;, or
lfmmu,s,,nridiu..\lt>1-1miln .\-..•u1.~• .l'i. 2&,.
0,11grt"l'S o.( PoraJilofop,y !Romttt, t. '.?39---:! ~O.
aa:; ,i. ~ tt.t.., ,1:1, tt.t-.. ,,uo. Sµurvg(•n~· uf l.t·uux,-co:ov,1 '1flJ
,ALU:, . /\ ..
11-; mu 1,1 ll, u ... 1v11J.tM1~ r , a \\ .,,.....,.ti. "1:1. ari·a, OmJthophUit Culi(uufcs
Htlfmop,ouus in si.mu1Ud\ ant! ccr-1topogon1dac .Jmi :1 h'\1 ""--d sµp. Dlptcmi: Cerutopc,J;omdtt~l frc,m :\:'!\\" Brun~'\\.1l'~. c..::innd.l . .snd
1:l11;~ific..1.tion r.>f th~ f-la~m1J..1pOridli~1a. r.a,wdim, Jrmrmtl ofll.HJIQS..'l', :t~. lrr,pllcDtiozi,. oftht-ir in\'Ol\'\'ntt-"1 In h~tmopro-!e!d wn.,mJ~!Qn
faur,m, of .\/t."tfi({1l L'nw11;0Iog;•. 1 t iOl-70 ;.
315-228.
(cr.uopol{imt-dn1.· th tntnm,.o.Jlatr ho,i-.
1:!8 f.AW>, A, ,1. &- tltS~-t'T. G 1- ... 19&1 M8 •~~l :,t 11.., .c ... 1-.\11m:. \ . A. . .t~ tt,1.t!S 1,..n ~ 19U Equln~ t:11/f1,·11itfn
(or HtU!mOprtJti!liS and other para_,}l(?. \ltil'(Uita .V~u·s. .?? . ! -28, 1.1,'J'<.,..•MilMl\' In Aotida: b!tlng midges colttc1~d ,n liJ;h11rup, n~M
h.,,,.._ \ftdu;(I/ mrd l',r,•,11111,v lill/m>111/1tl{\· 2, 129-IJ5
~ f:AWS. l\ ,M., J,\C.Otl:!O,. ILL. a. RA\'UOU.1.D. 1., .• 191'3. JJa.t-m.uto1oa in
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:1Jmm11/ Mtk,o/,/olos:11. F.pfd,m/all)/(II. I ln11111111oblt1/QI/II, 7{81, 24-28 1in .-. nouu,m, v.. 1985, O~rirnjnn nr H<'JµJlf1(J':$lis lct'inc,. nr\,·<1:11«it~, n
Ru,nlnnJ. para~!H: 01 f';t•ropus p()lifh"l!p):4/c,i from Qm.--cn..\l"and ,l\thtr.1H.1J••o\mmft.•.<
d,,• Pn.11u,·tolag(,• /lumumt\"l VJmpaM·. 00. 313-382_
r.. J\Of!TA'?... fa."
!-;"; .:,\11-.SlAb, LIi .. fLOCllf1', U.I,;., ~f•,\1.,\11:'\, J(A,,.~,.
k.P••
19;,7 F..>Stt!rn equine! ~ncerh11lornyt:lhi~ ,iru,: lsolatc:d from tb.u:c 'JH!c1rt- ,gG Lt ou, J,\\ HOCt L .,.1. a.1•:,ttu11:o. r.r•• 198'.•. Ecolag) of Urupuucht
of Dlp1era r,o,n Goorgi.. s,.,,,,,.,.
125, 39;;.39.;_ ,·1ru~ a Simbu grc.n.1porbQ:v1m!r> irom nnnhurn ~outhAmeru,."tl /u: I.ABUD.\.
M. & t:..,u.,uo. r..11 (NfS.j • .v,,w~.\SJ)r"('ll '" Hcotogy 11f .J,,rl)or'fru~~...,. ln~nttu.:
r.'8 ,-;..\\, H.lt .. 80RllUI\M, l•,f,f,., ()l'CJ, A,f.., It ~lA..''fllAST, ti.A.. 19;8, 8l(){)cj
of l irt>I<©·, /Jmtls/m-, 103-1 !J.
ftcdlng panc1n.<ofbitint1 mid~.. 1D1ptcr.c c,rawpngonidae, 01
Ko\\'4.tl)~nu,. C...lpl! York: P'"·n1muJu, northern Qucc:i~Lffld. Br1fh:ri11 of 197 ux. \ .n., 19;,. r~ohmt>n, )t \iru1t, f«im fidd 11npulallon~ a! Culicor.d/!$
l:ntomologltof /les,,nrch. I,. MS-149. i01p1cnu (:t,r•1optl\;011ida,•J m :-.IK•ri• /011ma/ of,\f,diml fmomol/J/(1',
16, fc,-;'9
,·.tt.. :-.iookt. n,1~, t-11C>N . n.a, C'\\J~n-, p.H. t. ~,un,•11)-. r •., ••
ft.lj >...t)t:P, G.J:... tfl·,
l97~ 1\01on).:m v1ru". ~ n,..,\ HJtahdo\iru> rl1'h)h.-d to ~tokol:, \"irw: or U-1,· ,a.a Llil,. \',H., t\\lSli't, O,l'f, (,l:.\flOf\L 0,1... ,,111. 8h11:tOn}:,"Ul' ;tlld rcl~1U!d vlru\~-
Rabies ,-;crogroup. ·tme!1(fm Juur,wl uf F.pidrr111ology, 93. 43-1g, ln lb•dan. :,.;,g,ria, httlotlon and prellmlnill') ,dcntific>tlon orrln,scs.
180 1.1-m.r.11.s.• 1$9-: .1/t/dlcol mrd V,1m,11>ry· EruumolOII')' L,1ndun .tnd
Am,:n<art/OUrtt(I/ oft't11•rlnorr R1.."S("(Ufit, :J5, 1105-t rna.
Sydn<~: Croom Hehn. rst, t, 1- ·.-.c:.• 011•.~f"J, (
"· & 1-uu. -: .u., l&nS, -Stu die, on !t.-ucoc·yto1otmo~b of

1th l.'ltAXCAU, c11.).1, 4i i..TTTI...E. n.~ .. ,cm. Tht.1C:1UrtmlN l,ntr<.•lllt" (l)ipccr.t:. .:hiclcsw. IV. a,~ ,•cctm u( L. am/Jrry1I 1n T:tl\,:u,. 1aiwmt /Qunu,I t,f
Vf'terlnary .\1ffhd11cm1d ·1nlmal 1lu,lm11d1~·. ? • .U-17.
~ratopngunldae, o(F.:ist Africa, 1~11n<a<1/o11r 0/1/r,• Hay11I
Et11omo/ogi<i1t .,,,rrrr1:of u,11du11. 123, 1-9:\. 200~Ln· , ......, 19:9. ~utrien; ag_ar mb.mtt."!i furcuJmre of fo<Jd tUJ.anhm,

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132 S1:et10, o\t; !\Spccts lllflucndng tJ1c occurrence or infec1lous dlse~scs

forpreda.:4ou, Culleortla ian,ae ll.l1p1cr3· Ctr~1opogonldatl Jo:,mal qf 1•ro1~<1lon of horr.c< from C:ullcmd,s lH>littm;s 10lp101;,:
,\!«(icoJEt1LO"fU(){~•. 16. 171-172. Cern1op0gnn,dnN, a r,,cerutr rdemificd, ll<'IOr or Aldan horw <iclmcs.<
20, ,.n.. 1~5. Wtin-gmidgc, <Oipttta:Cct.11upogonld,·w) .1,~\cctaf:'I; of
UNUl\',
81tlltti1111flitt1M1&IOf.iMI Rq;onrc/~ 90, SOS-SI 5.
nomirnl animal pa1hogcn,. Jorim<>I qJM,dlcnl Fmomn/r,gi,. :?2. 588-59~. z.t.t ,n l!,\\·1~)1;..( 1 1 H,. tl,U )"
1 ,,. 1,.,AhVSClli\,.X£, i,.- _.. MEU.Q.A-. PS•• 1999.

20-: U'.':L!.Y, l,M.£.-D,\\'ll .. , J .n,. 1971. S:mdOh.'i- .1nd inurhm ln f)orid..1 und thl' IJnpuhl.~h,-d d:110
B.1hamn, :md C:.inbbcM.n :m:a~Jot,mt1/ b) l;",•,momic lbuonmlol:)•, 64. n, Mti~·wt~KM ~,. HH"-\c i,,.:. t.t ,,. 1u~4 African hoG,-,.,kknes,
2&1-27ff. cpid,mlology- five >pecil!S oi C11/lroltl11J tOlptern: Ccr.uopogonldae1
103 u>O~us. E..c... Hus:..q:1r..ru.1Ln.•osnuNs. •·•· 1985., f:pidemlolog_ic ~rudtt-,. coll~tcd the behind 1hc<al')ond ,11 the dung of the Afri<otH•ltpMOt In
of blueu:m~ur.:,inbon lhC!Tefn rnnch. C..iHfomi3! \'("Cmr ho<L i.,iru). 1ho l,rngor 'lallonal 1'01~. South Alnca. D11tlm1,poorr Joumal oJ
:cbt!ciru,,h;p .... J,r. U,\ftl}(k, t.L /; IOOUM. 'L ,, OOsJ Blt1d/(Uf},,'"t' 111111 \'N,~ri110I)· H.tSMrr.it. 6t. tSl-tiO
Rt:{ntt!il OriJJ,.,Jru.u~. Xe\\' York. ,\!w, R Lb.-... »it \111!1>\~1'-lr:1'.I.. A. & o,·ct. A.L. 1qS,9. AfrolropicaJ Cutlro;'drr. Sy11l1t1lri1
:0-4 LOW,U(. R,C,, lUl'.IU( \,ti),~., ~ O)UUt'l., l.C l$'8. f)( \'clopinent of
1 lrJclr<·r, 1925. ro~urrc:ctcd a.. >uhg~nu~ tucmbr~,c 1osr1..~1..'"1. (l)IJHena;
Tc1rapmolonN>1t1 m1Jimt)stU1~ 10 t-hc inrecb\•< stage in Cul!t"Oid,!t
1 Ccratopt,grmid;w,. 01ulc>1.ttt-JHKJn Jm,rmll oj\'4.•trrl11ary R11rarrh 56.
lwllm.tis and c Jur,ni. Touma/ of Par,uitolaw, M. 100.l-lOO,. 147-163.

20S LO\\'n.lL 1\,C , OJlllltL T.C. & f.hLTIIIAM, ,,. t- ,,a:. Cull,CJld(~ l'(lti,'p~nni.~. n ~ ,u, ...w1,1a1.. k., :-..n•u 1. t,.,, & \ 1 ,11-1t. t..J .• 1,94 Vectors~ 01/fcmilrs ,:pp.
laborntOf)' ,-ec1or for the Ama;,;on form o~ ,\~an.f<mvlln o::arrr/1• .~m1--riCflll' In: r:n1·11..PM. t ,!'\.\\ nm,,!10~. u_n,., 1usn~. 1t,., 1.'Cls . fnf,nfous
Joumal ojTtoJJtc"I ,\ft.vlrrmr-,m,t H.Q:u·u,. 31. 166-167 Dr~,se, of L,w,111tk u•,1/r S11tY/al Heft•,...,,- 111!i<Jmlr,r,1 Ajrlro. C..apt;
2o6 L0\\1uti. K.C. tnfr.~.,,,. t hlPIL\kD~. ,u.. It >!Art.. )..t'., 1~8..,1
M(J
Town: 0.ford UnWCMt)" Pr~~ ~nuhttm Mri,:.r_pp, 68-$9
Ass~~nt~nl or /,!pt«ollDP• l""7tm.'tl1 a, i po:cmlo.J vector n( Jfniuo11,•//11 ~6 MEJ...,.,,,,aa.. rt.&l'AWU..:.\. r.1 .. 1998. The., 1998ou1brc;1}r.:ofhnrsc ,id.nh.~
=irrllln Halu.Amrrlcan /011r11<1lofTrorllrnl .\frvll<ln,1111d H;'!,</Mr. :12, in South Africa: \ no" C,1//cnitwsl.Jur~III• tC<'r.11opo1,'0nld~,1 <<.'Ct<>r~
)013-1015. Ab11,,,.-t Fu111rl, trr1mu,r111nnl <:Onpes. ofDrp:rrology. i:,bt,Colley,
20-; '•L\Ct-al. J,w..s .. 1 ~ OnM>me .Egyptla.11 UrJropogon.tda:t. Bu/lnin of Ox/o,d, Ul.'..6-13 ~plfmb,,r 1\lllll. pp. l45-14fi
E111omo/og/<t1I &si:11rd1. 15. 6Hl7 n7 ,su!.\,'Ji\nt, ft. &-P.\W~K.\. J.T.. l999. Thct 19'-.}8 outbreak oi Afnra.n ho1~t
aos ,w:m. r.w.s.. 19,<;. C.,ra10pogon1d~~ (rom ,he i\nglo-l~·ypoon ~<:di}ll ,-1d,ne<•. in th1.• '-':\<tcm ftct' St.Ht'. South Africa· ~c\,· jnsfg.hts into thl
Pro«M/11g; 0/1/12 llo}tll lincum11/n1,~tol Socwry ofwndmi ill;, I~. 1•2. 11pidC'm.nlogy nr the di-.t,1-\,~ /.'rnct."4.·1litfJ;J r>f'fl:.. ,\'1n1ti tnu1r11m{oJl/d
S)mJ>UjJ11m of r/f~ i•.:o,ld...wotitm"u of lr~tc.'fitta~' !Ahommry
2:09 MA<!Ht,. J.\\',~. ~t~(.ki\.,\f, , •• I ~ . Cl'tt.\Ul n~i:rle;,. ofin..«<t uu.~m \\c,.t
D11.1J;no.,tirlnnta11d OIF. 8/or«l1110/t,;:>• S.•nrln<1r. Collugt> $101lon. kw,
Afr!Cll. Bull~t/11 ofEntomological ~,.,nuI1, 13, ?~1-2$4,
USA Pr l.tt
210 """"· ,., .... M-\wn·, c.o. 1996. An cpldemtol<ll:fc an•l>~•~of 1hc icn ;.t:J.IJ >1o~wi-:.UL n. &-\·E..'"(ftft. u., .. H,su. 01,dt:~cpoort\'t'lt•rin~ry Jns1itut~.
pln.;ue<o(Et,")'PI, C/Mm,. 12. 7-2~.
Onders,cpuorr, 01 ID '<uu,h Afrlrn. t Jr11mbli!h•d datrt.
2n ,to~...:TosH. K•.,1.. 1981">. The-cpfclctniolnm, Mr1rthropc,d-burn~ \inb~, in
m )tlLLOR. P.~, l!Hl· Srnttlt-~ of 011t'/Jnt't!rr~ i'l!rtrittiliJ Rullllct~L I lcnry 1910.
~ut.hcrn Africn. o.s,. Tht-st,. Unhc~!ryuf Prewrla. I. Oncll«,:ft:t> rc.•ril{call$. lan.'Ut! in 6tifr1,·h hnf':Seli Journal Q/
2-~ Mlll\\'{l~l\l:.l a 198;". Affotrop1enl t,,llcold,~r a: redesc-.rlptton oft" IMrntn:l111/t11:)·. 17. 97·-110.
l.,kor,t,(1} Karmgm ~hama.la & ~t:nh.· 1971, rcan..-d frc,m dt:phan1 dung 1n
230 "' "' ""· r.s.• 1974 ~)udies ur(J11c/1<1<1•rtr1 ,wutrolls Radllc1 c1 Henry 1910.
ihc l<!Ugcr N•tionul l'ilrk. 51>111.h Alric• o,ut,mclJ(>Pfl /orm:nl of
I\. Bthavtour c>rtht' \'~tor CulinHde.r 1mb1..•r:ulo.ms in rdwion to lhe
Vc1crmory R~rtl,, S4, Sa5-S!W
:r.insmii,sion of Onrfutci:l't'a cctnfitlllis. Jt>umal a[/11.•bnintl,olog>', -18.
21.1 >11U•w11<. .L K.. 1989. Afro1roplcul Ct,1/rt>ule.: • r<'<lc:.<:rlplion .,f C.: 28:-S.28·1,
(Al!m,t/11/ 1111/c~/111'tclT<•r, 1913 (l)IJncra: f.cr.r1op<)gunu.lMJ\\ill,
~~• Ml'.U ott. l'.ti..., 1-9/S, Stud,I;', (.)( (),1<11,)(:('r·ca C(•n•i,·"lls Hailliet c,
th?nry l910.
d('...cr!p1lon of the clo-.ly :,llicd r (rl, l,n/itwo, ,p. n1>v. rearod from 1hc,
dJmg ot •ho .lfncan bu!l'oJQ, bll,e \\ildcbco" and conic In So111h Mrica
v Tht ccv~lopme,nr nf Onrhocr,rr, rr11 le11/ts larv:ie :n rhe \'t.'(lots.
1

/Ot,'1:tt/ t>J JMmintllolol:)•, 49, 33-12.

0,ul{,,sr,·rJ()()rt /01,r,u,/ of\'ctrctr;muy R,i:u·:11,:h. .56. ll--39.
~u \U!b\\1~>.1-L 11. 1~51. Afnnrupical c,,Uto/:lt.'.S. C..
(A1"fltltluJ mlumbo)p. :t3"2- Mf·LJ.on. , ••~ .• 19;G. lnftt1lon of Cutlcr.)fdes m(b«i1/QSt«$, Cullcol(lci ri~rht
a.nd,kdt.f ,,e,opti \\1th ,\trmtautlla o:::z.mdl. TranM1a,rms pfrltt: R(Jyal
nov.. • \\1de,;,re•d ,pccie< clo\-01)' allied 10 C 1/\.J lm/rola ~101Ycr 1~13
S<,crei,• o/Tmpirttl Mr·,llti11n<mtl //ygw,ru, ,0. ;153
<Oip,ern: 1:1:mropogonld~c), Oml,'11ti1JJll!m Jmm111/ 1J/Vvter/1mrr
/le.$\'(ttf/1, 58, 155-1 iO. at \!lU,()R. 1• , . l~)St,n. , .• H<\.\JRU:-0, ('." & OKA.HAM.!>,., 199q, b4>kltion~of

2.15 ,,a~w,,"t-l.L x.. :.~:t~.,fro1ruph:Jl Crdka!t/(':J! C c,w,.,,11/111/u.xo,lou,l.s ~P~ \ftt~n hot<<' "lrL'. nh,,. \.in,r.. (mm,,.~~,, lnt«11. nuu1t- rturiug thf' 1988
nQ\'•• • new membero! inc lmlcola group 1Dlp1oro: Ccro<ol"'l,'<>n\dnol cp1,o<>1.cir1 ~114!n. J:prd,•m/olug,·n,11/ /11/e-c1io11. IOS, Hl'-154.
a.o,!'i()(.fatt."'d \\ilh th(" Mrican «•lc.ph.tmt in ti\&.! l:rug(o:r 1'\,ttmnJI
~1~td.:, "\t.luth 23.1 ,nuo1t. r,s. ,.. Mt:c.\11;. J. 191.- Tht 1>robablc t<lU4-C of .,,,·ctr heh' rn
Africa. (),Wr~Upnt,tt /t>umal o/l'et,•rmtU)' Hcs.rArth, 59. 14f>-159 lln~lar1~ T/1,• \tcrm11t11)' R,-<ord, 95.411-115.
2:16 'lOSWl:o..'l.,.1.. ,t., J99.3. Ondersu:poort \'et.,•rinun· lns1itut<', Ondt't).tcpoon. 235 ~ff.U.Olt. r,.s... osaon..._L. R, & JL...-x1 ...c..:::i... o.M .. 198.;. l~olatlon ofb1uetonguc
OI IO SOurh Afrfr:L Unpublishod dO!a. ond n•l.1~d ,iruo;cs rmm C1d/(r)i1/e;,pp. In Sudan Jou11111/ of flJ'!;l,•n,,
.2r; ,u:1bw1~Kll11 pc., tffi. Afnmop1r;..I rulko;,I,·>-: tm.>,.;':'lcmat1c~ 01 the: C:uml,ri.-lt,.-v. 93. f;2J-628..
lmi<t.1fagroup, ~ubg,nu~A1'Qrim1 fl>lpi<'ra: Cera1opngon1dao1, \\i;h 236 ,,u1.0,t ,,.,.. & 1•n1.ou,. c.. 19:-9 Observations on brc{ldlng sil...s ilnd
special reference 10 tlle cprdcmiolot:) ofAfric:an horscsl<l<nh>- M.Sc. li8hi-trap toll<'<'llon, ofCri/lrord,., during"" ouibtNk ofblu,~ongu• In
!J\gri<.I The<ts. Uni\~r<il)•Of PrclOno. Prt1ori11. RSA. Cypru .. B11ll,1/nn/ F11rom1>!1Ji;i<11l 1/mvm·l1. 119. 2:!9-234.
ai8 \IFJM·•i~,::r c ~-, 199; Oi$CO\'t'J') of n C:'ull,out,., Jmiroln•!ree T.one fn ,23; ~11urr. P. 1.- ftu\:, o .. a,84 Unto nnuvellc-tiln.rca de l.t pl~. 1:'tlfilnru: J.:.all{ai
!>ou1h /\fric:1: prelimlnaey 1101c, and pcu<nu•I signlficon<c. n. ~;,. tm1dunln.ic) ct ,on dC,l•loppc.m\rtU che:t Cullroldrs mJbt.oeulostts.
Ond,·rs,~JPl;or1 /ounrul u/Veu:rm(lly R1.,.,r1rd1. ~. 81....SG. A.>uud(fi(ft J'(,ra.sltolog1t hunumur l'I CtlfflfJllrte~ 59, 177-178.
ii':! ,iu<, tlSJ."l..t
1
• 19,gS. The 1996 outbreak of :\fri<n.n hOr14- ,,r.t..nr:>s ii'\ 338 ):ZL Wt.'i', F. , &...\..~l).\U. : .• 1982 Culir:oiddS nuba·ukJsus. ~n cxp<'tim.c:ntal
SouthAfrf~- ihc c111amologi,al pcr,p<o:tlw . .-\lrh/,--, uf1 ·,r9!og;· ,·rccor cJ 0: new 1rypano.somt froJn a psim1dform I Psittacula ro$'Qfltai·
(~upplcrnm1 I-I , ~ - Tf)'/JIUUhmull IJ11.J.:;.•rf. new S.pl-Cit~ilunafes d ..• J>ara.titotogie lwmtnn~e:
.:.20 ~1msw1:r--"U'l... n, '- 9.-\\'US, ·"·· t998,. Morpho!ojtkal conflrm';\ti<Jn uith4.! ,omtxJft'(\ 57, t2J...;28~
sc;,:imtc sprci.,. <'tOlU< af (;u/u:o/d,. /,kuriliJrJ uudifld/111.< t>elfin,1do. ~9 .\tlLC~. f,. UNOAU. I. C;.A,..:,.rsc;;, r_u.• K()t)k\t,\,,, f. ""KM.l!.\ O,tt ..., .. ,g.76,
196l and C 1,1,, ,mtcoltr Klcffc,. 1913 •.t>iplff.L C.crraiopogonidati l/o1pn1rxi~ri.f dl· Mo.Jol<e. Ill. OcwlOJ>pemcn, d Flq111tl>C)'lt# ftr~1 chez
()11//rr,r,{1(><H1 /,:,rmra/ •I Vet,.•rlnorJ 1/,s«irc/1. 6.o. 9-16 c,,llrold~s m,l,.•r.ul&u.t ct C 1.•11tiipt·m11J.. Am:a/e.$ ,It.• PdfllJtJO"logi~
221 '.\JdS,"\\'b'-W... R M,'IJ!t. M. .S. LADU~CL\C~L i.... :tt)()O, StabUnt: and ch~ lmmnl.i-.t,nl ttmlJH~rn:. 51 2~302..

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 \ l'Clors: Clllico/de$ spp. 133

uo ~ULTG~. F... u.s1uu. I., RA1A.'\:o\WO-RABH..\~. :,. •• • YE,,'JIUTR.\, s .. lg8l. 2SS :-.1:-LSO-s. R.L ASCRJ\',\!\t, 11.r.19':'2. 1sot~lion~o1 :a.rbO\iruS('-S: fro~ parou_..
P1:1nlut1Jmo1)rDl1mJ rk.Jf.t.•rl n .. ~11,: g:amcttJ~onlc, t-t schl1.0J:?Onie ch~ mldg.e(.of th(' C,1llr"ftf1."'S 1·11riiµ&••rn/j'"totnplc:x. :1nd parous. rntt.>1 in bitini
l'hote n&turnl. l'si11(lt:t1!n m1n.11a de Th.tl!3ndt M)'J)Orogome population,, /o:,mnl of .\.Je.'licnl [11wmolo~· 9. 2;-:--2:81 .
r\.pcrime.nu1.k.- <"hez Cullcoldt.'S nubeeul0Ju1. lv1nalt'Sdt> Pa.rasf.:o!ogfe 2S9 .sr:-10\ 0
.\. :,;,\'. Dt.lL\'C.JI(\ • \ r •• C';0k1lt:1;\'.\. ?.,.£.. XO'STY\fKO\", :.l..A. 1989,
ltw,uum, e1 t'f>mr>uret. 56, 123--J3().
1,olanon ortssyk-Kul ,iru. Crom b!ood,sucl.!na bitins fii<> C:11/,ro1dcs
:Jt ~UUMt\, \., r.oro. v.• »:uuo. ,, . .s.. ioso. 'Ii .. 1988. holahon c,fChu:t.Am \1rus. a sclwl,:.ci F.nct:rldn In southtn1 Tad~hikts\411 in Russrtn1 • •\lttlic/11}kayt1
rtt:\, tn~mbcr u!1hc P"Jlyam i.ul>~n,UJJ urthc);cnu\Otbh·l1ux. lrum canlt: l'um.:JtultJ~)« i Pilt1J..UIW fQIC s,,1~.:.1ll. 6. 2..\-2~.
.ind Culltoitlts nx_\'$I011ur m Ju.pun, .~,wJrium /our,mf <>{Vl'wrim,ry
.!6o '"'"" t- \l, 1967. Blologkal >tudi.-. on some i.oulh .",mean Cuilco:d,-.,
Rtt:ri'tlrCh, t9. 2022..'2025.
,pccle, !Dlprera, c:crntul)ol!on,d~,·; ond the morphologyofthd,
242. ~,o. C,S... THO)!PSOX, WI., UO\t,\S, ),. 0\'1\i.OQ, \~ 'r.. C.AAJSf~. e.c.. immurure $ta~.._-,., M.S('. 11\j,t,n~.J 111esis. University •lr Prt:ton-a.
GOSt.\t:f.~. J. & SAC~/~ \I k., 199,1 , Uhli'.'tOnJ,l'Ue ,iru~ i~lotion-s from vecton.
261 ' " u.1. r " ,~fiB, A ,1,;nlflt,,nt new btc'Cdlng .Jt• or Cullcofdt.$
s.nd rumiJmms In CentralAnt.e11ca und the Corubbean.A,m•rfcm: /Nfllltl1p,•1111i, (:orrcr. tngr,111 nnJ M,tc11e (Oiptcric c..r.m,p<>80nfdaol.
Journal of Vc•tt'tttU11)' Resetlrch. 65. 211-21 S. /t1111u11I of1hr Smith /\frtran V,•t,,fluury• M.:dl,al M'IIXJmcon. J9, 6l
243 MUHA:\1Mti.:o. M,1!..11 .. )ll!U.C.)A:, 1•.s., 1SCtt1, Funhcr smdir.s on b1umon~ue :9ff9. ihc murpholo~ ot the inun~ture stagi:,: of some
262: :\'L\ tLL f.• :\f.,
and bluc<011guc-rclu1ed orbMruo<:s In the Sudan. li11ia,mlo/ogyo11d South MMe•• C111t,:o/llvH;"'c10, Diptc-nt: c:,•mtopogonld•oJ.
/11/«tion, 105. 6111-63?. QrulrNl~poo•t /1J1mm/ of\ 't•Jt•rilUll)' ll,•Uiln'lt, ,l&. 265-2'3';
24.: Mo,t-sou,:. ,.a.... 1,s,. Cullcotdr.s m,,,t't't1lo.tu.& ;.sc1g,1 <Oipu!ra; :163 xt, ,u. , ,, .. 19;,. Cattle ond Culltt>fde, biting !nldgh as po,,U,!o
Cer.uopogon,dne,. po~lble intenncd,3tt ho-. 01 the filnna 011chll<•ri-a o\tt\\'in1crin1; ho,of"' ,;( blur-tongm.. "inas. 0,1dttnutpoor1 Joum1.d of
r2ticulato Ot('.$. in the Ca1muguc. Q,mpres R1ndu,5 •.:irodemii! d,.-r.~ :kie,u:es \ '91'1ri11a,:.• RfflVll(/t, 1A, C~r2.
(Parls/.233. 10?-IOJ.
~<i.i ;,,.:t\11.J... 1.. ,: t978. nw u.-s-c of t~ttlc ;o pro.rce1 .shC\.-.p: from blue1on~c
2'!S )tOJG:>'OIJX. 1,11.. ,951 The Onch1>c•rca of equlchi<'. ,Itta Trop/c(I, 9. infNtitm. JournnJ of11,~ SOU/I: Afriaw \ t!tt'rftJt:f)' A..(J.U('Jn:frm. 49.
125-ISO. !2!1--1:ffl.
246 ~10R11, Y. 11 KrTAOKi'\, , .. 1968. lnOucncc ort~pcrarurc on 1ht -<;,orogon~' 26S ~nu I l \1 1...\StUJl!)QS, 'DOA,\.1972- tfO)l p~fc-rC."'lCCOi Cutfroul~,
ofAkllJ<, ca111/q1y/ In llu~• Cu/iroldts <pccie,,. \"m/onnl t,11ri1111~ of mJtf~ Oipu:ra: CcrJtopogonufoe-, in SoUlh ,\fora a~-dc-l~rmtned by
.4mm/ll H.alrh Qunrr.,rl;- t ro,,.,,.
R, 21 B-21~. precipuin t\"}1 .and li~.u tmp catches. 01ul11rstttH)Ot1 Jrmmnl of
u-; .\!ORJI. T.& ,a.-r,\O>:A.a.., 1968. Rclatlomhip b+!C\\'l'l.'n the.cour}leof \ ·e:.:rt.,u,,,,R<JMrt·Tt. 39. l-l"i-15?.
or
gamctOC)1e appearnncc llkl/l(t rm,1/,•ry·i ln the <hkkeo and spoiotnitc 266 ~C\ tu. L\f, 1..RA.\\1u-... B,J •vt.,n.R. (;.r .. 1ruµ,. \ "iix-ye-ar sun'e}' of
production In Cullro/d,s arnknuwi\ Vt11fo,u,J h11tir,11~ufA11/m11/ 1/1\1/rll ,irw;-cs 3.""0Cl'llcd w'nh Cull«>ulrs b\oog midges throughou1 ~uth
Q11t111crl>• /To()-o). 9. 20,1-209 Arnc::a Drp1(r.1: QrJ1opogonidnN In: ,,·.u.m~.-r.r~• o-ssu11s. n,s•• r<-ds::.
2..:8 ,10AJ1, r .. kUAOt.:A,., & J\,:W, .:." 194.i5. So1\a- fnvestigntionson the Blu11wn,:m•. AfrltJm UUNt•M,kn,"S.~ mu! Relim.ul OrhfllinJ.S(!S.. Ooc:a Ramn
<poro1,..,ny or1.,,u<"0<)·to:,,OJ1 coullfJ)'i In !abommry-«ar.-<l bi ring_ mldg,,s flot'ltto, CRC Pre<,, 1>1>. 31 ·1-319,
of four Culit.'(,:'t!M j.podie.s. Nmitmul /n.)·mut,.c/Antuu,/ I-IMllh Quat1trl:, :.th7 '-l \"11 J f ,, •,•1,,1nt, n.J. I O'... •UUu,,. ,,. ,,AIOR. 11.P.. 'tffl~\\-.,. ......J:"Jll, JI ~

m,i,"'J· s. 109-1 to. ""~ G-\S. 1.u., 198,$. Culicoitftssp,cc1t.!Sa~.soei,1tc:d \\ith fa·esmc:kin the
U9 \.fOHII. r .. \li\1'$UJ. T.. Ull:'>IA. T., fUIIXAGi,\, t .. tgS.i~ Jntectivi[}· of S1.d1enhuseh un:a nr tht· wC)i1em Cape- Pro,1ncc. R.epublic: orSnmh
Um'OC)'tot.0011 coul/uryf ~poro1.oitl'!'> devtlloped £11 J'itro and In ttlt'iJ. ,\fur• Dlptcni, C',er,nop1>1,omdoc) Ond,•mf1><l0rr /01,mnl o/Vttermol')•
/11temnrlo11nl}o11r11nl of />t1f/1$lrQl011J•. 14, 13!.-HO. Rr.<l'J1Nlt. 55. IOI 106.
250 '\101UI. T.. :,.'~J(A..'\n,Jfl,\, ,; . U'l.. ,.c 1111)'!.\, T. 4 11GJI, ):,. l986. Obsm>a.tions. :z68 "'tYIU u. 19':l!">- Ond~htepc,.:in \'e~.:nn~I")' ln.sdtute. Onde.,tepoon, 0110
on tltt' 'fo.iwanesestru:in n{ l,.•u,·QCYtomon cauJIU)·i Hfaemosportnn in l>ou,h .\fnco. Un1wbl,-hrd dota.
chi(kcns. 10,mwl ofProro:oology. 33. 23 t-:?3.. •69 M\'IU. u... mn., t., 1%3 AfrOtrtl)>lr.tl {.'ulk1Jl<IN: Oe-,crlpllon and
2.51 ,,uLU:'.'\:,.. ~ - ~~urz, o.A., !98,1 Ag~s1rm·,w<and survh"Orship or cumporb!on o( the pupae of ,c,·~n ,p«,e, or the ~lmlh,..,t>ctgroup
Cul,roi/1"1 r'tttili><•1111:< , Olpt•ni: Cer.i.topogonhfo•• '"ccnrml 'lit-,< tori. 10iptt:ra; Ccrllto,>0gunidth.~J~ Onda,repoort f,;mmal o_f \/e.urml11)•
Sme, USA. /ournol of.\fediC11/ F.,1u,molog:,-. .21. 1*-203. Rio,'llrrh. 61.s.;...100.

~2. .'-tOU.0:S, 0..\,, \TI.Tl:.._, ,u. ... <,Htm·, A,C.. 8M\'Ol)t.\;-.", 'I, Ir LNUJW,, K.C.., 210 ~tcJ1ou.~. , .. ..i 195,} The b1onornl.cso( Cull(oJdt'I nwwnt, vt'C'tor of
>OOQ. Feeding und sunivoJ of C11/lrotdl!J! s011om1sis on cattle ireau,d Atamh«llt11lan~m11 pt-rtttuH in the m1n-forc\.t ofLht British C:smeroon,.
Wtth p1."m1,cthrin or p1rtmiphos methyl. ,\Jt"<flcal and Ve1~rlrU1')' :md nlhf't .. p,'t'tl'~ whit-h ffli\)' hr' \ ·1'('1ttr,; 10 1h1• "'m,.;.u,.., ~mMl.1 nf
F.momology. M. 31l-320, Trnpi1Y1/ .\tt!f!irfmumrl P1t1AAfot()g,t·• .Ji. 1a;-206.

~ ,1u1,JJ 11, M,J. & sr,,.sora.,s1. Jt,A., 1')66, Vectors of ephemeral fe·re:r group 271 ,SICJtOL\"i, W,1.... J:1!.RMIAW. W ,L, 19is.; SlUdlt"$01l the int.akcn!
,iruSC). rn: sr,clP.Onc.c. T,o•• ~\Y. n.11. "m.oi,.. J,. ted,, ), Arlxu,fr;,s RiSicYl.rttl m0<10fi!arfao b1• 1l"•lr w,~un, 1hcirsnni,,.,J, and th<ir elrect 011 th~
m Ausuati(l Pr«a..'tf11l8$ F<ntrtlr Symf)()smm. '>-9' Mny 198G. Ori!ib.me. ,u1'iv.tl of their \'L"Clor~ Ill The lntJkeonlw mkrafi!Jlri~C of
pp. ~S!;,-2~1.8. t-l,wmluJ.4·111,flr,,wmn p.irstnn by '11/itqldttKmu1~n/ and CuUtol:lt.f
gmlumrii..-\111111/,, t,J'fmpkill .ltrlflrlnemlll P,1m,it~lag,·. ~B. 20l-20;.
J.54 \ilJ!il:11, t.z .. 1:,.,\. J .1'.\"\',, C.ltAl~Q, lt,O .. tllZ\""T,\. M.G. S..U:.L\, 'R.M, $- lo,,;\PMT).,
rw .• l!l-98. 0,//coidrs n,soc,ated wllh da,ry co·,,-, 01 Sebele, Gaboront. 272 ML-<tn\11.L. u.1,, uv tor I K.,., a~~r ln'-'~dgadr,n... Imo ;he tnnsmh~ton
80C$\-:ana. tr<11uc"J /\mmal Jlt"alilr nnrl P1mluc1m11. 30, 305-JOi. ortro~;rckru,,,., Ondcmcpoon during the se~<1111 1932-193:l.
Omft•ntcpo9"I Jm,mal t>f V,•r,,n,mr,· S<'lNrei•mrd Aufn:al Jndustryi 8,
!.5,5 \lU~VJv\, <t:-i., ,1u>·,,·:~"-D.1t.. u.wu-,;. ,, t,:tu:r. f",f. k MLu.,uit, P•.s•• aoo1.
213-269
Pwsden« of Cr,/1coides 1mla1/11 nnd other SJl"de> IO!ptcro..
Ccratopogonufae} at dght site, In 7.lmbobwc./oumn/ 0/1/wSoutl: 2il Ut~U ..Jl.\\ ,.(. A. wo.-..1 \'_ G. P•• ._&!_, 19fi0. 1'U/(lt4!mtn ut Rus~uu.1).
nfrium Vertritui')·Ass«·umou, i2~62-63. Ml'dga. Mo,Jc,.·a, ,159 pp.

~~ :;.~UM, R.,. MAbUt~tu. \., YML\S,HrTA. X ._ \",\.\1\,\10TO, T .• 19-sG. ~74 Ukl u.,. ,1.u. \flJ I UM.··~ Ki\WU't.>. ••. • l'kU, M , •• l!:J~8. The ~c-a.$0n0.:.
Studl~i on 'Ka;~n orho"Q In Holoc.udo. Ill. Annr<h on th• octu:tl and ~eoitap)uo tlhtributinn of Cttliroitlu., 11111.:0(11 C. 1111/ia,rl,itn>up
,m, of the dlocase In Jopane'-\!I. /11p/111o1H /o:tmnl ofWtenflnr;· And C. obsol,1111 group biting m,dg<"- In eomrol and southtrn Spt,.ln.
.~ult1,,,s <1/ l1r<,/oi:,•tSuppl•ment HI. 8.'i-91.
Resear,it, 4. 81~.
>.,".' ~,ntA~. M.o .. 1$1 Tr.snsm!is!on ol the huinon rilarlnl par:i.he o,s ,,ion, 11.1•, :98J·t9tl5 Ondors.tep<>on lc10nnaryln~nrutt.
.1rn,1so11,lla fl=rdit)y C11/lc'Oi1/et plt"'1,01onllu .Dlpu,ra: Ondc1'tCpoon. 0110 south Altic;,. Unpub!"hed dot.o.
Ccr.nopoi,:on!doijJ In «>11>1:tl n<1tth Trinidad. Bul/et/11 of E11mmologknl 276 v,,1<:m.1.T.r.. -8ft7, Acull::.pstble. ~m1•nutomntk <ent•(}-pe..em,rgence
Re,,(l.n:/1, 71.97-106. rrap. !uiHtb!..- for ~mphng Cul,coldf.S rn11m:n1: C4.•ratopo~onu:!a~) from a

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134 "':n,,- o,.,. .A,~p~cts inllumcing the occum,n~~ of infcctiOu$ di<ease.

v,ide mngc ofoabit•t, 011d,~</JOOfl Juuttrnl uf Vtrctl11arv RMeattll. S-l. !1-95 SIRt.E\'. U)." \\'l,l«XrA. 1J,:•• t98J. Su'M"tptibility of pi!kfng (;bUIS
9!)-101. pla1yrlzp1rlion and mw,.CO\'\· tCnirluu mo,relmtn'. duds(n
27":' t1 A8SONS.O:-.\ 1,M. IC' ~ ...owuo-... w.A.. !98S, ffluero~e. tpi1.ootlc
/ftremoprowus m'tllonis. Juurnul oflA"/ldl/fe JJ/J,-rul.'>. 20. 108-133
hnemorrh:lgictll(ta...,~ or d~e.r nnd rtln,~ \'ln1Ses'# current..sitc.:itlon Iii 2·96 .S.."1,"0BTSO\'.\, l"•.\1.., Olttl~L\~l'\<$KII. v.1 ... .SJOO.tl\'.\, C:..i\... S::11\TTO~S'S..\\',-\,
.•\U~tm.li.:\. ht: tu,.f11'tm. 'r.t.. ~ JOCJrtM. "·"·· icdj}. RlmJton.gun nr:tl R,•lmrd :\.0 .. AJUS'rt>\'A, \·,A•• S.C)\:IJkASHJ~A. I\.".,.., POU".\J.:O\'.\, ,\...~.. l-rJMS)OV, SH,~•I •
Orbivlrum. ~i.!W \·o~ Alan .R Ll*'· pp. 2r-35. tst1.t>uscw.1..,~. 1,-~uacm:~~o. 1·•• 1~z. R~ul" ofa \.'fro!ogicsl sur.t!)· of
th~ anhtupod \ l.~lu~ u1l lite tc, 1ilU•> vf1'u: Wuenia, H,olog/(l \ 1ruJo,
278 PATON. 1 •• 1863- Thc"horscsld,,,css· oidu:CopcofCood Hope:.
/.J.IOS£'(U(I), 192. J:i~-14.i,
t ',,rqrfn,1nru1. JU. ,i89"""494.
297 ,:.·t., ,QPMT, H,~. ti ~YCE. u., 198z, lsolatJ0n ot Thlmfri virus from
279 J,T., 2001. Ondcl"'>tcpoon Vett:rinary fns.titut~j Ohdcn.iepoon.
PA\VtiSX."',
C11/ko11!,s h1$tr<0 (t)ipte,.. ~rotopogonlda• coll,cted in 11011hom
0110 South \!n<a. Unpubh,hed d:lta,
Aufilmii.J. Jozmml a/.\frtlic"i E111ouwl11gJ•, 19, 2.12.
28o PAWfl..Mt.\, J, 1., OHtDf-~. <;.H.'"' J.1Eb\\1~J.U... ~ .. 1999. QndC"NtCpoO.rt
J9,8 >1:\ ,Oti,',-,1, H',i\., l>\-U., .\.L ~, (.f,()UC.f, T.f?- MlJ1.UR.- \q., 110U~KT't. ft.1-
Ve:"1111"'} lmtituto. Onder,tepoon. Ol to Sourh ,\ftica. Unpubll<hed
d•t•.
t:H. \JU.Ii\. 1.n. *nur,1cu. c .. 1984. lsolo1io11 of :ubo\'irwu:s from ln""l.'£.b
collecte~ at ll<•a,rtce HIil. :'iarthem l'emtory of .¼>tralia, 1~7-1-IS76.
281 1•Aw1;.<.,.. p. "vrsn,. G.1 •• 1999. Tht ~il'Ect of ttmpcrature .,Id ,l111tral(ttn Jo11r1111/ of8iologr(t1/ S<:/<11c.,. 37. 3:11-Jij6.
in~ubution p,:riod on rhc replication ofl,lt:;ionguc ,'fruo in C:11/fcoid~, 29~ ~C.\'\t)fMH, H,A.. \,OW.JI. "-1, 4,-Wll.S(')\., U.t>.• 198;\, \lorutlif}' nfCt,llt:oidVJ
imioo/o and C. bolltim,s iOiptcra: CcratopngMJdn,•). Proc,11vlmg,<o/1lt.,
b,.,,,,11r1/1fcd M ,111tl• :rc.ated wlrh h·emwc:ln. /m "'"""'-'·'· • 1or1m:.
T11·,,1t11 Co11g,.,,,.. oftiw li11/<Jmvlogictil Sorl'fl}•afS<>111/iem ,\fricn,
Porchd~1t<>om. 12-15 July 1999. p. 91
'L" (ed<l. Bl11No11g,"''"'ti n,,fnttrl Orblt-lrt1""1, :-tw \"or,: ,\Jan R. U;;,
PP 611-616.
!!8;t: P.,\vt.sl(A. J,t., \'t'-Tt n, G,f, & Mr.u.()n, P.tr., .1nn~ \'~1orcomp~10ne. or 300lo-"T.\~Ot'A:r,,T, HJ\ .. ~,.. r,lOM(l.f_, r.ti,. DOlffiATY. ft.L. & D\'C£, A,L. 197'5 Se:atti(e
South Arric:un C.1/lroidC$ species (orbh:c1ong1Jc ,iruHCIOl}-pi: I rl!TV· tl 1101 pro. ce1 - vfru., £iol;,1tion..c;. Rr.pori a/th,• CS!RO Diciisi-tm of,\nimol
\\ith spcclol re!ercnce to the effect or tc:nporaturc on th• mu, or \'lru~ 1/rn/JII ,o,,,~rrOJ, 6.2, 82,
repllc:ndon in C. Im/colt, und C. l10/lt/11os Mtdfcnl n11d ~,i.•rlnmy
Emomology, tG. 10-21 3QI s-runtto. 1.s.. 1933, ()nt'i10t'.trw rcm1ffaiis {Railllt': et Hcruy, t!HOJ tmd It~
d1.!'\-0iopm.::m in Ce1/lroidl!1' m,hrc11lcmu Mg. Rt.1UQrtofth~ lnultlllt of
~~3 Pm,t.PS. •·•·· 01.Aa11un;,;. ,:.~. ~ -:r.u.u. 1-, 1982.. C11/i<'t>ldr,s .Olptcr~: .t,1/mo//>arh~/OJO'. Unl\,'TS!ry of c::.mbridgc.J. 2,2-28-1
Cemropogonldac)c,uch<'S sntl ,1ru• i.,nl3t!on< f!"l,rn them In th<
MukwadtJ V311cy. Zimbabwe, Jo1m1n/ ofthi Emomolo;;irnl S/Ki1•r,• of 3r;2 :.f. ~;tul\Gt, T.o.. 1985. The '\.(larch for hluctr,ngu~ Yiru-.es in Au.stnlia. Im
lt\Rl'fll. r.1 kltX-U.f\.J. ,,.,:., cd:io Blut1tongunmd R,•JmCDOrlJ11'in.rse1.
~11t/1<mAfrlcn. ~5. 195-200.
:>:e" \,,rl;; Al,,n ti. L,.,,.. pp. ?!li;-305.
28~ R \IU.tA1', '-t..'-1.~•ttAlilA)tbWAltAI'\, ~•• 1991. holatfon nnd malnamance of
u,11rocy10,t>1m ttmllvl)'i: VRI $!min. J<lr1lo):siall v,·1r,i1111ry Joumt1I. 3.
JOl v1. c101>~,. 1.1,.. 199~. Bo,ineephemcnlfeecr. ln:ai,,.,.., J.a.w THO.\l·

l»-69.
..,,s, "-" ~ rusr1N,R.<.., eel, . lr,jtwinll.i n,,..,,.,,
of Utv$//JC"k with
~;,erial Ref•"""" rt, So111ll,•r,1 t!jrlct1. (;ap~ T0\\11: o.eford Unlse":I}'
2ss R.-o. v.fi. It RI)\ Jl,. t98~\. GPnl'ufc \"'atiatlon 01 blutioni,1c virus scron.-pt- Pre-» !'lt1uthe:rn Afrka, pp.. ~53-"562..
11 botntcd ·fron1 h~t ($hccp) nnd \-Cc!O:' CulltnlrlttS.Pllriipc•n,:ls'; Dl the.,
JOJ ~J.GlO'R(..L r.o.... O'Jtl~Sn. ru1 .• rtUPJ'IC)O. r ). r.A,tl.J1\", J,(; .. 1-9;9. The
sa.me~1ic. 1mc.rf«m /oilfnnl of\ftttarl,u,ry Rr:.t-tJU.reh. 4-l. 911 ··9J.;
!s.olITTfon of thft.'<' Shnbu g_-coup \iru.u.~ ,H~\\' to A~tt.lH~ Au.stMlUm
Z86 R..A,WUSG~ r .• C.\.PEI.A, R.• l'l'tO. ,,., .. C,J'11:C.A. \JJJ,~ ra;..NA, J,, fl:U8~0. c.. C..\SCA, /tWt'mtl ofP...,•pt11·/mt•11wl Bloloi(!•tmd M«ilcnt Scfentr. .5:-, j8 t-582..
A," \.ftJ 10Q, r.s...1.998. TI1e1e.J:uionship tt.1ween cJim.1tc wd tho
iho c:ontlnucd
JO;):')"'f. CKOflOl., 'r,O .. Cr1tt:..~. J),tLk ST.\Nl>l•J\ST H,A- 1982
dlmllmuon of C111i,oules Im/coin in lbcJin. :lrc//lvr.t of\110/(JfJ·
~tarch for btui?tOJlJ?UC rclau.-d \'irn~cs in AU!\traliL 1,r. '1', CJ?orcct~ 1~. -.
\Suppll'mcnt 14J. 93-1 O.!.
1:.w, "·"· frd•J. Arbt.wit11, Hrsrortll /Ji tiusuwlia. Procadln~s of,11~ Third
28i ~vu. w.c.. SCRl\'A.~t fl.P•• ~eLSOX, n.r.• tfAfU)Y, J,1-, R()l:'Efl.U.. l>,Et, (. Symp,1>iu111. 15-1, Feimurry• 1982. pp. 183-lfl3.
Buuom,ilto\.. '-iru~." nuw arbcwin,s isoliltc:d from
Slit.SON. ,u.. , 19;0.
.306~,1 ftl or.O-L. 1,u .. croc,s-1,.1. o.n .• Sl"A.'"Df'AST. HT,.. ~ ,su, c..r. ~
mnmnu1l, :t.nd Ct1liC1Jide., midges in Kern Councy. Cnlifamia..Amtrknn D1'UA•f'Oit1l\, A.f. 1983 The- i.>0hmon of five difTel\'nt v1ru:,e:;: of lhe
Jouttrt,t n[ltopiClll Medll"i11ra11d f/yg;,•11t. Ul. S<H-SSl.
cpl7.oot ,c h-acmorrho-gk d'tsea\'C' or deer sero.group••-\us1rollarz Ve1cruuu,·
283 ROUN.1(;2U7., , ..... HOOUIIULS, II, fl(..4~JAS°O, ,,., .. 1992.. ACrican ho~ Jounurl 60. 2l6-2li.
sickn!!Sl' in S1>oin. \"tt<!f11urryM/croJ,iof<r.,()'. l3. 129-1~2. Jo; ')r t.t<wiu:. T.U ":-.1uurrt. ,t.f.. &984. TI1(! i~atian ofa blucton8,.uC ,•iru.,
289 svn,~-tli\Nl, f., Mti.l~\, 1"'Ktt., ~-. ,:o~rntsi.:t. L.\l,. (arc:um..,n-so, ca., fmm 011/colde, br,~1·lrt1rsi1 ..~trftrnJl,m \-C'ltrlnr,r:,· 1,,w,,11/, 61. 95.
\.UlUJ)R, P-~l'I,, M,\1.,\0UO.\, A. lt. t,, c;,\SPliJU, e,., :001, :-.101<.-cul:rr 3()8 ~T. Cfi<'Jfl.tir, T.n .. sr.,~·or.\..,t, tt •.\. a- cvmxoo, o.u. :~7S.1sol41lon of
diflcrcnrintlon of thl' old World Cul/coid,. tmlroltt >'J'l'Clc, cumpt~, ,\i.about~ \.inJ, frum )~ntinalcuul~ nnd Cu/ia,fdt'S bri~vira1$l~ ~u,1mUn11
[l)lp1ero; (:cr.,toPQgOrridoe), Inferred Ll<ingrandom ompltfled V~ti"t'frth,y/<mtnnl, 5-4, S5&-S6t.
potyn,qrphic D;>;A marke~ Mole..wtar &oluir,·. 10. 1773-1'8/i.
lO!tST.C.tOfl,C,i,F. r.o.. Sf.\SOUST. H .., •• (:VHINSJ;I. UJI.. DYCL. A.t.. MOW.~- ILr .•
?9(1$lllliR<, lt.f. l'fil>GU\", D.L fi TUCIQ'R. )1,A.. 197.'• Po<~i~lc Sprc~d Uf 0011um. tl.l. .. f.;AHU v. ,.,•.• 1'1Uf\f'l(':tt, (',. & Fa\?.l[n c.1••• 197a, Th~
.\lrlcan hor,e skkr1o;., (ln ~,c 1,ind. Jo11m11/ o/11ygir11,. (~1m/Jridg.>. 7!1, lsol..irhJn or u hlutmnguc \'in.i.s.from C11/icoitft•t collected in tht:
:?19--297. ~-0nh~rn tNritol')· of ·\~l!'.il(a. \U$rmllnn t i•lf:riuon• /aur,u,1. 3,
2!}1 SJ;'.."\Dt)W.... ~V~W.:.IH, 11-.. t RAbMU:>, b.J... IJA.'\:\U~. I-'.\\', 1993. h,Qlo!liOn of !5J.-IS.1.
bluctongue ,1ru,, ,crot)"pt 2 l from Cu/ico/1/rs ,pp. In lndono~lo. :uo s,. GEoJl(;e. T..o •• ~u..,·Dl \~1. H •.\o., C'l'lU~s .. ,. D,11. FIUPPJOJ «;. ~ c;.\RLi\'
\leter/11af)· Micro/J!o/og)', SO. 34!1-3SJ. J,<i, is$>. P~1tton virus e oew Simbu ~roup ru-bo\'U\b ~sotatl!d from c;nd~
:t9:l ~HARV. :\,A.O., 192r. Ot:\·ctopmvnt of .\Jlcrojitnria pcrYtnni i:1 C:,lro!,k.-: ~nd Cuira,/tl~ IJll,ttirnrsu 1n ,,~1ntlfn. ,\usm1liar1 Journnlof8!0ToghY1l
gm/Jami/, a prelimlnar; norc. Tr11mt1.r//cru oftile floJ'tll ~(>((,r,·of Scitnt.V. 33. 2l!'r-244.
TttJplc11/ ,\/~licln~mul 1/j'giMt. 21. 70. 3U S.T C,O[UIWt.. T,i>.. ~-f.\~Vf \.S1 fl-A., ,nru.llk, 11.J.• 7,..l..~.E\\'Sltl, fl., CYBiX~~.

293 SH!\JU', N,A.u•• 1928. fll11rit1 pu,~11111~. 11.s dne!oprutnt in Cu/l.:al,ltJ o.u. lSI c1t,.,"i•1s. n.s~, 19:-8-. ttt0h1tlon ol arbo,1nlSes. from cattt\· and lnsceb
flll$1ML Tm11S(t,·1f0111 OJ 1hr Rt>)'f'll Socle1yo/Trt1plrol Mc'illt:i11, 1111d
at senu.,el s1t<"S in QUO'ffi-i.!and. Aui1mllu1 1979-?985...4.rbo<'iru;
Nygir,w. 2J 371-396. /nJonnation F.J.'l'lumg,,, Junt.-. lS9-Jli7.

2..~ S:t-UMf>$AW/\, , •• A..\tAS.0, A.• ~uxu,a. ~-. ~·U~OGA. ~ •• ):URIUA'R,\; t .. 1...:-our. l .. 3)2 ~UM\ll l.O, H,H,, Ml... ,u~. c;...., \tUUJ'".Jt, ti,, ):CG.nttn'. $1.B., om.11•• V,L. •
to iniE'Ct lird~ (.\(~t(Oliei 1mg"fc"TOtU$) \•;Ith
nl·'.\"ff,\ \ 1 1 D.A., 197,", ,-\1tett1ptS
°'oi-wxu,"" "011~11. 1 .. 1986. Hpf,.oollakgical o;;urva:yaf chicken
tcucocytozoono,t, and Ct1/lcoidnorn~"""e· 81111,tln o/1h, Coll,,i, lVi,,Ja-wrfn b11nCTrJ/ti, Onr,/:(J(('Ta, tl()lrrultJJ. l.aiJ loo- and 1\Td11s.on.-lla
of.-tgrlc11t111,,.n,1d 1·,rrrtnn~v ,\/td/ri,r,. "lhon Unhct>lty. 43. o=rdl JQ11mulof llrlrmnto/ngy, 51. !1?-13.;,
202-209. 313 "'''" V<l•i. 11 .. 1994. C!,.<slfi<utlon, ep!d<>miot~· and comrol or

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 \'ectors: C:111/coldl!S;;pp. 135

~pod·bomc\iruses. /It: r.ocrzr:.A. J...,\.w.. -Ho,:,.1sox.esc.~1us-1", tc,C., llj!l,1-1111p <Ollilcii.ons. Ond,•r,wpoorr Journal OJ \ i'rcinm;· Rt>Ml't'h. tl.l.
tcdsJ. f11f,x.tw11s Disau,,so[Lh'Clft(}(I: uif/h Sp.'Ci4/ 11.'f,Jtii•~ ;0So11wrr1 31S-325.
.'ifriea. Cape Town: O~ford Unhw$lty PT~ Southern ,\fries. pp. 103- t.),. ,&U,:-i DIR U~llli T,f;. 01' K.. 1&96. Geog:nphlcal
33,2 \'t,_..,n.R, 'lo,J.. 'l'\'lU.
120. dlsuiburion 4nd relo1he a1>11ndan0< of 510cl-.·associared 01/i1:i1,dcs
3U SWA~FJJtlhL A, c, 11:UCKBUFt.~. ~ ...... 19;6. A ne\\' Oll"fl'lbcr or tho Pa.l~.tm speaes Dipwra; C<-r,110po~onic.bel ln southem Afril.":a. m 1ela1ino to
serojtl'Oup or orblvirust..>s:. nu., Vtl.t!rinn')' R«r.rd. 99. 360. ,heit .poi«mt:ol ::i.s vtroJ \·<.'tlors. Omlrr,.1rp,1011 Jounwl o/V'tNrlm11)'
11,'ttartiJ. 6.1, 25-33.
:,is T.\ll..\(;.u:,ac;>,;., w.1 .. 1991. <;tnctic canuQI of o.nil :iu>e~pobility (>! €ulit:Qitlc,
1·1rri1pfn11ls-Whh h1uetongue ,•iru" /\m,•ritm, /oamol o/Trcmrnl 333 \'E!\'.Tl.R, G,J. '1S\'1U.., l:',M ... \'\Sorn UJ\:01;1',C, Ot. lo .. l ~ St-.tS(WU\l
,\fedicine and llyi;hmt, .as. 666--67' I. abundance md 1)1lrily of ,roe1<-,,.-soclated C11/lcolck1S',>i!c!e< (Dip1era:
C:i:rotopc.>g()nid:ie} ln different dl~ttc- rc:giOn'> fn ,outh~ Arriua In
316 t.,AAJ~'>lll+ J.:., 0\.,\, A,, OJ:..\0-\, T,. MA'f5U(), 1\., ~UMA, M, & ~UliUtffl, U.,
rcloalon to 1helr \'iml vector p010111l:rl. Ondmtepoort Journal of
1968. ,\lno ,;M. • new member or the Slmb1.:group of ruboviruses from
l-',•rcrlnary· Rr.,rarclr, G4. ?ful-271 ,
mos.qui toe~ in Jopa.n /OJ)(lu~~ Journal fJ/1\frtfitnl Stwnc.-1 ,,mJ Blo/og;·.
21 95-101. 33.; \tl!~'niR-. c••J, " r.>1t\\'F'~ J.t., 1999. llluc,onguc, :1 muhi-\·t.-ctord 'j\,~SC in
South ..\frlc;s /'tol:re1/i11g, o/til< 121/r Co11gr,"' oft/:.• Enromohlgrt11l
317 TU:A.'\11\~V. tL. f·UJt~AJ.:I, K., KITAOK,\.~, IStu, T. & l"UJlT.\, J.• 19"8..a,
Slldo~.o/So11llum1 Africa. Potchcf<troom, 12~1; lul; Im. p, ·30.
Cuhlvo11on or 1,wro,yro;:,,on ,m,lllfryi: purlficanor. morhod or
l35 \"E.~Ttt- G~t,, P.IU\"6Slt\, f..T, \'.-\~" n.rp.:.A.A., :.tii.1.L()B.. p ~-" r.UL\CW-.'1C:).., W.J••
!tporozoit~ for inoculum m Ol'O, Uullcriu o/r:1t!.\"1pp,J11 Ve1edmu')1 cmd
7.oorr<l:mcal Coll.og,. 33, -1-i5. 19:18 VectorcompctM<l' of Culirnltks bollrit:os and C. fmi,:,,/11 (Dipiero·
C:L-ratoposunirlac} fo: South :\irk.in hluc1ongul.' ~in.;" ...tmtypt•:ot ·t. l und
318 LU:.-\QKA, ffu.,.W1't, C., 8.-\rS, 0,. or.AT,\, )r. 4,, U.\11:A, M,. 199:). lfWCS~JIIC)n of
.a Mtdftaltnd Vcn..•rim1ryJ;·111omolr.,g.1~ J2.10l-1D6.
C.~,l/co11feHOip1era; c:.im1opogonida~, m rel,uon ro 1h~ trwnm1'5lon or
.n6 vr~,-u.. G.J. s. S\\U1'M.,~. (;..S:• .t989. Seasonal abundance :ind p:uli')' of
boV1ne onclloeetc::& and Qther nt31'iae In c~nuill K~-wchu. Jupnn. P,rmsll~
-/OtJmal HK. Frtmc. Pnr(l$irology, z. Je;7-3; _ Culi,oides bhlng mid~ a,socint.-d \\11h li'<'Slock ar R<>rno. Lesotho
(Dfp11:ra: Ct'rdtopogonida~ l. Or.tl('h·tt/NOl'tJIJurm~I ofttf:1~ri1itt.ry
3l9 T.\\U>lt, L.R,, 1962.1bt?abso!ut<; c-Didmcyo:'tnstct 1mcdon traps. Ammls. H,"8J.'tttrh. 56.17~~177.
of,!pp/led Biology, 50,40:H21.
337 ""'~tR. A,R. 1977. Adult li(c,pan ~d rcptoduetlve status of (.i11/co1dC$
32onm.utt. ,., 1921, Ajrlm,, Hurfi•Sld:11(1$ (Par.r. Eq{,orumJ. Stltntt- {D1p1tra-: Ctratopry,unid.sci in !.:-cn>"iL whh 11!/c:rem·\! to virus
8ulletln. Xo 19. Departm<nl ofAgriculture Union ofSou1h Afnca. pp. mmsmas.<ion. Bull,1/11 of Emomologi<al ft,•se<1rch. 67 W:,.-il 5.
1-32. 333 \\'UJCl'Jt A.R,., bOIW<AM. l',f.t... 1976. lllood feeding or Ctrliroi<lrs
~ nt£ODOKU>1S., ;\., ,-....:vn.r.. L.111,, u •.c., 1t.1 . .;,1 BO~<Ofll, s.r., J!J79, VJrust~ 1Dlp1<m: Q>ra1upogo11id11,,1h1 K•nl'!f In n,l;atfon 10 rh<' epld,•miologyof
isolnted from C1tllca(d,; anidll"S In S1>u1h Africa during unsu«es,lul blut:1tm,8\li.r and cph<'mc~ re\·cr. Rnllotl11 (ifEntomalogl"'' HC.."'-'mrh, 1,6,
ancmpts: 10 isolnte bo\1ne eph~mero.l (evcr\-rru~. OndtJf$Jf/J<HJfl/ourrml 181- 188,
o/V.r,rin«ry /lt-1(/1. .;6, 191-198. 339 \\'ALJ.3iR, A.rt.!. DA.VU:~. f .r. .. 19;1. A prl'.'1hulnotl')' ,uri.~~ oflhe
32.!:t 'TTI)WD.L M-"'· ~ TIUWtiU. ,1 ,., t98:?. Dcvtfopimmt of MonsmuJ/a or;;;o,dl cpld~mfoll>i:)· of bluc1on1,'Ue in ~enya. Jouma/ ofH;11i111,. Olrnbrid~e.
In Slmulfum mna:.oniC'um. Simlllum cvgqm,xm,,m nnd C1,Jicoi<lr., 69, .:;-6(),
insln11aws from Am'11Ana.... Colombfn :\mmcan /()umal o/ Tropical 3<0 W,\LTO!\", 1,\\, \\11.Dll. 1•--\., l:R.A"\1fJt. \\".L. (,\111 U, t",.t:. U,\\'IS. (,., ltnUUtOO~
,\1«/lci11,a11d/1Jgi•1:q, 3!.1137-1141 r.ft,. ,:001U..c.e1.. ~1ruaL r.1 .. so~~~. ru1. ftJt\."\'xn ,;.r..... rqa;. tpi:f:ootlc
323 u,~•t.. "· 18ll4 F.pid!!inlnlogl"'11 studfe, ohuttuuer ,'t(Cffill in lti:l•nd w,ic:1Jlor,1omotltl, In Colorado, USAI 1982: cp!demtologlc and
ponll!> Prukri,<:J,;, Tlnar::1. 65. 65~. entomologic s.1.udh:,),,.. •.\.1tu•rlc-tm /oun,nl tQTrop,'(.(.1/ .\Jfd/dnL•a,1d
ltyglMr. 3~.16&-176.
:iu ,·~, w,,,,,_ ...
1\!90. Unl,'C:'>it)'t)r l'n!loria, Ond,.,..,epoon 0110. South
3M \\~l:'.~lA.'"' J.IL ).IUKJ'HY, K- h'XDERSO,S". Ul., DL\fAJtTISr.1.<.:,, LOXGHUHST,
Africa, IJnpubllslwd data.
W.M . .,, co~ l\'Ol L.Y, c;•• 1979. Sea-:.unalphwofonce-.. pa1ho!ogy nnd
325 \ ,, Ak~. H. At \l>lSW"~>l- ll., 1q~ ~11b,an1plfng Ofl:U'gtl llgh1 lrdp lrnnsmhsion of 1ht· quail lwart\\Ot'm. Spltmditln/Uarin l'lfllfurnien.i:f.J. in
cruch,·> or C11/lco/J/"1 ntptcr.t: Ccro1opo~onldae1. o,1de1Ste(}t10n/011mal 11onh~rn Cal.ffomln U.S.A. 1.Xuma1odtl: fllaroldea~ o,,,artfrm/ourr:rdof
ofV.reri1111ry R;,,wn·h, ;ill. 1sa-189. 7.oUfotJ• 57. l!lil-(8:7
s..."'6 \ ,,.t."'· L, 1969, Culltold.:.s M 1ranun.hltf\- of di!ta~ C'i<1n!tt1 .t.tc,l!cndP J42 wn1.ar. ~,.~· D.\'l'U~. ,1 .. ftA\\'U"'CS. r. ~ MEUOK, tt.s.. 1996. Effects of
M•xlro. 99. i82-78i 1e.rnpvraturt un sur\'i\·41 o1nd nut ar\'Ul~ntsb ui Afric:an hor$e ~<""imc.."N-
33i' 't"t.YlDl, G,J.., GrtAIL\~1, ~.D. & lL\\UIUt\, ~ :!QM, .\frfclln horse fiiiC-knm \irusln Culiroides trarllpennis tanf)r~usi.s 1Dip1era: CerotopogonicfQ.e-:
<pidcmfulogy: vec,or comp<l1ei1ce ot' South African C1tllroid,•1<p•ci"' .tnd ib ,.jgn.ficuncc ii, rchuian io th<" cpid.1.'miology of disro.se. 8ufU1tin of
for Afrkan hors• stcL:n,•,< viru< wrotlll"' 3. ~arid 8. Ml'iliMI and f~11omolog1cal R'-'Si.·Qrcl:. 86. i15-i20.
Verorimil)• Emomol01ti, l·h 24~250 .343 Wl.Ta.t... lL, ~'t\1U.. f..M, &-Elt.\S..\1U:,, B,f, 1970. ~1udl1.~ on the tr.:osmbsion
).28 \·r.n,M, C-t,J,.f,Rbt~'l\\'ALD, D,:\1,. f>A\\&:~EA, f ,t . \'L~T0\1 l:!..IL II H(')WEU. of Atrkon horscs:<'knr.ss. Ontlnstrpoon Joumal.cj l·'..-r,~rtnmy Rt"~-..w~h.
P.c .• 1999. V<:<l1or cornpuu,ncc l>l s.:l,,:n-d SouthAfrlean C.i,Ucoidl/$ '.li, 165-168.
,pcc.,es for the Brya.nttO:l <crotypeof oquin• on<q,halosh ,iru, .\/fd/m/ .l-14 \\1n~1'U:11. T... sw.,"'''"~t..w.. IO~R. t'hawc1orl,.01il>11 o(p~tenrlnfly
m:d \'rt1trl11ary l!ntomoloio·, 1a. 393--lOO. (Ol?toaopfe l'alyi1m serogroup arbMrU>O$ '"''••eel In ZlnthJbwe. /011rnnl
J.2'9 \1i~ fUM, O.J .. Htlt.. l!l.AJ\f, Pi\JOR. 1.r.P." ~!\'lU.. J!. ,i .• 1991. Ille u.,..e ora oJC,m,rn/ Virologi·. 69, 2:.'21-1227
membranv fttding technique 10 dc1t•rmim.• the Infection nuc- of 3-4S \\'ffl.STt.f.Jt, T. S,\\"A:O;POO. 1\..1 T:.R,i\$)1V~. B.f,. 19"89. Charncieriz.•urort of
C,11/et,/des imlro/11 U!p,cm, Ccm1orogon!d:1t) ro, 1wo bl11e1onguc ,1n1, l>al}·am ,~ro;;mup arblvtnue< i3ola,rd in South \irfcn nnd <;L•rologt~
~e:rorypcsin South .\rrfca. 011d,m,poort /oumnl ufVNtrr/11ary Rtai!ilrr/1, ~vidt11cc for their \vidC",pfC'n.d th,.trihutf()n in the rountl'}'
58. :,-9. F.pld,mlologftrrl lnforma1fo11. 102. lfi-32-1.
330 \~'TF.H, C,I, "MU-.P.\'JNl:llt.. A., lll~A. Th1 1 virtu:iJ ~b1"1."nCe a( C:u/if:O(do, J,i6 WtEst:MSCffO,U'f'. 1-, \\11..SOS, W.C., I REXC:U, D,D.. 8,\Jt\,.t, .\. S.. fOll., t~D .•

imicolu (OiplNa: Cor.1tOJICl&Onldllc) in o llghMrap >'JI'\~')' o( th< colder t993, 81u..:-tongue vlru~ tn ~ccp and dlulc and C,ultcoftl~,J.'flriifN}uzU
high•lyini; •••• ofthc~tem ()rongc Pree Stair. Sou ah ,\frfra. and :md C. '1elllfer tD11>1er.i: C~:,nopogonllinoJ In L<>ulsl5n:t. /0111111'1/ of
lmplit:ndons for the mtnsmlsslon or arbo,i.'We>. 011d,meJHJtJt1 /uim111/ .\l,·rfh·nl Entomology. 30. 7 l9..72..J.
o/V.reri11ary Re,1•1m:h. 61, 32i-840, 34;, \\1UJ,\ \1ij, n•.L, 11..\U.. R,U. 8Rtl<:E. r\,B. ~ S(:llm I~ P.J.. 1905. /J1-oe.r10,k
331 \"£~'TtA. C.J .. MU~w1-sAtl H.• sl?Vlu.• 11•.,1. ~ Enh .\fU)b. M., 199b, CuUroides 1iluomQlogy.,,c,, York' f Wiic,.335 pp.
1D1p1tm1: Cc)111opogonldael assoclaae<l h1th I" e>tMl. ln the 348 w1nm.\'x, t'..J •• )1tu.0n. ,~. .& ?r,\'l'U\', ,, .. 2001. u~lng cUmatt d,tta to n1~J>-
Ondemepo,m Men, G•uttng Provine,,, South Afr!cn. ._, dt,10,mln,-d bi 1ht pownrfnl dl$1rlbutlon ot CuUtol{l,s lmi,ola IOiptern.

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136 ,•cno.x o"": AspeclS influencing the occurrence ofinfec1ious diseases

Cemtop<>gonld~) In Europe. R,w~ ~cnfit111,01 t«:lmi(/U,, 015cc 350 Y•n.. 1-'-· 1.0wn11!.. iu:." nt111~\1W. M,L. ts8: Dcrclopmtnt or
International d~s Epi%oo1io,;, 20, 131-74(•. Trtral)(!ta/011,mo lle1,·eJlynt to the lnfe<:th..., ~ · in C11/ic,;/des ho/1,nst,,
149-Y,UIA$UTI:--\, In t"JTA.UUR..\., U. & KAXA,,!VKA.. A., 19i'S. Studies on 'K3$Cn' Of
Joam<1I ofPa,asltoloK,•, 68. 293-!15?.
hoN<!S In Hokkaido iV. Researches on the punl;Jes In liol:kaldo "ilh 3.51 ,A).tU·\\"tf.t, tt. ti' cvm"'s~,. o.H., 1~84. rs.ol:uion of Klmburl1.1 ,~ru,. a
de,:oription or an.cw ;pedes./flpor:IISl' /oumni ofl1ewl11ary R...,,.,c1,. 5. rhabdo>i1U$, from Cul/coldts bt11vllM.<is. AuJtrn/ian four,111/ of
89-96. /'.xJNrill'Mtrll 8iologi•,.nd .\.ftdlr'11 Sc/mu. 1!2, 1,S-780,

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 6
Vectors: Mosquitoes
PG JUPP

Southern Africa and congenital abnormalities in livesmck64 (see Chapter 94:

Diseases caused by Akabane and related Simbu-group
viruses}.
Introduction
Rift \ 'alley fever. WSLand \\1;\! viruses are also important
Among the pathogens that c,iuse di$ease in li\'esrock in in causing human disease. while 1wo other mosquito-borne
southern Africa. onl)' a few viru$eS are transmi11ed by mos- viruses. namely chikungtmya [Cl UK) and Sindbis {STJ\'} \i-
quitoes (Table 6,J l. These and other anhropod-bornc ruses. only cause disease in humans. Their epidemiology
,;ruses (arbo,'iruses} characteristically multiply in both the and association with human disease have been summarized
vertebrate host and the mosquito vector. Successful tran$- recently by Mclntosh.• 6
mission is dependent upon the vims producing a ,iraemia Apart from virus isolation from wild-caught mosquitoes.
In the vertebrate. while the mosquito which reeds on the ver- the lines of e,idence incriminating a mosquito species as a
tebrate must de,·elop a salivary gland infection followed by vector are its susceptibility 10 infection \\ith. and Its ability
excretion of virus in the saliva. The arboviruses belong to 10 transmit. the vims concerned {vector competence) plus
several families and genera and their taxonomic categories irs relative density and ecological characterist ics. Vector
and host relations are discussed elsewhere (see Chapter 7: competence ls assessed quantitatively in laboratory tests
Classification, epidemiology and control of arthropod- while data on dcns11.y and ecology are obtained b}' field ob-
borne vin1ses}. servations. Relevam aspecrs of ecology are feeding behav-
Of the mosquito-borne ,1ruses that occur in southern iour. including host preferences. lnr:val habitat associations
Africa, Rift Valley fever ,1rus (RVF\) ls periodically respon- and biology of the egg stage.
sible for serious disease in sheep and caule in the subconti- Climatic factors influence arbovlruses both directly and
nent, while Wesselsbron virus (WSLV) causes a milder indirec.1ly. Temperamre conditions directly affect tl1e multi-
disease. affecting sheep in panicular. \\'est Nil~ virus (\,\/!':VJ plication of the virus in the mosquito vector. low tempera-
anrl Ran1i vim~ tRA¾''V). howrvrr. h:we only hePn t'xperi- ture." ' slcming down ,~rat replic;icion :md lowering vector
mentally incriminated as causes of congenital abnor- competence.JS. 101 I lo\,·ever, more importantly, climatic fac-
malities in lambs in South Africa. 12 In France. Wl\'.V has tors openue through their influence on the geogcaphical dis-
been documemed a~ the cause or cncephalomyelilis in tribution and prevalence of the viral hosts, both mosquito
horses?2• 33 and vertebrate. In the case of mosquitoes, high rainfall and
Orher mosquito-borne viruses of the families Tog,wirl- temperatures can lead to higher population densities of spe-
dae and F//111i11iridlle that do not occur in Africa. bm which cies belonging 10 all genera. particularly Aedes. Climate there-
cause imponam diseases mainly in hotses and humans out- fore has a considerable effect on che epidemiology of
side Africa include the alphaviruses of Enstcm. Western arbovirus infections.
and Venezuelan ence1)halomyelitides and the niwi\iruses Apart from some work in the temperate. moist highland~
of ~lurray Valley, Sc. Louis and Japanese B encephalitides and subtropical. moist lowlands of Zimbab\\'e, the areas of
lsee ~e chapters on the specific diseases). It is possible southern Africa lying omside Soulh Africa have la.rgely only
that Akabane vims. which is a bunyavirus of the family been studied for the presence of mosquito-borne ,iruses by
Bun_vtwiridt1e, is also present in southern Africa, although means oi serology, without virus isolation from 1,i!d-caught
thh still remains to be demoru<trated. This virus has been mosquitoes being attempted. There is considerable arbo,i-
isolated from C111icoides spp. and from mosquitoes in rus activity in the tropical region of Mozambique and the
Japan, suggesting il is also mosquito-borne.98 Akabane Zambezi t•alley. and in the subtropical, moist northern belt
V'irus h,as been incriminated as the -.-ause of abortion that extends westwards from Zimbabwe and embraces the
137

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138 <r -no, c.,.: Aspec1, influencing the t>C('tlrrN>ce ofinfer.tfou< dl<Pa,!"S
Table 6.1 Mosou,10-eorne
viruses causing disease 10
lrvesiock and/or humans in
I VIRUS MOSQUITOVECTOR Wl,0
VERTEBRATE
HOS~S SHOWING OlS~A.SE

soutnern Air.ca HOST HL:M.\NS LIVESTOCK

Togav,naa:
Alph3'l'i1JS C~ikul'~ur;,, a 4e (Die Jfwc,fe·
Sim,itlis Cx /Cux.) uni1,r:,Hus
flavimoae
F/2v111rJ$ Balll• Cx. (£um} robinows Accents
Wessetsbron Ae /N~.J circum/Jteolus i:!dents1
Ae /Neo Jmcmcrnhi
Ae (Ocn.1caCJ/lus!Jupp1
Wast Nile Cx. (Cux./ umM/BIUS
Bunya,1m:lae
Phlebo'llt~s Rift 1/;illev Cx (Cu~ J1t.eileri
ieve, Cx. /CuxJ 1om/Jaensis
Ae /NeoJ c11cumf,1eorus
Ae (Neo.J mc,nrosh,
Ae (Oen Jcal;allus/;upp,

a Disease produceo ate• inocula11on w11h \•:us ,n laboratory

Okavango Swamps in nonhem Batswana, and the Capri\~ deve!opmetn of a poor general state of health in sheep and
Strip and Ecosha Pan in nonh-easrern :siamibia. 13• 511• 61 cattle. Sheep may even die from loss of blood as a result of
A:rbovirus acthity in Sou1h Africa is more or less confined the continual biting by vast numbers of b1sec1~. This is
to the subtropical and cropical Lo1·,Yeld of the Limpopo and probabl~ che main reason why mosquit.o comrol in South
:Slpumalanga pro1inces, l() 1he KwaZulu-Natal coa,;1al low- Afric:i is necessary. However, effective large-scale mos-
lands. and to 1.he temperate. high inland pla1eau of the quico comrol on farms would be difficult and very expen-
Karoo and Highvcld. 'i'hcsc regions all fall In 1.he $ummer ~ive to carry out.
rainfall are.i. Th<> inland pla1eau has 1'inuallr no rnin from
)la)' to :November. while in 1hc ,,inter. from Ju11e to Augus1.
Virus isolations fron1 mosquitoes
night 1emperatures are often near free1.ing and periodically
faU belo1, rhis. In the Limpopo Provincc-Mpumalanga Table 6.2 list.s 1.he six mosquito species which )'ielded mul·
Lowveld and Kwa7.ulu-Na1al lowlands. v.'intcr,- are dry but tip le isolations of RVFV "·hen coll eel ed on farms during RVF
mild. epidemics among sheep and c:aule. The larges: number of
Records of rainfall in the ~umrner rainfall region of isolation5 on the inland plateau came from Cale:< rheileri
Somh Africa since 1910 reveal a 20-year oscillation be- (Figure 6.1 ) and in the KwaZulu-Natal lowlands from Ci:.
tween decade,, of above and bulow a,·erage rainfall. 111 The .:ombnensis. Fcwt!r isolations were made from aedine mos-
1950s was a decade of h igh average rainfall. 1.he 1960s quhoes because collecting <mly began once an epidemic
below average and the 19i0s again above average. During was already well under war. by which time the acdine popu-
thP £1Prioci 19:i, rn Hl7<J, rhP m;ijnr nurhrP:ik, of\\"\', <'Ht K lmion, \\'NI' waning -\111hr J\,>1/e.~srr gi\·rn in Tabl e 6.2 m,•
and RVF virus iniections in humans and domesric animals floodwater mosq t1 iloes belonging ro the subgenera
have occurred mainly during periods of above average ,Veomela11ico11i<m (Seo.) a11d Ochlerormus (Och). Aetles
rainfall. (,'\'eo. ) m.imoshi and .·le. (Och.) jllJJJJi are illuMrawd in Fig·
During tim<is of heavy rain certain pans of the inland ures 6.2 and 6.3 respecti\·ely.
plateau. not.ably the panveld area of the western Free S1.a1.e :\s shown in Table 6.3. 43 isolations of \\'$1,V have been
and northern EaMern Cape pro1'inces. ma} become so made from mosquitoes collec1ed in South .\frica and 15 from
saturat.ed that large pans and areas of pastu1·e are flooded. mosquitoes collcc1cd in Zimbabwe. Apart from two, all these
Such accumuhuions of water mar remain ~randing on isolations were made from Oooc,lwater aedine mosquitoes.
farms for several months. panicularly if there is a grani1.e However. RVFV isolmions were obtained from numerous
underlay as. in rhe case of the Kaap Plateau. Under 1hese C11/e.t as well as Aedes spp. Like RVF\' 'Table 6.2), the mosc
conditions large populations of floodwater Aedt>s mosqui· hnponamAedesspp. involved with \\'SL\ were Ac. (Seo.) cir·
toes are produced inirlally, which are subsequently c11mh11eol11.~ (KwaZulu-Natal lowlands) . .4e. (OC'II.) <'llix11/11sl
replaced by equally large populations of Cu/ex mosquitoes. juppi (inland plateau). Ae. (NeoJ mcimQS/rilluridus (IJ1l::1nd
\t such limes epidemics of RVF may occur. bur the pla1eau) and1!e. (.\'eo.) mci111oshi (Zimbabwe highlands).
extremely high densities of mo~quitoes and 01.her biting from 195G to 1980. 139 isolations of \\''.\'V were made
flies are also detrimental to livestock in other ways. These from mosquitoes in South Africa. Of t.hese. 136 came from
include reduced milk production by dairy cows and 1he rno~quitoes collected on the inland plateau and 1.hree from

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 Vector,~\losqui1oe. 139

Figure 6.1 Cu/ex rh1if/er1

the KwaZulu-\'\atal coastal lowlands. On the plateau. 128 of sion tests for six species of mosquito \\1th RVFV are given in
the isolations were from Cule.-c 11ni11i11m11s which occurs Table 6.2. Dara indicating the approxima1e length of the
tluoughout this region.8 ·l There was a large epidtrnic on,-:-,: ex1rinsic incubation period for each species are also shown.
tever in the Karoo and :'sortbern Cape l'ro,~nce in 19i l, On the basis of these results the order of vectorial impor·
when 1he lnfecrion rate of the Cr. 1111il,i11m11s population lance is Cx. theileri, Ae. c:irc11mlureol11s. C" ;;om!Jaensls,
reached 39 !nfecre<l mosquitoes per l 000 at Upington.90 Ae. 111cimoslziand Ae.Juppi.
The low incidence of \\'~V in the KwaZulu-Natal coastal Table 6.4 gives the results ior preliminary te~ts on three
lowlands is apparently d41e 10 1he p:ior susceptiblllry of rnore mosquito specie~. all of which had high infectlon rates
r.ulex nenvei 10 infection. :;;t C11lex neni•ei replnces C\·. w1it•i1- after feeds of high titre of \ints. and \\'hich subsoquemlr
1mus allopatrically in the KwaZltlu-Xatal Jo,,·tands, where it 1ransmitred the vintS. Comparison of the results for Aede$
is appare111ly the mah, vector. 36· 8:.1.11.>. 11 • 1111ide111m11s (Table 6.4 ) \,ith those for Ae. mrintoshi (sum -
Banzi virus has been isolated almost exclusively from marized in Table 6 .2) shows that Ac. 1111idencaws is a more
Cu/ex (£umela110111yia) rr1binotu.s at Harare in Zimbabwe efficient vector.
and at Pon Shepstone, Empangeni and :.lmbntuba on the l'ield studies during RVF epidemics ha\'e shown only
KwaZUiu-Natai coa,1. h1f11ction rates in this sp11cies have low inrec-rion rates in 1he mosquito vccwrs. ofum no more
often been high. 91 than one infected mosquito per I 000.;,. 89 In addition,
high vector competence evidemly requires high level,; oi
viraemia in the vertebra re hosi.45 81 • 89 Despite this. a strik-
Vector competence
ing feature of RVF epidemi<:1i among sheep on the inland
Values for the 50 per cem lnfecrion threshold and the trans- plateau oiSouth Afr lea rs the rapicUty with whicl) infection
mhsion rate determined In horizontal biological 1rnnsmis- spreads through Oocks. causing heavy losse~. To account

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140 m.-n11, os,: Aspects influencing the occlmenc:e of infecrious diseases

Figure 6.2 Aedes (Neo/ mcirrroshi

Table 6.2 Southern Afm:an mosqu,tospecies yield111g mul11ple ,soJat1011s of Rift Valley fe1er virus from 1953 tc 1981 number of ISOiations and results of
transmission expe:iments

REGION SPECIES NO 50% INFECTION TRANSMISSION RATE" REFERENCES

!SOI I\TIONS THR~SHOlD3 \DAYS POST-INFFr.TION)
LOG ,0LD.o/'ml
SOUTH AFfllCA.
KwaZulu-Natal Cx. {Cux./1omoae11s,s 7 C 7,5 3%, a1% 42%, [14 22 29i 56.92
coastal lowlands .Ae. /Neo, 3 C. 7,5 32-38•.e 116-31) 54.56.92
circumluEJJOlus
SOUTH AfRICA
Inland plateau Cx. (CuxJ rl1eile1J 22 ' 5.7 13-35~: (15-22) 22.41 75,84.89
Ae (Och,!JUPPJ 3 8.7 e~. 1111 BS
Ae /0Ch 1ca/Jallussl 6 8,0' 0%' (12. 13) 22 89
Ae. /Neo/ mcmroshid 2 8.1 17% (10.17) 89
ZIMBABWE Ae. (NeDJ mcinroshid 2 75
HIGHLANDS

a The. titre of wus neai!ed 10 infect SO per cent of the mosquitoes

b The propomon oi i11fecled mosquitoes transmitting viws to iamsters
c Experiments dor.e with Ae ceball~s ss.
d Reponed as Ae /inearopennis

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 Vectors: Mosquitoes 141

Figure 6.3 Aedes(Cah.J;upp1

for the high level of infection, biological infection by mos- that Cx. 1wiuir1aws is by far the most efficient vector, with a
quitoes Is almost certainly augmented bymechanical rrans- low 50 per cent infection threshold of about 2,3 log LDll0/m1
rnission by various biting lies. In mecbanical transmission and a high transmission rate of97 per cent.38·~0
there is no viral replication in che vector, only a passive Owing to the high infection rat~ of BM1\I in natural
transfer of vi rus. F.xperimenial mechanical 1ransmisSi(ln of popul(ltions of Cx. mf,ino(U$ ar Port Shepstone. it was pos-
R\'FV has been accomplished by various haemarophagous sible co conduct a transmission experiment In 1he field in
Diptera: Glossi,w morsir<ms. Stomoxys calcitrrms, 1-wzomyia which indh~clual hamsters were e;..-posed 10 16 groups ofbe-
longipalpis. 01/icoides uariipe1111is, and several species of rween 4 and 19 wild-caught mosquitoes. Banzi virus isolates
Aedesand C11/ex. 2t1, 55 In imermpted transmission tests from were made from the mosquito groups that had fed on 11 our
,iraemic 10 susceptible hamsters by the bite of /ledes aegypti of 16 hamsters, which suggested a high degree of vector effi-
formcs11s.55 cransmission was achieved by groups of mos- cienc:y.81
quitoes for up to 25 mlnures after the lnfecrive feed bur not
after 5.5 hours. After shorter intervals of 3 10 5 minutes,
Ecology of mosquito vectors
single
mosquiroes were also shown to transmit the vims on three C1tlex (C11x .) lf1eileri
occasions. C11/ex spp. deposit their egg rafts on the water surf~cc,
·n1e resultS of earlier l'ettor competence rests with WSLV where. depending on the temperature, they hatch within
are shown in Table 6.5. They indica1e that Cx. lheileri can be two or three days. These eggs cannot resist dl}ing. hence the
excluded as a vector. while Ae. circ11m/uceol11._~. Ae. cabal/us prefe.rred types of larval habitat are permanent or semi-per-
s.l. and Cx. z-0mbae11sis are potential vectors. manent ground pools, although temporary pools arc some-
Laboratory evaluation of all 1he common mosquito times used. C11/ex rheileri is 1he mos1 abundam and
species occurring in the Hlghveld as ,,ecrors ofWNV showed consistently prevalent mosquito throughout the inland

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142 » mo, u": .-\spec1, influencing the occurrence ofinfcctiou~ di<ca:<es

plateau region of Sou th Africa, but ii is usually rare on rhc to some exrem (endophagyJ.~1 Epidemics of \11:\l fo\·er are
KwaZulu-Natal coa~t and in the Limpopo Province-\lpu- probably partly a result of increa~cd feeding on humans
malanga Lowveld.4 h· r...i. u_q, 112 Thi.~ ~pecies feeds on goats. when panlcularl) high densir!es of the vector occur, as the)
cattle. sheep and hwnans, mainly at grow1d level did at Upington in 197~.1111 Precipitin tests on blood meals of
during the first three hours after sunset. ll also feeds 10 ll Cx. 11niui11arr1s colleeted on a cattle farm near Johannesburg
limited extent In the 1rce canopy a.-. it ls moderately ornltho- showed that 3 7 per cent of them were of mammalian origin.
phillc.1 3 ; 48• s, The larvae are round in a wide range of and 5.'> per cent of these mammalian meals were from canle.
ground poolf> ,'3rying from clear \'l~is wll h emergent \'egeta- C11lex 1111i11i11arus will milize temporary, semi-permanem
tion to pools highly polluted 1\ith organic 1va:;1e from and permanent ground pools as larval habitats. although It
canle.28• 34 prefers siws with clea.r water and emergent
vegetation.:!ll. "'1 Recd beds at the margins of Lhe Gariep (for-
Culex (Cux.) zombnensis merly Orange) River were probably utilized at Upington in
This species is common in the nqrthem KwaZulu-\atal 1974 aJld 1976 when densities of C:\'. 1111i1'i11arus were tugh
coastal plain, 18· 8 •· 86· 02 butrarer in the southern pan. 49 It is there. It appears to be important for C\'. 1miuirra111s adult~
mainly a ground frequenting species which feeds noc1ur- and larvae to enter a state of quiescence to overwinter, al-
nally on canle, sheep. goars. antelope and humans.fti\. " 2• ii9 Lhough Lhe available e,idence i.ndicatcs that adt1hs pass Lhe
Cu/ex zcmbaenslsseems more selective In its choice oflarval winter in a nulliparous ratl,er than a parou, condition.~; s~
habitat than Ct. t/1ei/eri and is probably more confined to
cleaner water ,1~1h emergent vegetation. such as river mar- Culex (Eum.) r11bi11ot11S
gins and permanent pans. C11/ex rubi1101t1s can be colle:-cled in reasonable numbers
0nly \\1thin or near its preferred habitat. This i-< dense
Cule.i: (CtLi:.) 1111i11ittar11s marshland \'ege1ation. always associated with permanem
This mosquito occurs throughout Lhe l-lighveld and Karoo l\'ater such ni. lagoons. rivers or streams. and dams or lakes.
regions and is usually one of the most abundant Cu/ex spp. Hank reeds. papyrus or grass are always prescnl bu, trees are
1aken in catches:18• 53· 112 le is predominand)' ornilhophilic either few or absent. Although this type of habilat occurs on
but sometimes feeds on humans and oLhcr mammals 1 17· the inland plateau, it is particularly common on tht! Kwa·
4 a. 53 It reeds nocturnally. partlcularlv in the tree canopi• but 7.ulu-.:-;;ual coast, and large populations or the mosquito
also at ground level. While this species is only slightly ru1- have bern sarnplcd at Mtubatuba and Empangeni.9 t The
thropophilic, It hai. been shown to feed on humanl> indoori. mosquito prob11bly ~L>ek$ out lhis dense vegetation be(';Juse
of its sensirh•iry to wind and low humidity. Laige numbers
have also been taken in mature sugar cane fields when these
Table 6.3 Nt:mbe' of ,soiauons ofWesselsb(on virus from mQS1lu•!oes
adjoined the natural habitat.51
collected in southern Africa
Cu lex rubinows feeds primarlly on rodents, to a moder-
REGION SPECIES NO. AEFEi\ENCES ate degree 011 domestic ungulates and rarely on humans.51
1SOLA· The feeding rate b high on hamster$ in mo,quito tra~
TtONS placed on the ground. Female mosquitoes confine lheir ac-
tivity almost entirely to ground level and rarely enter traps in
South Africa
KweZulo·Naral A~. /Neo.Jc,rcumluieof:JS 13 s. 77 105, 113 the tree canopy 5 t
coas:al Ae. /Adm./ minuws l 113 Larvae of c,·. rubfnows have been collected in reed beds
lcwlam!s k /Adm I veenia11 0
y in shallow\\'ater. 51 Hopl::insza recorded the larval habitats as
Mansoniw /Mnd./ 6t1.. ·3 being papyrus S\\'3mps and weedy or f>isrfa-covered bor·
umtormis
row-pits.
Cl( /Cux I nea,ei• H3
South Africa
Inland Ae. /Neo./ mcmroshi~/ 3 60 Aedes (Neo.) clrcumluteolus
plateau el/Jolhorax' This is the predominant member of the subgenus St!ome/a-
Ae (Neo/ mcinwshi/ 5 46 11ico11io11 t0 be found in the tropical coastal lowlands from
lu11dus northern KwaZulu.:'><aml in South .-\frica northward, Imo
Ae /Och.) c~/Jelluss.' e 13 cO ;\•lozambique. In Tongnland in northern KwaZulu-1\'ntal it Is
Ae. (Och.IJUPPitce!!ellus 5 ~6
,,mDabwe Ae /Neo.J mcm1os1r,• 10 7, 75. 77 often one of the rno.;,t prev(l.lent of all mosquitoes during the
h1ghlaods summer.16· 50• 74 • Q5 86 · 113 It has been collected in rnuch
smaller numbers in other parts of nonhem KwaZulu-Natal
at river margins or in marshes after rain, although it can ap·
a Reported as Cx 1C11x JunMuarus
parently become quite prel'alent :it Lake St Lucia. 17 This is a
b Reported as Ae /Neo.J lmea1opennis
C Probably All /Neo) luridus floodwater species which o,1posits at the edge of rivers a11d
d Could be Ae c;;callus s.s or A.e. 1upp1 pans 1,ithin the flood plain. ,\edine egg1i are resistant to

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 Vl>etors: ~lcsqultoes 143

Table 6.4 Resulrs oh~:or competence ,es:s v,,t11 Rift Vallev iS\er v,rus and wee mos~u to sl)EOes: mtec.,va feeds M \Maem·c hamsters but
transmission by the in vwo capillary metiloo!;
S?ECIES ITTIH' Of 111/FECTIVE MOSQUITO INFECTION TRANSMISSION
FEED N
LOG 0Ci'D~
PFU/mJ< OA"SAPER RATE' DAYS AFTER RAT:b
INFECTIVE MEAL INFECTIVEMEAL
:,a INeo / unidenunus 6.8-i.8 "2/1! 186%1 11 71"2 (53">)
J.e /Adm. demat<1s 6.8-7.6 11 2Si3\ 1gO%l 11 5'28132%)
17 6/8175%1 18 3/6150%)
~ /C1Jx.J poiciltpes 7.8 15 2s1zg ISO%! 16 412605%)
30 24/30 {SOo/ol

f Numerator ,. no. mosqunoes infected oenom,nator,. no mo~u,toes :ested

b Numerator" no moSQ;J:toes transm,ning. denominator= no niecied meisqu,toes feed,ng
c Cr,:opathic dose tCf'iJ~ Plaque fanning umts 1P'UJ

Table6.5 Results of vector competE!flce tests ·:mn various mcsQw:r, spec,es and Wes~:sbron wus

SPECIES VIRAEMIC SLOODMEAl INFECTION SUSCEPTl!!LE TRANSMISSIOfll ilEFERENCE

INFECT1NG TrTRE RATE HOST1S ATTEMPTS
HOST LOG,0LD5Jml
Ae /Neo / c,rcvmlulsolus Arthc,al 72 7/9 Mice 16/18' 37
Ae (Och.I caba/luss I. Mosqurtoes 1/1 1 ,1
I collecred 1r.
epidem,c
NT lambs

Ae /Octi.J C1Jb/Jll~s s I. Mosqu,toei; "1T M,cc 1/3i 1 57

collected in
ep1dam,c
Ae /Ocn.J ca/;allus s I. lambs Ni m Mite 212• 57
~ {Ct11.110mbaens,s Hamsters a.e 13126 Hamste•s 1/2' 8
C,, {Cux Jtheileri Hamsters 6,7-72 21/52 Mice 0/15 8

a Numeiator = no. successful transrn,ss,or.s. Jenorn,nator = no anemp,s wnh gr01;c;s oi 11osqut1oes

o T:,msm1ss1on ra;es
NT Not tested

desiccation as long as R high level of humidity is maimai11cd southern Africa.'·.; He named two new species, .-le. w,idcnra-
wilhin their mlcrohabitat under Lhe ~oil where, like Ae. ms and le. luridus. making a t<Jlal of eight species including
Juppl, they are probably deposited wilhin the top 50 uun.21 !le. circ11m/111eo/11.~ and Ae. linearape1111is. Prior to this re\i·
They do nor hatch until they are inunda1ed ac the next river sion Ae. rmickmaws may ha\·e been calledAe. liwwwpennis
flooding. Local rainfall apparemtr hai 10 be heavy and con· in Johannesburg and at Lake Chrissie for ex'ample.< 8 while
ccmrated lo cause surface flooding similar to that asso- Ae. luridus may have been confused with Ae. al/Jorhomx. In
ciated wich river flooding:50 19851 luang described a new species, Ae. (Seo.) mcimoshi.
,\ study at Ndumu in KwaZulu-~atal showed that during and shom~d that African mosquitoes previously reported as
the dry winter and spring momhs (June to October) Ae. cir- Ae. li11ecuope1•11is really belonged lo this spedes.31 Aetles /in·
<wnlureolus declined to veiy low numbers. bm it never emopPm1is it~elf was shown to be confu1ed to the Philip·
completel}' disappeared. Blood-feeding and ovarian devel- pines. Thus, reporis of Ae. /ill(:atope1111is in the literature
opment occurred throughout the year.;o Ae.(/es circumluli>o· after 1971 refer 10 Ae. mcimoshi bUt before 1971 c-0utd have
/us is a ground feeder reeding during the day and also at been e ither Ae. 1mldenrn11u or Ae. mcimoshl.
night from about half an hour after sunset. It feeds avidly on Aede-1 1111ide111ar11s is probably more common in the
cattle. sheep. antelopes and humans. 50 86• 90 moister eastern part of the High veld and lhe mjddle veld of
the Free State Prnvince. where it is probably the numerically
Aedes (Neo.) unidenta nis and Aedes (Neo.) 111cin tosh i dominant species of the subgenus.7'1 Aed~ mcilrtoshi I$ a
In 1971 McIntosh re,ised tl1csubgenus \'eomela11lco11ion in more widely distributed species adapted 10 regions \\ith

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144 ,r=r. ost: Mpccis influencing lhe llCcurrencc of infectious diseases

markedly different climates: it occurs in the Karoo. High· found in similar habitats 10 those of the subgenus Neomela-
veld, Limpopo-Mpumalanga Lo\\veld, and the KwaZulu- 11ico11ion, and often species belonging 10 both subgenera are
:'\atal coastal lowlands ofSoulh Africa. and in the Zimbabwe collected coge1hcr at 1he sameaqua1lc site on 1he inland pla-
highlands. In the nonhern part of lhe KwaZulu-:-.:atal low- teau. Rec:1?nt work at BuJtfontein in I.he eastern Highveld has
lands this species appears lo be dominated by Ae. ci'rc11111/u. shown that in the ca,e of eggs of At. juppi and t\e. mcimoslri
reolu.s. Tts prevalence rela1lve to ..ie. 1mfdemarus and Ae. in pans, the development cycle rrom egg 10 adul1 can be as
lurid11s in the Highveld and Karoo respectively is 1101 ye1 short as five days after inundarion:•~. H
properly kno\\11 because of insufficiem sampling. Large
numbers of Av. 111ci11toslli ha\'c recently been collected .u Viral maintenance and occurrence of epidemics
Villiers in the nonh-eastem Hlghveld. where it was about as
common as Ae. u11id11ntat11s. In southern Africa. the distri· Many years of work in southern Africa and in other African
bution of Ac.>. mcimoshi indicate~ that it is probably best countries have failed 10 provide evidence supporting a con-
adapted 10 regions ofinrermedla1e 1~rnperature. 1lnuous cryptic venebra1e-mosqui10 virus transmission
Aedes 1111idenratus feeds both by day and night. with peak cycle chat serves to maintain RVT-Y during the inter-epi-
biting commencing abom half an hour after sunset in Jo- demic period. Evidence has, however. been ob1ained In
harmesburg. It feeds voraciously on goats, caule. sheep and sou I hem Africa and Egypt that transmission by various
humans. 1• 48• 74 Aedes ,nci111oshi has the same host prefer- mosquito species, notably Cu/ex spp., occurs during epi·
ences but its peak biting in the eastern Highveld at Buluon- demics in sheep and cattle.a· r 89· "2 Virus isolation at·
tein in the Free S1ate Province was from late anernoon umil 1empts over 20 years from some 40 000 mosquitoes trapped
sunset, wilh a lesser pe;ikat da\,11. 16 in R\'1: areas ia KwaZulu-Nam! during the inter-epidemic
Larvae of Ae. rmldenraws ha\'e been collec1cd near Jo- period failed 10 incriminate mosquitoes.2
hannesburg in temporary rainwater pools in flooded pas- Cu/ex rheileri and C,:. zombne11sis are the important
rure.34 Both Ae. unidemams and .4e. mcimoshi larvae occur species on the inland plateau and KwaZulu-!\a1al coastal
in large numbers in flooded pans and. 10 a lesser ex1em. in lowlands respectively. There ls strong evidence chat Lhese
vleis In South Africa: species transmil RVFV biologically between sheep and carde.
and that mechanical transmission by several mosquito spe-
i\edes (Och .) juppi and Aedes (Och.) cabaflus cies and other biting flies is responsible for the mnintenai1ce
~ !cl ntosh made a taxonomic reasS8$Sment ofAedes COchlero· oflheepidemic. Humans are also thought to become infected
rams) ca/Jal/us and tound that mosquitoes that had been on occasion through transmission by Cr. rheilerl and C-1:.
identified as this species before 1973 could have been eilher zo111bae11sis, although some human infections during epi-
Ae. cabal/us s.s. or a new species, Ae. juppi. or a mixture of demics are undoubtedly caused by contagion during the
both these species.16 From the limited number of locality slaugh1elingofinfecwd animab.
records available for lhes-e rwo specles 7• ii appears that Ae. In South Africa. uanso\'l!fial 1ra.nsmisl>ion tc.s1s and 1est-
juppi is adapted 10 temperate conditions and is endemic to ing of mosquiloes reared from field-collected egg~ ha\'e
the temperate regions of southern Africa. It probably out· been undertaken to try to incriminate \'ariou~ lloodwater
numbers Ae. cabnllus in the High\'eld, and seems to be the Al!<le.., spp. as reservoir vectors of RVF\'.21 · 45 • s<J Such studies
only-species in I.he western and coastal regions of me\ \'es1em ha\•e so far been directed mainly at A juppi and have indi-
('.ape Province. It is sympauic with Ae. cah(l/11,s over a large cated that tl1is species has a low vector potential, that it is
are-a, including the Karon and southern Free State, bu1 the only a subsidiary epidemic vec1or, and that it Is no1 invoh'ed
relarive proponions of the rwo species require further eluci- in venical transmission of the virus.
dation. Aedes cabal/us is apparently adapted 10 higher The pauci1y of isolations from floodwater Aedl!$ during
temperatures and a lower rainfall than Ae. J11ppi and is epidemics is panly because sampling ofrhe mosquito popu-
absem from tile coastal regions of l<waZuJu-l\a1al and lations for virus assay invariably began late in the course of
Mozambiqut.'. an epidemic. Despite this, isolations have been made from
Aedes juppi and Ae. catin/111~ have very similar ecologies. Ae. mcimoshi and Ae. juppi on the inland plateau and from
Bodi feed avidly in the light and just before and after sunrise Ae. circum/Uieolus 011 the KwaZulu-:'\atal coastal loll'lands.
on goats. sheep, cattle and humans. ' 3 • 53 • 8'.l In the eastern If one or more of these spedcs should be responsible ior
Highveld Ae. Juppi has hs biting peak from laie afternoon transo\'arial transmission in the way lhat Ae. mcimo~lli ap-
until sunset "i1h a lesser peak at dawn. 16 L.arvac of tie. cabal- parently is in Kenya (~ee funher onJ, such in(ected mosqui-
/us s.l. have been recorded from small or large temporary toes probably occur only in a few endemic foci where RVFV
pools or pans.9~ More recent studies have sh0\\11 that the erupts to trigger a new epidemic in sheep and cattle when
eggs of botl1 species are laid in sedge or grassland within climatic condi1ions are favourable. Figure 6.4 summarizes
low-lying depressions (pans} and ha1ch co produce enor- 1he known e1>idemic cycle and 1he hypotl1etical maime-
mous numbers of larvae when these pans fill after heavy nance cycle for RVF\' in South Africa. Theoretically it might
rainfaU. at. ·12 Hence. larvae of lhe subgenus Ochlerocams are also be possible for some infected Cu/ex spp. to over-

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 Vcx:10r$: :'>IO$qultocs 145

Comagious route

foii!emic cyr.le

\\'inter and inltia1e transmission the nc.xt spring, especially On the temperate South African plateau rhc Ae. caballus-
as small numbers of C,;. 1/11!ileri adults have been collected juppi and Ae. mci111os/1i-lllridus-t111idema111s group of mos-
in 1he \,inter in Joham,esburg.3.~. J« quitoes are vectors. Along the tropical. eastern coastal plain
Recently RVFV has emerged asa small outbreak in African of northern KwaZulu-Natal and Mozambique, lie. circ11111lu-
buffalo (Sy11cer11s cafferJ a1 SkukU7.a in the Kruger :\ational teolus is the most likely vector. Because the tropical areas of
Park. Mpumalanga,23 and the reuospective testing of buffalo northern KwaZulu-Natal and :-.'IOzambique receive a higher
sera rrom this game reserve has indica1ed rhe presence of the rainfall. and are therefore moister than the drier temperate
\in.is in mis area at times when no epidemics were occurring areas of Somh Africa. Aede-;5 spp. are probably active there
in livestock in South .-\frica. 30 This sugges1s that RVF remains over longer periods. This would account for the higher level
ende1J1ic in the resen·c over extended periods. of\ \"SL\" acthirr in the tropical areas.~;
Antibody sm...,eys in humans and livestock ha\'e shown The only evidence for the: possible existence or a \\1ild ver-
that WSI.V is more comm<>n in the moist tropical part of tebrate host for \ VSLV is the single isolation of \irus from the
somhero Africa - nor1hern coasral KwaZulu-Natal. gerbil (Desmcdillus nuricu/aws).0 Howe\'er, the absence of
\lozambique and :'\amibla - than or: the inland plateau. CrtJc>.,; ,·cccors suggest& that viral maintenance would be
the southern K\\·aZulu-Natal lowlands and the Cape tluough transovarial transmission from one generation of
provinces of Sourh Africa.58· 59• so. 93· 105 noodwater Aedes 10 the ne:-:t by vertical transmission, the

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146 ;t~,,o~ c,~,: A$pec1s influencing the occurrence oflnfcc1iou$ di~cas<'S

Eo:demic cycle Contagious •oute

Viral ma'n,enance
cvvenscal
11ansm1ss1on?
/

Figure 6.5 Transm1ss,cn cycles of

WesselsbrM virus ,n soot~e.n
Africa: maimenance of virus
;~rough 1he dry w,mei 1s
r.ypothetically bv means of vertical .....
[,ransovariall vansmission by . ;.,
Aedes tNeomefamconion) or Aedes
tOchlerotatus) mosquito sl)ecies

virus surl'iving dry periods in the eggs or these mosquitoes. first developing in the a\1an popula1ion. Tho:> exposure of
Only one attempt has so far been made to show !he presence sentinel pigeo11s at Belhulie in the FreeS1ate PrO\~nce indi·
or infected floodwater Aedes mosquitoes among 1hose cated 1he occurrence of such epidemics in April l 970 and
emerging from an inundated pan.47 This was unsuccessful. April 1971.82
probabl~r because insuffic:ient mosquirnes were tested. Serological surveys for ncurrallzing antibodies in live-
Domestic ungulates are important incidental hosts with stock. ha,·e shown a pre\·alence of 16 per cent for w;,-y in
\\'hich the vector could probably maintain the \irus for quite sheep and cattle on the Highveld generally. and rates of21 10
long periods in 1he absence ofa primary vertebrate mainte· 47 per cent in older canle, oxen and horse$ in Johannes-
nance hos,. Livestock, particular!)· adult sheep and cattle, burg.61· 8~ It appears. therefore. thac lh·estock .ilso become in-
are of1en inapparently infected. Humans can be infected fected 1,1t11 WN\' in the field. although calves show only a
either by contagion through handling infected carcasses or trace of viraemia afcer inoculation 1,i1h \'irus.3 Such lnfec·
by mosquito bile (Figure 6,5). tions mighl be expec;tcd sometimes to lead 10 1he de,·elop-
During each summer on the inland pla1eau of S0u1h ment of abnormalities in rhe foetuses of pregnam animais. 12
Africa, WN and SIN vinises are maimained in a feral cycle Culex rubinows maintains BAi\!V among wild rodents.
between Cx. u11i11iua111s and various avian species. The That rodents ure indeed the feral vertebrate maintenance
same mosqui10. Ct. 1111iuirrarus. also transfers infection co hosts is dear from aspects of the studies referred 10 above.
humans.38• H . 48• 117• 83· 90 An outbreak of \\11\: fC\'Cr in the Addi1ionally. rhere is the iSo lation of \~rus from, and the
human population will depend upon an epidemic situation presence of antibodies in. Mastomys 11acale11sis.19 plus the

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 Vectors: \los11uft~ 147

,iraemic response of wild rodent<; w moculation with the ing meteorological sa1elli1es whereb)' data on green-leaf
H336 strain of the drus. 73 Thc oct·ucrence of C:r. m/Ji1Jo1t1s in vegetation d~11amics. ground moisture and rainfall panerns
other parts of Africa. with associaced viral isolations, sug- can be collec-.ed over wide areas of Kenya.66• 117 It ls hoped
ge~ts chat il is probably the vector of BA.'\V over a large part such data will enable predictions 10 be made as 10 when and
of the continent.24 • 95 where dambos (pans) would be flooded. This would allow
!>erological surveys in the Kwa%ulu-Na1al coa$1al IO\\· vecerlnary authorities time to vaccinate liw~ock before an
lands showed that neutralizing antibodies were present in outbreak ofRVF occurred.
canle at nlles of2 to 16 pen:ent and in sheep at racesofO to i
per ceni.5~ This e,idence. together with the moderate feed-
ing of C:r. rubinows on domes1ic ungulaces, sugg<'$1S that Sub-Saharan Africa, excluding
pregnant caule, for example. could become infected. par- southern Africa
1icularly if thC)' were 10 browse near the marshy recd beds
where C:Y. rul1i11otus is pre,'lllent. It would seem wonhwhile
Tntroduclion
10 examine foecaJ material arising from natural abortions on
1he KwaZulu-Natal coast for the presence of 1he 1iru, and JnAfrican countries omsidesouthern Africa and ~ou1h ofche
developmental abnormalities. \.\'hether bo,ine infections Sahara, RVF has periodically caused epidemics in lil'estock
everat1ain 1he level of an epidemic is 1101 known. Banzi vims in Kenya and in the Senegal River Basin of nonhem Senegal
has been isolated from humans and is known 10 cause at and sou1hern Mauritania. The mos1 recent ombreak In
least mild illness, io; but the extent to which it causes mor- Kenya was a very large epidemic which occurred both in rhe
bidity in hun1ans is uncertain. Garissa region of north-ease Kenya and in Somalia in 1998.
which subsequently spread 10 1he plains of the Greac Rift
Valley and norchern Tanzania. 11 These outbreaks have fol-
Mosquito control
lowed unusually neavy rainfalls excep1 in the Senegal River
From the point of view of comrolling R\'F. priorlmmuniza- Basin, where damming 1he ri,-er caused widespread flood-
tion of livestock would be more c:ost-effeccive than vector in1r During active sun•eillance for arbo\.'iruses in mosqui-
comrol. However, for other reasons mentioned abo,·c, it toes in Wes1 Africa and the Central African Rt'public.
may on occasion be ad,•isablc to institme some mosqufw multiple isolacions or \\'SLV have been made (Table 6.7) al-
comrol measures 10 reduce the degree of ex'µosure of stock though the mosquito species concerned have vet to be in-
to mosquito bites. Since the peak period for biting Is from criminaced in !ht' transmission ofWSL\' to cause disease in
just before sunset umil ~uorlsc. and because mosquitoes sheep or goats. Onl)• occasional isolation of\\'SLV have been
are always ac1ive at or close to their breeding sites. stock from mosquitoes i11 Uganda, 25 Kenyati and the Cam-
should. if possible. be moved away from pans and vlei~ 10 eroon101 that have not been associated with any livestoc"
higher ground before nigh1ra11. ln the case of dairy farms. disease.
the milking sheds shou ld be made mosquito proof. and Tho only other mosquito-borne ,1ms tha1 nppcars 10
perhap, Culicoides-proof, and monthly sprays or their in- have vecerinary importance is the Semliki Fores! Virus.
terior walls should be carried ou1 wich a residual insecti· which is an Alµ/u111irus(Toga\'iridae) that has been lhe cause
ddc other than an organochlorine. Organophosphace of encephalitis it1 horses in Senegal. 14
Insecticides, such as malnthion. fonthion (Baytexl. fcni-
trothion (Sumithlon), or carbomaces such as carbaryl
Virus isolatio ns from n1osguitoes and their
!Se,·in) and propoxur (Arp rocarb). can be used a1 rates or
vector competence
1,5 to 2.0 g/m 2 •104
Control of adult mosquitoes outdoors would probably be Table 6.6 !iscs the eigh1 mosquito species which yielded
neither effective nor economical. Omside comrol could. mulLip!e isolations or RVF\' toge.1her with the results or
however. be direcced at larvae in the main breeding pans. vector competence experiments where these have been un-
\'lei.~ and dams. Lanicldes such as Temcphos (Abate) can be dertaken. C1111'.x pipiem from F.gypt is included as !Is level of
incorporated inco sand granules or µellecs and sprinkled veccor importance appears 10 be similar to C.r. rlieileri in
imo the breeding pla.(:e In the case of pans, a dry pan could South Africa It is likel}' chat the epidemic \·ec1or involved in
be seeded with such p·e11e1s just before a nood is e.,1Jec1cd so 1hc 1978 Egyptian epidemic was Cx. pipi('J1s 1110/em,s- 1his
that u1c inscc1icide is released imo the water as the .tl!dine ~uusµi:<:ic.,, uoL uni) fted" on th e~tu<.:k. uu1 is abu hi,t.hl~ im-
eggs hatch. An alternative lanicide is the bacterium. Bacil- thropophi!ic, which could account for some of the human
lus rl111ri11gie11sis ismelemis, whosf' spores can be lncorpo- cases that occurred. The other species listed were either col-
iated into pcllers. This may become practicable ifits present lecced during epidemics ot' RVF or during imerepidemic
high cos1 is reduced. The emphasis should be on a control periods ln Kenya and Sen~al. 1\vo of lhe three species im-
programme with measures that are selective and integrated. plicated as vectors in Kenya, Ae. 111cillros/1i and C,:. zom-
Recently a technology was developed using polar-orbn- bae11sis. \\'ere the same as 1hose importanrin somhem Africa

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148 ,n=, o~,: Ai.pcct• lntluenting the cx:currenc<t of lrifectlou, disease;,

Table 6.6 Mosquito species from African countries ouiside soutnem Africa vielding multble .solauons of Riit Valley fever wus; number cf 1solatiQns
and results of transmission a~penments

COUNTRY SPECIES NO tSOLATlONS S!l"o INFECTION iRANSMISSION REFERENCc

-l{r!ESHOLO (DAYS
l06, 0LD15()1'm1 POST-INFECTIONI
Egypt CX. (Cui1,J p1p1ens 3 <6.5 -17-12• 27 S•
Kenya Ae (Nel).J mc,mosn, 10" <5.7 -171 15, 70. 100
Cx (Cux.J amenr.atus 3 <5.3 -(1-211 70. 110
Cx. (Cux.J lOm//aetJS,s ii ND ND 70,72
Senegal Ae. (Adm /daii,eh <I ND ND 20
Ae (.4dm.J fa/lier, C. 6.3 +(71 70, 10!1
Ae. (Adm./ ochraceLJs 3 ND ND 19
Ae. (Adm./ ve.~ers . 10 NO NO 115

a pools of !err.ale mosquitoes collectea d:ret!ly in tlie f:eld

6 pools of ~ d female mosqunoes
2 pools of reared male mosqu110es
All isolations from other spB(jes were from female mosqu,roes
ND Notdone

(Table 6.2). Of the l O isolations made from Ae. mcintoshl, Table 6. 7 Multiple isolalJCl'S of Wesselsbron viw; from moSQu,tnes
one was made during an epidemlc15 while the remainder collected in West Africa and the Central African Republic (CAR110
were made during an interepidem1c pcriod.70 In the la11er
COlJt-ffRY SP£C1E.S NL ISOL.\TIONS
study. eight of the nine isolations came from mosquitoes
reared from field-collected larYae lsec Table 6.6, foomote). IVO'\' Coas: CAA Ae /t.dm.J ebnormalisgrp 11
Cu/ex c111re11nc1ws. the remainin{,? species. has never been Ivor\· Coas:, Senegal Ae /Adm I daltleli i3
prevalent during outbreaks of R\'F In ~omhern Africa. Tht
vector competence rests done on Kenyan Ae. mcinroshi and
Senegal
lvorv Coas:. CAR
AE. (Adm I mlnu1us
Ae IA:Jm Jtarsa/,sgrp ,.7
Senega' CAR Ae /Aam Jv,uaws 3
CT. ame1111ancs showed they were both highly susceptible to lro"I Coas'.. CAR Ae /S1g./ aiflcanus 5
RVFV. The higher susceptibility of Kenyan over South Afri- Ivor, Coas'. Senegal Ae /Sig I /ute«ej)halus 3
can Ae. mcimoshi may indicate that the mosquitoes teswd CAR Ar: 1Srg I opok 3
from the t\'l'O different countries represent different specie~. Ives, Coas'. CAR C, ;Cvk) vertvscus 3
This would seem likely as memberi. ofrhe 1\'eomeUmico11io11
subgenus are morphologically very similar. ,\11 the isolations
of RVFV from Senegal were obtained from floodwater Aedes
mosquitoes belonging co the subgenus .4edimorphus
Ecology of mosquito vectors
(Adm.). J\edes vexmzs is apparently the most important of
tl1ese vectors, although none of the three species concerned Aed es CNeo.) 111ci11toshi
has l>een evaluated for vector competence. However, an- Recent studies on the ecology of Ae. mt:i11ll)$/ti near :-:airobi
other ,-1edimorph11S species. Ae. fowleri. has been shown 10 in the Kenya hishlands have indicated a behaviour panern
be susceptible and able to transmit RVFV. which ,;uggesti; similar ro that of South African populations. Larvae hnve
tha t other members of the subgenus could be shown to he been collected in very laf!le numllcrs in dambos (pans) on
labora lory vectors. bushed grassland.68 Adult females feed avidly on catlle and
,\!though \\'SLVoccurs in a number of sub-Saharan Afri· humans with peaks jus1 before sunset and just after sun-
can countries ourside sourhem Africa, only single isolations rise 69 -, The dispersal distance from a dambo wa~ found to
have been made in Kenya from a pool of Aedes (.4t/m.) de11- be 0.07 and 0,15 km for males and females respectively. and
u1tus mosquitoes95 and in Cameroon and Guinea from vari- tl1e dail) survival rate 8.82 and 0,85.6.'
ous Aedes spp.1 08 ~fuhiple isolations of the vinis havi:.
'hvwever. been obtained from nine dilTcrem mosquito spt:· Acdcs (Acdimorphus) spp.
cies collected in the Ivory Coast. Senegal and the Central /\J.- Lirde infom1ation has been published on the breeding places
rican Republic 10 (Table 6.71, Out of these 59 isolations, most of the Senegalese Aedimorp/1 rts lloodwacerAedes mosquitoes.
(45 isolaros) came from AedimDrf1h11s species and it is ahhough Zeller and co-workers state tha1 Ae. dalzieli oc-
thought that these noodwater mosquitoes parallel the curred in small ground pools locared in the nood plain of
Neo111elnnicl)11ion and Or/1/eroratus noodwruer .-ledes spp., temporary rivers. in the gallery forests. 11 ; In 1he US.A. , Ae.
which are the vectors in Sourh Africa. £11!.mns larvae also occu r in I he small ground pools as well as

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 Vec1ors: '.\losquitoe~ 149

in the larger bodie;. of waier fOm\ed in 1.he flood plain of riv- area,, are able to propagate Lhe virus in a mosquitO·lO·
ers.29 Apparently all three species of Aedimorph11S 1hat have sheep lcanle :ransmission cycle 10 maintain an epidemic.
been incriminated as ,,ectors are mosquitoes that breed in ln Senegal. IVes1 Africa. some of the virus isolations in-
temporary ground water. although whether they mHize the cluded in Table 6.6 - those from Ae. ochraceus and Ae.
dambo. or arc confined 10 small poob. i~ not known. They Vexans. both Aedimor,J//us species - were made when 110
feed on cattle and sheep 19 and their beha,1our is generally epidemic or R\'I' was in progress. This evidence. together
similar to that of the .Veo,n('/aniconion and Och/ero({/russub- with other e,1dence from serological ~tu dies in livestock led
genera including a very short development period from egg 10 Zeller and bis colleagues 1i; 10 conclude tha1 there was some
ad ult. Fomenille and co-workers observed that ,le. 1te.· mns de- endemic maimenancc of 1his virus in 1he Sahelian region
posited eggs on the soil or temporary ground pools in the dry and that these cwo Aedimorp/111s species should b!! regarded
Sahelian regions of Senegal and that adult mosquitoes as lmerepidcmic vectors. The) considered that these spe-
appeared only four days after che rain." cies fulfil this role In West ,\frica. which the ,Veo111e/a11ico-
11io11••~e. Mt:imoslii. fulfils in Kenya. They poimed out.
howe\·er, that while epidemics in Eas1 ,\!rica are asrnciated
Viral maintenance and occurrence of epidemics
with increased rainfall, Lhls same relationship has 1101 been
As mentioned above. RVF virus has been Isolated from mos- established in Lhe Sahellan areas. 115 For example. the RVf
qui1oes collecced in Ken)'ll during an imerepidemlc pe- activity described in southern Mauritania in 1982 to 1985
riod.;o These included eight isolarfons from adull Ae. occurred durmg a period of drought. and the 1993 epidemic
mci1110#1i (reported as Ae. liT1Mtope1111ls) that had been col- was not associated with e:mins!ve rai11faJJ. 103· 11-1 It is note-
lected as larvae or pupae reared co adulthood in the labora- worthy that z.eller er al. 11 • made no isolations from Cu/ex
tory. These immatures came from ne"·Jy nooded dambos in spp. during the imereptdemic period as it stren1,<thens the
bushed grassland near :-:airobi. The only oLher isolaiions concl~ion 1ha1 C\'. rllei/eri in South Africa and Cx. p1pie11s in
from aedine mosquitoes were from wild-caught Ae. cum- Egypt are on.ly epidemic vectors and probably do not haHi a
minsi adults and another from wild-caught ,le. mci11IO$hi viral maintenance function.
adulc.s. This important finding pro,ides eviden~e that RVF\' There have been no field studies on \\'SL\' outside south-
tan be maimained 1hroughout imerepldemic periods i11 a ern Africa that Implicate particularly mosqulto vectors in the
state of dormancy in the eggs ofile. mcimoshiand can 1hen maimenanceo(ihc virus or in rhe transmission of virus dur-
be transrnilled Lrat1S<>varially Lo the nex1 genenulon of mo,- ing epidemics. I lowever. it i& lik.ely that the Aedi111orp/111s
quicoes. Aedes rncimoshi ls therefore considered to be im- flood,,·acer At•de< contribute towards maintenance of WSLV
portant in the generation of epidemics b> seeding prim30• by transo\"arial tran.smiss,on. Experiments need to be car-
,irus in caule and sheep and susceptible \\'ildlife species. ried out \\ith Aedimorphus species. ,;uch ~ Ae. dal::ieli. 10
Subsequemly. ocher mosquitoes. Cu/ex and Anopheles spp .. see whether such transmission can be demonstrated in the
colonizing Lemporary and pcnnanem water in che ..-pidemic laboratory.

References
-'~DDWS. n .. 196i. lico1og,.cal Mudu...")on Sindbi.> and w,.-.s1 ~lie vlmlit'i 10 .,-...o, :t98.i" \mum/ ll••p<1r, oft1r~ Ptt$frttr/11.s1itmq, P.O. Rox 220, Dak.1.r
m South \friO&, IU, Ho~i ;>:cfercnc.'4:~ r>f mosqunoc~ .is dt!termint"d b~· s~nei:'11
1hc pret'lpmn ttSt, .')(Jw/J -tfr1(a11 JournoJ of ,\J,•tlrtal ~ic1tt1'S, 3~. 3-1-J:9. u ~"" 1998. An 0111hreal.<11 ltJi1 \i1ll~rf=rln Eos,~m.-\irlco, 1997~19!>8.
2: ' ·"o°", \Sj.3-lg.81. Ari>o,,ru~ Unit. ~at1onal ln~tUH: for\-irotogy, 11,,·kl.1· Er1/1f,mlnl11girnl Jkmrd 13, 10.>-112.
lohanne.lmrg. lJnpubH,h,dda1a. 1~ a~,,n.,,.~.J.1f. "voc;rs. .r..r l9H6, Flu,;\iru~ in South .\i'rlen:
.,~()\, ,~au. Arbov1ru~ Un\c. "\ational Jn,.,1irute for, frol<11,,")', PouhogcmcHy !or \;hL-cp. Oud.usu•pocm Journal of\..rtl!rmary Rt$<'<lfflt •
lt>hnnn••hurg. Unpubhsbed dlit• 53.1.U-238.
l..l st.\Gmua.~ x.~. 1,, w,,r\.,U"Ol.L "-· tg,So. An inn~strgat-lo'n of fln,i,'fn.r-.
,1 ·'"n~. 1~,G-1v88..-.\rbo,~n.1, l,h1i1, ~:ition;tl ln..;lluu.• for \lirolnb') ,
[nf,-.,1ions ofcnute In Zlmhahw,• Rhndei!o wllh pa!llcul"f r~!o1c11c" tt>
lohan11c,hurg. Unpublished data. \\cs,.eb,brnn -,rus, Joumnl QJ J/_1-gt,iie, l\:., 1-3.1,
.; >.~u:,:, 1!t(i;, ArhmimJ Unit, ~t'ttiomd ln~titut.: for \1rolcg:.·. u BR-1:-s p,_, 1 ~ lmp{lt:t of ,1rbcwlni1;rs on human and iimmnl lll:nhh, hr:
Johannc..,buri;. Unpubll,hed unrn. :..101,·Anf, 1.,.. (td1l, f lw Art,r,1:intSt!s.: Rvld,,,,1,ofotJ· ,md l:tofogy. Vol 1
El A.XO'\:, Hi68.Arboviru."'" Umt. '\'-atlono.l lnsdtutt.'I for\lrulogy. Bot11 koton, 'loridu; CltC l'rt». pp. 1-18.

Joh(u>nL"burt, Unpubli,hcd data. 1.s o,\\-U:S, r:.c: . .& 111t:.1no, M.n.. 191\r,. Possible \'ttton: of Rift \'!IIILi (ever in
KCn}'Q Trnm1.1crim1.{ uf rh,· Rc>yrll s«icl)·o[Tt'Op!Ml .,t1t't1-irlrl,1 a11tl
; •:-o:-. l91i'!. Arbo,'iru, lln\l, :-otlonol rnstl:uto for 1·,rotoll!'·
H.,-gi,•11e ;4_ 815-316
JohannL-.bu~ Unpttblisht-d ®"'·
16 or >.stJt.LO~
•·· 1~~n H.,o,, H f,4- ,tu$M\a,n. t. tgs; s,udi~on the
a A.XO:-., 19r1.Arbovi.ru., Unit. ~a:t1onat ln,-U<utc forV1rolo~·. mhropod·bom•· eiru,,..., ofTt>nfl"lantl. II. Not.,. on lh< nio,~ common
lohonnesbwg, tJnpublisl1cd data, mo,qulloc,,, SQmh ;frr«m /011ma/ o)'Mf'ff/"11 Sd"17r.,;. ?l, ,;;-s3.
, ,w:<. 196'1.Arbo,iru, Unit. Na11omll lnru,utc !or \1rtil<,gy. 17 198,.1'otion'11 lnni:u<cforTmp1l""1 Oiknse.. f:show~.
l'.ClWtD. 1>.\I
luhllnno,bu'l\, Unpublished data, l'erson:tJ coinmunJc::ilfon.

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150 SOCl1<l'"·'c ,\Spcctsinfluencing the occurrcnt<! of infectious diseases

18 G.CK.\RQ. 0,"M, \\OJrnn.NGTOS, v. ~ JUl'J\ P.C: .. 1988. ~1M(JUilO S!)tcie~ 38 tuPP. r.c., 197,1, t.abonuory StudiC$cOn th~ tnmsmission o!W<is1 Nile
collec1ed •• Rich.um ll3\\ Natnl. fo,m,n/ ti/ti:~ £11111mol1J$MLS<JC1uty of _,ru, b)' Clllll• !C1<fexl 1111Mrro11,sl11cobald: f:lcrnrs inJluonclng the
Scu1/:cr11 ,1[rfar, Sl, 140-1-13. 1ronsmtS.Slon rare. Jou11wl of.Wedie(,/ Emomolog,i. 11. ·155-458.
19 t:Q.Nll!SIU.li, 0, TR.\QJUM,.\~tT7.,1".:,\ ;',t,, 01\LI.CI, )l,, 'TllQ?l.;'\Q~, J., DlC"~UTIF, as tUi'P~ i>.G.. 197,&.. Funhcr srudies Qn 1.he ovemimenng 5tngcs of Cul,~t
1,P;f, nu.ER, H.(:., 1998, :'l:c,,· \C('lQ~ or f.tft Valfc}rfo\'Cf m We~1.Afr1(a mosqulwes (Oiprrr:,: Culicidacl 1n die l llgh,c!d r.-gion orsou1h Africa
Emt'ti:111f/ /11/1tC1101,s DI.sea...,·. •I, 389-393. Joum(I/ qfllw w11omotog1((IISaclnyofS0ull1m: :\frlm. 38. 89-97•
2() l'-0:ff'l!NIUE. D .. TR.i\ORl!·LAMJ;_,"_.,\:,;t. M .• 1.nu-s. n •• M0.'~00. )!.. l)l.\UO. ~.,. ~ .io rurr, r.e .. 19;&. Hit ,u..:<pllbill1y or four South :\fri<at\ •Jl"cle. of C11t,-x
<>ICUUTr>, /,P., lU9!1- Shon
6cpon: Rift \".Uey (c,'nr In Wos1cm ,\frlCJI: 10 \\'ccs1 :'\Uc ond Sindbls ,irusos b) "'" dJ/!mmt Infecting methods.
isoln1io11> from ,lwfr. mosqu1coes Junng an ln1ercpi,001lc pcnod. \fqsq11itoNet11$, 36, 166-173.
,1111,tlca11 Jqumol o/Trnpieol M.,lfcfne and HJ1!irn,. 52. 103-l04.
.:1 1uv11• 11.t;._. 1983, ~ath:m:tt tn,tjrtH<" forVtro!ogy, JQh.snnc.'!-tburg
:!l C'.AllCA:o-l, T...P., JUP.P, P.C. &!\"0\IM:, R.J., 198&. P.an\'f!ld at.: O\'ipt,,illon .SilL"\ of Un11ubfl•hed do1a.
Ooodwmor ..i,,ttr, moS(lulloes amr ,m~mpis co show 11-anso,·•rfal
,42 ,us-1•, 1•.t,, 19a1•.~iti<:,r.,,1 ln~titute for Vim1Cl8), Jr1h1:mnC!-sburg,
troo,mls-;iun nf Rift Vallt)• fcwr ,in» In Snmh Mrlca. Medlr.o/ n,:(I
UnpubU,hed dala.
\fererfllnry Emomology, 2, 231- 236.
43 1u11P. P,r,., 19,,88. \tntional lt\~druu· forVi rc,lo~·, Joh1.1nncsbutg.
,.r. & ..:ors):Y. n.. 19;1,,;.. Rm \"al.le..'}·
22 GEAR. ,,1-1.'$•• ot )!EJ.UO:'\. a .• u. noux.
llnpublbhod data.
£over in Soulh t\frl~ a study on the 1953 ou1br,ak In !ho Orange Free
S1nte with special Tt'fertmct: lo the \'~cu)rs and ptil.:5ibll' rt'.M!rW,it ho~t'!I.. J.I JUrr. P,G,. 111,,c..-t1uJt:1o, :,,;·,;.: .. nt0MP$0S, o ..l,. ¥r )U-E.'\:8HAX, r;.e,i,s •• l986.
South 1\frlcn,r .\lltdlcnlJo11r1111/, 29, SM-518. Shtdbfs a.n1..l \\'es1 .~n~ vi~ fnf1.-ctio1\s In the\\ 1r.,·o.tt:t:SrnodtPrc,orh1:
tf!lion.Sout/1 ,\frirrm .\1/!diMIJoumfll. 70. 218-220.
:tl GROllRnlMR, o.c. ~ M~"CI$, tt•• i999. National Pnrks Bonni .a.nd
ilepnrimont 01 Veterinary Services. Krui<t :-lotl<inal Park, S~uku,.., 13.0. 4& ,orP.. r-.c. 1, i.OR!lof.J... .\..,., 1988. \'C(;101 l.!•Jml)l:ttnct t~u w1ih Rift \'~le}
UnpllbHshcd data. tC\·er \1'."U.$ -ind fh-e South ,\Jrican spec1~ of mosqunocs./auma/ of lhi•
tmerlctm M,,sr1m10 Ctuurol.r\sst>eft.u ltm, 4, .i-a•
..24 umollll.SON, e.c.~ MCCRAC.A,.W,R., ):.IA)'A, t .G.. S°l;.~ii:\JIJ\IGI:.. \:, -0.::~l':.\lP:U.I\,
s.o. ..: .. 197:!. Arbo\iru.s tpizQOtits 1nvohm~ m3n, musquhOC$Gnd :i6 Jua•P, i•,,, . "Xli)1r, .\. 19!)8,. Sn:dft"~ on nn uurbn:.s.k c>l WCM<:l.sbron \·!cu.,
\'Or1cbrates •• l.un)'O, Uganda 196$.. \mmls QJ 7'roplm/ Medlcw~o)ld m lhl' freeS\nt~ Pro,incc. ::Wtuh Africa. Jo«mal ofthtt\merf<nn
Pomsltology, 66. 3"3-3S5. .Wo;qulro Courro/ ,tssoc,'01/0t1, 1·l. 40-15
::S H(~'iJb8$U~. ll.f·:., TUJ.::1:.:1, V,M,, LUU~ >J., \\'! ST, R, ,11 ..MUJO~IU.Uo, L, tS67, ~; 1u••· M•, "~l!.\lP, .,.. 1998. '/utlon:U lnMltuto tor, lrology. Sandringham
.,\rOQ,iiru~ iclentific:atfan smd.ics: l~::,1:ulcms from mosquitoes, W!-t 2131. South ,\fric,,. Unpublkh~d da13,
Afn'can Virus Res:Mrclt fo.~rlwtt Rt1JKJrt. 1; 22. .;A JUl'J•, P..C. & Mc;1'·ro..<H, a.11 .. 1967. l~olo.gical ,tudle.. Cln Si::ulbi:- und \\'t·)t
26 H()Cll,A,L., M1tc.,:<. ,.r.,. 11.-ULE>', C.L, 19e;. Mcch:u\lcal 1ran,ml»lun (l[ :,(l!ntruses In South Mrlca II, ).Josquho blonomle> So111/1 tl[rfc"''
lllft V11lll'y fcw"iru, b)' hcma,ophagou, D1ptern.,1nwrlm11 Journal of Jc11m:al o/M,vflc11/ $€/e,,c,.,, 32. 1:;.aa.
Tropicnl Med/clr,~11nd lln,~•·11•. 34, 183-1~3. ~9 ,UE>P. P.G. at.)JCIVTOSU, D.:\I., 19i3. National Irutf[Ut~ for Vlrology.
:r.' lc(')()GSTRA,\L, tt., )1£(CA:,:, 1,M .. Ktt\UL. G.~~. &. ADI1A~1, F.J!., 19;9. lht> Rift lohann»burg. Unpubll<h•d d~m
Vlllloy rev~,tJ>i>ootit in F.gi1H 19n-78. 2. F."colngiml nnd tntornologlc.,I :;o SUPP, P.c;. tr MCISTOSU. a. .:i,L. l98i',J\ bionon\iC SlUci)' of udull AW~"
srndll)S. Tto.,wm/011; of1/1~ 110)11/ :,oc/e1yofTro1,icol Ml!(lfclnumd
l.V,"011u:lun,(1J1Jlon) ('fr~umlutL'Clusln northl•tn Kw:,zulu, South 1\frfca.
Hygl,11,. ;J. f,24-629. Joumnlo/1Ja1tlmrrkn11 ,\!u.t.quieo r..on1rt,l:UJotliUitm. 3. l3J-1.3.6.
:,8 lfOl'J.aN'S. C.11.£,. 19~ .wost/UitOf!.S()flhi! J:1l1iOpim1 Htg{OIL I. l.tJn,nl
51 suvP, P.c.., 1'tc:::i~TostL tt:i.1. t.A~t>F..R..'iOl\', o .. :976. Cula rErm1elt11mmJ,'it11
IJJon()mfcs ofmo$tfulrr,c$ n,uJ uixonom,·ofcull",ur lnrt.101.t. 2nd edn
rriilf110111s ·t11eob.ild as vector of Bami. Germ,ston Md\I 11wu1ermmd
Londo,~ llritl.sh ~luseum {;'1amml IHstot}'l.
,ire~~ !V. O.bst:JVationson thebio1ogyof C. n1Ui1ww.i... Journat a/
)9 UORSFAU.. W.R., FO\\'LEft, tn. U.\1,'., MORETTI, J..J, & TAU$£.~, J,R... IS73, ,\/~t/1(111 li'llll)I/IOWl.(l·, 12. 647~I.
Biouomirs 011!1 F.mbT)'Ologyoftlre /nlm:d Floodwarer Mo;quW,M><lcs s2 1u1•P. P1r.o., J.·a;1~qo~1. -.:-.t 1c 81-At~un..... -....K.. 19Ut>. E."<p,er,.menW
vexans. Urbana. Chlc•gu. t,)ndon: Unh·e:$11)' of llhnols Press. ;l)Se~5mcm <,r,h~ve<aor,:ompc:,cm:c of (:ulcx(Cul£wJ nea:1tl(l'(Tht(l~fd
30 nQwm.i.. P.G.. l999- Faculty of Veterltu,lryScicm<"e. Uniwn.ity of Prc1oria. \\ith West Nile and :)indbh, ,iru1i.t$ in :,omh J\!nc.a. Tfnnsactltmz oj ch.~
Onilcrstcpoon 0110, Un.pubbs)led data. Royal Sx/cryoJ Trc,p/col .\frditi!M and H;,'tiicn~. 80, 226-230.
31 nu.\~G. v.,r., 1985. A n,·w African ,pe~,s <>f lletlt< (O!p1cra: C\llkldnel. ~ JTJPP. r>.G.. '1Cl::\'TOSll, n.:u. ~ ~-m·1u. U t .., 1980.A survey of the mosq\lito
""-"U/fl(M .\)'$lrmntlr.<. 17, WA-12n and cu,,c,nd,,s f:aun.\.. at C\\'O foc::i:dit u:i In the ~:ttOO ,c~on of Sou:h'
32. JOuerk'1. L, t,9~,> L':irbovlrose \\'Ml :'\'-flr• .t6ono:i:~ du midl ..\.fnt{I \·,llh ~)me obst:nauon.~ on b1onoinics_ (J11<ter~1q><~fl Jm,nml <,f
ml!dlterrnnecn de la Fmnc~. Bu/l~lin dtl:~rt1dOmirNmlnnalt1,I~ Vfu·ri1utr:,• HCSJ.Ytrth . .;7. 1-4'.>.
MMlt/m>, 159, 495'·503. 54 JUPP. P,e;.,. Mn,"fm.H. u..,,~ .. a. TtlO~U-$0S; l'-t..• 1983, f5ol&tfon Of Rift
33 JOOn(RT. L. OUOOR.. , .. 11!\NXO\'~. c.. IU:.HOUl', lJ•• Ct>JiSIOIJ. ti.. auru.o.,;,, \ .:UIC) foxcr virus from.4ec/u.,; tSr!Qm~loniron;Qn) nrc11mlmeohts
J.C.. tt PASTiltEt(. n .. 19iO. Epid€1niotoqic. du v,ru,: West ~Uc: <itudc <fun andior lmt!D!atera!is coUt>cttd during nn outbreak fn cattle in the
fo)~r en Cnn1a."gll.e. 1,·. La menin~o-ei1c~ph:tlon1ydU1edu che-·.i. coastal region .,f:-3caJ, South Africn. So111/1 AfricnnJ011rnnl ojScie11ro.
..J.Jmol& lnstlml~ P~lst11ur. t 18. 23g...~7 ;9. 3.;;,
3.: JUPP, n.c-., 196;. UU'\1\lh3bi1n.t s of culkinct mo:squimru,; tDiptMtl! 55 1u1 1r. P.G .• )1<:Jllo'tt.i:SH, Jf,M • .e.. fflQMPSC'I:<, u.t.., 1964.~\ie('hanit"lll
Culicidaol inn se\\~geemuem disposal :th!~~~ me Souti1 Afr!Cl>II tran...m.ssion orRlh \'alle;- r.evc:c ,,ru~ b)· mo>qultoc~. Sm,th Afrlcm1
!ilghveld.Joun"'' of,,,,
E111omologfcal Sorwtyo/S0111h .-\fffro.. !IO. /Ollh!(I/ o/Sc/q11c,. 80, ?:6.
242-250. ii~ );O,a:n..,vT. 1\.11., l l!T!.t..\:'\S, c....,~. MUS:PR.\TT. ,. ~ WQU~HOL\t!I, s .... 1937.
3S 1uv,•, ~1.<;,, J969, PrelinunaJ)•.$md1e, on 1he o,·erwincc.ring~tages o! Cli/i.').· Studk, on orthropod-bom< ,1ru~.,. ofTor.gol•nd. V. J~ol•tlon of
,. mo~u11on (Oip,cr~ CuHcid,u:) In the Hlgtwr:td r~s«on o( S0u1h ,,fri<'ll, Oun)·on\,:crn nnd llift \ 'o.lfcy fcyc-r v i ~ frornmosqultoc!S, ~~11tlr
Jo11m(l/ of,,,. Emomolo,:ricnl Soc•t1> of!0111hm1 A[,·1et1, 32. 9Hl8. .-1jrl,011Jo11mal o{M,'(lical Srlrn<es. l~. 71-,9.
36 10.i•. r•.o.• 1911 The u,xonomk :<1ntusof C11/1U'(01/e.rl 1111Mtum1Sin 57 .:Qt,;J>k,"OT, tt,J4., VMI JbO~. u.r.. • m \:1111uo, 11... 19..,\8, Snidlcson 1hr
Sou1h /\fricn. /011mal <1/1il, /iJ11omotogiml Sl><'My0JS0111h,•m ,\fr/m, 3.J. craru.m ,~f<m orWc>Se-t!>bron ,iru!. by Al!d~ \0chlemrmu1' cobt:l/111
33!l-35i. Th~). Somh Afrlam Ml!dlc«IJovmal. 31 5~1>-5-18.
37 /UPI'. P.G .. 19;,. fMdstudk-son ui., icoedln,;habit1ol'mosqul1oesln the ;;8 •o>:tRM)T, It.IL, s>1m1au,;o; .•.c., C,,\.,'t>,\ll.\, , r. MCl~"TOSI., B,M. •
I lighvcld rogion or South Africo, SQutlr .-Jjrlrn11 /011mai of.Wt<Jleol ,m.,.u.,;s, c.s.. 19Go. Pro,-a, d• n~111rall2n,ao com1,0ros d~ indMdu~
S<lencr~, 38. 69-83. r~,,denu:, t.·m M~ambiqut c:oncrn tlt'terminados ,•frus i~ol~dos c.m

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 We tors: .\losquitoes I 51

Afnco. 1runsml11dos por amopodC'5. ,lni,i~ D<> lttlffwto ~ Mtrli<ino 71> ,10,i,"1'0~11. B ,,•• 1973. A ti\.,onarruc rrassc·ssmenr of /t1,,.vf~ .Ochlcwrouu;
Tro/1<(11/, I,. 201-2'30 cabol/1,s rnieoboldl (D,pttra· C.ulicida~J including a d•scnp:ion or a
nt-w ,p«l('S of O<ftlcrotnwJ. Jo1m1t>I ofthl! fmamologtcnl Soc:e1yof
S9 ::.~smuiu""'· x.t. +· nur..t. f .• 19fi1. Xeutrolb;ang
J.OAl:P::\OT. R.tt ..
Soztlltt"1'$1 VriM. 36. 261 -::!ti.9.
an1Jbod1e. agoin,t arth1n1l0d·b<>me 11ru>e$ "th~ teraor dmnesuc
quadruped,; mni;1111: ,n Tongal;;nd, Umon nf ,outh Afncn ,-\11nt1ls of ;; ,1r.1:-.. rD!\11, a M., 1~tc:SCJ, 'fhl· cpideimolog} or ru1h1opud-b,,mt.1 virus.t.·~ in
Tropl,·al Mrdlriur muJ JAt1mJltol<>i!)', S5. ~ southem \fm:a. O.S<::. Thcsi~ Uni\·,~""'1rof Prcroria.
6o ~0):tHSOT, le,H .• )MUfl8UH..,, "'.C:" l' Hllk\O!',,., tt f. I( OE Mfllt()S, 8,, 19,tk) ;~ ,,n,,.u~H. B>.1,, 19ft(i. Ma,quJto,bornc \irus-dJsc~sof m:tn In ~outhcm
funhti- tsol:Hionj. ot \\"~"olisbton ,·lru\ from mo.,quil(><.~.Somh Afn'ta11 ·\fr!'-"', Sa111/111fr/r1111 .\tcdl,a/ Jo11rnor, 70, »9-72
.\fer/1,nl /011rnnl, 34, a-1-s,4, T'9 !\1C.l'.\~11. a ,,., ni<:t.:Ja-XW~. IUJ \1.1!.0'l!UA...~. G ..M. 6- DOS :s,,~-IOS'. I.S,l.
61 ~eutraHzang
t:CIAEkStJ'f, R,11 .. 4-\HllltUI~, • .;.c.;, & \\'1-NHltEX. ,1,1••• 1956, t~';&. Culr.~ fb'unu'lonom,·Jn1 rubi1101ut11u:ob>lld M \'ttt.or of'Bantl.
antlbodl,•, 10 1u1hapod•bomc viru.c, ;n lwm.in hclngs nnd t\llln>ols in Germbwn und \Vitwa11.~rs.rnnd \'inlM:~- 11. htfecdoru; In -.cnthtd
thu Unloo ohou1h Alricu. /ourunl of lmrt11111vlo,o,. 7i. 313•3:?:t h~n1>1"'< ,i11d 1,,1,1 r,k1':n,~ /n11mnl ofM,d/tt1/ Ei:mmo!ogy, 12, f,4 f,4;-14
6~ l«'l~f;k~OT. Ii.ti.,"'l.i.A.'"· l!.l., U.\1trr. I,,_ Mr.t:.. tfbU, lt.M., &965,. S!UVI?)~ for 80 )ICC~fOSH, B ,1 .. DICE~Sitl~. D.R,, l(;lRArL'-;1, f ,T,.. rg.61.AntlbodlesngohlSt
antibodl"1! agaln,i enhropod·lmrru: ,iruS<'• I!! the ,crn orlndlgcnou~ b..~a.ln ,.ubor \l'im~.s In ttero from humnn beings ru\d dam&sticnnim.rl~
re<ldents of 1hu Caprl\·i Strip mld fk:chu;inaland l'roll!ciorat~. lrom the South .•\frftin High\cld. Smuh AJik4n/our,wJ o{Mttdicnl
Tro•1$n<1io11Jof11te Ro)11I Sorlcr1'0/'fmpl(n/ JMJ/do,,,.,d /~~gien<'. ;9. Sci~n~·t.tS. :r:, a-:.. 9,J
iS3-562.. 81 ,;c.a~"ll)Sfl. ~ '1 •• DOS S.\:\'TU!' l5~L. • Mtf.Nl:KA"'\'. f,.M., 1976. o,rex
6J ~J.Al.tt, f .<,., J(:)UA:-.: ••, .. MtLc,,~. J,M,. C":"Ot-'.~.'-'.(J, N.U..., \\'IU,OS. ~1..L, Pot.H~. (1i111n,lni11m,y111I rubino111.Hh(!,;)b:1ld as ,-eciorofB.tnZI.G<:muston •nd
c.,.. 01(iol,J'n-t.. J,P•• (;IUI-L\UD. M,. ,u-.rueour,. ~.(). & ltlUl:tA,, F. ,: .• 1g.89. Rift \\1m;.ite:·ir.a.nd UNk.~ nr. Tranimh.,,on of \irU~ to h:11m:tersb)·
Voll<.")' f~ver omong dom<e>tk anlmrtl.s In 11,r r«ent We;1A(rie., ou1h:cak. \\1lll•c:iugtu 1111ectecl <. rubtnarut. /oumtJI OJ MedWII Emomo,ogi·, 12.
R=r.-lt lu 1'/rq/o~· HO. 6i-77, t;.;:,....646,

n.1 ..UllOCI. u ... l:S.\Ji..\, \".. coro. \·., \tlUA,,\. '·· 1 ,>:..,•t,hlU, 11.. S.\10,.>,; 0)10DJ, 1, S.2 ,1c.1, 10,H, fl ,i. • JUPJt. P.G., ;9f9. Jnfef.t10rlii ,n sc:minc.ol pigeons by
& ,1.,roMOTO. " ·· a9;&, Sctl)(ogtc ~idtnc<" (Orl!uologfc role of Akab:mc S1ndbt" and Wes, f\lh.• \"ITU.~1.·s in South ,\lrtca. \\'it}) ob,ec'\·attoni; on
,,ru, in optzoottc abortion - anhrogl)'po,is-hy~rancne~ph•l) n, <~uti•.t ;Cult•x; rm1,,mnun iL)iptL·ru· Cultcfdai:-1 ~mrae1ed to c.hCR birds
<"Onlo in l,1pan 19;2,74. ,l,.-1,1,..., ofl'ir<>l1>!(), ~i. 71-23. /0,,,ru,111/M,'1111·11/ F,iromt>logy. 16. 23~·U9
65 USlllltUM. k.J •• OASLb'\'. C..l.. bAVICS ....... & ~lltU. \., 1985. Ob:sen':ttion~ &a Mt;1,to,u. n .,..,. "11111•t. t•.(~.• is&-.1, F..cologft.ll o;cudy on \\'c,t \'ilc vtlll) In
on th,• dispcr~.il nnd sun"t\'tll nr tt pOJ\ut,Jtion ofAt'(J't·$ liiWtuopcm:ii ,anthem Aincn 19C>S-J9a(). /"; ~T. fit.mt<n,, T,n . .k ~,,,. P.H.. (edsJ.
tludlow) IOip<ero: Culicidoc•) h1 Kcni;i. Brill<!t/11 of lin1omolob1r,1/ .-trbQNn,~ Rc:$('(1trt, m #\uun,Jla. Proceeding\ of 1hr 3rd ~yrnpos,um of
Rt•Jt¥Jtrh. i•t. 661""6i0. the (.umrnom.,l-uhh ~P\'nlifk nnd R(';Sc;ur.h Organization and 1hc
Qurenslund ln~ti<u<c for \lcd~AI ttcscarch, IS-Ii Fobruar)'. 191!2,
66 u.,,·mctlM, };'.J •.• BAIUY. c.t.. rua.t.'"R, c.1 .. \~GU.Rf~(':tl~ 1).H•• ('.A.~S<>S. , •.
Bn<b~n•. Au.11nl;:1.
1.0G.\.N, 1·.:.1.. <il8B~. P.tt, k ~1r.K1:Sel}.. J .. 1991, Towards reti1 .. 11m~predlc:don
of Rifl Vull('\' fo\•cr epidemics In Afrii.:..1. Pfl•J·1•iul1,14• \'cn:rlnnty ,\ltdh'imr. 8..1 -M<;1:\..,0.'Jll1 1.1 ,1., 1u1•11. . . . c;.., ~xut.K.\Ct~. t>.& tuo.r.,i,"$.0~ o ..... lS7J, Rifi
11. 32!>-.'l:!1. Valh:~· !WiJI':~. AtU.!inpU,.10 uan..ml1 \1CUlo \\11h ~""\'en 6~CIC'.l al
n,10,quitQe,...Joumnlo/tlrr&,uth :\jrirmt V~1nfrwr;;A.~1>octatfo,1, 44,
6;- Lr:O."llllC:UM, K,I,, 11,\IUY, C-L, TUC.Xt:.n, CJ .• \lfTCIU:.U,. l<.O., 1-,0GA;,;, T,\:,.
57-60.
l).\\1~S. f.G.. J;.i,,L,o, c..w., TJIA."'1;Df ••.(',. &. W\C.\T(lf. PL. 199(). ,\pp:icanon
of polar·ofbiring mctC'o:ologiml c;ateUite dar~ to<fca.""Ct fl.oodjng of Rm 85 :\1t1, l!J:-.)1, fj , ,. im•1•. r.e,.;; nE. ~US.\, J.. 1912. Furlhl•r b.oJ;uior.'i of
V:tlte)' fever viru\ \'t,."CIOr mosqurto habitats In Ktn~-a. Mi:dlcul and ,ubo\'iru•t• Imm 111n,qul10<'• <otl<..-1t!d In longal:uul, South Africa.
I 'cteri1111ry 6111omo/ug;. ·l. 433-438. !96/l-!968 /<)11ri111/ o/.ltll((i"'I f.numw/,,gy. 9. I S5- IS~.

o8 u.,-noclD1, ...-., .• u,\\U~ »:.<.,, ~rtn·, c.L.& ):,\fit(), \., r9&.1. Mo~utto 86 :\fr.,:--.-toli.u. 11 ,1., ,u,•P. P.c.., or ~UM, 1.. 191..t. ~lo~ul~o,:,~ f~t.."dingat two
species encountered In a 11;:,ock,l grils,land dambo m Kenyn••I/0<:11<i1a horixomal 1t-,.cls in galfcr,,• fo11.-st 1n "\;,rnl. South Afrk.i., \\oith rtfen•ncc to
St~'1J . .J..I, 22~2~2, po>-,ibl• vC(lor> o( chllungunra ,1ru,. Jo11m1,/ oft/it EmomologiMI

69 u:-."TTun,,,. K,.J .. t'H\'l~. f.G .. a"'"\11'0, \., ,994. Obscrv~monsofiht.•b1ting

Sc.:i,•r;·11f!iut1tl1em A[rfCII. J,. 81-90.
ucti\ity or mosquitoes..01 i, flooded dambo In Kenra. MoJ.Qui10.v,,,,,~ +i 8i ,1C1'\1"0:o,1I. I). \l •• JUN•, I•.~•.• t>:t.X{,-:.O:o.. OJS., ~C!aUJ\1"\'. C... •, :. 6

;11,;-m. ~wu,.,~,1. J .1967. licolngklll ~tudJ..-.. on Sindbi~ and We-st ~il~ \.'fruses
In c.;outh AftJ(ft. I \l'frnl3.eti\itY 3~ rc\'t.1a1t!d by 1nfocrlon ofmosqultoes
'7Q 1 ISTHICU'.\,1, t;,J P.Wll:j,,,, r.G, l.;A14'0. "· • Jt.\JU~Y'. c.t... 1985. lU(t Valley fo\~( and ~·:uln~J fowh, SQu,lt Africun Jm.11,ml o/Mt!tlldll Sdtm:cs. 32, i-14.
,;rus 1fnmlly !lun;·,n1ndnc. gonu, Phleboli=l. l<olotlons from D1ptera
88 \!Cf:\ IO!'oH, a ,1 .. IUVP. ti,lt, 4.. nn~ SA'nO:o.. , .. 1978. TOl\.'"CliOI'\ by Smdbls. and
colltttod during an !n1<·r·opl1<101k ~rlod ,n l;on)'l. /our1111I ojH,-giur>e,
D:lmbr,dge, 95. 197-20\l. West 11:ile ,inise< In wild papulluions of C,J.l.,_t 1C11/tXJ 1u1M111Jtt1S
n,eobald fDlptern: Cullctdatl ln ~ourh Af1kn. /011ma1 ofrhf
-1 U:\"TltlC:UM, l.:,J., J..\ffUIU.\, H l'.A,. UI\VU -.., f.,(., ~t INl>r.)Ul,T, ~.f,, 1985. ,\ Emomafo1,.'i.(d/ St,r/~ty ofSOmlttTII Afritil, ,1 I. ,;7-61.
bloodtne~l analy;I, of engorged mr»qulto<> f<>und In Rifl \'all<')· fc,-.r
89 \1Cll\l~I. 8 ,: .• IUl'I'. P.n .. UQ,, M~"'fUS, I, f(: U.:\BXAJ:ID, U.f.ft .. 1960, VA!CtOr
cpl~ooric a:r~s In l\enrn, Joumnl oftl:e ..,mertcnn .\tosqulro (:Ortrrol
s111dlcs on R1f1 VaUe)' fever In ~outh 1\llico. So111/J -~Jr/c1111 M'1dicnr
-!$o0rw1/on, t. 93-95,
]QUftl(f/, $3, 127 132.
; , t()(';AS, l \t., U~T111C:U~~. k.J .• O,\,\'ll!~. "-~~. bl'HP.\t. \',$, t,. lt(Hll:14.h. C.R,,
90 ;i.u.·1,ro.'li11, u ,,., ru1•r. r.u.. t><r:, .)j\~'t~ 1. NMU...'iuu,i,. c;,:-i .. 1si6~
1!r,Ji. boltttion of Rift \':illey re,•cr ,iru.s Crom mo"lullo.,. (Dlptern:
F.p,d•mie> c>I l\"'t 1'11• ,nd S,ndbis mu.<•• ,n SouthA!nc.l l\Uh C11/,.r
CuUctdac) col!t-eted dunogan outbreak !n dCtmeiuc animafq; 1n l\cnya,
rC.-r,lc,t) mm mm,,~ Thtob,dd ;is \·t!c:11'.ir South "4.ftie<m Joun1al a/Scitnrl".
1

Jour11nl o/M,:;/lc11/ trrwmul<>J1Y, 28. 293--395

f1.29~300.
"J \lr.1s10>11.11.M~ 1961. Su,ceptibilhy ofs,;me .\ftia,n \\ild rodfm,10
lri,!~dun wfch .,,·,u\ou"' ~nhrnpod ·borne \'lR1~,._ Tr(m.~~:tfcms ofthr
~t \tu,10:-u, u ,1•• 1uN1, p.<:; .. 00., ,, ,..,o.;., ,.~.t. ,..
~u,ett...~. r,.:,1.. 1976.
Cult:i t;·umdanmnylat r1,bi,rows Theobald M, \"'-'CtorofBanit.
Ro,11/ S,X,1errof l'TOJ}it:tlf .\letlt<·ittr (Intl H,-g:,nt ;;;;. 63-68,
C\!rmts.zon rutd \\lrw.m.-~r.md \iru).~'>. I 1'-,Qlaticn O(\.i."U, from wild
nw .ledlnc $ut,v.enu0;.\1tom~ln11it011lon :\C\\')lcad
- .; '1C1:'\'iO>U. a.:., ... 1~';"1. popu!•uon;cifC. rublno:u,. JourMI of,\lrrli<al linromolog,r. IZ, 637-
rtulicld.ie. O:p1cra, 10 /,,0-\lthttrn ,\(d('a w1;h d~p1lon~ of r.,"O ne\\' 6-10.
,p,.-clt.'$. /1>1m1nr of rli, Fmomo(ogl<11/ Sor/tr:,· •f S4111/Jrn1 Africa. 3<.
~~ Mtl"<10-.u. s M•• 1urr. ,.G... uos ,,,,,sco~. 1 & ttO\\f.. .,.c.. 1983. FitJd and
'.l!S-333.
laborato" ••fdcncc 1mpli(•tlng C11t,xcoml•111m;/.; and 1"'1rs.
;s MC:r~10,1t, l:\,M., 1972 Rm \'allt..i f<.'VCt I. \'(:CtOritU<Jlcs in tht: field. Ctft'1.m:1t,tc10,1m a, \'<."Ctot) o('FUft VaJIC'y fC\'l!t\'iru~ in COil)l.;1l South
Jo11mnl of,;,,, Soll/It 1,/rit:rm 1',•r,•rinu,y .\$$<)('/J11i,,11, 43. :!9!-J~S. \!rl~.. $0111/1 ,lfn'r11n .1/e//ical 1<,11m11/, 79, 61-t;.I

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I 52 ,,mo~ ,,-r. Aspects Influencing the occurrtrm.• of infoctiou.s di,ea.~es

93 ,.,c1N"tosu. e..M .. s.t11.,1-1s1,. r. 1.. mc:1,.,-.,w:... ,>.a "wus-snr,. ),t..J.t•• l$U)1- 10.; ~"11tv1t:L. ,,.w.,. ,~,so• .\ Guide !tJ.\(tV.llc<1l l:'momql~·- LcmJort: \'facmlllan.
1\ntibodfts ugalnst cenn.ln 11tbo,itU$CS ln ,,,.. from hum.in being, 10~ .,.,unrnuf',, ...(;,, >:OKt:fl,OT fUt.. tttYMA'.;i.. c.~. t.1.\'0~'fl,f\,f,X. M,J• .. t9S9·
r~ith.rnt in tht coasmJ-area.s o( southern ~at-..tl and e~h.•m C..-ipe ,,umllt7.mg '1ncib()dl0i for cc:nain ,,n,,;.t~ in lhl' scm of hum1m being>
Pco\>!ne<: or Sourh .-\fric.i. S,mth -~frim1t}Qtlf1tt1/ ofMl•d/MI &/mu,. 2,. residmgm nonhem ,a1:tl. '-Outh Afr/am ,lltt/11:nl Jwmol,3J, i,5$-561 ,
77-88..
106 "-"tmlHtJk~. Jr.,C., .t.OUMO..Ot 1t,,t., \'.11:\IUU:~ . \t,P, , . Dl 'otl-lUI)\. II. 1:93,i
9J \1EE(J-A!\. I, ,1. )i:11.WL C:..M .. UOOGiTI\.\AL. H, & ,t'>UA \I. t,,t,... 1~1'1. holatlon of\W;,tl~br<>n ,·1ru, from~ m1tu1..!I) lnfl<:tt!d humdn being
f-'J)Cchucu(df tnutsml~iuu ,md f1..1J bul,tU1,m ~lUtllt:t 1mplleclllug C11to and fran \tdt"1 :fkmbmrl/J1: <1r,11mluttolm TI1eo. "<tmh Viira,1
ptpicns ns. u \'ee:l()ror Rlfl V:tnl'\' r~:tr._,nb in Egypt.,\m... /tJur,u,f of ,:,,w /o:,mnl r,{Ml"llktll St'iC-lt(?I.. 22, 113-120.
fropk<il .\'frdicb:e,uul flygfr11t... 29. 2.;oa-1-110.
lo:" C.)a1 M»'- o,, ).,C•. J•,\nfbO-'. )l.l., ULV\Vt..,s-. L.!,_ .. \\1' 1 tJl. ••A.I'),. 19-59 ...Vl
.93 '\UTS.t.f..'-"', I).. HE.,il~SO~. n.f KlftYA, G.B..
runt, r,M.,. Of.. Ot.u,. A., agent t1.io.ted to Ugi'Lnda S ,1Mb !rout m:in and mo-,qldtoL~ in S<>uth
191.i. Jsota1Son o! .utu,·1~ in k~:nya. 1:.366-1971 Tm1lS<,c1ion.s of tlw Afm."'.~\- Stmtlr Afrlttm .\!t•tl!t-alJvurJ1ttl, .;ti,, 9~~Ytu
Royal Soci•Q•of Trop:ral Metliri11ra11d lf;-g11•11,•, Ii&. 111-123.
108 ~,.. \.'1(,,1.l(U 1.. H•• 19-38. \\'C'\.'i..Cl,;.bron \.in~ dts.CAS<? In: MO:\",\la. 1,P•• tcd,
9b '1U<,l!A1T , .. 1955- Re>carch on South \frican <:ulicinl llllp<cr-•. T11<·.~r/l(Jt'iN1$<'fc £µ11t,111fol(//r,•and l!.riJ/fJ&\', \'ol. l Boca RJtan. Rondo:
Culleldoe), Joumnl oj 1/w l,i,wmoluglr,1/ S,x/Qt)' ofSnudwm Nriw. 18, enc P,cs.. pp. 31-;,,
149-207
10, TVfU'iLl, \1 J,. f,\l"I.\:\ \1,li. COlt"'l1', M, i< n.MLC\" CL, Ss89. \ 10C10f
9i \1USPkA'rr. J.. ss,sm,aun.:-:. .:.c.j PA1Tfl..O~. Ul1. f. >-O.:t<M!\'01 , M,H.. 19:;; n,c compc«ncc of Scn~cs.o A,'(/e;fowler. 11Jlp«ni· C11hcld••J for Rtl<
l•hnm1ory tr,n,m1ss!on of W..scl,hron l"iru, b) 1he bite of ,\rda \'ollcy tnTT \'1m-.. /rmm(l/ of ,\lrtlirttl F.momology.15• .lt)2--t00.
[&,11/:siilellal ci,.,uml111ool,uThl-u S<J1ttl: A{rlcn11 /tJlm:ttl of .\ /.-.lira/
!.ci~nt:l'tS. 2..2. 12.1-126..
IICI TUMf'Ll ,1 J,. PRE!,U:,. , .,1,, 4-."\0,A.:,.1,, UJf'll ,.t..
OOH~,. U.f,, MUIUUU.. J.C.
._ \M'IHIUt, H,Jt.,, It)~,. Vec:;arcamp ....,cnc:e r,(Fgyp1i.in mQ~qUUOdo for Rik
'98 O'l"o\, .\., 01:V.:O:().. T .. OC'r.\T,\, T .. l:.00.\\'.\4-fll. 1.4 .\1A1~UY>AM,. T•• 1.961. \'allty !t.-vtrvln,.!o. Amtrknn /our,111/ of'l"mpl,:at ,\ltdlrm11uml llygft!mt,
Akab:tne. a nuw 3Ibor virus isolati,d in Jnp:m. JntMme.w• Joumol t,f ;~. 136-199.
Mlcroblologlrnl Sd,·11cen111J /J/l)IOl:l)', 1,1 101-108.
111 n~os, P.d .• 1~-;tt Hr,,fnfoll chM~e\li ()\·er South Africa durin; 1h 1.1· period or
!19 VA 11±.JtS,O:-<. 11,1:!.., BJ\0:0.SL>f.:\'", , •• L£:\1TI', ,. ~ \~Oltl II, c.;.,H .• 196.c Sonw mt-t~owtogic.J rt...•curd Ju; \\'u:rA. ~,~""- [cd.1 111()$.i.-ogmµ:,y "'"' F.tnfogy
cuU<:ine 1110.<quirn,,s !Dlpt<tta: Cullddoe) at Ndumu, Hcpuhhc ar~oulh ofSoutJ:,m Afri«L nie llu~ W. Junk.
Afrk~; a study or their host pnrfercnl.e\ :ma hO!'l't r.sHgc ••,rMi~'11
f>ron'l:ding,,, 10. 183-1.92.
U2 ,.M, orntJMJl't T.C,Df Ir.., Ill Will'. 1•.11., \'A~ l•UT/J~ ..... KOK, 0.1 .. HmKIL,

198::. ~pcc1i:s nchness and rc!:.th·c abundanet

:,... \!0)1"IflT, t'l,f. 1c xm.. .,.,
:oo rATR,ICI\.', t,..\. r. H.AltEt', c.t.., l98~, lnge:)tton of iinmunc bloodmca.J.s and of fematc. moS<1ufio~ :u a sm.! in lhl• w~~tcm Orange Free.Sm co. Joumol
lnfeorion orA,desfowllfri, Aeda mrlmoshl ond Cu/ex ,,,,,1,11.1\\irh Rife atrh,, F.11ron:a/1Jgf,'lll jq(;1,•~·0JS<JUll1t•rn Afntd 45, S1-67.
\'alley fe\'ervira:s:.•.\menw11 /OtthlOI afTmpr,"I t\f4:tlirt11,• rmd It;1Zh't1t',
lt:i wUMift, c.n .. ,wrut~o:--. n..t (( oc Ml!tU.O:". as .. iabi.. The fncidcn(e ot
40, ;,3,1-540.
nnhropod,boml" ,in,~ m J pupuhulon nf culk:Jnt:. ml)5-quitot•:,, In
101 ftl;f.i\'1·",, \\,(,, Bt llA"1Y. ".t..
tm&. A,r. ,e. (,C"JU\'.\~1. K...... l.964, \l\.llyffi of ,he l'onJl<l]ond. Un,on of !>Outh Mr.co (Janual). 1956, through April, 1960)
clttucrutance~ leading 10 .lbonlnn ofa Wc,1em equln~ Mr~h;llftl, :\rnn,amJoumnl o/Tm1m:ot MNlrdm•m1d Hyg,rt14', lb, 583-Sit:?:.
cpltfcsnlc, ,"1.marlcnn Journal of JIJ1,.rfcn,·, l.sO. lO-l-?:tU.
11.! /.1!.IJ.tn, U,G,, ft,\, M. \!, "'- ,\Ml;l~ I, A,l .. lt'}'')S, R.ift \ 'ttlh:)" t''-<\ C'f t."J'lX.c>lUit ln
10:t S..\L\HS. J.J., RIC:kE:'\llACH, /t.., OPC~. t1 .. BRO':"'f(~ 1I,, ~l>RM.\J-..: '1.. E.OUL.\~. ...mall rumln:inl( Jn ..outlum, Mtturirrml;i (Octob~r 1993): riik of
J.P. a. r0t.RAM, 1•• 1969 ~ 11rbo\iru, bolf,s ~ parur dt! mou,.uqut\~ au t.!Mtn~l!O\llhrcali.~ ~1~wnl1nfSt>tlcl)"l?.t:IK..• \/t.'dtdm.• Tmplc11I. ;;;,
Camereun.. Bullt•rin oftl:c World 1/t."Olrla Orl(/wisntftm, , I, :.?33-2,; t. 13:,-1,10
103 s.\Ul7.ZO, J,f,, f)l(.()VTfl;, f,P... C:lL\RTIJifl, C \URl'1NtZ, I>, & B:\Di\. R,, 1987, 115 11 U.rR. 11,(.. , fO~ll.SltJ.S:, U,, lk.At)H£~l,\)U'l\,.\, '1,. r1U0!\C.:-\~~•. \' k
Focusot Rift \"tt.11£:y fo1,·cr mmsmiu!on in southern M,1utita.nla f,rmtc1 I. Dmoun-t. J.P.• 100::-.. hn1.oolir ,1clivlty l)f R.ifl ,·.11(4,.~ fovt:r in StnagJJ.
50~. .-\,nj•fienn Journnl o/Tto;1fr11l .\.ftttflcln&?.mul llt;i,m". 56.. 26.S...272.

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 7
Classification, epidemiology and
control of arthropod-borne viruses
R S\\"ANEPO.EL

Introduction monkey~. lly then, ,;everoJ arboviruses of "eterinary signifi-

cance had been isolated, the firs1 m·o being African horse
,\s recendy as 1he lare 1960s the cerm arbO\'iruse:.. a partial ac- siclmes~ and bluetongue viruses in South Africa. Although the
ronym derived from arthropod-borne ,iruses, \llllS used in a exact dates and autl,orship of isolation oi rhese two ,~ruses
taxonomic sense 10 denote the existence of an inferred un- are debatable. they had been maintained by subinoculation
derlying relationship between vin1$c:< \•.1th similar mode:. of in hor,es and ~hecp re~pecti\'ely, long before they \\ere
transmission. After morphological and physicochemical shown 10 bt· filterable agents or were adapted to laboratory
charactcrizarion, it became evident thac the \~ruses a,tually hosts:•,.;:, 16<. 1-,;,21G. 21,.2i,,
belonged 10 a range of widely disparate t'amilles. and since By 1939, 16 arboviruses had been isolared (Table 7.1). al-
then there has been a tendency to stress the diversity of the though Lbi~ flgmc is somewhat misleading as it does 1101 take
,iruses, at tJ1c cost of overlooking shared epidemiological Cea- into account 1h::u. African horse sickness and bluc1ongue vi-
lures which dictate a common approacn 10 the inves1igarion ruses are each made up of mulrlple serorypes. Of 1he 19 ar-
of arbovlms diseases. indeed. the study of the arthropod- b0\1ruses isolawd between 1940 to 1949 [fable 7.2) eight
borne viruses has ah,11ys been a specialized branch of virol· were human pathogens from various pans of the world. but
ogy in which there is. nevertheless, a need for co-operation I l were l'iruscs of unknown significance that had been iso-
berween such \'ll.l'ied ln\'esrigarors as\irologists, medical and lated from mosquitoes and a sentinel monkey in the course
veterinary clinicians. entomologists. 1.oologists, ecologists of yellow fe1·er investigations in which tJ1e Rockefeller Foun-
and clima1ologis1s, among others. Except where otlwrwise in· dation Virus Lilboratories RF\1.) of New Y(>rk played a lead-
dicaced, the tollowing outline of the subject is drawn from a ing role. During the Sccond World War, RFVL concemrated
number of genern1 accounts of arbo1~ruses. 22• 109, 2211• 240 on the in1·es1igation of )'ellow fever. but during the l950s it
For Cl'nlllries it wa~ ~u~pP.C'IP.cl rhm rPrrnin rlisea~<'~ \\'PrP sPc-nndP<I "t:iff to l;ihormorir" in rnany pans of 1he world, in-
1rru,smittcc! by anhropods. but it was noL until l8i8 that cluding what is now known as the i\:arional Instilute for
Patrick ~lanson demonsuated in China tha1 rno~quitoes Cornmunicablt> Oise/lSes in Johannesburg. to establish and
~rve a:. vectors for the filarial worm \\'11ch1,1reria IJ<IIIC'roftl. In foster arbo,irus units which were intended to continue op·
1893 Smith and Kilborne established that ticks transmit the eratingas regional cenues \\ith locally trained staff.
babesia, which causes Texas redwater fe\'Cr of cmtJe. and be· The ne\,·ly established arbovirus unit in South Africa. ini-
tween 1895 and 1898 Ronald Ross performed investigations tially under rhe leadership of Smithbum of RF\1.. conduc1ed
in India that pro,·ed that mosquitoes transmit the parasite of surveys for antibodies to a range of arboviruses on sera
malaria. Shonlythereaf,er. in 1900, Waller Recd and hisasso· collected from humans and domestic animals in South Af-
ciates in Cuba demonstrated human 10 human transmission rica.62· 12~· i:r. 11u, 1~5 •213 :0.!01.ambique.1i 3 Angola. 11i; Namibia
of the virus ofyelloll' fever by mosquitoes. as had bcE-11 posru· and Botswana. 1211 The highest prevalence of amillodies was
lntcd .earlier by Carlo~ Finlay. Not only did this constitute the found in ~era from 1'ongaland, n subrropical area occupying
first proof of transmission of a virus by anl1ro:pnds. but it was the coastal plain of the northern part of K\\aZulu-Natal
also the first occasion on which evidence \\'35 produced co in· Pr0\1nce In South Africa, and consequent effons to isolate
dlcate tha1 cm agent causing disease in humans, as opposed ne\, ,,mse, were concentrated mainly on this are,1. In the
to lower animals. was caused by a 1irus. However, 11 was not decade from 1950 to 1959, members of the Onderstepoort
until 1927 that Smkes and his co-workers in ~igeria sue· Vererlnary Institute isolated \\'csselsbron ,;rus from a lamb
ceeded in maintaining yellow fever l'irus by serial passage in in the,Vesselsbron district of the Free State Pro1ince, while
153

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15'1 «nm~ c.": :\SP<,'Cls influ~ncing tho occurrcnn, of lnfeellDUS diseases

Tabl& 7.1 ArthrallOd·bomev,ruses ,solat.1;! throughout tha wo:id p,,or to Table 7.2 New artilropod·!:orne viruses ,soia:eo throughout th~ worlo
1940. lnformat,on 2cap:eo irom Karata!sos 11985i1os from 19~0 :o 1949 lniormat on adapted from l<arauatscs 11985)109

VIRUS ORIGINAlSCURCE YEAR VIRUS ORIGINAL YEAR COUNTRY

SOURCE
lliuetorigue Sheep blood 100:·
Afncan horse s,C'.o(lleSS Horre blood 1sos· Wyeomy,a Mosquuoes 1940 Colomll,a
Afr,can swine fe•.cr Pig blooo 19;0 AnoonelesA Mostiuitoes 19.:0 Colombia
i /a1rob1 sheep oiseaSe Shespbl1iod 19'.0 Anoi:l\ales B Mos~uitoes 1940 Colomoia
Vesicular stoma,,t,s Cow tcngue ep,1hellum i925 Semlik fw.si Mosquitces ·g42 Uganca
-Indiana Nia·,a Mosounoes 1943 Ugana2
Ya110111 fever Human blood 192i' Cal,for~,a encephan1is Mosqu,toes 19,3 USA
l.ouping ill Sheepbra,n 1929 8unvaom.ve•e Mosquitoes •943 Uganda
Ritt Valley lever Sheepblcod 1930 Sendfiy :ever - S1citv Hbmanblood ·s,3 Italy
Western encepllali:•s Horse brain 1930 Colorado urk fev01 Human blood 1943 USA
Eastein encephalitis Horse bra,n 1933 Dengue\ H~man blood 19t4 Hawaii
St Louis encepnalit•s Humanbra,n :933 llt:aJS Mosquitoes 1944 8rwl
Japanese ericepha it1s Human brain ,c--
• .1:1 Oeng,;e 2 Human blood •944 New Gi,:nea
So!Jon • • Humanbra,n 1937 Sa~d'iv !ever - Napies Human blood -944 11atv
Bwam!la Human blood l93i lika Sentinel ·9;,7 Ugalldo
Wes; Nile Human blood 1937 monkey
Venezuelan encephalitis Horse brain 1938 blood
Uganda S Moscw11oes 1947 Uganda
See text for derivauon of the dates grven here for Jsolat•on of Omsk haemormaglc fever H:iman blood iS-7 Soviet Union
bluetongue and African horse sic~ress \iruses Ha~zalova· Human b1a1n i9'l8 Czechoslovakia
Sof1in = pro1cwpe Strain ol Russian spr ng-summe, enccphaiit,s virus Tnvittatus MosQu,toes '946 USA
Neg·s.~1 iiuman ·s(a Japan
terebrosp1nal
'lu1d

personnel of the arbovfrus unit in Johannesburg Isolated IO

r.anialova = first wain of :he virus ol Cenval European encepnal,lis
new viruses from Tongaland and four from elsewhere in 10 have been 1sorared: i~d•s1ingu1sha1!le oy cross•neut1ali2a1ion from
southern African (Table 7 .3). Most of these ,iru,es were the vir,s or Russ,an spnno-summet e.1ceohali1,s
originally isolated from mosquitoes or birds collected in sur-
veys and have not been found m be of great medical or ve1-
Classification
erinary significance. alt hough a few of the same viruses were
later Isolated from the blood of persons with febrile disease. The system of C'lassifying arboviruses on the basis of immu-
Since 1959. there has been less emphasis on surveys and nological relationships. which was largely de\1eloped by
only three new 1iruses have been Isolated from anhropods. Jordi Ca5als of RFVL and its successor. the Yale Arbovirus
although several potential veterinary pathogens have been Research Unit {Yt\RU). has been borne out b1• modern mor-
isolated in somhern Africn (Table 7.4). A number of arhovi- phological a nd physkochemic.il characterizarlon of the vi·
ruses that had original ly been isolated elsewhere, were also ruses. Serologically related l'ini;,cs were first placed imo
encountered in southern Africa. These included Rifi Valle>' groups A and B.35 and then later a group C and a Bunyam-
fever and Akabane viruses and human pathogens such as wera group were added.33 J.l 36· 3 : \\'eak serological cross·
chikungunya. Wel>t Nile and Slndbis viruses (Tabl e i .5). reactions which 1,ere obsen·ed to exist between group C, the
There was a need to collate information on the wcahh of Bunyamwera group and certain smaller groups led 10 the in-
newarbovirusesisolated worldwide from 1950 onwards and clusion of these groups in a Bunyam\\·cra supergroup. 7
in 1959 the RF\ 'L organiZcd meetings that led to the forma- .Many other unclassified arboviruses and rabies virus were
tion of the American Commiuec on Arthropod-borne Vl- found to be rod- or bullet-shaped and to share certain physi-
rmes (ACA\1. In 1960 a Subcommittee on Information cochemkal properties. These were then placed together in a
F-~change (SIE, produced the ·catalogue of Anhropod- thabdo1·irus group (rhabdos Gr.=: rod).~~ 100· 103• 11 5 • i;s. t"3
borne Viruses of the World'." hich is cominuall y updated by .'\s African horse sickness and bluetongue 11ruses were
1he mailing of registration cards for new viruses. However. found to be resistant to lipid solvents and had a similar mor-
many of lhe more than 500 registered viruse, hO.\'C been phology 10 rcol'iruses,28 · i;;i it was suggested 1ha1 they
proven 1101 to be arthropt)d-bome but are nevertheless re- should be placed in an orbivirns group.2ff named for the
tained in the c.i ralogue as they are likely to be encountered ring-~haped c:apsorneres seen on the surface of virus par·
b~ arhovirologists in the course of testing material, such a~ liclcs (orbi$ L. =: ring).
organs and sera from small mammals and birds. The title of The overvd1clming majorit}' of arthropod-borne viruses
the t:atalogue has been altered t0 • Imernat ional Catalogue of "·ere thus placed into A, B, rhabdovirus and orbivims groups
Arbo\~ru$es Including Certain Other Viruses of\'cnebrate~·. plus the Bunyamwera supergroup. The 'groups' were initially

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 Clas<lfic,111011, epide111ioloi,,y and control of arthropod-born~ ,'!ruses 155

Table 7.3 New arthropoa· VIRUS ORIGINAL SOURCE LOCATION YEAR R£FERtNCE
bome vir~<Ses 1sola1ed ,n
sout.'1etr Africa i1om l 950 to Wes;elsbron lamb liver Wes~elstro~. RSA· 1955 264
1:,59 Pongo1a Mosquitoes lake S,mou RSA 1955 128
Slmtu Mosquuoes lake S,mou RSA 195S 262
S;,ordwenl Mosquotoes lake Simbu. RSA 1955 12S
lobombo MoSQu,toes Ndumu. RSA 1956 22
Sanz, Human blooo Ndumu, ASA 1955 2g
M,~delilufg Mosquitoes M1!ldelbc.rg 1957 117
Cape, RS.\
Nyamanini Bird blood Nvamanim Pan. 1$57 228
RSA
Gemi1S1on Mosqu,tces Ge1m1ston. ASA 1958 126
Wit1~atersrand Mosquitoes Germiston. f1Sl\ 1958 152
Usut:r Mos~ulroes Ndumu.RSA lSS2 228
Tete . 811d v,scera Naumu. RSA 1959 SIE l97o··
Mos;u1il Mosquitoes Lumbo, MQZ· 1959 i23
Ndumu MOSllUttoes Ndumu.RSA 1959 i20
lng·,,a,;u"'.a B1rd viscera Ndumu, RSA 1959 153

:,SA: Republic of Sout.'I Afr,ca. MOZ: 1.1ouimb1~ue

SIE. 1970: Subcomm1:-i£e on lnforma:,on Excr..mge of tile American Committee on Arwopod-bome
'.'truses. i 970

Table 7.4 New acthropod,oome VlRLS ORIGINAl SOURCE lOCAilON VEAR REFERENCE
,,ruses ISOiated 1n soutl!arn
Africa subs..oquent to 1959 Sho,we Mcsqu,,oes Ndumu. RSA· 1962 jq7
01.:antsvlei Mosquitoes JOllannesburg. RSA 1963 '09
Cascara Horse brain. viscera K,m~r,ev RSA 1967 70
Ap,.s Riw;r Cattle blOOd Ooderstepoori RSA 1967 266
Bovine ephemeral fe'le.t • • Cattle blood Onderstepoori. RSA 1967 253
~afflll Horsl! hlood Nahoomsoru,; RSA 1971 71
Prei1)na Argas,d uc~s l'letoria. RSA 1973 50
Nrabi<a Cattle foetus Nyabira ZIM· 1973 m
Kaa!plaas Horse blood Ondersteooo11 ilSA 197~ 71
Kyalam 1io1se viscera Onde,mepoort RSA 197d 69
Gweru· .. Cattle foetus Gweru. ZIM 1976 265
Brvanswn HorSl! visc,,1a Johannesburg. RSA 1976 71
lani;ebe1g Horse foetus Pary5 ASA 1977 69
Marondera Cow viscera Marondera. ZIM 1978 265
RSA= Republic of South Africa. ZJM: z,mllabm
Virus Isolated in 1967 from cattJe blood stored 10 1958
Results of recen: te~ts on unidenufiec viruses In storage at the Natio~l lns111u1e for Communicable
Diseases. indicate that Gweru virus ,·.-as 1SQlated from Culicoidesm,(!ges in Sou1h At11ca in 1SS9 and
1970. p11or ,:> ns racogn1ti-0n as a new W\iS ,n Zimbabwe ,n 19762' 5

raised 10 the status of genera,5 · 269 and some were later des· and those of hog cholera. bovine virus diarrhoea and equine
ignared as families as the general classmcation system of,;. viral arteriti~.
ruses evolved, On lhe grounds that they possessed similar Bunyamwem s11pergroup viruses, and serologically un-
morpholoi,,y with lipoprotein envelopes. groups A and ll grouped \iruses with similar morphology. were placed in
were initially included in the new Togar•irus genus (togn I..= the ne\\ family Brmyai•iridae.73 17'1• 181 \\'hich initially con-
cloak,\, 5 but "'ere later designated as the Alp/tar,irus and Fla· Lained the single gem!l> Bunrm,irus. but 10 "'hich was later
11iuims genera of a new family Tog<wiridae rna"i' referring added ;\'airo11/ru$. Pltlebo1•ir11s and U11k11t1in1s genera
10 the yellow of yello" fever \'iru~. the type ~pecie~ for thl' (named for Nairobi ~heep diseasa, phlebotomus fover and
genus Flavivirus).7' Subsequently, the elucidation offLmda- Uukuniemi viruses respectively).24 116 .\lore receml)', the
memaJ differences. Including differences in replication genus U11k111'irus has been dropped and its members incor·
strategy. led to the ge11us Flcwi11irus being placed in a new porated in the genus Phltbo11irus on the basis of biochemi-
"°
family Fltwiviridae. However. the genus 1llpluwirus re- cal simllarities. while a new genus Ha11cavirus (named for
mained in the Tog<wiridae along with a few other genera l lantaan 11rns). which includes a group of non-arlhropod·
containing non-arthropod -borne viruse~ such a& rubella borne. rodcnr-associaced 1iruse1, producing haemorrhagic

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156 '.'if(. 1UJ\: 0\1,! Aspects tnnucnting I he occurwncc or infocrtous disea5os
Table 7.5 ArthropoC·barne VIRUS SOUl!CE LOCAi1Q~ YEA!! REFE~ENCE
~,ruses orig,r,ally iso1atez
elsewhere and swt;so.,lluently R,;; 'fa !el' fevei Cane live· Joha1nesb"urg. RSA· i9S' 3
1sola1ed ,o ~uihe'l! ,:.,t:ica S•r>eo•s Moso,,to<S Springs. '!SA 1554 252
Ban-,am\1.era Mosq.,to.s Ndijmu. ~SA l<l''
v~ l 18
Cn,,~ngunya Human clood Le:at:a. RSA ·g~ 84
West Nile human 0,ood Ndumu. RSA 'S:a 119
Tah·ina Mosqu,to6S Lumoo,MOZ' 1959 122
Sllmlik1 Forest fl.'osqu,toes Namaur:a, MOZ 1959 156
011arani1I B1rdv·scera Naboomspr~,t. RSA 1960 22
Crnn1iaa Argas,a :,cks HartebeesPoOrt. ASA iSSG 147"
S!!aamomta M,oges Onoemeooon. RSA 1S67 ,~r·
Sab~ Canre blood Onders:epoort f!SA lSS7 56
Shu1i Ca,tle blt»d Oncs•Hepoor. RSA 1gs7 56
Gcmc\a .. • MoSJ;.;.toes Ha;a•e, ZrM· 1969 25
Th:ni~, ~;,a:ges Lake Ctmss,e. ?-.SA 1971 25
Arumowo, Mos:iu:ioes Por: Sne;istO!\e. IISA ;972 150
~taaoo Algas,d 1,cks lambern Bay. RSA ;973 147 ..
Ahad,ra Canie foetus ~yab,ra,Wo/ 1976 26~
fsra,i Mkev men,ngoer,c~phatms Turkay ovanes Krugemfaro. RSA 1978 15
Bagaza Cau,e foe1us Ofav,. NA~1• 1978 16
Ugllll<!aS Mosiiu,ioes Pon Sheps.one. RSA 1979 147
Cnrr.ean-Congo haemO!r~ag,c lever Human blood Sloemhof RSA 1981 224
Akaoane M1ctges Ma1owe.Z1M 1982 26

RSA= Reouc11c o1 Set:tn Africa: MOZ= Moumbique: w: = Zimcabwe: NAM= Nam,n1a

Addi;,onal info·:r-ar,cn :rorn unpuohshed labc•atorv iecords. National Institute fc· Communicable O,seases
Gomoka and Tn,mir1 v,:uses we:e recently 1dent,fied in tests o, un•denlihad ,sola,es in s;orase at the
National lnst:tu?e :or Commumcallle Diseases

fever with an associated renal s~11drome in humans has '-:ames of arboviruses with a long history of general usage
been added to the 81111,vcwil'idae.9,; were ac<:1!p1cd u11aJ1ered for registration in the catalogue. ;s.Je"
The famil~· Reoviridae. as originally esrablishcd. induded viruses are nnmed by the authors reporting the isolation. it
arthropod-borne members in a single genus Orbil'im;. but being common usage to name arboviruse~ after the geo-
later it was found that Colorado tick fever and related ,;ruses graphic loc:a1ion in which they are firstdb:covcrcd. Geographic
differed from orbh~ruses in the number of KXA 5e1tmems in names are spelr \dth capital initial letters. but arbo\'iruses
the genome and they were moved to a new genus Coltivirus. named for disease~ are not capitalized - for example: !lift
Anhropod·borne members of the famil~· Rltabdoi'iridae \'alley fe\'er. African horse sickness. and bluctongue.
were placed In a genus V1?.sic11/0t1im,~ (named for \:esicular Viruses registered in the catalogue arc accorded
stomatilis virus).-4 abbre,iations derived in a specified mann('r from their
Altogether ~97 of the 534 viruses (93 per cent) rcgiswrcd names.6
in the working catalogue aL 1he end of December 2000 be-
longed to live families (Togm1iridne. F/m1i11iridt1e, B1111y1wiri-
Ep ide1niology
dne. Reo11iridt1e and Rhabdo1•iridne) (Table 7.6) and.
although not all of these viruses are known for cenain 10 be Arboviruses arc defined as \'U\J$CS 1hat are maintained in na•
arthropod-borne. ve1yfew oflhe rnmaining37 viruses in the mre principal!~. or to a significant exiem. through biological
catalogue are considered to be definitely or probably anhro· tran~mission berween su~ceptlble venebrare hosts by hae-
pod·borne. ;\cw viruses are continually being registered in mamphagous arthropods. TI1ey replicate and produce ,;.
the catalogue and it is believed that only a small proportion raemia in at least one ~pecies of venebrate in order to
of the arboviruses exlsting in nawre ha,·c yet been disco\•- facilitate infection ofhaematophagous vectors. Follo\,ing an
ered. Although it can be assumed that virtually all viruses extrinsic incubation period la.s[ingseveral days and involving
lhai cause major epidemicdisea~cs in livestock :rnd hum.'lns replication of the ,iru$ in the tissues of the arthropod. arbovl-
are known. It is possible that many arboviruscs that cause ruse~ are transmitted to new venebrates by uw bites of the
less obvious 'erosh·e· dfaeases such as sporadic abonion and anhropod vector. B) contrast, m.echanical ,ransmission of ar-
teratology. remain to be disco\·ered. Furthermore. it bo\iruses im·olves the direct subinoculatlon imo a second
occasionally happens that pathogenic roles are discovered vertebrnre host of infected blood contaminating the mouth
for viruses registered in the catalogue and which had no pans of a biting arthropod during the feeding process. Robust
previously known medical or veterinary significance. tabanid flie~. whirh arc painful biters and are consequently

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 Cta5'-.ification. ~pidcmiolo,zy and romrol c,f arthropod-borne ,iruses 157

Table 7.6 Taxor.om1c sra:us o: 534 viruses reg151e·ed in the a \~ms should I.le regarded as arthropod-borne include:
·1n1ema1,onal Ca1a1ogue o' A1bowoses Jr.cti.::i~ Ceru11n Omer VirJses a: • l11e frequency of iisolation of the ,ims from narutally ln-
Ver:e~rates·. at the end of December 2000 fecu:>d arthropod~:
FAMILY GENUS ~Uf..lB:!' OF NUM8Eli OF • demonstration of natural transmission of infection 10
SEROGROUPS VIRUSE.S suitable ,ontinel anin1al, exposed under controlled con-
Togavifldile Alohav,r:1s 27 ditions;
lforrr.erl1~roup • demonstration of 1rnnsmission by natural!)• infected.
A aioowusesJ wild-caught arthropod~ after an appropriate interval In
Ur,ass,gned Uog·ouped 1 captivity 10 exclude mechanical transmission;
Flaviwridae Ftav1v,rus l 68
• demonstrauon of biological transmission follo\\~ng in-
Jforrre1i,=group
Aa,bOvi,usesJ fection or pumiive arthropod vectors, either by feeding
Bvn'{ilviridae Bun;1Jvirus 18 138 on an experimentally produced viraemic blood meal. or
Na1ro>1rus 6 24 by direct inocula1ic>11 of ,irus inco 1he arthropods;
Pnleoov,rus 2 43 • demonstration of virus replication In experimentall)' in-
Hanrawus l 7 fected arthropod~. including trans-stadia! persistence of
Unassigned 8 41
lie<;;,1r:dae Orblv,rus 15 infection during the life cvcleofhemimetal>olous-anhro•
59
Co!111,r11s I 2 pod~ such a~ ticks: and
0:111(]1eov1r11s Ung•Gl!pa:I 1 • demonstration ol lhc ability of the ,:irus to produce vi-
Unassig~eo Ung:01.ped ~ raemia in laboratory animals. suitably exposed semirn!I
ilha!):fovmdae lyssa11,rus 3 16 animab or in naturally infected. free-ranging animals.
Ve=lov1rus I 18
Ur.assigned 6 37
4renavi1idae Arefl8WUS l 13 S11ppi>r1.111g evidence includes tn.xonomic rela1ionship 10
Arreriwidae 14nenwrus l I kll0\\11 arbo,in1~es and strong epidemiological as~ociatlons
Asfarvmdae Asf1v,rus 1 1 with ;:irbovlrus diseases, such as seasonal occurrence of in·
rilr;viridae Filowus 2 2 fection tha1 coincides with 1he biring acti\'ities of poiemial
Nod:iviridae Alphenodavirus Ungrccped 1 vectors.
Orrhonwxowodae Thogotowus i 1
Ungro~J)ed Ungro~-pe:I 2 Negative evidence which renders a virus unlikely tO be
?;;ramyx.ovlfidiie Paramy~ovirus Ung·oLped I arthropod-home. includes:
i'oir,iridae Unassi9~e~ I • po,,t/ssion or physicochcmical characters that ar11 not
Unassigned Ungro1.ped 2 !'}1>ical of ~nown arboviruses:
UnCl~s1f1ed 3 12 • fnilure 10 replicate In a banery or suspec1ed arthropod
534 vector species, including failure of infection to persist
1ra11s~tadially In hemimcmbolous vectors: and
• isolation of the viru,. from salivary gland, urine or facet's
frequently disturbed during feeding. constitute good of a wnebrute. since this implies 1he exi,tence of an
mechanical vectors (see Chapter 4: Vectors: TabanidaeJ. alternative mo-de of U11n~mission.
Arthropods That have been incriruinaicd as biological
vc>c:rnrs of virus<', in a11imals :incl human~ 'inrh,nP clip1Pro11s On The ha,h of nvailahle e\'idence. <;EAS classifies each
insects of three families: mosquitoes. ceratopogonid midges registered virus as cllher arbovirus, probable arbovims.
•Culi,·oide.~l and phloboromine flies (s:mdflies), as well a~ possible arbo\1rus.. probably not arboyfru~. or not arbovirus.
ticks (see Chapter I: Vectors: Ticks, and Chapter 5: Vectors: Of 1he 534 \iruses registered ln the catalogue a1 the end of
Cttlicoules spp.). Of the 53'1 vin:ses registered in !he arbo\•i- December 2000. 214 (40 per cent) were rated as definitely or
ru~ catalogue at the encl of December 2000, 263 have been probably arboviruses and 33 (6 per cent) were rated as
isolated from \\ild-caught mosquitoes (predominantly culi· definitely or probabl) not arbovlruses. In essence, this
cine rather than anophelinc mosquimcsl. 117 have been means 1hat the information available on Lhe remaining 28i
iSolated from argasid and Lxodid ticks. 45 from sandflies. 36 viruse~ (54 per cenc) rated as possible arbo,'iruses ls 100
irom midges and l; from miscellaneous other arthropods. meagre 10 allow meaningful condusions to be drawn.
The figures proVide an indication as 10 the relati\•e impor- The concept of threshold intenshy or ,1raemia for the
mncc of the varioui. groups of nnhropods as vectors. but infection of haematophagou~ anhropod~ was csrabllshed
definite proof of biological transmission in nature is lacking with ins1amancou~ feeders. namely mosquitoes. Threshold
in most instances. 1lowever. much can be gleaned from has been arbitraril) defined as 1he level ofviraemia al which
analysis o( supplementary e\~dence on virus uansmi'ssion, 1 to 5 per rent of arthrnpnd~ taking a blood meal become
and for this purpose AC.AV cs1abllshed a Subcommittee on infected 1\itr thl' virus concenied,39 and the concept has
the Evaluation of Arthropod-borne $rams (SEAS) in 19;0. important epidemiological implications. For instance, the
Criteria used by SEt\S when determining whether or not detection of antibodies in a ~urvey would indicate ,,·hic:h

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158 « 010, "" ' \<p;,c:1s lnRu~nclng 1h~ 11ccurrcnce of lnf~-cliou< di<ca-.•s

venebrate species become infected with a \~rus in nature. mission by arlhropods2s1 but differences in geographical
but the performance of experimental infection~ would allow origin or limited pa.~saging of vim~ strains in the laboratory
distinc1ions 10 be mnde between \'enebra1e species in which do not necessarily aJJect vector efficient}. 1118• in, 2M
the intensity ofviraemia exceeds the threshold for infection Holomeiabolous \'CCtors, ~uch as mosquitoes. midges and
of anhropod vectors and venebrares which may become in- phtebotomi11e flies, undergo marked metamorphosis during
fected yet fall to develop sufflc1en1 lmensi[) ofvfraernia 10 their life cycles and the immamre l11s1ars exist and feed in en·
fnfectanhropod.s.' 13· 114 · 178· 193 \itommmts. often aquatic. that arc very different rrom those
It is a general propcny of arboviruse, that 1he) can r!'pli- in which the adults exist. ~losquito larvae can be infected
ca1e in a wide range of arthropods af1er being directly lnotu • with ;irbol'lruses by ingestioir11-1 9 but the epidemiological
lated lnto the haemocoele. but that they tend to produce significance of this phenomenon is uncertain. ai1d in general,
natural infection in on!~ a limited -ange of specific vocton,. holometabolou, vector~ must gain and rransmit infection
The proportion of arthropods that becomes lnfecred \,i1h a during the adult stage. Female mosquitoes may take a blood
vims generally increases as the inte'?sicy of a ,~raemia rises meal e,·ery cwo to four days and live for sel'ernl weeks. so tha1
above threshold value. but for each virus-vector relation- af1er an extrinsic inC"ubation period of one 10 1wo weeks. an
ship there is a maximum proportion of arthropods that be- Infect('(! mosqui10 is able m 1ra11smi1 infection to several ver-
come infecred. lntense viraemia with a particular \irus may tebrates. In hemimetabolous vectors. such as ticks, 1he im-
produce infection in a maximum of 30 per cent of one spe- mature instars resemble and feed as adults. so that infection
cies of mosquito, 80 per cent in another. and yet fail to pro· can be gained as larvae or nymphs. and d1en u-ansmined in
duce infection in a third specie, of mosquito. ho\,ever the succeeding insrars as nymph, or aduh!i. Lxodid ticks at·
intense the virnemia. The exact determinants of specifkity tach and feed O\'er a period of days so 1.hat determining
in vector-virus relation.ships are imperrecrly understood. threshold levels ohiraemia for infection of these ticks is more
but intrinsic factors in the arthropod are genetically con· complicated than for instantaneous feeders such as mosqui-
trolled,91 • 187• i06. :?:?6 , ::i;., and appear 10 operate as barriers 10 toes. Furthermore, ixodfd ticks feed otu}' once in each s1age of
infection at the level of enzymes. membranes and rel! sur· the life cycle and a period of weeks or months may elapse be-
foces.07• 99, 199 · 2oo. 2•R. 259· 200 Following ingestion by an ar- rore the tkk has mou Ited to lhe succeeding instar and is ready
thropod. mostarboviruses infect and replicate in cells lining 10 ~eek a further blood meal. so that transmission 10 a second
the mesemeron. before penetrating 1he basal lamina to be nmebrate may occur long after initial infection of the \"ector.
released into the haemolymph 10 set up further cycles of ln- Argasid ticks tend to have multiple nymphal instars and to
fe<:tion and replicaiion. In 5ome il1s1ances. arboviruscs may feed rapidly and at frequent imcrvals, so theorerlcally an ar-
penetrate the mesemeron without replica1ion.23Q but the gnsid tick could transmit Infection to numerous vertebrates.
concepts of a mesemero11al- or gut-barrier to infection and Ther are also able co \\ithstand starvation for mon1 hs or el'en
a threshold level of virus in the blood meal to overcome the years, and can conscquemJ~, barbour infection for protracted
barrier, remain \'alld. Further barriers to infection appear to periods.
exist at the levels of organs such as the 0\'8ries and salivary ·n1e phenomenon of u-ansovarial tran=ission of vimses in
glands. By definition, a virus must infect and replicate in the arthropods has imponam Implications for the perpematlon of
salivary glands before the ex1riJ.1sic incubation period is suc- the lin1ses i11 nature. Early in 1he known hiswry ofarboviruses,
cessfully concluded and the arthropod is able 10 lronsmit e1~dence was produced to show that viruses may p;1ss from the
the infection by bires.:ui Arthropod~ ma)' gain infecti<m with ovarie<; or inf~-ctcd arthropods. through the eggs. 10 the
great efficiency from the ingestion of a viraemic blood meal. succeedil1g generation of mosquitoes. 142• l 4:t i ;o Licks.58.. 112 or
yet fail to transmit virus because the Infection has railed lO sandflie,. 111. i.9. 2!18 bm im-es1igators were unable to confirm
spread lo ti1esalivaryglands. Hence. it is important in vector the occurrence oftransovarint transmission of virus in mosqui-
219
studies in the laboratory lO distinguish between anhropod toes.6 '- 82• a;, Although some workers failed to demonstrate
infection rates and transmission rates. the phenomenon in ticks. other~ obtained comincing
Replication of virus in arthropods is influenced b) ambi· e,~dence that it occurred on a significam scale with tick-borne
em temperature. and, 1vichin op1imal limi1s. increasing tem· viruses.~· q;, IOI. IO;i. 1:lO. ' '" · HO, 18<\ ,~, . 192.215
perarures ha\·e the effect or shortening the e;,.1rinsic The belief tha1 ttanso111rial tran~mission either did not
incubation period and increasing vector efficiency. 17· ~0 • r.o. occur \\1th mosquito-borne \iruses. or else that it occurred on
S8. 89, 10.,. 131. 175, 25 2. :1.55 It was formerly believed lhat infec- an epidemiolo¢callr insignificant scale. prevailed for manv
tion \11th an arbovirus wns harmlc!s$ for the arthropod vcc ycars. 3 1• • 1· ti6. 182• "32. 256 However, the bunya,1rusel', and pat·
t0r, bm i1 has been observed ll1a1 infection with certain riculatl) the members of th~ California encephaliti$ complex.
arboviruses may produce morphological changes in thesali· were mvestigared most intensively and in recent years many
vary gland cells of mosquitoes. reduce ability to refeed. and investigators ha1·e succeeded in demonstrating trnnsovarial
reduce the longevity of mosquitoes. as well as prolong larval tranSmission or these viruses in mosqui,oes, with filial infec-
de\"elopmem ume after transo1-a.rial infection. 18· ":!. 88 uz. 169• tion catesgenerally below50 percent.6 · 1o1 l!l-i 1. z3.w• .u. 1s. 51 · 5s.
z1e. 22;. 230,250, m. 207 Viral genetics ran also influence mms· l i ':"'"" :l. l:tl, 134. 136. 1$8. J:;9, lf)S, 167, 206. 2 30, 231. 2.;;-2,ff; , 251 ~ T. ::38

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 Oass!tkation, cpid\'m!oloi,:y and control of anllropod-bome,iru5'!5 159

In more limited observa1ions, dear e\1dence has been ob- Tesh:!3"! cxammes the latter possibilicy and dra\\"s an analogy
tained that u.msovarial transmission with 101, filial Infection with inherent 1~mses of arthcopods. in particular the sigma
rates, ge1,crally below I per cent. occurs among Oa1~\iru~es in 1'inis of the fruitfly Drosophila me/1111ogaster. Inoculation of
mosquitoes. 1 t~. s2. s... 63 . .e. w. 88. 89. 102. 111 112. 1,,. 1ss. 16$-190 fruitflies 1,i1h sigma 1irus ge11erally results in ·non-stabilized'
Filial Infection n:ues as high as aboul 20 per ,em were obtained infections In which there may be transovarial transmission of
with one navMrus. 81 There Is some evidence that \'ertical 11rus 10 a limited proportion ofprogen)' ror a fewgeneraclons.
transmission offla1i11nJses in mosquicoe.s may involve the in· Occasionally. so-called stabilized lnfec1ion arises in which
fection of fully fom1ed eggs rather than 1h<: passage of 11n1s there is infe,tion of germinal cells (oogonia) whh sustained
Lirough the ovaries186 and ii is not known whether this phe- mmsovarial 1ransnfr,sion of 1~rus by cytoplasmic inheritance
nomenon occurs with other fmnilies of ~,mses. 1ovinuallyall progenyo1 succeeding generations. Mos1 in fee·
Liule evidence has yet been produced 10 show conclu- rions produced experimemallr in laboratory sllldies of vec-
sll·ely that cransovarial transmission occurs with mosquito· tor-virus reiarionshlps ha\'(' probably been of the non-
borne alphavinises and phlebo1·iruses:'.· 110• 13• lntriguing stabili7,ed C)1>e, but It appears that stabilized infections may
evidence has emerged to suggest that certain mosquito- have been experimentally produced in mosquitoes with a few
borne viruses may be maintained b)" uansmission in ticks, member<; of the California encephalicis complex. with sus-
including transovarial 1ransmiss!on, but the full significance rained filial infeclion r'"dtes ranging from 71 to 90 per cem.1'16·
of this phenomenon has not been deiermined. 27• 14 191· 225 238 219
The existence of such subpopulations of anhropods
Transovarial transmission of viruses by sandflies has been \11th stabilizt"d infection, in nature would serl"e to perperua1e
well docurnemed.~· 67 197. 233. :!!14-237 arbol'iruses. even though 1he infection rates in the 10tal
E~dence of mmsovarial transmission is obtained eicher vcc10r populations remain 101, ..-\rbovin.1s diseases are char-
by testing the progeny of arthropods infected in the labora- acterized by:1>6· is-. •ez. ~;-211 zu
tory, or by isolating ~rns from eggs or immature arthropods • a 1endenc)' 10 occur within specific geographic limirs.
collected in the 1,ild prior 101he taking ofa blood meal. It has ahhough spread 10 new areas sometimes occurs \\ith
been observed that a period of days may elapse af1er the in- dramatic resu lts:
fection of female mosquitoes before infeclion of 1he Ol'aries • seasonal occurrence in periods of vector aetivicy: and
occurs. analagous to the ex1rinsicincuba1ion period for infec- • a propensity to produce major epidemics periodicall~·.
tion of salivary glands, so that virus ma}' be absent from the
progeny of the first OYarian cycle following infoclion. yet ap· Primary factors alfeo1ing the distribution of arboviruses in-
or
pear in later progen} .10· rn,. z1 o Lack of awareness this phe- clude latimde, altitude. topogmphy. geological feacures and
nomenon may account for the fallure of many earlier workers clima1e.11iesc determine the nature of the vegetation cover
to demonstrate transol'arial transmission of arboi~ruses.232 and che availability of surface water. which provide habitats
Since male mosquitoes and sandflies do 1101 1ake blood with suitable rnicro-climate~ for the breeding and ~urvival of
meats. isol111ion of \irus from 1\ild-caugh1 males has also some arthropod vectors £see Chapter 6: Vectors: J\losqw-
been interpreted as evidence of uansovarial transmission. As toes, and Chapter 5: Vectors: C11/icoidesspp.)
an extension of this observation, it has been shown that in- Within areas or foci. arbovlrnbes are maintained by cir-
fected male mosqLLitoes can rran,mlt infection veneroally to cula1ion be1ween vertebrate hosts and arthropod vectors.
females. and that the females can subscquendy rransmit The ,·crtebraws involved in maintenance transmission
vims tmnsov:iri:illy en their prngl'ny.1;s. 1;; 185, 2-i-1 24S cycles are often bird,. small mammals or. less con.monly
Transovarial lransmission would seem 10 represem an large mammals. Possibly as a resuh of Jong association with
ideal mechanism for the perpetuation ofarbovirnses in na- the viruses concerned and the confcquem subjection 10 se-
ture. However, mathcma1k.il models based on filial infec- leo1ion pressure. the vertebrate species involved in mainte-
tion rales observed in laboratory sllldies indicate that the nance transmission trdes frequently appear 10 lack
1iruses concerned would 1101 persist in more than a few gen- susceptibility 10 the pathogenic effects of the virnses, ycc
erations of arthropods ir rransovariaJ transmission were the they develop sufticieml} intense vlraemia 10 serve as a
sole mode of infection.;,, 77 source ofinfcccion for arthropod vectors. In the va~t major-
b1 an incisive review of transovarinl transmission, ity or instances. the ven<>btatcs are 11011rue maintenance or
Tesh 23~ poims out that the above finding suggests two pos- reservoir hosts of virus in rhe sense that they develop
sibilities: chronic infection with persisrem ,~raemia. Jnscead. the ver-
• a:..postulated for some virusc5, tntnso1·arlaJ transmission tebrace and arthropod vector populations collectivcir main·
permits lhe \1ruses to survive periods of vector inaetivity, 1ain virus in rransmlssion cycles in which the vertebrates
such as winters or dry seasons. bu! ampllfica1ion ofYirus serve as amplifier and dissemlnator hosts of1irus during the
is required through the infec1ion oi vertebrates during seasons of vector activity. Such basic I ransmission cycles
periods of vector ac1l\~ty: or usually remain cryptic unless susceptible venebraccspecies
• laboratory s tudies have not gh·en an accurate indication that exhibit disease, including humans and domes1ic ani·
of what occurs in nature. mals, impinge upon them. i\n example is the circulation of

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160 >U."11<'~ a,r: Aspe<:1$ inlluendng the occurrence of infectious di~eascs

\~ruses ofthe tick-borne encephalitis complex of rhe north- is vertical strarilication of habitats in fores cs as weU as hori-
ern hemisphere in fores1 ticks and rodents. It is generally in- zontal distribution. and it has been shown in Uganda that
ferred that voriebrate species 1ha1 are prone to developing AedR,afr;r-Cmu~ mosquiroes. which feed in the canopy layer.
disease have. in evolutionary terms. compararh•ely recem rarely or never bite at ground level.1 12 These mosquiroes,
experience 11ith 1he virus concerned, and that they are dead- and monkeys in the canopr. can maintain sylvan yellow
end hosts that play no part In funher dissemination of the fever which may be communicated to humans b)' the ar-
virus- in otherwords. the occurrenceofinfection ih them is thropodan link host {Aedes si111pso11i). which bites at inter-
incidental or 1ange111lal 1q the basic transmission cycles that mediate levels. People who enter fort'Sts co hunt or fell
serve ro perpetuate the viruses. This is particularly true of timber acquire infection from Ae. simpsoni and themselves
1.he sporadic diseases most commonly produced in humans serve as venebrate link hosts by conveying the infection to
by arboviruses. bur livestock more often contribute sign'ifi- centres of human habitation where tta11smiss:ion cydes of
cantl)' to the perpetuation of pathogenic viruses or to the urban yellow fever may be iniliated between humans and
generarion of epidemics. For examp_le. sheep and caule are the peridomestic mosquito CAedes ae~7;ri) which enters
thought to play a key role in the dissemination of virus to dwellings. In southern Africa. Riff Valley fever is transmitted
mosquitoes and mechanical veccors during Rift \'alley fever by zoophillc mosquitoes. and h umans acquire the infection
epidemics.93. t-19. 247 prindpall)' from comact with the infected tissues of live-
The roles which vertebrates and anhropods pla~ in the stock. 154• zzi Consequendy, human infection in the subcon-
epidemiology of an arbovirus disease are compounded by: tinent is encountered almost exclusively as an occupational
• the nature of their infection: hazard of veterinarians. farm workers. slaughtermen and
• their beha\~oural pateems; and Others engaged in tile li\'estock industry. On the other hand,
• population factors. in rhe first outbreak of Rift Valley fever 10 be recognized
in Egypt in 1977. amhropophilic mosquito vectors were
Thus, the duration ofviraemia in a vertebrate above thresh- invoh•cd In the spread or the disease co humans of diverse
old intensity for the infection of arthropod vectors affects occupations_lGI 1~ .Many haematophagous arrhropods,
the number of arthropods which are exposed to infection, paniculadrdipterous insects, tend to bite a1 set limes of the
while the duration of the incubation period in the vertebrate day. Some arc diurnal feeders, 01hers are nocturnal. while
and extrinsic incubarion period In the arthropod contribu te crepuscular insects feed in rhe twilight hours of dusk and
to the length of the intervals between successful rransmis- dawn. Ir :,; imeresting to note that long before it was known
sions. Once a vertebrate has undergone infecrion. ii is usu- that crepuscutar Culicoidesmldges were invoh·ed as vectors.
ally immunologically 'dead' t0 further infection \\1th the it had been observed b~ farmers in southern Africa that Afri-
same virus and. for persistence of infecrion to occur in the can horse sickn~ss and bluetongue could be controlled by
area. it is required that the rate of replenishment of the ver- stabling animals at night and releasing them after the de1\
tebrate population 111th susceptible individuals should be had evaporated in 1.he morning.
commensurate with the rate of transmission of virus. Birds The transmission of arbo\'iruses ls seasonall)' incer-
an.d small mammals such as rodents offer greater opportu - rupted by the suppression of vector ac1ivlry by rhe cold win-
niries for the circulation of virus as they may occur in large ters of temperate zo11es. or by the dry seasons of the tropics
tlumbers and are liable 10 popiJation explosions under and subtropics. Prorracted dry seasons occur in much of.o\f-
favourable circumsurnces. Small mammals often have re- ric;a and both cold and dry condftions prevail during wimers
stricred home ranges wirh considerable overlap between in- on the inland plateau of southern Africa. Viruses transmit-
dividuals, and 1his leads 10 high population dens!Lies,90 ted bi· arrhropods may sun~ve dry seasons through the
which favour transmission of virus. Furthermore. they at- transmissicm being sustained by limited numbers of the vec-
tain sexual marurity rapidly, give birth to several young ar a tors which cominue co breed in restricted sites. such as in
time. and may produce several liners per annum. so that the tropics where mosquitoes breed in water conserved in
there is rapid replenishment of the population \\it.h suscep- tree-holes. 83 Alternatively, viruses may disappear entirely
ribie individuals_ Migration of birds and mammals can also from an area during,\incers and dry seasons, co be re-imro -
serve to replenish the susceptible host population of an duced larer by migrating or wandering infected vertebrates
area. or 10 inrroduce (Or$easonally reintroduce) \'iruses and or arthropods. ·mere is evidence. for instance. that under
ectoparasitic vectors such as 1ick5.91• 99 Habitat preferences certain weather conditions involving favourable tempera-
and manifestation of nocturnal or diurnal acti\'i.ty arc vcrtc rw:es, humidity :i.nd sustained \,~nds. infected rnosquitOe$
bra1e behavioural characteristics which influence their ex• or midges may be carried vast distances at high altitt1de to
posure t.o arthropod vectors. be deposited in a viable state at a remote site where they ini-
Haematophagous arthropods manifest host preferences tiate outbreaks of infection and dise.be in i·ertebrares.201 It
and other behavioural characteristics that serve as mecha- has been suggesred that vectors of ACrican horse sickness.
nisms thar isolate chem from panicular vertebrates while bluetongue and Rift \'alley fever could be transported in this
bringing I.hem into contact with others. For instance. there manner. and that mol'cment to-and-fro on a north-south

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 Ch,,<ificmlon, Cf)id1,m,olog)' and conrrol of anhropod-horne ,irusc., l GI

axis in Africa occurs in associaiion ,,ich sea.~onal movement the harbouring ofwes1ern encephalitis virus by\\ild birds in
of the inter-1ropical convergence 1.one!01 - 2~ A variation of :S:orth America. ,11:11-iowe\E:r. cattle may harbour blue1ongue
lhis mechanism, in\'olving movement of virus only. i, postu- virus in this manner. 139 A variation of chronic infection is
lated for the perpetuation of yellow fever in the vast tropical the suspended or latent infection. which appears 10 result
forests of the Amazon River basin of South America, where from the low~red body temperature and metabolic pro-
large popula1lons or mosquito<.>s and primates constitute a cesses and delayed Immune response, \\'hich occur In such
continuous substrate through which the l'irus can mi- animals as bats. hedgehogs and reptiles that hibernace.:;:i.
grate.208 Within a gh·en area. epidemic infec1ion soon leaves r3 i.iJu. a.H-2 ' 3 The infections are activated and run their nor-
most primates either dead or immune, and yellow fever mal courS'e \\'hen 1he animals return io a.he active scate.
virus is perpema1ed by Lhe extension of infection from the Finally. allowance nmst be made for the possibiliLy a.hat
periphery of the focu~. to circle back at an interval of years \'intse~ may persist in hit!wno unsu&pecccd arthropod and
\\hen the slow-breeding prima1e population is replenished vertebrate hosts. aquatic organisms. protozoan and mern-
"ith susceptible animals. 1.oan endoparasi1es or even plants, some of which may
Between the extremes of coml11uo1.s transmission and occur in the food chains of currently accepted prim('lry vec-
comple1e disappearance of vin1s from an area, several tors and l'ertcbrate hosts. 18? There al'e examples to indicate
mechanisms are poslUlarnd for the oventimering or hiber- 1ha1 viruses may use more than one mechanism 10 ensure
narion of arbo,~ruses through the occurrence of persistent ~urvil'al during perio<b of inactivity of prim~ vectors. As
infection in either arthropods or vertebrates. lnfected ar- an example, mention has already been mode of the possible
1hrop()ds may enter n stale of diapause as adults, and hiber- trnnsmission of mosquito-associated ,~ruses by ricks.
nate in protected niches or resting ~Iles with favourable In many instance,;, arbo,irus activity does not merely
micro-climates. /\lternatlvely. they may diapause anti hlbcr· abate for the durarlon of a \,inter or a dry season. but the
nate as eggs or ocher immature instars. Ovcrwincering of di~el!se ~eems to disappear from an area for an interval of
,'irus Is readily explained in ricks, which e,·en without dia- many years. only Lo reappear suddenly in epidemic form.
pause may spend weeks or months away from hosts be· Rift Valley fever is n classic example in which there is dcbrue
tween the feeding of successive instars. For female as to whether the viru,; persists al an uncecognized, low level
mosquitoes to sun'i,·e hibernation, LllC:} mm,1 cease O\'arian of acthity in those an·ru. where epidemics occur, or whether
actidry in autumn and deposit reserves in the fat body in- the, inis di.~appear~ w be reintroduced 10 the areas during
stead - a physiolo~cal state which is triggered in a propor· ep.ldemics. rr.119, 2<1.,. 221
lion of the population by being subjected 10 shortened day rar1ors which precipitate epidemics include climatic
lengths and cooler temperatures from the larval stages of events such as the occurrence of partlcularly hca\'y rains
de,·elopment 011wards.66 Failure 10 understand thcphysiol• which fa\'ounhe brccdingofar1hropod veccors.59 M~.:m or
ogy of hibernation ma} have contributed 10 past failures 10 unduly warm weather whkh shortens the extrinsic- incuba-
demonstrate the phenomenon in the laborotoJ)'· tion period of vlruse~ in arthropods and increases vector ef-
le is a prerequisite that uanso,,arial transmission should ficiency 106· 107• 151 Furthermore.good rains may be followed
have taken place If virus is 10 hibcrna1e in eggs or immature after an apprnpriate interval br p1)pulation ex'J)losions of ro·
arthropods. Flood1,111er-breedi11g aedine mosquitoes. dents or 01hervcrtebraw hosrs of cctoparasites and ,in1sc.s.
which hibernate as eggs in dried m11d. may be involved in as a result of the availabilit} of incr<'ased food supplies. The
the o\'erwinrrring M !lift V:illc.>y fever vln1~.~· t:I, F:iilur.- in eli<•r1~ or fn,•1\\1t11hlt• ~r:i,nn~ nn vim~ rr:insmi~sion mil}' h1>
demonstrate \irtas in hiberuncing adult or immature o.rthro· enhanced if imersperse'd \\ilh a succession of unfavourable
pods in the past could have been because ,<irus wa:;. only or drought sea.son~. creating an unstable situation thmugh
present at low concentrarions or in forms ~uch as tempera- the lowering of 'herd immunity', Although lick-borne virus
ture sensith·e mutants. \\'hich arc not readily dewctable by clisea~es sometimes occur as epidemics. the long imen1nls
convemi011.al methods.6b It may be necessary to ~tJmulate between tick blood meals and the utilization of small mam·
the preliminary replication of virus by warming hibernating mal hosrs with rapid reproduction rates often lead 10 rhe
mosquitoes and allo"1ng them to cake and digest a non- situation where ciclc·borne diseases occur sporadically in
infected blood meal before anempting to isolat.e virus. 13 It fixed foci which remain infected indelinitely.2oa B}'contrast,
has even been suggested that, through the mediarion of the high biting frequenci• which occurs In ombreaksof mos·
reverse transcriptase. the genomes of R;l!A arboviruses quito-borne diseases usually leads to explosive epidemics
could,.bccomc incorporated in arthropod host cell D:--:.\,170 ,,•hich ~ub~ide rapidly once most of the ,;usceptible vcrle·
and, although there is linle e,1dence 10 support this hypo- brate population has undergone infecrion.
1hesis.J2 iL fuels speculation that some arboviruse-s had their Human activities frequently have an over·riding influence
origins as components of arthropod cell~ on the occurrence of arboviru~ disease,. and, of these, land
There are few examples of thronicallr infected \'ertc· millr.acion probably has the most profound cffecr. Primary or
brates cons1iruring true maintenance or reservoir hosrs of Indigenous forests tend 10 have a rich and varied flora
arboviruses with persistent orimermiuem viraemin. $UCh as and fauna. which become progressively simplified wirh

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deforestation and dh·ersion ot land usage to secondary for- Two examples im·olving Rift Valley fe\·er suffice co indi-
ests. pastures or monoculture of cereal crop~. until a single cate the impact that arbovirusc~ may have:
species of rodent may be the don:inan1 vcnebrare:~ ln the • according 10 one estimate 500 000 ewes may have
proces~. haema1ophagous arthropods and arbo\1ruse& may aborted in the 1930 to 19SI epidemic in South Africa and
transfer to new ho,tl\ with potentially serious Implications. 50 000 to I 00 000 adult sheep may ha\'e died; 198 and
Edge habitats. such a$ Junctions of Core-SI$ and fields. ha\•e a • the official es1i.ma1e was 1ha1 there were J 8 ooo human
pankularly high potential for I.he interchange of \'ecmrs and cases of !lift Valley fever in che 197T to 1978 epidemic in
vinises. 11 Egypt. with 598 deaths. but unofficial estimates were
The final lngrn<lient needed for the occurrence of out- much higher.
breaks of disease is the introduction of susceptible popula-
tions of humans and livestock inro rhe ecO:.')'Stem. In the Outbreaks of arl>o\·irus disease are tcrmi nared when:
past. major socio-economic uphl';l\'als. such as \\'ats and • most of the susceptible vertebrate population is eithc>r
.
laod resettlement or rcclamarion schemes. have somecimes
provided the influx of humans and livesrock which triggered
dead or immune lthat is. when there is so-calico immu-
nological exhaustion or the population :
epidemics or led IQ the emergence of prcviouslr unknown • fac!Ors whkh precipitate or facilita1c the outbreak are re-
pathogenic vinu;es.'lG. :to7, 210· 221 Large cMl. industrial and versed narnrally. ~uch as when the onset of 1,tjmer frosts
agricultural engineering projects. such as thl' building of in southern •\fric:a terminates outbreaks of Rift Valley
dams and irrigation schemes. sewage plants. power stations fever. bluc1ongue and African horse sickness through the
and factories "~th effluent waste water haw also brought suppression of vector activicy: or
about cha11ges in the juxtaposition ofhuman. li\'estod.and • d1ere is effective human intervention. such as the admin-
wild vcnebrate populations. a11d pro"ided breeding sites for istratitm of vaccine.
arthropod vectors to trigger outbreaks of disen~e. Air, ma-
rine. and even land transport provide the means for the
Control
worldwide dissemination of arbo, in1s-infected vectors and
\-Crtebrates. including humans. The planning and institution of effective comrol measure;.
At the levels ofindividual humans and households. occu- require a proper understanding of the factors in\'olved in
pational or recrearional exposure to arthropods, livestock or the epidemiology of an arbovirus disease. The complex in-
wild vertebrates will inctease the risk of acquiring zoono1ic terrelationships between viruses. venebrare hosts. arthro-
arbo\!irus diseases. as does living i11 a rural en\'ironmem, pod vectors. the environment, climatic factors and ac1s of
particularly in housing with poor insect-proofing. Humans human Intervention are amenable to logical analysis 111
gain infection either from the biles oi arthropods or from and mathematical modelling. 76• 208 Frequently, disea;e
contact wl!h Infected blood and other tissues of domestic represents the end product or a minor and abermm path·
and wild vertebrates. hut \~ruses which are pathogenic for wa} for the virus concerned, created or facilitated b}'
humans do not necessarily produce disease in other ani- human action~. It has been stared that for the epidemiolo-
mals. For example, Rift Valley fe\'er viru~ causes disease in gist. in contrast 10 the clinlcia11. the occurrence of dbease
sheep and caule. while Crimean-Congo haemorrhagic fever has no significance other than the implica1ion ni its
virus produces mild or inapparent infection In these ani- recurrence. 141 ;\,teasures for the c:ontroi of arthropod-
mals. Both ate important human pa1hngcns. Livestock un- borne virus di~eases may be applied to the susceptible\ er-
dergo inapparcm infection with rick-home encephalitis of tebrate species, venebrates which serve as maimenance or
Europe. bm serve as link hosts \·:ltkh transfer \~nts from link hosts of the virus. and 1he anhropod vectQrs.20R
cryptic circulation in ticks and rodents to the emironmcnt tn many respects. protection of the susceptible verte-
of humans. They also serve ro amplify and disseminate in- brate species through vaccination appears ro represem the
fection through human consumption of infected milk.208 most practical methods for controlling arbovirus diseases.
Pigs are link hosts for Japanese enccphalitls virus. They in- Mowever, despite the availability of vaccines, arbo~iruscs
fect peridomcstic mosquitoes. which in rum infect hu- such as Rift Valle) fever continue 10 produce massive out-
mans. 19' 1 /\rbo\iTuses such as dengue and yellow fevers. breaks of disease. A major pan of rhe problem is that it is
which catl be propagated b)' circulation between humans difficult 10 111011\· ate and sustain regular use of vaccine dur-
and mosquitoes. can pose a threat even in urban siruations. ing !mer-epidemic periods which mar last many ycars.221
parricul;rrly wh<.'rC mosquito breeding sites arc afiorde<l h)' ~22 and ii ic. equally as difficult 10 predict the occurrence of
poor drainage or water held in discarded containers. outbreal(s of disease sufficiemly early and accurately for im-
Tite populatic111 explosion and drift towards urbllnization munizauon 10 be applied effectively in inscanct:s where it
ex-perienced in the developing countries of Central and South has been neglected. 1l~. zzi. 22i Moreo,er, arbovirus disealies
America, Africa and :\sia over the past few decades, has proved sometimes ail pear unheralded in areas where they ha-.:e 1101
fertile ground for arboviruses and the literature abounds \\ith previously been kno\,1110 ot·cur. as when an explosive out·
descriptions of massive epidemics,1°, ~,o. 216· ~ 1t break of Rift Valleyieveroccurted in c~1)tin 1977. 1t; 1 Some

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 Cl,1,.,itka1ion, epldeminlogy and tomrol of anhropod-bol'f'~ ,iruse< 163

,11ccines have inherent problems or efficacy, such as the dif- Control of vectors include;. selective clearing and drain-
ficulties encmmtered in obtaining immune re.~ponse 10 all age ofland to ahcr habitats that favourthe development and
c,f the serOt}'J)es of 1·irus included in multivalent vaccine.-. suf\ival of haemntophagous arthropods. Laf\·icidal m.'llt·
like those used for bluetonguc and African horse sick.nt)ss.'~' ment may be applied to wacer where vectors breed and in-
Some vaccines are only partially atten uated and can cause secticides may be up plied to resting sites of adult vectors in
problems wllen used incorrectly, SllCh as live Rift Valley bulld!.ngs. Acaric,de~ may be used to treat 11,·ei;mck For ticks.
fo,·er and Wesselsbron vaccines. which can be tera togenic However, care should be taken 10 minimize pollu tion of the
and abonigenic when used in pregnam ~heep. 16 A further enviromnent with tc>xic chemicals and their u,e should be
problem i~ lhat vaccines ar<" not universally available for all combined with ma11agemem systems de~igned to limit vec-
arbo,;rnses of medical or veterinary importance. Jn some tor populations. For example. in1ensive grazing uf pasnires
Instances. such as Crimuan-Congo haemorrhagic fe,'l!r of over shon periods, eombined with effec1i1e acaric[de treat-
humans, lack o f potenrial dt•mand inr.lbiLS the product ion ment of llvesiock, can be used 10 reduce or crndicaie tick
of vaccines, particularly in view of the e~enses incurred in populations. Howe, er. efficient vector control may lead to a
meeting stri11gem safery and purity requirements. potentially ml~table situation through loss of herd immu-
Palliative measures for the protection of humans from nitr. Biological methods for the control or \'Cctor:. should be
anhropod-bome viru~es include insect-proofing or the considered. such as the use offish or bacterial agents co con-
home and the use of mosquito nets and insect repellents. trol mosquito larvae in water.
The ~ame principles can be applied to livestock.. and pos- Finally, measures 10 c:ontrol arbovirus diseoses include
sible measures include 1he housing ofnnimrus a t nigh1and 1he promulga tion and application of regula1ions 1hat re-
rhe use or highl} effective insecticides. During epidemics quire the produc1ion or health and vaccination c:crtificates.
which are caused by \' iruses rransmiued by mosquitoes or ectoparasire treatment. quarantine and ln$J)Cetion of im-
midges. it may be beneficial to move livescock from tow- ported vertebrates. This includes nor only susceptible live-
lying marshy areas 10 high, windswept ground. stockspecies, but also animals such as cage birds which may
Vertebrates " ·hich are clas~ed as \ermin or pests. and serve as reservoir host~ of viruses affecting humans and farm
which ser\'e as maintenance or link hosts of arboviruses. can animals. /.lo\·emem permi1s. vacci nation certificates and
be subjected to specific population control measure,. Th~e ec1oparasi1e control are also common requirements for the
include the proper handling and ~torai:e of crops and feeds translocatlon of pers and livestock. within coumries. lnspec-
in the vicin ily of homesteads and stables. Livestock which tion for. and control of. arthropod vectors and vermin , may
serve a, link hosts can be immunized to prmett humans also be applied to aircraft. ships and ~u rface transpon
against zoonotic diseases. an example being the immuniza· vehicles on international routes.
lion of milk cows and goats agamst lick-borne encephalitis
in eastern Europe.

References
\fl'Jt.r,. T.tt,("... il'<H, R 7' Rf \1', M. ,-,, •-:n"'"'· t, 1~f fn1thiWIHfal ,qr.q -\rbrnina. n,)"mr, A,,w•frn,, Jru,mn/ (lfT"Pflf.~rti \trdiri"rdt)d
1ran,ml,>lon of ;,·llow fe,·e, ,in~ by mc>"(juil<><'> ktfr.a,-g,111[), ll;g,tn<. IS. ;JJ-i34
.-lmrlkiJ,i /011mal ofTrop!Ntl .\lnllrl11, ,md l!Jtl,•,11·. ?R, t 1g..z1z g A'-OX\'\.!Olh, :97n. Subt'nmmht\'t• ,,n lnfcmn:iucm f..'<eh:tni:t~ u! dtt-
'2 .,rn::r~. T1ll,C'... \\()(H>>\U... 1.1,.. ,\~fH1+\Dr.•.ui.~....... ~,.\nrrtl, ( .. 5, "illOf>•• Amtricar. Commtu~ on ·\nh:opi,J-bc)me Viru...~. :9':0.. Caiolob'tlt' o(
n.F•• 1~a. P:u:ul \itu:),_ phlcl,nmmilw Oh,."'\ .md ...m.ttll mammal~ in Unuil. anhropod-bn,m.~ \'int~""' or th~ worlJ.•.\mttrlctm J1,ur1ml ,~}° TmplL·nl
Jn c1>ldcmu1i01,,fe-.il study. ~m,rltmr Joumal of1ropico/ ,\ft'dl(i11c n11tf .W,'tllri11rm1I. l/lt;11•11,•. l~{~uppJ.,. 1081-1160.
f/vgir11,. 24, 358-36- 10 \.....:0\:\ '1tllJ'.'I. -~,a1.1, YcJlow ((l\"t•J in 198i Weck(}' tpul,·•mfolog,tnl lb.,'tOttl.
J 1,,nx,~r,lln. tt-,\.. 19.;t. Rift Vo.11r\' fever in rh~ Unr0n. fournal of:lu SIJ1111, G.i.J;-13
.,t[rit.tm \.'ttlt."t'ir.nry .\!Mfet.11 .~lllr(o,, 2.l. 10~112. 11 \troY. J.a.193& lhc toC'.lli;,,.ation of di.,casc \i.ith special rc-r~n:nr1: ro the
.\l.C(ASJ)LA. n..\.., 1957. R{ft \'all~ re,·cr. Pr(J(HtlfnJl$ of:Ji,, f'uuflli
Mt~i!r.g lOOno~ Tronsac1lom of 1f:1• Royuls«,~t..''O[froftit:nl .\frdlcith· and
ofrht lnurr :\fricm1.Ad1•l.u>")· Cnmmltlt.'t on Epo,otl<" DiSM.'tl'$. (Jt1kdr. Jf_vgler:, 32 Jllll-328.
JO II ~lay.
J~ \UOY ''"· & H,~1usos, ,,.,. 1-g,51, A re\1tw of inn:1-ugattons on mne 1yphus
£ .,'\nRt\,,, c·.H .• 1910. Generic; names 0C,1nr~c-,..n1 vcntbnurs nro/o,r.·. m aurm.2 .and' ~3Jnyo. l'l.tS.-1930, 1ra.111ac1/IJ11$Cf 1J1t~RO,\'(tl W(Ny,;j
-JO,lOi0-10."1. Tt'i/lit'lll M1"tl(Om:111ut f/J-g.ft:nr. f-i. 3; J 395.
t ,\S't"ftl \~ \\',~... PIOWt.n , 'N.A., \\()N(;.. v.w .• m,)Pt~f.Y. o.e,"' lf.\UI rJt, \\ . j,, l.!5 MIU\', t:.L, t...lHUlH,t.. IS.l•, 11\\l.., 11 ll., WAl'b-, t),\l,, 1'M\tM\UUtl'), U,t h
19~ 110,larmn oru,,imu\ti \'bus from l.1f\'ilc ..ind aduJLs rc~rc.-d from l>\IJ<>'\IPI.E. 1,>1•• 1978. Jsolorlon ul ~, Lou,s cn<-0pholl1l, 11rus from
fiold•collc.:tcd l•r.= 0(:1.,,J,., rrfirft1nw, <IJiptom Cullddae). /ormurl tJ/ o,~el"\,'lnttring c:uk.tpl/nt·11$'mQ,qllho,e;s. Sc1c1iC~ 199, 114'r lJt9.
Medico/ ~ntomol(lfJJ·. 13. 699-iOI. u l\.\ADO~. , .• U'l'R.\ J. • ttUI.\UR, l- .f. OLU~!C"."l1K, I, 1,9:'8. \1rologic:d
; ·'"ow,,ou:,. 196; World Hoalrh Orgt111!uuon 1967..~rllwrn,.li',, 1111d rxa.mimuion ormo'-.qulfo l;m.·ae- from .)outhem '.\h1r.1\i.:s. Ftitf(I
Huma11 Ofanu.'I. T«hniml R"I""' ~..,.,, :-.o.Jt'9 Pnn1.1!t0Jog1(11. ts. :S.-78,
,. A"0''l''H'tU'- , ~ Wl*:1<'.'.-1" Commiu~.1.•011 ,\nhropod-bomt- \'iru .. L"'t ·~ ftAR~\RD. a, H 1.. "\I\'\,"' ti .. rm l 1f;EE%.. J,11., Gftr.\'Ll~C~ '-'"· 1w \1, ,~•. 11.,..

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164 'l'~ to~ o,i: A<p<!Ct< lnllucnclng the o,·C1Jrrcncc of inli'ctiou, di,eM~<

l{)80. Turkey menfngo.cnceph~Htl-i In Smnh Afn.ca, OmkrJU!/J()Ott JO C.U.\t.s.. l. •·\\111n1t\,, t.. 190(). \ nt.\, anti~enic group ofuz:hropod, bom.t
/OUrtttll of V,!lfflllD'}· RtJ4.'lltdr, ,$';. 89..g.; \'U'tb-C1". Thit Bul\\Jm..,~r.a group. :\mvrlctm /omwll o{Troofc.·at j\Jcdldn~

n, DMlS-ATtD. e.1.>1- ~, oct:s. !>.r .• 1986. ffa\1,·ln.i.)C> in :,outh Afnca: nmt 11.t·Fir'lh.'. ~\. 13-ii
diaATIOShc procl-durt:) 011d,·r:•te11oort Jo:mm/ CJ/Vtti!rma0· Ht"St11«h. 5.1. 3i c.,'-\U. ,. "'\\ t11r \1,,x. ,~ 1961. GiOUJ) (.:. \ OC'\\ !'l.(rolog,cal poup c,(
181-185 hnhc-.no undt•st·til>t•d .inhtopQd·bom~ , , ~ lmr:1unulogi4.·uJ ~tud1es
n '"' .,..,,., M.,.. ROC.\·O \HCl1', ,,•• 1'946, rhe dc:~elopmeni o!llw \·lnb ot t.cl!0\"1 .'4nh~rttm1 Jo1,r1ml OJ Trosu(tJ/ .\lt'd1rim! anrl Hygft•l~. 10. ?SO-Z58.
rei:cr In flu,m,w,gus mq,quitorl, A11wrt«111 Journal of lroplcal J8 ('.JL\\J..fiOWX, H.\\ , t tlMllt..,l'A.'\, t.c. 6-. 1,Jn-'". H,J.,. l'.tS4· 5tt:dicir; on the
.\/,d/c/11~. 26. 585-u0.5 :Sonh \mcrlcan anhropod-bomc cnc,·ph•l!ude~ The e>trln>k
18 BtAn. u.f .. nsu. Jl.ll. a. ul1'is. 1 1u;,, 198~. l rtt.lU0\'3.ri3! tran~.mt,'".:on of fncubat tin or t:a.""tcm .md \\c.,t... rn-t:"quinet•nccph:1JU1:o. m mo-squiiOl>:.
)t.'lln\\' fo~l\r \'iN-i In Stt-gtull)"Ia mo~quil06-. \mttfitwl Jot~rtw! o/ Amf'rir<m Jmtr,,(l/ ofJ(;g,,11,•, 60. U~7
r,,,,,1r11l Medlc!nr m:d 11,1;1,,,h,. 29. 125-:3: 39 O.f:-\\ Ultt.L\I~, fl\\',, .)J).b .. ,>.., ~l!L<C,,'\', O.ff.,.,, ":IJUU. ~,.o.. :954. S.n:Ci~->
19 aMn.1u. ~ n .1o~tPSO!\, w.11., 1975. Emertc.ncc of l:I Crosse, iru:. 1tom on 1h• :<orth \mrric.u, anhropod-bomo cnr~phahll<le>. QW\nm•mc
endemic foci . .-im,1rlcar: Jaurm,I o[Tropi,11/ ,\f,•tlu,.·lne mu.I Uy,:l,mt!, 24, dett'nnln.a.cioih or ,ini~-\·~ctor n:lJtion,hip-s ..-\111,·rirno /mu,,aJ of
685-69 1. H,rgi•1"'. 60. l78-~8S.
20 eutv. RJ. & nroMPSON, •., .u •• \976, Df.,1im·al1Qn ortn Cros~l,vinn in 40 Cll,\..\lBUL\I:\, k.\\,6-!~UDL\. w t>.. 1915. tlTl"Cte. tJ! tem:ptr.1.1url' npon

de;\·~lop1mm1al stag~ of mm~1>vamilly mfeCh.'ClAtvfes rris,•ruuu,f. extrln-.ic fnn1h:ui<m ot" l':J.;H:m 1.·qulrtu unct!phahd\ In mt')t,quit<>cs.
...tmrriran Jo:,mul ojTmJJfctll ,\lt'Clfcl,u,,,,,c: Hygftm•, 25, :iU:t-512. k.th!ri.c,.111,ntrmll of u-~g,,,,,,., n2. 295-JCQ.
'21 GJ;UO~<:tl:>..~,. f*OUU~. t.. )lAfnl, j\., \UBI'-. \., •\U\·t:t... \f ~ JOU!o>IT, f .>. .. .i1 tH.,~t1hJtu1,. R.\, . •.su01.,. w,o.. 1961 \t«han1,m uI :.ran,mlssion o!
1981.,\etl\11} afCcllfomfa <'t1,"ephol11h llJUUI> virules in tmrel,< ,1ru~t'S b~· mcnqufl0t~~.Annu11J Re,·ieu•o/t.:nrtJmol()J,r,~ 6. ;i/1-390.
{Province of Quebec C.nadro. ca,111d/o~ JounUII tJ/Mlcmll/0/<1~·. :?ll, .;.: CHO\\", T.l,.,QIQ\\ . f.tt.""·"-SO:'\, RP,, 195-.~ Morphot<Jgyohesicu!ar
Si2-S79. .stom:m, ,·1ru~. Jv;rrm,! o/&«t,·rtoft,;(J·, 68. 7~,1.::2u
2!! ftt:Her, ·r.o.. 19':'S, Jiuttmarfo,wl C:O.tolosutof.Vl>otJ1ru~. OHF'.\ u (Jlkb~ ......., . "·"·· tlU\H.d \\' .\., WON<i, , ..,, .. UOH,l.l u.c.,i IL\U'.iUft,
PubUc.itlon N<1, ·.CDC! ;;;.a.101. 2nd udn w.,,.t,lng1nn, l>.C:a U '" w.1 .. ,~,R I ..::aboruw11- "\tUd~). oi trn1isnv,.ufal !tit.n,miv.itoo oftr1V'dHarn~
Gov,,rnmtn1 Printing Oili'co i,iru, h) A,y/,•5 rrlm1tt11us. •4m,,ritdll Journul o/Tru1,foal M ,'t.l~·i11, mul
-23 ltUUC:Y, A.L, LA LOXl)A WlilGEkl, tJ.I,, C:..\USJJ[~. C.H•• PA.JbOSS, \1,,\. ¥t MJAR, J(tg,,•n~, Zi, HU-tRfi
t':.T•. t9T,. E\'id~nce for tntn,.Ma.rhd tr,msml~sion of J:nnesmwn c.,n~·on ;..t ClUl\l \'M)\.", \l,J•., PrtllO\".\, '!>..I' .- W '\1).\1.., \'•., .. t'}-;';. ~,ucf~ nf uhri.a1.irL.:.).
\'1tu.s in Ohio. Mo.,;ffuiro Seu~. 37 .;94,-;St1 ,n,ephalhl<. VII •.\rtlfftal .,daptatJon o! :ht ,iru, c,f lit~ ond J~p:in=
:U 1115HOI', J).)f.i.,CAU:,11l£H. c.u .. c.:.\S:\U, 1, (.J(U\t\li.{J\', ,,.P.. GAJOA\1Dnc11. tnctphalld't m \·:srio::, -.pl'<lt.>> of tick.., of the fomi!\ J,cc,dJJ,11."
S,Y,\,, llA.!\XOU:"lo" C L\'O\ p.)i .. ' ' '"~it.UL, t..O. o~~-nw~t ..... ,\!i'tlltlln1Jt.(ly" PtJrdutofogiy11 I Pmh:.i1u1n:...,• &l!r:.r:t, J.I, IU--24
Pll111f<SSO~\ A,F PORTUU-JW>, IS.., RU5StU.. P ..... .)1t()PI. JU 4 WE!-1,\W.\1:. ;;5 (.L\RJ;;.(i..(.,,, rnrrut..\ H l... ROI-UU.R, \\.11., lt\fllfOH, \. , J,\);'tJ8CJ\\'S£. r ..
t.c.. 1980. Bun}·a,irid:ic. Jn1,m•Jroloi:;• 14. 12s-1'13, 1!NJ:t. PL·~"'l,l\,'OC',, at 1,a ~ ) t o vinl~ tC,11ifprnJ:s l'nl·uph,tllli5>---.0tO"droup ·
::5 tu.ACt.HtJJt~. s.,-., J~q.. N,tti01utl ln,tlrutt: of\ ·1ro1uto,. Priv~tc &~ X-i. in \'orth~Ccmml lllinol~ ""'"""'"' Jrmrm:I of I ttmlnrl ,\INfftill•'<11id
l>.ln\lrlngham. Johunntshurg, 'iouth Alrka P~rs<>n:tl c0111munlcmiu11. ll,,,iw11,, :12. 11,.18-1,
i& 1n..,c-~uu1t~. s.~. "'- ~f--\>U.t<. L. 1935-. \1ru!les, i~J:itl"tl frutn C:ufto,;t/t"f ~6 COl''t/,U•, I,\ w, & U \k.~.\t\O, b.f .H.. Jr.i, J-ly<ffOJ,S lllllllli ll\ JIJ,1,.ie,r
.Oipt~m: C,.mopogonldac1 e,,u[th1 at 1h, ve:eritUI!}' resea,ch I.um. tt.~),f)cinr~d \\ Ith 1,ydr11n-..-n('l'phaly anu 3rth:ogrypo~i-.. ,,,ilh \\{"\,t•{o;bro:t
)!""°"""· Z1mbabw•. /01mtnl qflh~ Enlon:oltlg(rn/Stxt,•t)·t>/Sau,h disc~ iltltl 1-tifl \';dlt>} f1.•\·l't \iru,.c:s ,1~ :JcltuluR,it.:J.I ,1io:t't:!s Oml1.•rs1.:;'1()(nt
.~Jrlco. 48. 331-336. Joumu-Ju.f\!t•ll'rinor.· /ltSl'tlrrh. ~.,, ! 19.
~, .BL\TrXUI. R.J. ~ H£\'.SS. l'.\f,. t(UJ. B!ood,RJcking \'et.1:Dr.i ol tnr~ph.tliris: .t; COLU~5 w L,. 19\>2, Tt.tll\•.,1.1dlal ptbSO{tC oi 5t Lou:> C11Cl"J)hUUtls \1ru~ ;n
F.'l)cnm<:nt:iJ tran.<n1Jso!on of St l.ouJ, cneeph:iliti< u luhh,rd stra1n1 ,o !•ul.es r~pu mosquitoc..• lmqrfm11tmmull of Tramm/ \/wlt,11,ea,11/
,, hiw S\..iss m!ct.t bl' thl! A.JncrkWl dog tk.b... D<mnaet:mor t•tJtlnbllo Sa~. HY)tl,,,tt•. 11. l3!>-JJ3.
Joumnl ofE::qx•rfnumltl/ .V!t•dlrlnc1. 79. 43.9,.....;5.;_ ~ COU.J\) \\',f.., t!ii)J. r,ao.,ml~1ooofS1 l.ouh.<'o~ephalill(, ,in.1s b~-
2H BO'IU)l;~. ti.C... SUOM', Jl.E.. MUftPlU , f,A., :g;i. Phr~ko·Cht'ntlcaJ and larvaJ-mfocwd Cuk.rq11mqudcw:w1usrnu"Gu1roes . .\,,un!so/tlu·
morphological n,lntion~llp of sonw :mltropud-bomc t<l ,;,= Emnn;t,fOJtJ' :)()f."(L'fJ.' <>f A.rm•rf,:y1. 56. 2Ji ·239
blu,:tonxuc ,1ru,. A new wxan<>nut i;roop. Ph)·sko-<:lwmic..1and ~9 cow~,:;. ,,.,.,..• H \IOU«'>\, ,'\.I· • ,uMrhtt. J.Jt. t;fm. Tran~mh,Jonof 1.1as,icm
sorolo&Jcolstud!es.Jo11r1u,/ o/G<•11rrnl n,olot",·, 13, 261-2;1 t'C]tlln,• r111'·r11h:dflh \'in:, hr ~pt/,.( hi,W'/Ui jotivtl'ff h)· f11n~l 1•\"J10\.tlrt""
29 noKoM l-'v\. R.t>.;. Gftl.)bt.,n. P,lt., 1~ft6, \in.i~A·L-ctor·host r~httion:Jllp ... nl ,,nd meml.u:uu.• (t.1.'d1r.i;. .\fu:J1/ui10 .\11,(,/j. 26. 3~-36"7.
.<\L"rleJ srlmulnns and Jomc.,to,..n C:nnyon ,,ru-' fn :,t. N'onhcm lnd:ana 10 t'.()M 1 lt\l. J,J)•. unnr..~T-ML H .. M(HJ~"'· \1.1., '-"'"'-U>. I, ),j U,1<f,R• .M.~ ..
('OZOOllt' f~u~ ••o\mtrican Joum(l/ ofTmpfroL .\leilemenmt /f.1,gi~f:.·, 33,
l&ii. Pttu,rfa \10:" a nn\' \l.ric:.tn .\f;t'Ot In 1h,• tici,horn«.· Otoni Gh~J
12ffS..1295, Kh.m lllGl.1 urnu1> 3nd anti~,•nlc rolodon,hlp, \\1lhln the l)GI." group.
30 l\lUn\~, P.. 1986. The cla,i,iftc.:11lon und nomcr)dutun.\ c>f ~irui,,:...: Joumnl n[.\l,•1//~nl Em~mology, 12. 202-20;.
&ummn'}' of l'l'\11IL, af r.u•,;1i11g; or th• lntcmntionul C:on11nlu~ on S1 coa....:nt. u.:;.. tttllltHJ'o,. \.~.. lt\.Y~. t.N ,. "t:l:',1, 1.n., i.qao. Cncht..·
T"s,mom)· o(Vlruses in Send~!. Sl'J>w1nb.r, l98~. 1111,tvtroldg1•. a. Val!cy "uus: npl.'rim~ntal in!t'ction in CullJ".:UI mor1101t1. CtwodJm1
141-143. Jm1mnJ1ifMirrt1//lo/1,g., 26. zs;-290.
31 8VRCD01-lfEJl, \\'.4_ \'AR)l\i !lt.t.n_, 19tl;". T,,uu.-itadlal ;tnd lr4ll~O\.lUfat S2 C:OA.~M. \I,. M()UIS. \ .. Hl \U. , •.\D.\M, e. fU.' \Ut>n , .• ,. \L\Dt, .\I, ,.
dtve11,pment of dliea.i:.l! ~cnts In .irthropodi.~A.mmfl/ Rtt1lf't1' of E\it,,o. •,i.. 197q line pou~ tpl7Mtiquc d" ftc\Tl' j.aunc.: .....-l\-'tltlquc uu.
Et,tomolo-1.0·. 11 , 347-376. Se-ne1ral orit.1uat. I\O!trr.:cm di! \iTU.\ dt rots de mou,11que, adulH.".So m:llt..~
):! e;..umrno. c.,., 6 ' bPAR:r..., , .• 19ii'. !ttductioJ\ of\ 't"nczuclan ~qulnc et {emt'l.:s.. Mt'ilttcin~ rt .\!afm'lln l11ft'(ti,:11.~'$-. 9, til-f,~.
(•nctphalitis ~·luc3mbo) \iru"- b)· ludodt..-OX~'Uridinl' ln c-hronlc.ill\' 33 .,;;g"'"""f,\:'(, I .C., l.\,,JOn1'. l,C. I•, f. 'l)IITfl, D.~- J9Jtl $U\Ccr1dbilnro:
ui!-..-"Cu.-d 'cured' cultured mosquito cl'IIS_/m,,11•Jrulogy. 8 .. l93-:?03. ba1s ,n tNlaU\ cnc:rph"1i\1, viruses.. h-duurum P101,·t'i.'rl11~$. 15, Sb~.
!3 CAS<\l.S, 1.. 195,. The arlhropod,bamc grm:p 01aaimlll \1rlli4'>,
.sa coz.1 .. num "',C-OH~n M.1.,Ko111:--.,.. 197ft lr1tni.mfs~an
Tm11sncti1J1uqf1ltt Xuw York .4.,"(ltf,.imy o,'Stli!111·cs. 19. 2t9-23S. ,run.,ovltfcnn\' d·un fl~\i\iru'I I!.! \·inn: hqulJnJW chtrJ: /u,"i/1."1 ,1,'iflPll l l I.
3,1 CA>-\l.S, 1.. 1961. or
Procedures for i~entifioarion anhm11od bomo ,iruse<. 0 t'-{Jm/JIC r.,ntfu de/ .\rodt'"1il· dt"S $(if111r:1."'I. 2.S.'l t>i.. 109-1 ]{.),
Bullt•tfo oftlze t .,·orltf lt.:alrh Orgo,1iH1rio11, ?4. :'23-i"J.1 .iS CA.\:\."'} I.,, I .. J r-;u,,o,-:trf:ll irurKml\.,j()n
H HU. J\.L .. (.\U)Hf·H, (.U. l!f-
3$ 1. lo'. a.ft01.,·~. 1.,., 19~ llcm,,ggluunalion \\ith i\nbmpod,bnrnc
C..\.,\1_.i;, ot Cill1!om1.;. l·ncepha.lttb ,1ru:,. tn tht.• mo~uuo .-l, ,Ir, durt11I" at Bh..1e
,1ruses. /ounrnl of F.:qxrlmentol .\fwflww ~9. 4:!9-1~9. la~f!. U:a.h. 1\{QSl/ll/c') \',!U.")'. 3; ,1~1fl1

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 Clsssification. epidemiology and control of onhropod-bom~ viruses 165

s& tM <:O<tA :.1tN0-1·.-. \'.,,_ ,~a., Th1..~ i~ohult,n J.nd impo~t:ut<'~ ofSlmbu .:. 11~11. P.t.,1,, M--;.. Ve.-cuu" i'lnd \en1t.1J 1,an.,mlfi,:$1Qn: an tpld(.'mJC)log:C!II
grouJ"in»c> In ,outh ,\!rica. M, \kd VtL "'hc,1~. Unh1'riit,· or pt'r\{>\•,,,\·~. \nr,al~ JJ/tlz'" \'..,u, l-Ort U:mlf"lt1)'0/Selma-1.. 266. 1';3-19.;,
Pretori•. ;6 .F1s~4 ll' 1 \t,. t98i. E1>idemtolo!!tc,al pr!nciplt'S of \'CC'tor-ntcdi-.H('(i
s· DAsn:J.O\ ,. ,·. (. H\-n,. J•• 1r,9. I .;thar,nnry dl".mon::uutlon of uanriovnrfal tron<mh,ior.. hr: \H.:..J:t.L\'&,V. '"" u .. O.Okl.t>Gt. a.,. a,. \.t.\kA\tuK('I\U'.1-1, ....
1~n--,mb..~fnn oJ ll1h~n3 inA,'(/c·!,,lu.·an..<.md ,,e rolco(th~ :ed-s, \t,"t'tar, of l>L'(,v,&\' -~~,w
lm,,nr,fons u•Hit P/()1111. Animr.ls and
mL'Ch.3J\iMn In o\1.'1\,1n1cnns of thh• urbu\1rus. Fo/111 P,1ra1fro(o:g.fr,1. ,\1tW ~C'W \'o,k: P1,u.1tC( Pr..~~
2li. 361-JbG, ouc.
-r; fl XE, r. [_,.,, & 1.1 ,.w.
1q ..S. rowMets a quanliuuivc und.t~!'mlnding 01
58 o.,usstA. ft,;. uuu,n"\, l,N .. 19.:11. X:\frobl ~h1.•,•p dh,ea'ie. Pnrn.fitology. l3. the rpjdrri\lolom ot· ~c;·.,.torie ,1n:t. In lhl' t:ssrem U:\hed Sutt~.
,;o;-52.1. Am«itrm Jounu,I o/TrtJpital ,\t,:dldm•mul Jlygiem!. 27. 3~2-338.

;;9 DAvrns. 1,.t:;., u."nucu\1 li..f. t1. J"rn"", A.O,. 1.9~s., RainfaU and cpt:oouc RJft ;8 f AA!".('.l·. o.n.. RUCII, W.i\.., .\10'.'l!TO\';\, M' .• r.\'.C:Llo;;U, o.s. ~ O.OU~·. "·"'· 198o.
\ 'nlfcy lc\'or, /Ju//11:/11 n(tht ll'nrld f/M/1/1 Orgn11/i:mlon, 63. 941-943. f rJ..Osov;uiAI fTUllilinli!o'ion nrst l.ou1$ CflC~phaJhls \1[Uj by <;u&!.t p1pit.'ll:J
60 O,\VJlli, .,.c.. l!tl:l nu: et1CC:t or ,·.uiou" 11;1mp1,,-rature1 ln nu,dl~1ng the
romplt•x 1110~\lllO(·~ Am1.,r/('m Jou ma/ n/Tmmcal \ fe(l/clru:1md
//),g/M,, 30, oll9•iOS.
<:<uin,ic incub,11,un period of the ;rllow (.,. ec ,;rus in Aode,«'IIJ.'P'I
,'1mcrt,mr Jourrtnl of l(tgtem'. l ti. 16.l-l ;'6. fU rRtUF'I. p -.«11t-u w, i.e..• 1984, 0ml :tnd nal'lSO,-a:rial mm.smi»hui of r..,
61 P-'\'l«.. :--.c,,. s1-1 ~"~o.~. 1t.c•• ,~30. ·n,,..
loc.1tlun of yt•tlow iC\·cr \'inz.s in
c,o,.~e \int<t by A1YI<.'$ a,mp(ltims. ,\m~rlc,v1 Jouaurl u/Tro,,rca: ,\frtlfcrn.:
mu/ Hyg,en,• J:1. 71)3-71-t
infe<tcll mo,quiloc,4nd the po~bUi~ ofhc-wditar)' 1roru.mK,!on
,\mPricnn Jouma/ of ll_l'l(IM<. 11, 33;..34 I, so •mni:c. u ,. Mhr~ 1, 198; \·cn:ic:iJ tr..in<ml<slonoi dengue ,1n1.st:sby
Ii:!. t>t«;Ji:ISSO°', Q_ft., \t('.GU.U\'Jt.-\\, a.,, .
\1('1Xlt)~Jf ft ~- 1,,\\1-sTER.. P-.,\,n•• 19:$5,
mru.quitc>c:.. of -1h<- 1\,vh-,#ut,•Unrls group. Amaricnn JoumnJ ofTropical
.ttrdlrtt:,•t'ltUI Jtn;I,•,:~, J;, 64()..&1;
Andbodirs. .;g3.ln."t eonllin nrbo\'i.fm<"i in ,(!ri {mm human bt11lnp J.nd
dom~ric :mim:c.b from the.- ,ou1h,we,,.<f..'m.1nd notth·w~um, tc.-giont c,f th rttrlnt. lL • t\O\t 'J, L. t9M. \\•nical tr.m!lmlS$IOn of d~llJtU~ vi~S: b~
the C..pc., Pm,incc 11rSclu1~ ,\Inca. Soll/h -~fr/Mr. J(l11m<1/ of \/,d/c<1/ Atlli"J m,·:!io, f1wmi..:t,m•r1(im Jm,mal n/Tro,,1,ct .\f«fltinca,;d
.l<'h·11n•, au. 11. ll)'/11"1~. 39, 218-222,

63 DUTAA'\ h.f r. I.I- DUC- J.W. 1.981. rr:\I\\O\'Url.11 m:m,mlS$iOn of ~~Ua·,, 82 t KUBl\lUR. M.. l>A\'l!,. ~.(; I. )11~"\~o,. tt,L., 19.ll• On.the lail.Uft" ofyellow
rL"\ \'r \'frus
0
br ,I "Yl\-.1rif; \'4'('IOr. /ltllWltJ.R.1'gOs ('t?ll(IIU~C. 11'fmJm;tfon.s oftf:t.• r,..,.., •.1rus ,o pcrsm In ocolon)' ofAed,s n('lr}p:I. AmmcnnJournal 01
R(J)vl ,;,,,;/1'1)' of1'rop:<af .\J.-llrwetmd U;,,<i,nc,,~. 128. ll}'g!,•n,\ J.1, r.;2-J.16.

6-: ,-01..:utu", ., .• t90<l, South \tlicull hol"<'S:<:l.ne<s. It.< p,11holo10· •nd 83 c;,\U:,orti0, r., 1958. munomk,i-o; Sr.lxtthttJthlOrQp((:rus I tumbuldt. n
m~thud£ of 1>rotcct.lW immun1;,4uion. Jouma: ofCompara1fl;,. Pathof,1!?>' vc-ctor of ~yh·dn ~ cUow !c:-vt't in middle t\mcrlca .•.\mrncan Jo11mal of
a,,d J11ett11><1<1fc.. 1:1. 200-231. 281-300. Trop!rnl ,\Jrdi(illl' nnd i{)Jic,re, i. 1?9-410.

6S tm"\,,10:\-. ,,., 190.1 runher note- on 1hc correllHo:i of SC'\'trai di;;e-J~h 8,l t;MR, 1 ,.. All:>, n.1 1.. 1~17 fh~OCi.'\Jrftru;c f1f u Jengut'-U.kc f~·c1 in the
occurring t1mr:m_g "'1od. in S<.>uth Afrie-..i /ournnl o/CDJnparmfw nunh,t•:,-.trm lmn.sv1MI. SomJJ.-\Jrlran Mediaal /0{1nml, 31, 2..;..1-2"5i.
PailrOIQgyfind J11<ropemia.11, Ml-141 sr. r.n.u n. J.l").• TCC>~. h .\'t'.. OIC)!.G.\\',,A., IL-\nOn\\, A.J. -'°Jll!\\'l'n Ll••• 1950.
ti-11 cu,tttnn,-. 19a, h ·ctnr mruntennn(ot ol pniho~cn~ In nd\'erse F.xporinrnnu,10 tc.~1 the p<h-..!b.!Hty c,,f trtu,sovnriot t.mnr;mis,ion of ycUow
m,ironmcnts :with specml n:t~rence to mosquno maan:en"nc,• of ft.."\cnirus- fn th•~ rnMq_uito A,.'<-18 !Srl.'~Om),ftt) ofrit\'mu.,-ThcobaJd..
;irba\liru'I.Cf In \:<;-KU.\'[\, lft,. J.J.. lWMIU(Oil. P ,f·." M.\MMOJUJ'>C.H. );., Au no.ls of Tro111«11 Mt'(}.t,•ui, and ParosJlo/og;•, 4.;, 3~2-350,
·t:d.s?, t ·~:ron-ufDiit'tt!il!. :lg.mt~· fm~rcwltom u-;th 11/nm,t \mma!sand 8-6 t,;NIMH\O, N,:t .. Ro~.c;., o v. ii-< R\U~. (;,-.. 190,0. ·\e<iM 1r1:Un.a111.t fOip1er.i-
\fa,r. ' ""'' York Pri.\1..~µe1 l'rt.~(i C:ullc1d:u,1 :and L'1 Cros.c •inli. ~lodlfreauon o! mOS<JUIIO focdi11g
is; f'-Oltlt~, ft(,,, \'OU~G. r> r.. N 11 "lfl, h. It. 19g3, l n.rbO\'.lrlilt tran."-m&s.s.lon of brluw1c,ur b)' ,'iru--. infL'Cliua /01,mnl <>/Mcdirnl Emomoto.so·. 17. 1-";'
Rio Grande vin1s tffuny~\i:,d,u:. /lhltboz"in,s: br the sand lty, Lut.:tJm)'i.a $7 H \RO')' u..• uou~. t.t, lr..ft.\\1t.h., .n "' "-EIWG,w.c.. 1983. fnutn~it: fo..ctori
amho11hom Am~ncan fo:,mal oj rrovir-lil ,\ft.tJu;uw rmd Hygumct. 3~. nffc-c:mg YC(lor compe.tence of mosqultoc>fot ubovfruscs.Annunl
8~-ll&I. Hei·;,1ro/F.m(1tnolt>I:}·, ia, ~g..!~.

t>B Ut.,~,w,;. 11.,.. 1!1;-8. .\ new :ipproaeh :u pol~ dottuJ unmun~atfon liSgw:r\5t 8a 11.\RO\' l,L.. hJM.;\. L. JmA.)Ttu, l..b , 1•nr-«l!R. <..b., ,:O.:HMO\'Ht. u ••~.-J;
\fnc.m h"™"'k~n.,., , /n: ....,.,, , . f.T. "Gtull!. n. ,eds). Proa,"'1111!:5 of TU(-lntu. ,,.,. 1ti80. flf('C"1nf reanng 1cmpcr:uure ~n uan~varlal
1he-Fmir11t /ml'T"nt:ounl CQn/rr••,,r_.o,, F.Jiuin.-• lnfr.c14011s J)Js,·a~•·t, ~c:w 1ran~ml~1Jon of ~t Loul~ tm.·1:pl111.1itl~ viru1.o by mo>oqUitot"s..-tm,•rica,i
Jers'*~ VetcrinJf)' Pubiicauons ln('- Journdl o/Tm11/ru/ llffli<'IWnmt /(1-g/m<, 29, 963--968.

,;g f.ftA.'(~;11.,.. tt.J .. 1~u9, \ ·eot~rinil1' Hl~et.iu;:h !n!iintutc. UndC!lttepoc,n. ~ouu, 8!:I ll,\Jl1J\ IL .. Hll"'I , , l, tl..tl:\"I:~. \\".\.., M.tlJ\'A.,1. IU-',<61 J'ill~""E"R. ~U.., ~8.t
\fri~. Unpul>li,hcJ <l•1• Exp«inu•mol transa,~rl:Jl tron<ml,,!on o(St lours encopholitl< ,iru< by
~o l.R.\S-)1\Jfo. 8..1 .. ,\.Ul-..L\Afl l-f. ~I\UT1 J.IL UC.\1't.S. , .... 'U).\t"', r .. 19;0. nw
o,ft..t and .•ll"t./c·i- nu,,ttuilOl·~~.-tlill'rfmn /t,um1tl hf Tm1>kttl ,\f('i/r'd11raud
1/ygl,•m•. :il, ltl6-t75.
h,ul:ufon Ar\d ch.irucwri7.Ullnn oft•ttUlrw c-nc~hulrni~) vim,- Rul/1,tm
OJ]irl'lmm1nri0Mf 1t,., lip/!hl)t/1:$. ,~. iSl-789. so it.\R.RJM>:\, ,J-.1950. Rang..:ufmovcmem of some Malasan tat, Joumttl
&/1\l«m11UilOFJ•, 39, 190•20fi.
71 F.Jt.\SMU!\. A.J .• 80~11nn. ~.r. ~ ,,wn!flsi: 1 ,,1... 19;8. The tsutaifona.nd
ch:,mcu,."fiY.-HiQn or \!quln~ t'ncc1>hulo...t.. uncJ !tirologicall)· rcl:ut'd 91 Utin,. C:.<, P'\~111, n.11.. O.\Q,\fl, ~. • .u1M1..u. 1 .. 19,$.: Gcn-c::1c vam'Hlonfor
orbhirw.fh. from hoN-o..'-S. In:
DR\'AX$, ,.T...
Gf~ffrR, H • (e(f§;J. PtOCtt"ilbJg,t \\'e.._q '.\He ·,•m1~ W$C:1:fnih1lit)' in Cut~.f triwcmorl:p:rhw. Ametlrrm
of1111· Fourth TlltC'hUUfo,w/ c,mfrn.•11t,, au cf'/1:inr 111.fi'i'tfou.~ Dfs.,j/(UM, Jo11rnr1J o/Ttop,ml ,\ 1t:d/rim•mul Hyg_:N1t• .33. ;15..72.;~
'<,~, kr~ , ·~wnn,uy Puhlltallnn, Inc.. \SCI 1~"10(..lri, f, jlj ,,u'\::)Ul'\', J,R,. l!,j:I..,; l.aboratOJ)' :itUdib on
fll n \'tU ~. l.,fl

*2 J::\R\.,. \.T.L. nrRLU. ~,., •• JICl\tO)(K. , ...., .• IIOlfTlfJt. Jl..<~. t:tntt-s. P.IL. 1hc t~1~ml-"'"hm Qf we-.tcm equine cncC'pha.hh.s, UU$ by !)askatch.c,\---an
v. v,,o,. r .1 . ,. n.uln. c.t JCi8~. RtthJccd ...unw.1.1 of adult Cult.-.x pipl...•ns mosqulrocs ;Culr., tnrsull• i::,madta11 la11111nl oJComp(tro1i,~.,1e,//<i,;,.
lnfoct<d with !lift Voll~· f,,.,,.,
,1ru, ,.\l,utr/m,, /omndl o/Tmpkal 36.83-88,
.\'1cJhc11te nud Jlyg1t1J('. J:", ~O:t-40:9 93 m>Cu ••,.L. r.,Jte;"''11. , ,f•. ~ k \JU:,. c:.t.. , 19Ur.,. ~lttha.nieal trJru.m1sslon of
T3 n ..~su. r .. 1976, CJ~ilieltiun and nomenc!inure of ,1MM:. Second Rift 1·011,y re1~r ,-!ru, by hemo1uphogou< dJp1cro . ..lm,•rftnn Joomnl of
Repon of1bc ln1,•n1atlan:sl Cornrnluce on To.,onomy of Viruses. 1'r,;pi,Yll ,1/!!dldm• tllt<f H,V//t,•11~. 3; 188-193.
Jnt.:n.'imlog,,. t 1-J Hi 9.- HOOG'i>"fll\.U- u .. 1966.ncks in cc.,Jntfon 10 human di<i.C~ cau::cdb}
4

;.: l·f"-MR F, l't'HUR.\. II.Ci S'()flTTRfl(lU, I,~. JQXUK, \\'.);, & OOW~lf ,\.\\ ,
,•fn.i,;~ A,m:llll Rtt11/1u, fJff.i:ttm:olog:.·, t 1. 2.61-3.0&
l974 Fam iii and g<nctrl< n~m~ !Or\ ,ruse, nppro\'cd by lh~ 95 11oor~TRM1, ,r. 19;g, The cptdrnu.010~1 of U(').,.b(»rr..e Crtmr:.m-Conso
lnwm,11onlll C:ommilt<'l'on T..onorui of \1ruses. Juno, 19;~- humorrh3~it fo,·t,r In A~i~. EUTDpt' and Al'ric~.. Jrmrm1T of.\ftdical
/m,n·/ro/1>11)', 3. l 93-!!!8. F.mmunto;:,·, 15. 307-1 Ii.

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166 ,11J«·, c,.: Aspects influencing the occurrence of lnfoc1lou~ di,~a~t·,

90 un<x:;~1M-.\.\t.. ti.... 19t11. Ch.u1gin,; p,mc.m"' of tid:,bumc- di4._..,.,.L.,. in ~"iufut"-d fto.nt ul-d~ mu""quito~ during. an ~pW>Otic in sheep in th~
mc,d(!!n 4>txitt}'· At:mllll Rei•ff:lu q/Entcm:t1logJ.1, 2b. 7>'94.'. ~.1.Metn C,tpc- Pr()\'in<'<-, ~1111: .Jijrit'Du Joullutl o/.\fr,Jiral Sc.L•m·"
l2.
Hd:..:frnm Eutop1.•.1n°i\Sl0:UC bud,;
9';' UOOGSlJCA.\L II." );,,\JSf'.11. ~.~ .1961 .
HS-I~
rnigmtin,:: through F.g:~ pt into Atricn X,mtc. 133. 277..2i8.. u8 ~ma,R,01. k,U •• m Y"it\.., ,. -"'· :\Hbr•run.. J, .. ,1,·ol!-n:-muL'1t.. h •• 193,7.

~ flQ()v)n{MJ. U ... M~LA, ,'11.S .. TI\ATI.Olt :\i,A .. Ui\fil n. ~. «<,kUlM)Y t. 1981.

'tudil"~ on arthtopod-bot11e •, iru"'-'' in Tong,.Uand. \'. b,oltt1or1 ur
Tick,, fl.rQtloitliN.1 on blrd> migraung from \fncJ 10 Eurap:e, and A.,M, £hm~1tnl\\\.lfll .1.ntl Kilt\ J.ilt,\' fo,·cr \irus~ front moi.qu1t0o_ Sowl!
811/lf!.lm ofth~ ~\"()r/d Ht't1/fl1 OtJV~m;;.,11un. 24, lY7-:!J:? -tfnrtm Ivumal u/ ,\fl•dfral S"U:na,, :a. 11.. ;~.

~ HOU);. t .. J., .. AA.',trN, L.O•• IIARO\, l,L.,. rR.t.~UI. "I IJ., 11t86. An lntl"r..p<-clfi-c 119 >:Qun,ur. rut & :\H 1, to:-u. u.•,:,. 19~9. t,:ofacion onv,.,.i '=ilc- ,·,ru, from
muM1ui1u mtH.1t:I ror Ll1L tnc.)Cntcron.d infoctlon harrier co ,-.1..~a.·rn
1 :t Jl4turi!lly fnlcc-100 hum:in ~ing and from u bird :,y!iwrca h,1lut-i.!1t.•
equin<' cnct·ph~lumyt-Jhis \'fru,. {Cu!ct,1. rarsahs ;\Od Cttl•'·' pf171rn1:i '\'1emot • Som!t ,l[rh"mt ,\h·tilt:t1llt1um11l. 33. 98i-~
.\m,·rlc'(ut Joumnl of Ttr)J)/ml ,\1,"Ciicm.-(wtl H;t,1fl4'tu", 35. 632...6,; \. 12-0 ,.,;;<HJ '"OT, ".n,. ,u"t\'TO~u. 1t ,:. , \\·OltlJI. c.a. 19M. ~dumu \.lfU~. u
100 HO\\Afl«>s. A J .. 1970. V1:-.sfcufar S10ffl3tfti, and td.11etl \'iru~e~ _~\dl'/Ul(eJ hith1.•no unknown :,s~(.·n:. 1)o()l,1t1.od from tuhc-.m1.• mo~.qultOt,'..\, collected lO
/11 ~7tur R&-a,,:/1, IG. !9;",-iS6. nonhem '\':m,1J. Unum o!Stt'lulh :Urica ..·1merit(m Jaumol u{'fto1.m.·,,r
.,1,<1cw1.-n11tl tt;g,,·11,, 10. 333· 331l,
101 mt.UtD. ,, .. c:o.u-,n. I.P,. Of.RMAf\;, ,,. .o,. f ..\\jlt:,\.,. J.t•• 19i'$. Pa.<-SaR,l"
trttn..~Jvaricnnt du ,,rus Dugbe chcz 13 ttq~~-\mblyommll '"'"f8<'."'m 1:tl li:O~l.i.. '.\OT, fUt., M(;l:'\ ro::-K, Ii.,~. W(lkOI, C,fl: i. Dt wu ... \, 1.. 1~6;;:,

•rJbrldu~J. Hullclln dt: In Saci',•u•rl;.• P,:tiio!QJ:!.U• £\·otfrlttt' ,11,tt·~ J,•_· t'°Jat!on at ,fru~ from mq.o;quhoes coUtctcd ..:i; J.umbo. \fc,;a1mh1qut.•
F'iJUJJ/ts 71 19-~2. II. ~lo,"">l.til ,1ru11. o nc.•" vlm\ tsolated CrQm 1hc-Cul..·x /Cut~ 1,tlrm
\\1t:dcmimn ,;ruup . .t\fnera-·an Joum(il Qj1rup:4·u! .\Jt¥1n·mnuul N;,g:i ...,".,
102 HULL."·· 'fiMSlNVH, J.'., DJ! $0V1.A, M. ~ ~~,\HUSU,, H., 1~84. ~:nur.d
11, 683-68-1,
trnnsovan,ll 1r.1n:im)~~ion of dengue -t ,·S:us an At'tll'J"'®1Jtl, m Tnmd,\d.
Am('nton Jc,ur,utl o/1 ropic1.1/ M;:dr,:f,r.,... a,ul Hygw,w, l3. 1248-12:SO. 12:!' ~U"Hl'\'Q f, ft.It,. \ICl.l\lO"lf, U.:-01 \\'Olf[H, ~.\\·,, \lrUI\L\~ L• WU~tlfti..S,
,1 , .• lf\l.\al'°l.ltiun u; vbu'-'.") frori, m011c1uuu1....., C(ll!t:C.h..~ 01 t .umh<>
1u3 UUMMJ'UH, .... t97J, Sulh!t.,.s.h.npt.td \1fU)4,.'" Jr,: \1<\R\:\101H)MiJI, ~.
:'.h>l,i.mJlqut'. I. I .t.1mhc.l \'IN..,, ant•\\· viru, lwlatt!'d from ,\,'tit:. rSktts~r 1
a- Jii.'VH~At.:, E. (ecb,) Co111pnm1i:1< \-'1tu~,·. ' 't:\\ York.: -\nscfomk Plt!'o>,
J11°mU<t,•11st,TI1cobt1ld.,-\lnl"rlcw, /ourn«t of rroplwf .'.l;'t.fitttt1• mul
104 HMt.atn. H.S:.., 19i3, rht.'- c!foct of t!.mironmenr..tl tcmp..·r~tturc upou tht ll,1i;,,•11c, 11, t , ~ .
tr;in~rni.-...iaft nFSt !...ouls <'nr.c-pb.alitt..: ,w, b~ ('ul,~ pfp1'i11u
123 '-.Ol\'1;R'.'l.'UI. ff.ft .• ~\111HlfU"~· $:...C.., G.\X1Hfl.\, ,\.1;,., ,,ct,tn,11. O.M , 1,,,
,:,,l11q11;{t:ula11u.Jo:mt11/o/,1/i:,l/wl l,111•,mn!r,IJ)', 10, t-12,
nn"'"' c...<., 1i+Go. Pn,ma-) dl• ncuua1Ut.tcao com ..oro,. dt" lndh1duo.,.
105 IL..E~KO. \°.~. vOROll-l\.."(K.l~A. t.s...,. .S~t0RDD"~l~\". ,\,,',,. 1-)70, ~!:un rt..-.:kkntts 1.:n M()('11tnh•qut• \"Ctntta dtt-mntn.ldO) ;iru" J>0~::uk~ em
paucms o! tran§.Ov:ui;tl trar\Smi:;ifon ot uck•bomc tncvph;ttfli, \'irus. h~1 Africa mm.,mhldc,h l'Oi drttopod1.~. .\ru1u d<h r1uti1111,-, ,\1«liC'lm1
tick \"Ct'U>~ . .\11..•dfum..d.,'Q)'t1 POmtiw,'o.J.iJn l Phtt1-:.imm,,• Boh1: .11, 39. Trop,u 17. 20!-?:\0.
Zo:3-26;'
lU ~O).:U\t)l, IUI.. '\Ml1H9Uft!'o., \'...C. f. u.uca.. L. lSOL ~....uu.all.tmg
106 JlWI', ,•.c;,1 19;.: t.aboratory.smdles nn ,he tt.imn·u!;.S100 01 \\'t"i1 ~He .1ntibixfl1."\ .,~illnM ,,nhropot!·l)()tne vtrusc~ In 1hc -..rrc1 of Uom~uc
\'lnr,o b} C.:u!,,.t ,CJ,!~J uni,1imuus TI}e-01nld: ta.ctont inOu<incing 1hc <JU•drured, rnngini: In longJlond, Union nf '-oull, \lricn. A111mls of
tr ,sn~nd,,.ion r.u~.Joumnt 0/.1.\f('dfcnl E>t·unu>l()gy• .f, ;ii-;3,'8,, l'mJ1lt:'t,t Mrthrlttt•mui Ptmttlmfugy. S5...~5.
H>':° f111 1l•. ,-.r.., tHACli.lJUh!\, ,,r,,,. THO\li-W~. U l. -, ,11 l ~l.fll.\~. G '1., 1986. ~ ~l.,J.:Ul,Ol .II.II .• .._,111tuWJt\. \,;.<:... \lll!,f'Mn I I,. JIObGW~. b .• l!)5"7.
$imlb,~j)nd \\'c"'t ' '£Ir VffU>-i1tfl>clh,m-.iJ1 d~t.~ \\it\, ..1tltf"1fltnd •Pr~:uri.1 ~mdlt"S on tuth1tJpod·borni! \'fnN.'~ ofTong~fa.nd \111. Spon<l,,1ent
regjon . 'w111h
. .4/ricftfl M,dlcol /011ma/. ,0. iiS-220 in1,. ,m .is~·nt ptt.'\·!nu.ik unknown h,c1l,ucc1 from 'I'antlothJ•m·Juu.
Suppl~tnt to intc.1"1Tm1lqnal C'11alngu(' of
10s ...,ttAll.\TM1', , .• 1~i78. \/a1:w11iottl,,.J 11111/mm«Thro, Sn1111! .Vri<'Jm /011mnl n/,1,..fllrnl
arbo'.itl.bCS lnduding <cl'Utin other" lru~i-o( Ycnc,,l>rau?, ..Am,•rh·tm Si.·i1•11no,l2, 10S· It!
Jormml fJ/Troptml .\1«f(clnrti111d Hygi~nt, 2; '.\i2.-l SO
l2.6 ~(l~t·R~Ot. ,., ..,. !)\.tlf HftUk'-, \'..( • M'l"atSO.~. tt.~. 4. ,1crvro,11. ti-'.1.., H'lt\O,
11"9 K.\FtAtlAr~~ , .• !98&.. lnrtrr,uuion/llcn:11!ttg11l! ofarl.KJt1in.u1:.1 inrfodfn,: 1.sof,ufon ofGcrmh,.10 \'ir~ hi:.hen() unl>:.ncw11 01~n1. ftom CuJidnc-
c,111aln orltcr dr1ue.sofl:'l!rrtbtatcs.. S.tn \n1onir,, ·rr,:t.<· Am.t•tica.n !'llO\qul:,1t·""', .l1,d.; rt:J>(Jrl of lnt('(:tfon in two laboratnry \\t1rk.eN-
Soclt'ty of Trop,cal ~ledldne and H) ~,en,• .Anwricm1 Jotirr:al ofTrc,pir.al \ft"llid1u,an,l !ln;h~n,-. 9, 62·6..CJ.
uo MY. o.u- 1982. Thro-e in(>({c...-. of trun\llu'l;.;1011 o(Ro~i R.i\'cr ,•m1~ by. \tr1ri i::r;" XOlr.l Jt~OI IIJL fl\Hl HIJtJJtS, ),.,C. k \\H~klUt\:, ,,.P.. l95S. !\tUlrallling
&ofgJ/{J;.\, Auirr(l/11111 J<mmat ofJixp.:rirn~mal 810Iom·1md Afrdtcul Sti..mtt, nur..bodh.':!. to JithropocJ ·bomt' \inl~t--s in hurruu1 b1,.1lni:.'i and •.tnhJ1Jf!\ in
60, :i:SS-..:1-~•. 1ht' un1,,u •1£ Sou1h Africt.1. Joumal oflmm:mnlug)', 77.. Jl.3"'3l3.
111 t..w. (tu. s<,:.\HU 't'. t,c•. 1980.1·raasU\Jriui rran\tnl~tun of Murray V~llcy ;:us >.onn!IIUT. IUL \MUHIUIM.~. J,;.,C. W0~8~ \1,1'- $.. Dt \It U L,O, . 8,. J9!i,i,
i:11Ccph.ilith v1n.J.. h) .i.:<ll!I 01..'f1J'Pll {l. .~litlrnllrm /(1llf1:td ti)" s1udic, <>n u11hrnpod-borne ,iru,.. o(Toi,galant!. \ 1. hola1!011 or
J:Xp,•rlnhmtn/ B{olog,· <tnd Mcdi,c.l Sctt!n<,, 58. 601-SO·I. Pon~ul.. \il'\b fromtl,:Yft, (Bcml·sfnc,Jlaj c1('.(mnh1«."ii/u,n1e0. ~u111Jt
u2 ~m;\·. M,:\t -t. cIf..\.', .i.;.\.. t983, l'mn~,)\'iilrfal 1mn,mi<...Jon o1 tfon!_..'tle-2 \iN~ Afrl-,:-011 Jom,u,/ 11/ .\lt"1ti·al Xf,•m.•1.•. 22, 81-92.
b)' ,l,v/,s "o'/rlVli in 1mure. Am,Yiro11 /011r1111/ oJ'Trop/MI \lid1ci111•111ul 1.35 KOKUtXOf. fLH •• S~L\\11' f.1 I l \'Jrl", J. A M.C.l~-rollli, U.~I .. 1,6S,.- ~Llf\'C~ :or
0

HJ?;il•n.-. ~:?. .i~O-~~ t :inubodll"') agnin.s,t an.hrapod·borne ,1m.c'- ,n the ,.;or-a of ind)ge-nl>UJ
llJ .1,.:;;..,u:,.G, 11.E .. 01.\,UBDU.AlS, J,t,\\, (flhO~. :\l,I , Sll:11., k.t. Ac .IUCC.:.\,_ \J.\._, : .....,:dent, ur du· Ci!P,rin Strip •md lk.'-chuar..:.fand rro,c.'<'tOr.t1t·.
195.1. S1udtes. on the , ~on.h Arnerll":!n ;i.n.hropod--borne rn,cph.tliddc.."S. 1,mJ.R,aion1 oJ 11t,· Royal So(l~· ofTmp:,·al ,'.Jtdlctnr and 11.'tf,•ir.•. 3-."f.
Lncrm r,quinc ~·ncq,h;lfh1s Ul ho"1~ .••\mut,,m /01,r1u1l ofUygi.\•t:t'. 60. 553-56:?
237-2.';(I, 130 ).()SUie , ... 1U,\'A, ~-S f. nUPPO\Tr>. ll,\',, 19;0. fn(l"CtlOn rateot /xl)(u•
n: i,a~US(,, n.L.. ClL\.M,Wf.KlAIX. lt.W•• :'i1kf5 •. ,..~ IJUSOS. ~u. 19S1. Srndih 1,m.1<f(tttuJ lick..~ nnd ccr.aln questiun~ of tr'1nsov.on.tl 1.ran.\mi\,1on unt1
on lhC' ,'-:unh \mcrfa.n ar1hropod·borm· (!nCl·ph:illtidc.\.. F...1,t1Jl'n <..-qui no dOSDi,t4'<f iMccuon with tick,borne ~.ncl!phnJiti~ ,,ru., \'(JJ1,;u.,,·
c:\tcphn.l.ill"io in wild.bir:ds. ~1m11n'co11 Jo:mud t,f JlyHft•,u•, 60. 251 <?65. V11t1sato,:it, i 5, 703-70S.
IVi ~~un:;qx, u.,., 19:-3. Rha.b<lq\•i~. /oumfll oj'Gtmrral Viro!OJ{l'. 2.0, t3l J.R\Mt R J .r,•., lL\1iD\' f.1 ...
rN; ......W. :;..a .• 1983. F.th....::1 0(11:m~r.HUh· o(
10$-1:10. t:Mtin,-ic ,ncub~uoo oil the vector L'Ompetenc~ of Culr,\ tar~lb fnr
Iii• fii;()lo!J fl Mn, lt.H .. L\~C.\, \ _:l,l.f(., \\'l lHKI:'.'\, ,t.l' Iii ~1rt:,,1o~H. tl.M., lQbS,
we,-1em l'qUiur,mccphJ.lilb \'INS Amerir:uu JuurnQl of lit1J1te<lf
,\lf'fltrir,l*lllld lfy,;t,·11~·. 3'.!, 1130-1139.
Sur.·", for andbodil..., ..:.gain:tt anhrop()d·bonw vfn1'CQ in the "-ur.• of
'"dlgcnc,u.'ij re:,ifdt:nts.o!Angola. Tmn~ac:ionso/1he Rqynl ~etyof 13?. LA.).10"1.. 1.. -...• 1g.sij 13p;;nc!'-c 8 cnt..1>hnlfu,. in hnh durinJ; '-'lmulith.-d
r,o;m:<,t M~Yli(lfJ~ a,,d fl}'61fn,•. •'\'9. 563-.)70. hih(-m.,u,on . ..\m,•nrnn /ournt1/offf.i'Ji••1u-, 67 101-IOR.
117 KO,-:at~Ol. fl.ff., Dl- '•U IU.0;\', lt.... PATrR.-.0'.\, IU.. 111.Y\lA~\., ~ N 13-3, u. nu, .. t.\\ .• 1:t~J
Th1.• t'1.·olo~ <tf C.11.&fotn.fa group \'l1u-.t..,., J,mttud of
',..\ti 11taUtc:\, Jt..t>., 1957 '.\1iddelbu-rg vfru~ \ hllhcno unkmw, 11 .-S};t"f'it .\:r"CUc.at Fnwmot,~·.16. 1-Ji

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 Clnss.ilkation, epidemiology ttod control of arthropod-borne \iruscs 167

Uit uiou~ f.\\'.• ~ll\'D\OTO. \\., !il.D~OCl. 8,F- RUssnt.. P.~.4 ll;\RR, ,\..P .. s9;5, fovN in hum;insin South ,\frlt,t ~·umh Afritult .\!t,1f(t1l Jour,ud. 58.
E<olng)' or C1Hfrirn,a enccphlllill~v!rusHon inc Oct Mar Vo Peuinsllla_ 127-132.
11. Ocmunstratlon or u·.s.n"'.warial 1raMmi<-l>Inn• .Am,,,1am Joumttt Qj 155 ).tCJ!\'Tn!sn. B.:..,., SI JL\FJS-1, LT.. OIC.~~'60~. D.1!.. &·\\T.l~URE..'1, M.P., 1962.
Tropfcr,/ Medlcfne,mdHygiene, 24. 124-l26. ,\n1loodl<'S 11qilin5< tenain.i!bo\1rum in '<)ra from huinan beio~
:.E t.tE. \'.U. k J.."".D1r, c-;.r... tg;o. Coi,go ,1ims: t'-lCptiriment:iJ infccllon 01 ro<1;lcle-n1 h1 tt)ll~1a1 nr~~ ofSomhcm ~:nnl :ind EtL,tcm C3pc Pn.,\incei
H;'t,f()1t1111a mfi1>Mn.nd rrnn,mh:~fon too c;.df. Bt,Ut!tln oft/Jr of Sow.h Af.dro. S0111// Ajri<,m /01mu,/ of\/l!ilicv/ Sclmc,•. l,. 11-36.
e,tomD!of/1(:tt/ SOcftir;•••VEJ:ttrta. 2. 233--r33. lSO .Mcr..'TO:iJf, -0,!'1.1 .. \\ORTH. c..s. t:! KOXCRSOT, n.11 •• 1!)31. fsohuion of Sen1lilti
l31) l,t,.7JHCLrM, ~.J,, 6AU.E"\' C.L., [),\\"l6,<t;,, 11,<l. f, ru<.nJ<, (;,J,. l!)it], IJ.ctl'Ction of Fon.."Sl ,irus tromAMfli (t\("{llmorvlms) tlTgdt1wo1umcuuu~ (Thaobnld.J
Rift Va.llt'}( ftvcr ,·ir41 ~,mi1yfn Kcny:.t by~'"'Uit~ remote se~ing rolleued m Vor1uguese J;{lq Africa. 1'mu5acuo,u ofrlrt· Huy'fll ,'ioriezy of
images)'. Sciefl<"e. 235. 1651>-1659, Tropical .~fe,litl11rontl /{1,gimt. 5-5. 192•1!13.
13- U.."-,.IIICU~r. K,J.• Di\\~ .f.G.. °" ~mo, A.. l9$S,.Rift \·,1.Jle)' (6\'Cl't ,in.ih 1,;; \1<:J ri,;,. D.~1•• t9~. AtbO\lfU~t::-. In: fUIOIJt.!o, •\.J, ~· \'A'' KOOn~. C.f...,
lfGnlil)' llunya,iridnc. gcnm, Phl,•IHJl'im4 .. l"'1:,1lons from Olpr:or• (ec!,;·,, T,x,tx,uk tJ(\'irolo;.'V Wtltlrun, and Wilkin,.
(ollccted during an in1«-tpr1.0otic p,:riod in ;;,nyo.Jo11m"/ of Hygr""• 13$ ~1CU.-\~. D•.M.. antQ)t\X, !t I:..\., (iOUU,. ,\,P., OR.-\)3,. ,.s.. .MIUJlJt. )LA. . .
95. 197-209.
•~MtT, ,.,. ,~;;. Callfomln 1•n1·cphalhls ,ims pru,·alcnCi' mroui;t,ou,
138 U~ZA. \1.A., DE FOUAM'. G,A,. \'tJJU., T.~t- .£. Jiw.\ki\.~Ol'NOS, M,C",,, 191'"' the Yu~on 1·ctt11ul}·, 1s-1-1!t;~. A111frlr1111 /011mal o/Trupfrol Mt:d/<111,;
i'Tc,ulencre ra1c, of I.a cros,e,1m< (C:allfomfa encepholltls sroup) in mid 11;-gl,m• 24 676-6111.
lnn,;ie from ovol\,in1n,:d egg, ol ,lro,,, trl>i:rfon,s. Mosq11/ro Xe<,1, 37,
159 AICLl'.'.Jt><:l.:, l,.cunrn·, P.:t., W,\(';~rtl. Ft,J .. LOON<-, \Lk",41\'t.JI.AL~. 1.0•• 1976..
.~750,
J•oletlon of ,now,hoc ha,• ,in11 from,lallt:S lmf)lfMrus lal\'>:io 111
l.39 LUEOltE. A.J., J-OCHlM. ~L)J. & JOXES. H., 1969. BluNo:igue ln tattle!\1tc.-nthL Sa~k.itchc.•\\3n• .\lo111uitn \';m·,. 36, 2:l3-'2S7
Amerlto11 Joim,111 o/Vcr;rl,v,1), Rt'$«1n"l1. 30. 511-Slo.
160 '1EtGA' ,.,... 1919. The Rift van~rfo,·c,r ,·p~,..,,,.;c in F.ll)Jil 19,7-,9.1.
4q LV()V, x.1:,, Tl)tOrM\'A, A.A., (;tc()}l1~511ff\'S,Jr;I. \',l,. Ot.1$8VOV$,:Y, \',f.. Description of thtl cpizMtic :lnd \.irul<>gicn:I sHtdl~s. 1'rn1tsal'tinnltJ/Jltt,.
GROMO\', ;\.I,, TS\'.RJ;JSj nr.,1.. '"')<.nurni:...~KC), .\,G, Ii; i.0).-rYAI«>, •• :,.. •• 19~ Ro)'fll i>l>rlel'f ofTtop/CtJI .\fedlcl11om1d Hygl,11~. -3, 618-623.
'Salchnlhf ,in,s - • n~w a,bavirus isolawd from l:rod,,s (Coratlsode;,
161 :m:i:c.J\~. , ...~ .. UOOGST'R,\.U., It Q ,:oUfjS,\, ).1.(.. 19;9. -~ cpfY,CX,ltC()fRlft
pums l'ick.-C\mb. 1978, collee1ed on Tuten I)· Island. Sea of Okhots-~
Valley fr,..,, in Egi.1>1 ln 1977. T/1~ l'r1,w1a1J, Rttord. 10:;. 124-12;;.
Atcl:h1 ftir gesamn,. 111rnlforsd:u11g. 38. t:U-l38.
a6;l Mtt.<,i\S I- .., .. "thU )l, to,,~ tlO()(OSl'R,\,\l H, ~Afl.HA:\l. f,.K, 1g8o,
:.:1 "''" r.01>, J., 1~6:LTld,s o.nd dlse~~e iu domestic s1ocJ::-Creo1 tiri10,n. l:xpcrimemal transm~s~lon and t:e4d 1so1atlr.in ~tudle> imphr.ahng c:,t.~x
,_.,,rposl<l oftlw 7.nolog,r"/ Soi:iecyof l,111da11. 6, 29-50.
pip1Ms~, a vec1or or !Uft V3Jler fe\<'t Vlru> in Ei;-,1>:, .-\111,•ri(a11 /o,mwl of
1-12 \t,\R~":J1oux. £~,. 61MO~n. J'•L., 1905, L.'\ tran»nlS.sfoa hcreditalre du \'{,u). 1'roJ1tet1l ,\frtlicinr,mul /-(\-glfmr. ~. l405-1410,
de I• fievn- f~unc che,. !c St<'1jOtnyfn {<1.>dma, l))m111~ mu/11 dr In s«t,r,
163 MLL.s:c..i.~ t4L4: )1CnlMI"- M -.. 19fifi. aa,'>iRcatTon nnd nomrncl:uure of
rft Bio/ob-it, 11'«rlJI. S9. 259-260.
a.nimnl ,1ru"~~. l'tDg~"l in .\ltvllt-111 Virv/t>~'. 8, 100-.09.
~a ' L-lRl.uoux, !I".~ st:..10~0. p .. L. 1906.. etudes surla. !il'"\"re inunc .•..\1111a/s dt
114 ,-i·Mun.,,. r, 19t>o. Ahic':in hot<eskkne$..). Journal ofComJ)irmtit1~
J'/111JlitUI, Ptl$t,!ur, 20, 104-tUS..
J'atJ·mf,® m,,t 1111•rr,~uua. ll.1-30.
?44 )L\SQS,J.11,, r.ot.LS. f,U.WJ\. ,..Al F.xA.SOEte. R.A,, ;940. Culdvauono1
16S )ct.11.ll\. a.II.ff.. Ul!ATY, 1',J • .t. LOUR~S. L-1-t•• 19-82., \~atiOll Of l...'\ Cross.e
bluctong»e virus in fertil1%ed egg. produced 011 ~ ,i1amin-<lol'icic,n1 dio1
vfru.s Jllial in:CCr:on tattt~ In g,e<>graphfc:-stmin:. of .·hlflC'S tri;~t'rillWt
\'at,ire. 45. 1022.
tDfptcra: €Uctdae:). }tJUfJJlfl o} .\lt•dict'll E.mb11Wlo,5•. !9.:? 13-214 .
...;s ,.s.\nO~tOTO, ~. 1970. R.11bic~ vtrui ..A:dt1m1c...v In Vlruf !lt9:'<lrch. 16.
1ti6 \ULLUI, B.'R.,Dr JiO.U,Un, c-;,R, 1r. \'UIU l',M., 1917, \'cme.1hrensmis.sion of
2.5;-JOt.
LaCros1e •,in>< (C.ili!nmi3 ~n«-phllll11,gmup1: uansovanal ond fihnl
.;6 ,1~nut,W:., 1(.£.ll,, 193L Thoc:~1,:fi<;atloJ\ und :,omcnclaturc oh1ru~: info<1ion rn1<:>in ,lctltii truermus '1Jip1crn: Culci,fa~,./oumal o/Medkal
,ummary ~r re.suit> ormec1in~ o(tht- lncern:-tlon31 C:ommiue~ on FJ1ronwTogy. 14, •13i-+IQ.
Taxonom) of\~ru ....-s in ,irnsbourg. Augus1, l'ISl, /t11,T11lrolo;f;,, 16,
53-00.
167 Mlt.LIA a.n•• Q.I:.FOUAftT r..R,6 \"'\JIU, T,r.,., 1979.,;°i¥di'S. 1riscnt11ttSnnd ta
Cros~\1rus: lark 01 lnfec1io~ m eggs from th~ tiJ!lttwiltian crd,•
i.; n,e upldemlolog:, of a,lhtopotl-borne ,·lrusK In
" cr1'rosn. o.>i .. t980 Collo\,,ng or.at in(etlllm o(1cmales Amvri,au Journal ofTropi<nl
,ouLhornAfrica. o.s,_ Tho~!$, UnlWrsliyofPretoria. Jitdlc/11~ "'"' nygltn,. 28. 89,-901.
,..:8 ;•,fn~TOSll, IJ.,;,.1,, PICt:a,so;-.;. o.a:.. Sf.R,,\ftS.I. [,T,J. D( sous.~ J,, )96:?~
168 ;'dlU.F.K. Jl.ft. IX \flT(lfll!., C,T, 1$81:>-. fl~ge Of)cillOW r\"\'Cr\'lrU.<t II~ ~ect
,\nribowc:1 ng.i1nn ccncin a.rbo'virl~ in j(:rQ oihun~n bc-tng, snd on lnft.'C::.iu·n and 1111n,ml,~iun r.:u6 In /\c"i/"1 ~·pd..-\mtrir«n }Q11nia/
dom..,lic animals from thr South African ~wld. S0u1/J Aftkon ~fTropicnl 1r,'11Jrl11,•,111d 11,l'IJ"'"~· 33. 1302-1309.
Jouma/ o/M,Ylitt1/ :kil]J1t-e_ 21. 87-94,
169 ~u ,,... r ...-' .. 1u,, ,i,h tit\! \l\'iHAu.. 1,0.. 1966, Cy1opathlc ~fft!'tl orStmtHJ
...;9 :.i<::r:-.'TOSH, 8.M. 1- JUVT\ t•,t:... l,8,. Fpidcmib!ogi~ aspcruofRih \'aHt)'
Fon,si \inh In ll1• mosqui10 ,ltd""''l!i)1Jtl. ,\mm~m /ounu,/ o{Tropfcnl
fen.-r in South Afnco \\.-!th [cference tb \"'eCtO:-$, CrmrrU,,a,onR ro ,w,,,11r1111.•m:d J~ygJ,•m•, 15, tiS- -s-t
Eptdemlt,logyufltl B/om,1!111rs. 3. 92-99.
170 !\ti 1.\'\lUN,\, t. ).IJAOJ.:A. '\). \\ \t.\..~\.HIJ4S.... li0S0t, I.. TIL.,us. §., sou. o.,
~o ,iaxTOSII. 11,M .. Jt,JfJJ.', p.r... 1)0~ .~\;\"JO'- t.~.l 4'MU'~l:.ltA~. G.M•. i9';6.
X.\G.\c.,·1.,. "- JO; 1. t.. sut\UY.u. '-t .. 193.9, Wcltcrt" Unu:uuchuns::,-rn ubN
Cu/ex (Ewm•ltmo,m ill.' mlm1uh1s'tlieobaJd ,1.1o \-CCI or o(Ban71.
di• ll~lwrir.lj!Un~ dcr Jap;inhchi,n cp1<fomt,,:h•n Fn,.cphalhi,durclt
Gerrn~lon nnd \\1~\"i\tc:snmd vfruse-.i. J. Jsolado11 orv1,u~ from wild
Mucken. 1'r1111.<t1Crio11toftltt Soc/Ny/'or 1•111/,c,to~. '''I'""· 2!l, !12-105,
popuhttlon• of C mblr.01115-/01m1nl o/Medica/ liJ11omoloir,, 6, G37-&40.
171 MOUO.:..O\S.t.:l.b.D•• tH0.\11" \ . .S ..\ .. XOSSl'l;A, V~D.. PA\,.0\'.\. Ii.A .. U\'CIUTZ.
~I )l0~'T0SH, a.:i.t.. lUPP, 1',(;., DO!'-SA..Vl'OC., t, IA. '\1f:1'~1-l1Al\, c;,,1,. 19';'6
J.1... Pat.:. H.J.¥< 11nuK'r1:cwA. v.t•.• 19.3;. Studif:.s on papp.:lt:\cl fc-.cr. ll.. On
EplderniCll of\\'e<1 ),'lle ~nd Sindbis \'!rule< ir. S~ulh Africa \\i1h Cttl~x the main1tno.ncc of tho :1pp,mci fever llil'U> in w,dfile,, htt1ch~ frum
{Cul<-x) 1111l1•/11111usTheo~ahl as YL'CJor, SourhAfrlcn11 /011r11nl of'"rlence. cgg,.faid hr tnr..ctc'<l fomall-S.,\/t,/ftr/nsk/1.1'11 Pllm:irolog/)YI /
72.295-300. Par«zuamye JJole=n.1. 6, 922..'JJ:'
:,s2- \IC.1:0."TOSU, 11.'1,, lrr.D>u!'RX01, ft.H, &PATER.SO~"'-· 1LC... 1960~ W it\'ltlWtsrand
1;2: ~:Q:,..·rc-o,1uv; 1::.. l911. On u Urk~bom~g.u;rrocmerws or shc,cp and gos~
\'lrus~ nn apparently ne\-. \·irui s~Jatcd from rulfctnl! mosquhoe.:J... YJu,1> occurring in B1'i1L<h l:.i.'1 .Mrka. /011rn11I u{Cnm1>a1t11i1 .. Pmhol,:,gy, 30,
,Vrirflll }0111>1t1/ of.\/ly/i""I Sc./MC,. 25, 33-~i'.
ZS.57.
:~3 \ICJ!\'TOSII, U..M., ,tCC(UJVIQ\', G,M. • J)l~S.SOS. D.• 1965, 1ugw·.wuma
r ..c.. '\l"IOf'E., R1f'.. & 1-LA1lft1SO;\, \~\..... t911, PhysiO•
f73 \tl-Jki"'Tf\' f,.\., ROAOfN.
,·hus: an a.bO\'trus i~Olatt.od in S0u1h Afric11. Somh -'\frfttm Jo:,nmi of ch~mical ~nd morphulogic•I rclauon,;hips of somq arihropod·bome
.\ltdl<alScicnc,.3-0. a>-70.
\'i.tuks ln Qluetongue ,U\.:.s;-1 new t:rxonomic group. El<tciron
154 \lO:O."lO:SH1 ll,M,, RUSSal. o.. oos SA~TOS.. ,. fl Gt.AA.. ,.11,s.. 1980, Rift\'alley mlct05ropicsiudles.Jo1m1<1/ ofCl'11,raJ l'iroloK)', 13. 2:-3-288.

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168 'if<:nc>, 0\1.: Aspect$ ln!luendng the occurrence of lnfec1io1e, diseM~~

1r.; \fUIU•ff'\', f,A,. lt.o\lUU!SO~, .\,K. & \\'TITTTIB..D. $;(;,. )973, 8Un).J\irlda.c.: ~nhropod-borne eacephalhides. lrmoductlon. Conlribution< of n~w~r
morpl1ologi~ol a,1(! moll)hogcnctle slml!nrities orbunf3l11W~ru fioid·l•b<•rato,y opp:o•ahc,. ·lmu/cn,1 Journal ofHyglt11t, 61), 231-236.
krologia supergroup ,·in,...,s and 'l<',ora other anhropod•bome ,iru..c>- l9,1 "'OU:Ytf'JI. \\.F•• DUl?SCHtSt t..~ ,. ~t(,'(;1,,0Rt.. u.t.. 1959. Ecofogfc sn1dio:s or
lmcrvirolol{)'· 1, 29;..310. J.ipanc,~ encupholhl~ \ina m Jnpru\. V ,\1,,ian fo.i.:tol":lt AM•r/c(Jl1 /oumal
1';5 ~.-.VAK, J,K... EtO!>(S,L.b-K.~IGtn . J.W.. 1986.Lxptorim~nraJ \l!tlk~ o/Tropleal MtJdldnt-and IIJ,gi,mr. 8. 689:·-69i.
,musmi«lon ofS.,int l.0ui•enrcpholitl1; ,1n1, by t'lorida 1M<(!uiLON. lgj SCHERI..L W,f•• \\'r?..\\'tt\+ S.C., t'AYl.()J\. CJ\•• (.i,PP, [.\\'., DIC);at~ll\~. n..w•.&
Aml'ricmr Jo11rnnl o[Trop1M/ ,\1e1(1<1Mtllltf Jlyf/ll'llq, :1;, 1~%-1:101. RU In Ml, n.11 .• 196?. Vector compcwnce of Cul<'.X 1.'<trlm1oc,:m1on)
ti6 s1t.ScuuL1.., a•• 2932. OvN d.t inft."C1It ,~n ).tui7.en mtt her vlrus der WN~lOPUlfOr .sllo;nunc ond cpi:i:o.cnic VcnL.--Xu~lan l!'quinc
/.uld,t\frikaansrho PoucWokte, Twl.<cl,nfr ,wr TJilfrl(V11<'<•iklJ11rfe, 59. enccphllamyc-Hti5 vifw.eJ_ Amf!ri.J..•an Jaumal of TrQJ)icnl Medicinea,:d
),133-i~45. l{>g•tll<, 36, i9,H9i'.
rn rouMtT, r..a.. 198;. ,~;:di!$ rris,•riatus-und Lu Crosse
PATiltCJ.\S~ t -,\, & D"E 196 SCHM.\Uotrs-, c.~ .. 1l\t-TY, ~.r.. fJ\Lk\?1,tp l.fi, f,M, 1, Ll. l)UC, J.w~ 19~
Vtrus: simU;r \•encrc;,J mfecrion rates in LL'mttles given 1hcfus:1 ,\mlgenk nod ge11e11c propenle• p~,ct,intscs Un~ed to homorrha,:ic
bloodmeul immed1a1ei)' before maung 91' sc,·ernJ day$ nfm mitlng. f1.•,·c:r wilh rtnal .):·ndromt: mto a. nt.•Wly,dcfmcd ~rnusof6un;,..1,1rid.a~.
;\marf<an Joumnl ofTr(Jp/ral Med/clmrcuu/ ll.l'f.1CIII!, 36, S.~2. 5<;1'11CI', 2'.!7, 10,11- 1044.
l':"8 PAfRlf'.A~, I .. \., Ul! t'OtJAttr, ~.n..&YUIU, T'\J,.198:;, Oral infection and 19; ~01111r. 1.•.. ~a11,uor. \l.L .r,, \ID, M.t .. 1911. Phlcbo1omu1 rc,c, In
transmb~1on of La (;so~:.-:c vtrus bpm ('ll1,00tfc $1t31n ot Atdt-.,: mwriutu,<: E11ypt tio!3tlon Qf µhlllboromu.s fe\'erfrom Plrl,0/Jmomu• p,11111rasi.
teed Ing on t"hipmutU:s with a rang~ of vm,crn1a lcwns. A1naiaJ11 Jt:mr,wl Am;rlra11 Numa/ ofTroplrnl Medlr/n1t-011rl l{•'glffl,•. 20. 48S-190.
of rrup,ca/ M<Ylici11ea11rf Hyg1('11r, 3-1, 992-99!! ,t,11 <rnuv., , ..... lll,I, Rifl Vall<)' i<l\.1>rtn So111h Afric~. S1=lnllr,pon So,
119 P£TR1$0Ul.\';\, P.i\.. ~ ,., V:,,IOY, .i\,I,, 1938, On traU):OVari:il t:111\.(mi~Jon or ,5151 t>/Mfl)', 19.H,Joh.inncsburg: Union Ocpamncnt ofl 1¢alil,. Plogu~
vinL\ of paJ>pa1ad fcscr by >1>ndfli,-.. ,\rcl1i£,.. oftllu /Jlotoi:lrnl S,xff~>', R~ch l.abornrory,
~3. 138-1~1. 199 .c.orr. 1.w.~ uuruw;t. u,., 19114, R~pid Infection of Sllllvn,yglancl.9 m
180 PLOWntCHTt \\'., PtRR\'. C.T. ,,,_ Pll,HrF., ::.:.,\. 19:0. Tr1msovndft1Initttion 01./1.s.:t(J mi!lanurtJ \\ith L'tt~ern ~ufn~ cncti,h:slili~ \•frus:. nn oh..~trr.m
with Afrk:in \\vine re,,,.
vin1>ln 1he.~tgrud dck, Or1111J1orfoto1 mo11ba1n mlCR)Scope ~ruc.1v..,,,,cn·cmJ Jt111rm1l oJ7mpfcnl .\lt:dic-im.• nmt llygium•.
porcinttf\Valton.. He.search fu Vuteriutt·ry-~lcntt, 11 , 582-SS.l. 33. 961-1)64.
181 J)()$t'n!R.t-1E1.n. J,S,, OIS.\l.S, J •• <:11U:,1.AJ;O\'. ,1.J••• r.A Jl>AM(WttH'. S-"A,, :2ooscon 1,\\1•• 1m>1u·1u.,s,w. & oM·n, it.J., 11)8'1 Th~disnibutionand
HANKOU~. 0. HOf.,,tf.$, I.ti,, HOft.l{Sl:K. ~J.C., \tV'SSC,\Y~ \\. & MUSSl-:U. Jt,)L de\'rtapmem Qf u,u:m equme l'nceph-:lfitl:- ,·fru!- in us-enzooth;
llr.4, 8unytl\il'u$es and llunyUvirfdde. //wrv/rol<>gy, 2, 2i0.-!72. mo~quilo \'ector, Cu/i,se;n mPin,wrtL Amttrfl:011 Jouma.lof T"rDpical
.,r,,i.'riMmut HJ'l/f•nt. 33. 300-3JO.
182 l\GllVllS. w.c.. t9,J Overwintering of urbol.iru$(!;. Pr1>1,re.u /11 .\lclflcni
\/irol"i)', !,. 193-220. 201 ~•11.tR.." R.F•• 19,Sq. \\"eu1h~. host and \"CC"tor- thffdnt('[l)loy in the
,p,ead ofinse<.-i-bomc ;mimnl \'IN$ dbe~•<!',/011m~/ of H)gf,iw. 85.
183 IU!P.\"t,s, \\',C,. JIUT$()X, Co.A,, RFU~\lt\'. ft,f.~ SCRJ\fA.!\iT, k.P., 1nt. c:hnmic
oS-JO?.
latcm infections ofbitds \\'hh Wf.',,tt!fn cqumc c:m:c1>halom:•elit;s viru~
Pr«eeding,Hif ti:,• ,\n<,ny/nr F.vp,~rim,m1,1I Hu>tnc,•rrnrl 1\f1•dirinr 91 ~ ,a.tflru,., R.r., QIUlS, LP.. HI R"-1\t..\:\', ...., •• vrncu•.-, n,r,.. •TUtf{l'Ul. '\$,R..
T&:l-736. 1~·9. Possible origin of tht blu<1onID<C cpldemk In Cypru.,. August,
19;7.Jtrm:nl offl>)lf.mt. 8!1, 547-555,
184 rt,RJtACU:, 1.. 1962.. Tnuu,o,..1ritd mcn.s:mls.sion t>ftkk•hr.:irnc i:nr"ph~litt,.
,;ru~ by-lick~ .,rm
\1rologla,. G. ?20-121,, zoa S.tW 1\5-, R.J •• PtOGCLE\', o.t Ai TlJO{I n. M.J't .. 1.97'. Po~lble $.prtad of
\fricanhon<cs'iclm,~, on rho •,,ind. Jo/Jf//0/ ofH,1gie11,, 79.2i9-298.
185 1t0, u,. L, L96; Sl.'l:ual trMsml«hm of d,nguc ,;ruw-, br ,·lwl,,
nlbo1>icw<. ,lmtrfcn11 Joumnl of 'f'ropkm .lfrlllcfll1• 11111/ Hyg/"14'. 37, 2<M ~•u.1 Rtr-.- "4f,, Pl!.l)(j.EJY n.1 t,. TVc.;JO:n. M,tt•• 1978, Po-.siblc.• wlndbc,m('
398-402. spre4d orbluowngu• 1n Ponug.t. Junc•July, 1951>. fm,rn.nl of H;)/ien,•,
81 189-196.
186 ~OSl:i'\ 1.,, 1988_ funh1,4rob ~erv~tin~ on~<- n·u:thnnisin o! \'fflkal
t),(i, ,- TUl';1;Dt. )1,1\., l!*S2.. Rift Val!~) fovcr Egypt,
::.05·:t-r:LUR""' lt,1'",i i-CDGU,i.\'
1ro1mnls.,lnn ofllnvMruses b)' A,•t!t$ n,o,;quhtx.'1i, Amtrlttt11 /01mwl of
'Trop1't:J1J Mt(flcine 1111</ Hj'8/em• 39. 123-126. l9:7: Ob,c:i~e ~preod by \\1ndbamc in:wcr \·cc tor)? The v,1orlno1j1
Rern,d, J 10. 73-11.
18; l-t0$1!N. I.•• JU')~flU)0\1.1.-r .• cut.CR. o., .. W..\: ,.~.&(;JC,\SIOi l$. ns. 1985,.
206 ~11a<.1\n. o.A.. 1~··rnm10\·,.uiuJ m,,fnrcn:inc\! of San ,·\n·gclo \'lnH ln
Comparoll"' wsteptibllity <If mosqui10 spedesandstmln, to oral und
patcnud infocrlot, \\1th den~ue and Jap:snt:;e cnr.cphaliti~ \.iru.w'". ,cquen!lol gcntr:nlon. o;',llllfl'J nt/Jo11i,ws..,meri<'mr Jwmnl o/Tmpl<n/
.• \1m:rlct111 Juum~J a/Truplttll ,'1edfc111...• tmd Jlj'8lt.•,n•. l-1, 603-()15.
Mcdirlrwtuul H,vglNU•r 1~. O:M-417
20; ~>~1PSO:'\. tu.11 .. 19-72. Arbo\''lrur,; <.ll~cnscs. Brllf,,;, .\!(11/fcnl #ulfcrin. 28.
1&; r(oS~. t.... sHHOYI.K. v ..,.. • u.E.,.,\ 1.t:.• 1S5c,, Transovarinl tra:nsnltsSfon of
10-1.;
Japan.($.e encephalitis virus by Cttli'x 1ritatniorhynclw'Smo"'quft0t."S,
:\nu,rlca,, Jou ma I of1iopt<,1l M4!dfdmumd HyglPur, 29, 71 1-712. 1=:1ctorli m the1.ransmissl1:,n o( virus !n(ec:dons irom
20Ss;-.im1, c.1:.c.. 196;
anuml, 10 n,un. Sc/rm/fie Bali.< ~fMftflch1<• ,lmrual lli'111't•.•· 8, 12S.-l ;o.
189 HO.Sex. ~ SIIRO,T,M. 0,A,. ltiSII. fl.fl,, FR.aal. ,.t. & u r ~. ,.c•• 1983.
Transo."'W'hl! t"3.Mml~1on ordt:-ngue \i.ru~ by mo!-quiioes; •.;..,,tfe:s 209 (Mrm. c.t.(,,, 1963. Recent nckanccs in arbr)\'trus fl':S\"arch. Ab.Jtm,rs 011
a//xJpicm<and M1tcsaq;Jpli•.tlmU:cn,1 in,mwl of Tropical .111!1ficl11t 11111/ 11,-;zlr,,r, ~3. 139,-1,:3~.
H,1ogia11e. 3Z. I I OIH I I 9. >JO ,)llTII, c.e.c_ <97L The role of "lru< dfwa"'< in uop!cal public hc~hh.
190 KOS£...:, t... l ES.H. a..e•. ut:..~. J.t'- &CitOSS. t,fL. 1973. Transo\1tr'4l Tran,<11,1/0111 oft//t llo;nl So;fr·tJ o/Troplcnl .IIN1fc/n,1111d I !ygfNlf'. 65,
trlln$fllis..•,:ton ofJap:inc$r t•nccphnlitisvirui. by mosquitoc-.s. S<t~·ncc, 199, Supp!. 73-32.
909-911. :au >.Mffil, c.t.o. tn;"- Ch:lnging pnaorn, of dls~ose in lhc U'Oples. Hrl/lsh
191 ~'\.LUU(), 1,1-.. HHtvl', f,P,,MLAU~. l~J., G0.'.14,N, M,,COflM:T. ;,fl',, C,.:\ J\,'\IIC\S, ,\/,t.ifet1' 8u/ler/n, 28. '.3-9.
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J.1... HfMe. G. RQBL~, Y,, 1980, Cnrncte~r£que~des ~>uc.h~,. du nrut- de ~ S)JtTIIDVMX. K.C:., IL\000\\', \.r, 1,1.m,,~nH~. \\'JUt..• 19.;9, Outbreak of
la fo:\'n' jnunf.!' ts.oleesde, oe.uf~t>t des l3n-c, d'ur,e tiqwt t\mb{:,'()mma -syl\.)ll yellow (ever in U<:3nd;i \\llthAcdes (nc,wmyUIJ afri«flUJJ
11111'feganm, recohe~ ,uc le bciuil • 8;\ngni IC•ntra!igue). .~1,n{l/l'S dr 1hoob>ld .. princtp~ ,·«tor un<I m~cc:t host of\iru"' ,111r.{I/., o[Tropn'al
Viro/c,gie, 131E. !5S-.l6';, ,\/('dicinennd l>ata.,110/0,:;-. -13. 7-1-8!1,
192. ,:i,\_\lt)ll,0\'i\, T.1 ~ 0,1\SJlOVA, G,M., 197.;. The tltk l;i.-r){Jl!J
riclmlJ. il'S .Q ~13 :,.,~cnumnx, ~.., .• X 01"Ftc;>:QT, fl.tt .. lU'.'VM.ANS, (....)•. W'EISB1U:1~. M ,P• .Jc
Uukunicml ,i.rus \'CCtOr. 11. 1'raMmlssion and iranwv:irnu trammlss,on ,..r..., 1KOWSKY. o.. 1959. ~eum1Hzing ontibodi,, for cen~invil"U$~\ in the
ln ;,dult dt\"'Clopmenml state. hr von'.U.O\", , . .1. ~ ,·vcw, o ..... eds .. sero 11fhuman bring, l'l!Sidlng In nonhorn ,'-awl. .Soutlr ;friwn M4'dl(o/
Ec<>/og)' of l'ir11sesA<s()(/n11,11virh Bird$. Mins!!. for,r,u,l 33. 5$5-56 I
1'93 M:IIAF"Ffl'.11, ,1.~ Aft~Ot.D, E,11.. 195-4. ~tudif",. on the Nqnh .-\m~-ft'An

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·\mt!rl,m1 !mm:nl or 'fropfrol
,lj.rlr11/111rr, !rn111,.tt1ljlir IPIU-//Jll,, 110-121.
\l,tlid11~tlltrl fiYl(l•llt.8. IOl-164.
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l?lit-122• • ,\lrrl1rt1h· an(( 11.ltrfenl', Z':". 18;'-l!h,.
::::i.:- 1,\KAJL\.',1U, M . J. -SU'LU);"J~ .,,. t9':'9, lapanc~ t'n~plt~t.litis \itus tn rnosqu..uo ~.16 TM,m~. r ,ir111 .. •· "'"'" n. •· t9;tl, Cnlifornl3<'1\C1'phallri, g.'1>up
still\ ary Jll•nds. ,1mmt<>11 /011,11<1/ 1>/Trop1(n/ ,\/rdlti1u>n11d li};,'Wm•. 28. \iru:-.(•t L-.ol;.1h.'Cf (rum mo~uhor,<c,Uct"u..·d in ~uz.h"'~m An:11.c ~on,o1y
122-135. .-lrm Pmliotr>iica 1•1 \lftrol1iutogl(,) S<i2m//11"1cit, ~eci. R, 86. 3~-3.JI
~8 1 nunr. tL ,1 .• l9tr·. r..:,;Llloguc or~,r1hmpod·bo-:nr ,,ms.es oi du: worid. A u; 1u1ut1 ,,1. e.,n.u•. C.L "nft(.q, c- ,. 198.;. lnnc.:i,;cd mos.qw,o feedingun
coll~11on nfdarn on n,gi<tNt'd ar1hropod·bome onirnnl ,iruse,. U.S. FMt \ alh."\ i1.•\'(•: \1n1,,fnft.-ctt:J tamb-...-\mtrl1..·wi /r.mrnnl o/Trop;rnl
Puhf/r ll-.•«1flt .,cr:.fCt' Pub1ica11on \'c,_ l:ti(I. \\Uihmgton, D.C: U.S. '.li-r/h·i1t\• mul fl~1·gi1m1• :ri. f '?32-1238.
Govcmm,.mt Printins: Offil'c. \\'a.sh,ngtori. O.C
2.J.8 n.mn.1- M.J .. G..UtG\ ....:, r..v.. •11'AtU\. t. .t~, 1~84 Hf'plkatlon;,nd
;e..".>g l.\\"l.OA, R.M.. 1111"1 nu I. fl, ..... WO~. I.Jt >i1,r.~~os. >· t\, '¥1 H(t()f.~M\L, JI .. d'(... ')f.'nlln,1tl"\n ofR1h \ :111~· , .....·~r \'IN~ In n,tru 11i1n,.ns•.\mtrlrvm
1:i.Gfi. ArbO\'lnlS'L."S hol:m,>ti uom ~·· Udo, ;n EJ::)-p1: Quar.;uifil. Chtnud~ foun,al of nu,,t,~,t 1\hV/1(111,-m:rl l-fygl1·11r. J..'$, 1i'--1111~
.:md Xyrun.inini _.-tm,nc:1111 Joirrnlrl o/'l'ro11t<ol .\/1't/icitw anti Hygrem!. 15.
;(>.86 24g ruREtl .,u.• 1un1>'1'. f.l A tcHn~ \\.t ~9&v..S1.,hilfz<-d i.nf\."(tlon of

C?llfo1 nl,1 ~n,:cphru.ltl-' \')ru,,. in..\t.vht-· 1tor<J1/lt and 1r, 1mpJkatfons for
ZJOTmr, R,IJ.. 1$UO. ! .'tpcrimcnr;,.l "1udii.'t on ihc U..Ul.tO\.:irfal tninsmtsslonof \ir'.tl nui.Jttcr.m1cc: ln m1ture ...\,111•r-J,:n,tJtm11wJ o/Twµitt,1 \J<"tiletm.'mul
~unjin ond S4n An~e-lo nru,c~ in mo,quitocs.Amc-n·ra,i Journal fJj f!)gl,,i,.JI. :!.52-l2.;9.
Tropi(t]I Mt"1/,;,1wo11d l(.1-gw,1e. 29, 65;~.
25() runt.LI_ \I.J.. t, tvr.~. W ('_"II \Hf)\', J.I .. 198:,. Trnnsn\"ari:,.J ~r.d
Zll rn1<. "·" . 1981, V<nical ltnn,mls,lon of anhrcl)<ld·bome vint><'> tJf rt.1n"1o,,1:idlttJ trun"inlioC:<ion of f.nJlfomfo NU:t:J')ha.Utl~ \iflB 1n :\t'des
\·ertc.-br.llt"\, /n: ~t<.~'TL\CT, l,J,, (tl'))U(,t n.t. , ....1AHM1CJ1tl.)$Cll, K, it:dc.'.,
dursnli..,· and .\L'dt·~ mt1lu1amou. -~u,,,!t,m Jour,ur/ c,/Trop!<:tJl ,\1otftcl,:r
VtettJrs tJf 1)/.·w,JS, :\g,·m.,.. '.:i...\' \'c,rk,: l'n-11.-gcr f'ubli~h~~. audli.\-'KW'u•.31 . tO.ll-102~.
23:1 -i~H fut,. 1981. Tranwvartal tu11\)1ni,1'h:m of ubovinrn,'tli in their
2:;.1 ruru:u. ~1.J.. •tin-~. w.<· k H.\RO\, J.L.. 1g82 fa-nlu:t:ion t>l chcclfsch:nC\·
tf\\Ctt~br-.ne \\:C-IOr, /11: U.\J.lkl~, t..r. ....d 1,. Cttrtatltl tuplCI UI h~lOf ofrTIJ.n.'liO\.t.rltl. lftin(mttr.J.-ton nfc,Jifom1J Cn('t.•phnllillo ~. tta.1 strcin~ in
r~ffl1rd1. \'nl II :-;.,,, \ or!. Pmoiµ,r Publl,h~1<
Aet!cs dlJl'JU.lL< uttd ,h..,h.·.s nwla11{mo11. J\11h'ri,m1Juumal of Troplatl
m n.,,n n.n ~C':UAM011, »- , .. l97!t. Tr:m.SOHtrillJ 1'1Hll'lmis,ion nf\iNSCi by .\l-.:tftt-111,• ,md ll."1f"'""· 3l, J&:?-388.
ph1'cbmominf! S$!ndlli~ _.4111w/.t uf th<' \'au• >'or!: .4Ntdt~my of5<tr,1tt"'$,
~ TIJRUJ ,1.,. XO..'$), C,.\, ._ 0.\10,, C..L. 1~5. Effl'Ct o! itttr".n.S-ic intubation
266. 125-1:~. h.'mP4,::ro1:tur...-nn thl' .1.blllt\· 0 14•\,:d•.., JjJpiPnJ. to tt,,n ..mu Rltl \"ttU&)· fohir
23.i n:st.t. 1t.8.. t..n.,~1011s. -..s. & 1011:--·~s. K.M.• 19:~. \'t"Slcular .stomttti1' \irus \il'WI ;\11:t!rllrm Juurnul 1,.fTm11it-nJ !,f,vflril,rnnd f ft.1.,;l'ni• 3 I 1211-
•lndi.1n11.scrotype1. Tmn!>-,)\'Ari:11 tr.m-,mlj~!OO of phfcbl'IU>minc 1~18.
~:indllle-s. Sc.i.t'JJC<, 17.S, l,t:'7-1.;79.,
2.;3 \"\S UFR WESIHUIZC \, H- 196:. 5tudi(~ on OO\i.n1.~ ephemeral re\tt, I.
235 1"£511. 1(,..,., C){t\;o..!Ull~. 11...'\:' •• t'lTI;\LT.\, fl.IL 6 IOJI~~'.\ ).,\1., 197.;. Ecology of 1;.ol.ation u.nd pn•Umtn~i~· cMrn\'.teri7,c11lo11 <tf ., ,·iru~ from namml and
viruses Cs,ol:u~d from P'..1nomnni:sn ph1cbotomi:ne Sii!ndfltes. Nm!rtcm, t-XJJl'rirr.(~nrn.Ur pmUut,)Ll t.a<~ or btaYinc ephcmer.11 fe-t•l'r.
Jtmrnbl c,f rro1>Jral ,\ltvllti11-.•uml JIJ'l:h?lt'. ~3. :?.:i.8-269. Oml,•Mti•ptH,rl Jtmnt(}/ 11/\ 't.·tl'rlntJI}" Rt·.ttnMh. 34, ?..,4-;0.

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170 ,,cu" o-.:i: .-\>pect, intluendng the occurrence of lnfectiou, dl~:»b

::3.1 \\"AU.Ji,.. ,,.P•••\fl ~r,. I ,H,C:.. Ut!An. JI,., .• l,01\,L'\"l, L, •.\'/,L\lO, t,.p. '-' A1111·r~,,.~,, /011nml (,f T1,,11frftl MMflrim~tmd l(\-"8Wm.·. 35, 6g...623.
u.t1..\tu~1c1rr;. ,,.,.. ,~a;; Sc.•it<hon (or -,U~(c.tptibillty and 1~fr.1i:1orin~~ c,t .!6-: \\ ll~8fU , . ,1,11 .. HE\'.\1.\,:\, C.S., 1{0):E.lt.'-iOI, Jc..H". ir P\'fllRSO~. 11.C- l9Si,
.~1.'1ii'J d;.1!)'JJli to o:-1Unfcc1icm \,irh ~·t"Uu,.. k\'t!r, iro,..Amer,·tan /011mnl Mmfa.~ on ,tnhmpc,d bonH! \.1ru~th or Tong.tla.nd \'II. S:mbu \UU!>."
4

t)jTropiral .\lr'tlicmctaml Uygir1h·, :l.;. t!Z5-J?31. hllhc.-:tc unlJ10\,n -1gcm1 1.. olJh:d frnn1 ·kc.ft'$ '"Banl:Rn,•llal
25S \\ \Tl~ D, ,,_ bUR.l;t. o., U,\JUU!tiO~. JL\. \\JITT)Jllt!. !t.E, 6. \l~W.t,;, \ , tftt·wnl:Jtt'Olm ThcrJ ~uth .1ftt<tm Jourll(J( of .\!1.·diml Schmtr. 22.
198:. f.O'f..~t uf h.'h1p\!t1Uutc t')tt lhl· \Ll('tOrtffiCll.'t\l"}' or At-'l'1,~$(l~·ptt Cur ~3-102.
cfen~ut.~ :i! ,·uui.. ,·\fttt"n,"tw Juur1tul t>/ 1ropt<al .\Jetlttuu;1 ani.l .H_1gtif1lt". J6, :63 \\"t.:~am-... \Ir, )iQKnt~o,. R.H. -i. J.)UTil81JRS. K,C. 1,l6.Slt31fl4i.Of
l{'.I 1,2.
'£.im.ibis·lU..c \'lru'll 1so1u.h.-d rro:n C\11i<:uw :;:c);;qwtod in the Union or
~ss ,,,An~. o.M, ,. tU)RJI><,L. IU •• 19'i'.;. frilnsr,·.ui.ut mansmi.~Si•Jn uf !><>Ulh .',inc.,. So1t1lt ,VrlCtnl .\tl'tli<l>l Joumnl. JO. o31~36.
:itb(l\'IJ'\1$C> l>)' n\Oj(l\lilO<~ ,\ ""'"' .\f;dir,:/ BiDfoi()', 53. ?'.' 1-2:ij. ;:O.i \\ l:L!o"', fi.t... IIAI(,. !;)..\. ti!. .\U~<\..,U.Eff, fe.,J\., 195,:., \\'es,.elsbron ,'ln.-$ - ~
257 \\''\TN,, O,M,, r \'\/TU\\'\T\~,A. s.. 011 .-ou.,n.~. c..n.. \'UIU.,,-,):,,. ffiO\t~;o... \·1ru, ncn prcvmusly dL"'-Cflbt..-d. ,s.~~lc1u1ed \,ith iJbon,on tn Uomc."'.!i,tic
\\ .Ii .. 197'3, t r.m~O\'ilnnl lrnn'>m,s.s-lon of Lu. Cro:-sc \'iru) (CaHfomu amniul, ()111/,·~rt/J<)(lrJ /oumal o[V<1rri11aryli,.,<1,.,1,. r. 18}-195.
eoccphrul~ R-te)up in du: mo~1mto••·\i"-_ics 1rM~rco1111, Science. 1l40-
;n;.3 ,,.111..,-ru1t. r. &.-:twA,s1~'1(>1 "" 1stU.t Chn.rnctcrtlaiion o( potcml.atl)'
IHI,
(oc,1ouup!c t•olyam ~erogroup orblnru~ i1,01ated in l.in\bab·wc. /tJumnl
~ \\
1
AT1$, f), ,, •• tttm,Hk,OS. \\,II,, \'IULL r.,, , Oli fQLtMn. G.lt. & U.\~>0'-. N.J•,. of(,'t-nc:UII \litQlu~·. 6~. 2;?2 l-222i'
1974, Ovam-inwring of La rro,.w \'irui ln ..U.-><lts rrlierlow.,. Amrrf~m
2b6 \\ttL,1 l l'H I,. ~\\'"\~I f"OEL a.~ l:JL\";.\HJ:,. 11.t.. lt,J$9, ('.ha1ructt:ri1.:iU.ion of
Joi1111t1r t>}Tropl<<tl Mwll<lm• 1111</ H;-g1wu. 23, 694-700.
P;ll)·uin ,m>gmup crbi,·fru;,~ !,qla«1I in South Afrlro •nd serologfc
259 WtJA\/Tlli, S-.<;•• 19Sl~ Electron mh::rP'rop:t .1.naJ~,i> onor~tlcm patu.+ms c,·ldcnc:c for their \\'1di.·,prrnd J1 ..1ribuLion ut the ruunny. Fp/dt'm/111,,g:.·
(orV~llC"lUC.1nn tquinc \..'fltcphaJom~d.itl, nnJS in the\ ector mr.»quho, and lnfccrlor,, IQ!!..\!':-:UL
C11/u (Mt•lm10<.r,n/1m) latmopus .~n11•rl(ar1 Jcmrll(1f tJ/Tl'c>plrol .\lidfrm('
:r,; WHJTTIW>.....c., ,1u11l'H\, • \. >. ,u1>: ,. w .. ,.o. 1973- SI 1.oul~enccphru.1.1.i,
01111 Hyglt11,. 3;_ Ill-Hill.
"''N...__ .sn uhrJ-.tructur,1' -.,ud) o( lnfocdon in a nitKquhtl ,«cor•
.260 \\'£:\\'£A, s..c.. SOCt;lQ.:.ft, W.f.. tUt11', E"~W. ,.. C.\Snu.o. 0 ...\.., 198.:t Ba.mm Hl \·itO/t)~·, .\(ii, 70--87
di~min3unn o!Venttuefon ~nccpha1lt\ \1NS1,.~ In chc, mfdi.th.•
168 '"mn,,l-:11~. 11.r, .. t.:f!.! , Thl· •~NIQh>~· of phlt-botomu( (~:er.Journal of
Amerkon enzooti<" Vi.'<'IOr mosquito Culr., 1MrlontJC'.fmtcm~ ,ninfoput.
:tm,ri<'tm Joumnl ol r,oplml ,\1<,Hd11,•,mcl J{11{ium·. :n. 953-960. Suu..· ,\J1•ditin.-. ~i:!. 461-:Ei<J,
~f,g \\"Jt..1>\, t•.. 197i. CltWljt(afi01! cmd \"omttrct,,w,.._•oft'!tu,ts. Sa.sc1: S
~l WL\\'1°ff... 5,C., .SCIIFJllR. \\'.t,:. TM10Jt, ('" \ .• t.t::,.IW,0, I),,\, l- CUP\\ e..w.•
t986... L.lbonuory vecror compeicm« of Cul(..t <Mr!onor.orlion,' <<d«d for 1'>11:<'t AG.
~ympauc: iand allop:ulic Vcnozm.·Tan 1.."({t!in~ ,~<.:epOO!om}·clitis \'iruscs. ~40 lUUA"'O\-. \ ,,1 .. 1a-;,;. !nttgradon ot \iral genome!. Snwre. Z:-.JO, .,n-;i3
1

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 8
Special factors affecting the control
of livestock diseases in
sub-Saharan Africa

~}-1. PENRITH AN D G R THOMSOi':

lncroductlon Studies In this area have generally shown that returns on

such in"es1ment are highl) posirive (see Chapter 9: The
Although livestock production, panicularly in 1he arid and control of infectious diseases of livestock: Malting appro-
semi-arid regions of sub-Saharan Africa. is of vital impon- priate decisions in different epidemiological and socio-
ance a, a food source and for 1rade. it is also an lmegral pan economic conditions), 1o1 although there Is a l'lew that the
of social custom. Lradition and wealLh. and therefore of cost of eradicating the 200 I foot-and-momh disease (Hv1DJ
greater value than can be reckoned in mere fi nancial term~. outbreal- ln the UK wa~ grearer 1han the benefl1 it achieved.
Furthermore. because epidemic diseaJ;CS are more diverse The argu ment Is that a cheaper. albeit more time•consum-
and prevalem in these regions, animal producrion and Ing. approach would have been more appropriate. 15 The
household food securlry are mort! cornaraincd by dbeescs second aspt!.C:t rela tes 10 who pays for control or eradication.
rhan elsewhere,4 as many of the chaprers in this book atte.sl. In the modern \\orld It is generally accepted that the ·u~er·
The fact that many of the Office International des Epizootic, - more accurately 1he 'beneficiary· - pars. This presents
(OlE) I.isl 1\ diseases (i.e. those considered mos1 dangerous two problem~ because (I in sub -Saharan Africa most c:om-
in terms of international trade) are indigenous 10 Africa. muni1ie) dependent upon lll'c.stock are impoverished. and
often makes it difficult for produc1s that originate in Africa m (2) when it comes 10 effective control or e radication of epi·
gain acccprnnce co imernational market~ \\'here the hest demic dh\ease,. the! b~neflciary is not confined to live.stock
prices p revail.2 A result is the perceprlon -one hopei. erro- ownet'$. It is generally ai;cepted thererore that the State ha~
neous - that dis.case comrol is genernlly doomed to lailure this responsibillcy but, once ngain. many states in sub
in most parts of the coorinent. Saharan Afri<:a have rarely been able to afford large-scale
Successful comrol or eradication ornves1ock diseases re- comroJ1eradica1ion programmes. The eradication of conta-
qulr~ a holistic approach that in,·ol\'e, appropnate animal gious bo,·ine plcuropneumonia (CBPP) from :,;gamiland
husbandry practices. careful surveillance and control mea- (nonh•\\CSt Botswana) in 1996197 is a remarkable excep-
sures such as zoosanilary provisions 1har are enforced \'<IC· tion. \\11en the disease appt'ared after an absence of more
cination. chemoprophylaKis a nd chemotherapy. To be than 50 years, 1 the governm,:,n1 of Botswana devo ted more
sustainable these actions must be cos,-effec1ive and pracrt- than USS 350 million 10 1ha1 cau;c. At presem. internatlonal
cal. and be recognized by livestock owners as necessary and donors make very significant contriburions to the comrolt
appropriate. Ideally. livestock owners and animal healrh au- eradication of epidemic animal diseases in sub-Saharan
thorities need 10 co-operate in endeavours aimed at more Africa but these are not sustainable in the long term.
effecijve disease control. Unformnately. In sub-Saharan ,\f. NevertJ1elcss, donor-assisted programmes such as P:\RC
rica, this ideal b rarely achieved, which is arguably the major (Pan-African Rindcrpe:;1 Campaign) and its suc('es,or. !'!\CF
reason why animal disease control is generally less cfl'ectin• (Pan-Arrican Programme for the Control of EpizooticsJ.
in this regio1, than In most other parts of the world. both of which were funded br the Food and .\gricullurC' Or·
Another well-recognized constraim is tinance, to which ganization, have made \~£al contributions to the eradication
there are two aspects. In the first place, there is the question of rinderpest :rom large parts of ·\.frica as well as to im1>t0\ ed
of the remrn on the investment in conrrolling an imal health surveillance programmes.
I 7l

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 ,_
1-., " mo, " ": Aspec1s Influencing 1he occurrence of mfec1iou, dl~c3si.'S

In the C\'Cnt of epidemics. the measures and approaches group frequen1f~ live on either side of a border and people
necessary 10 comrol/erac!ica1e t!iem and prevem 1rans- and hvestock mo,-e freely from one ~ide co 1he other. Ylos1
bo1mdary ~pread are generally prescribed by international imponant. however. is the requirement for pastoralists. \\'ho
convention based on e~pcrience and approach~ acccp1ed live in a wide belt acro,s the continent. to move long
in 1he de,·eloped world. panicularly western Europe and distances. lrrcspec11\'e of c:ountrv boundaries. in search ot
"lonh America l n prac-1icc. i1 is olten not po.sible 10 appl) water and be1 ter grazing. Any auemp1 to s1op h~toric irans-
these measures effec1ively In sub-Saharan coumrie, be· human practices ls usually interpreted by pastoralists as an
cause of financial and/or logistical reasons. This sometimes attempl co ensure their demise. IJ1 addition, even within
leads 10 1he arcusa1ion tha1 the mea~ures applied to comrol coun1rie~ there Is often Jiule differentiation between differ·
an outbteai.. were not properly implcmen1ed and hence the em land-use areas (e.g. wildlife and farming areas), and fre·
lack of success. However. It is of1cn 1101 recognized by do· quentl) livestock are miu1aged on a communal basis rather
nors and international agencies 1ha1 th11 methods prf!· than by individual farmers.
scribed have no prospect of b~ing successful in the l\nerc outbreak1< of highly contagious dise-c1Ses are diag-
environment in which they arc applied. 17 There is therefore nosed. their subsequent control/ eradication ideally requires
a pressing need for control nwasures c,f epidemic disease;. in immediate deployment oi people and resourc~ to effec1
sub-Saharan Africa to be more effectively researched in movement tontrol so 1hat the affected popularic,n is isolated
order Lo develop more practical siratcgies to o,·ercome the from adjacent susceptible populations and transmission inter·
special conditions that prevnil. rupied. lloadblock,, arc the most Ob\ious mean$ ro this end.
Even in high!} developed countries. the possibiliry that Ho\,'C\·er. in mo,1 sub-Sahar.m countries our experience ha~
conventional control measures may not achieve the de.sired been that roadblocks have been esiablished vei:y late in the
result without excessive financial losses has been demon- course of the outbreak. if at all. and ha\-e not been effective in
strated during the outbreaks of classical s,,ine fever (hog pre\'entlng 1110\,mll.'nt of animals and live.stock products. 8\l·
cholera) in the :'\etherlands in 1996 and f'\ID in the UK in reaucratic delays and slow mobili?..ation of the necessary funds
2001. Failure to recognize disease early enough co pren•111 are at the n>ot of tl,c, problem. Although most counrries ha\'e
spread usually lies a1 the root of the problem. ,\frer the dis• permanent police and/or anny checkpoints along major
ease has been diagnosed. logistics. socio-economic faclOrs roads, these fall under ministries unrelated 10 agricu.Jrure. The
and politics can all comribute to the failurn of comrol/eradi· mt-chani,m 10 allow them to act as veterinary checkpoints
cation measures wherever they are applied. However, the often docs not exist. o.nd complicated negotiations at ministe·
nature of the logi,tical. pol11ical and socio-economic prob· rial level arc nec~sary either to allow a ,·eterinary presence al
lem5 differ~ berwccn developed and developing countries. t.he points or to a111.h1,rize the regular persannel to search for
,\pan from the fim111cial constraints discus~ed above. live<tockand for products. The establishment of 11 pem1ancm
other factor~ conrributing to the failure of control, eradica· m1er-ministcrial committee tC) respond to emergenclesolfe~a
tlon programme5 lncluclc the lack oradequa1eoradequatel)· pr;ssible solution. l lowe\'.;r. even if it is possible 10 establish
trained and motivau:d human re~ources, lhe lack of infra. \'eteainru:y checkpoints. !hey are seldom successful in pre\·em-
srructure. discipline. polirkal \,ill. and, often. the lack of in· lng movement 0111 of tht' infected area. There are :.everal
formation. In addi1ion 101h.is, sub.Saharan Africa continues reasons for this. In t\fric.a. movement of liv51ock is not
10 he affiicted by civil strife, rendering e1Tec1i¥e animal dis· restricted to roads, e"cn if t.he movement is across
ease control impo,,sible in regions where insecurity exists. intemationnl borders. :>·!any people on foor and on bicycles
Enm when pence prevails and the resources necessary for 1,ill b)'!)a,s any checkpoints placed on roads. particularly when
comrol can be supplied either by the government or h}' moving Illegal material. \luch trans.boundary movement
donor organizarion5, there are or1en cogt>nt reasons why takes place in t.he :<ame woy. p.irricularly in an.'il.s where the
conventional comrol meosure$ cannot work. border. which ma) be a road or a river, cuts t.hrough )ertle-
mcms in which pi;ople are related 10 one another. Thorough
Control measures ,earc:hL-"' of trailir pll!>sing through roadblocks are also imprac·
1ical. \lany roads c,my heav\· traffic. much of which consists of
Quarantine and movement control hea\ily loclen public and long·dis1ance cargo tnnsport. The
fn the managemem of comagious diseases, strict mo,11ment lime requln:d for adrqunte warches would bring traffic to a
control mu:.t be <mforccable for quarantine and other ~tandstill for unacceptably long period~ and would in fac1 en-
zoosanilar) mea~ures to be effccai,·e. In this (.'011te:>.1 it b im· courage .hC' u,c of mltim rnu1c;,. f'unhcnnur". uffidals man-
ponanl Lo remember that movement control or li\·es1ock in ning roadblock~ are generally underpaid and poorly
most parts ofsub·Saharan \frica -some countries in south· motivated, and may not confisca111 animals-Or their product~ if
em Africa being an exception - is difficult because the bor· offered inducements to ignore chem.
ders berween most countrh:s are 'open·. i.e. there is usually n,e pre~surt> 10 Indulge in Illegal mo,·emt'!ntS can be very
no indication on the ground as to where the border i~. and ,;trong among poor llve~1ock owner--. Lhestoc.kof1en consti-
they are often not controlled. ;\!embers of the same ethnic tute, 1hcir only rc~ource. and 1hey han• no reserves o·n

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 Spcclnl fac[ors atrcc,ing !he con[ro! of lh·es1ock d~ease:s ,n sub-Saharan \(nca l ~3

1·.hich 10 fall bad, when the marketing of th1>ir anim2!s i; necessary products ;md ,uilklent fioondal means. e i1her on
prevented. Entire communiiie,. are often affected. so lhal no the part of go\·ernmcnt or 1he owners. 10 purchase th.em. Jn
money other chan whai can be obtained by selling animals areas far from ci1ic~~ and towns. or, a~ is che case in many
or Lheir pn1duc1s is available. Depending on the ex1em oflhe coumries. in di regions orher than the capital city, product~
outbreak. this situation may be protracted. r.urthermore, it are often ~imply nm a\-ailable. This is fur1her complicated
is frequently beyond 1he power or volicion o( govern menu, wh,m produces. usually 11?ccines, require an unbroken cold
in sub-Saharan Africa to insist on a complete ce:;:;ation of chain in order to retain \·iability. lnabili1y 10 read the in-
trade for the duration of the ou1break. particularli if no structioru supplied in order co U$e the produces correctly is
compensarion is po~sible. !'or e,xample, when markets ore also a pmhlem in m,rny areas in the region. El'en when ou1-
closed by official decree. the)' are of1en reph1ced by unoffi- h:t:. for :.1orage and ;1ale ol' produces are pro,ided. either b~
cial markers that spring up in the vicinity. the su11e or b} the pril'ate sectc,r, siorage facilities and
le is arguable that tht.' inability of most :\frica1\ countries to back-up ad,ice and sen-ice may be le~ 1han optimal.; In
effective!~ comrol and 1race animal movement when it be- thb re~pcc1. drug, - hoth conl"entional commercial prod-
comes necessary to comrol ombreaks oi C<>ntngious dist.'ases ucu, and traditional remedie.~ -are ~ometime, more ea,ily
such as CBPP. African swine fe,·er (/\SH and F~IP. is 1he obtamable from unofficial traders than through ve1erinnry
major reason for their 11idespread occurrence in Africa. On or pharmacel.tkal agende~. Thi, meam d1at the authentk·
!he other hand. methods Lhat ha\'e been applied 111 Lhe del'el• lry and/Or stO'."Jge ol lhe products supplied ma} be dubious.
oped world or e1·en somhem Africa are not practical in most which can res uh in animal owners wa,ring their hard-
regions of Africa. TI'lis is particularly so in the \·a~t pa~1oralist uamcd mone)' on inappropria1e or inetTcc1ivc drugs. To
:u-eas of sub-Saharan Africa Unless a solution 10 this comm- some cx1em this problem is being countered by commu-
drum can he found, effective control. much less eradication nlty-ba!ted animal heahh workers (sec below) " 'ho ha1·e re-
or major trans-boundal'} diseases will remoin a problem. Per- ceived some hasic training in animal husband11 and heai1h
haps any real progress 101v-.irds add.res~ing the problem 11ill and \1ho are employed bv the animal 01\11ers in the commu-
necc,sitate fra11k acceptance of ics natu1e a~ a first srep. nities in which the) opNate. 3• 8 In many cowuries this ap-
proach i5 large!)· being adn)caced and finnnred by aid
Stamping our agencies, and it remains to be seen how 1iable the init!a1ive
In poor countries, and increasing!) in developed ones too. will be in the long run.
the concept of·s1ampingou1· (i.e. 1·omp11lsOI) slaughter and In the event of an ou1break of disease. ring vaccination
des1ruetion of animals i11 Lht! infected area) is accepted only mny be i ndicaced to prevent spread. The expense of vaccina-
with difficulty by livestock own11rs and the public at large. It cion like tba1 of comp,·nsation. mny pro,·e too great for 1he
is 1hereforc doubtful whether thls i,; a \'fable alwrnarive In gowrnment to be,1r. E1·1m when donor funding and Yaccine
most situations. h has been applied successfully in are m·ai!able. lack of infrastmcture. such as passable roods
Botswana 10 eradicate CBPP and also in Cc'Jte d'll'oire and in10 1he area. serl'iceable l'ehicles and the whei:ewithal to
Gham1 10 eradicate t\SF. In the laue1 1wo countries it is maintain teams in the field. may delay delive11· of vaccine
highly likely that the ,\SF problem \\ill recur because both and result in the cold chain being broken. Maimainmg suf-
have largely uncomrollcd borders and many countries in ficiem trained pcnsonnel in the area to administer the vac-
the region are infected. cine also has cost implications that may not be fea~ible for
The ottl)' war to ensure co-operacion bv owners. particu- the state to cover.
larly ii movement control is inadequate, l, to pay market- or
The complexllics and interaction some of the factor;,
related compensacion for animals slaughtered. When state outlined abol'e arc illustrated b)• the problem that ha, arisen
budgets do not penni1 such payment. :ir whore pa}ment is wirh respect to rinderpest eradication in eastern Alrlca.
retarded. 1he entire operation is doomed 10 failure. Con- Since the early 1~90~. 'mild rinderpest' hu,. becomepreva-
certed efforts to avoid the forces of government are llkclr to ient in some pans of 1ha1 region. pariicul.trl)' in the Somali
succeed. and may resulr in even more rapid spread of dis- eco-region 1hat enC"ompasses pans of Somnlia. Kenya and
ea,e 1.han would olherwi$e be the cn~e. \\'hen compulso~ Ethiopia The viru,o:s concerned have so t'ar all been mem-
slaughter does take place. strict monitoring or the dispo~al ber~ of 'lineage 2'. rn There i~ usually lit1le If any e\'idonce -
of carl'asscs is necessary. and the carca~ses ~hould be ren- otherthan residual amibody - for 1he presence ofthe infec-
dered unusable.; In poor countrie-., de~1rt1ction of large tion in cattle while wildlife. chieR} African buffalo {Sy11cert1$
amou.nts of edible mea1 i, a very difficult conc('pt, nnd exhu- C"njJer) ma} have 111orc Ob\'iou, di,;oa~o 11 The result i$ un-
mation of b uried carcasses is cC1mmo11. ccriainry as 10 how to go about comrolling the problem.
Limited ring l'accination I~ no1 advocated because it is diffi-
Vaccinarion and chemoprophylaxis cuh robe sure of the exact limits of spread ol inapparent dis-
Prevention of both serious epidemic diseases and erosi\·e ease. Nlass \'accinmion O\'er a \1ider area is also problematic
disea,e, of lives-tock ofwn depends on either v11ccination or because it becomes t>iqiensi\·e and difficult to persuade lil'e·
chemoprophylaxis, or boch. This presupposes access to 1he ~tock 01·,ners that such resources be dcvowd to a disease

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174 srn,c, o": hpecrs 1nn11cnring the occuncnce of iniectious dise:iscs

they do not recognize as serious. At present, there is r\o stood and applied hy th«.' live<tock owners themseh·es. since
scientific evidence that mild strains of rinderpest virus are thel' h:wc the rnost to gain from succes,;ftil control. For live-
able to revert co th!.' ,inilem form. although it is an ob,fous stock owners to play• either a leading or supporting role in
danger. disease control, Llwy nl.'ecl 10 be informed about the discai.es
and trained in methods of preventing their occurrence.
Surveillance and monitoring of diseases These m~1hod, 11My demand modificarion or even complete
:-laimenance of adequate ~urveillanct! In order 10 detect OC· change or their husbandry ~ys1em.
currence or an increase in the pre\'alence of economically In ~ub-Saharan ,\frica livestock disease control measure,
imporcam animal diseases at an early stage is vital for cffoc- are required for two complct<'ly dilferent levels of produc-
1ive disease control in any circums1a11ces. However. w re- tion. Commercial lh·es1ock farming. which is undenaken
tain an effective country-wide ~urveillancc system is entirely 10 generate profit. ranges from small· 10 large-scale.
expensive and lhe benefits are usuallr long-term and no1 aJ. and fncludes dairy farming, feedlots. and intensive produc-
ways appreciated by those who control the treasmy. Ap· tion <Jf pig, and poultry. These enterprises have increased in
proaches 10. and minimum standards for, surveillance are number during the la,1 decade. Explosive urban growth ha,
currently under consideration in many African coumries neceisitated higher levels of food production on less land.
and constitute a spedalii.ed field in it~ O\\~l right. There are which favours ,ommcrci;il produnion.
a number of texts that deal ,,ilh this subject that should B) far che greawr numbers of JiveMock are still produced
be consuhed for derailed infonna1ion (see Chapter 9: The in traditional systems. where animals range extensively and
control of Infectious diseases of livestock: Making appro- arc given liule or no ~upplemcma.ry feed. In 1he more arid
priate decisions in diffe.rent epidemiological and socio- pans of Africa, livestock keepers are often nomad ic.
economic conditions). 1" ,a.
When an outbreak has occurred. surveillance and moni- Disease control In commercial production systems Db·
toring bolh during and afterthe outbreak are essential to es- en;e co111rol b more easily applied in commercial. more
tablish che extem of che Olllbreak. decermine how effective intensive production syscems chan in extensive systems. It
concrol has been, and a~cenain it, origin, Ir b frequently is also cmcial. because i:oncentrations or confinement of
necessary thar resources in addirion to Lhose or the ,·e1eri- animals parricu larly ,vherc large numbers or animals arc
na11• service of che country concerned arc made available involved. fovour mpid sprc•ad of contagious pachogens.
during and immedia1ely after a serious outbreak oi an epi· Additionally, h igh morbidity and mortality it) commercial
demic disease. The major additional requirements are usu- s,·stems may prompt panic selling of animals through out·
ally trained personnel. tra11spon and equipment to mount lets lhat reach a wider market. and can contribu te material!)
an effective campaign. However. maintenance of surveil- to the spread of the epidemic.
lance activities beyond lhe outbreak LO ass_ist in preventing Commercial emerpri~es are more amenable 10 effecch·e
recurrence and 10 enable early de1ection is often extremely disease control/eradication for the follo1dng re.isons:
difficuh . The fact that donor resources are us~1ally a,'ll.ilable • ownen, are often more educated and Informed:
during outbreaks has t!ngendered a perception that trans- • owners are generally more willing to spend mone}
boundary disease concrol is an imernational rather than a on higher-producmg animals;
national responsibility. This has serious implications for • access to \'eterlnary sel'\ice:, ancl other resources
sustainability of control programmes. Is general!)• beua?r:
• quaranrinc is much easi.er when animals arc restricted
Alternative approaches to disease to a defined area; and
control/eradication • marketing of animal~ and p(oducts often takes
When considering approaches to disease control. it b im- place through formal outlet~ where some form of
portant not to lose! slgh1 or rhe reil5ons for control, facile as 1·e1erina11• inspection is available.
thi~ may sound. The<e may vary. buc with the e.~ception c,f a
few countries in 1he region. the most imponam reason for Additional advantages arc that the number of animals is
control is 10 enable production to conrinue at a local level usually known. or easy 10 count, and animals may be indi·
rather tlian to protect inrcmational trade. If Jlvescock dis- 1'idually ,denrlfled. Records of purchase~. sales, births and
eases can be adequate!)' controlled at local level. commer- deaths may also be kept. Owners ,,ill often seek veterinar:,
cial producrio1\ has a chance 10 reach a level at which formal assistance \\'hen animals are ill or unusual mortality ol-curs.
cxpon becomes a reality. Control measures should there- This encoma,ges rapid reporting of disease emergencies
fore be designed primarily to protect the interests of the live- Owner com pliance 1,'ith control measur~ 10 deal with dis-
stock owners and not to destroi· their enterprises to the ease emergencie-~ depends strongly upon con,indng them
e>.,em that they will be unable to resume them for a Jong chat che advamage~ of con troll eradication of the disease
time by, for example. wholesale stamping om. The most outweigh the shon·tenn advantage of e\'ading the control /
successful control measures will be those rha1 are under- eradication measures and selling as many infecied animals

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 SpL'Cial facto~ .nm,.~ting the con1rt,I of liv~tock di4-ct\Sl'<: in ~ub ...Saharan Africa 175

as possible. The key factors in successful disease comrol / doubt that considerable knowledge e.~ists among those who
eradication in the commercial secror are therefore: possess traditionally kept animals. However. the diagnosh
• organizaiion of the industry: or disease is often a problem. as is the a\-ailabillty of suitable
• awareness. information and train 111g: remedies.;· n 20 Community-based animal health care sys·
• a holtst ic approach to on-farm biosccurity to pre\·ent terns aw most ~uccessful when based upon existing know-
outbreak$; ledge and when there is an ob\10us gain for the
• ens~1ring that when outbreak control ls necessary. dam- communiry. 3 · ·1 Because community-based animal health
age rcsuhing from control me.u;ures is lim ited by assur- care lncrea$es awarcn6s of and inrormation about animal
ance of market-related compen~ation for compulsory dise(l..Ses. and concenu-ates upon prophylaxis. inc:ludlng
slaughter and/ or other measures that encourage 0\,1,er vaccination for serious diseases. it can make a vital contri-
compliance: and bution to impro\·ing disease comrol, particularly in tradi·
• participation of owners in disea~e control surveillance tional systems of animal husbandry in remote areas where
and monitoring. there are curremiy no od\er options. The contribution or
community-based animal health care programmes to con-
Disease control in traditional production systems Dis- rrol livestock d ise11ses depends largely upon the perceived
ease cont rol is challenging In traditional production systems importance lO the community of tbose disea~es. a~ well as
as these e nterprises do not have ready access to \•eterinary the approach adopted. Rinderpest control \'ia community
services.Although rhe animals themselves rcpresom weal ch. animal health workers achie\•ed considerable success in the
their owners frequently have little or no ready cash to spend Afar region of Ethiopia and in Sudan, because livestock
on animal health care. Because production in these systems owners had no doubts about the effect of the disease.3 On
is low. large numbers oianimals may be kept to offset higher the other hand, tsetse fi)'·Comrol programmes that were
mortality and lower reproduction rates. and the value of the rheoretically community based enjoyed \arying degrees of
herd is often measured in the number rather than the quai- success, which was at least partly ascribed 10 the fact that
l~· of the animals. If animals are sold. marketing is usuall)• the priorities of the technologists and the communities did
,ia informal outh.•ts. While cattle owners. in p11rticular. are not coincide.·1 Only when it was recognized that tsetse Oy-
often highly knowledgeable and skilled stockmen. their comroi provided tangible benefits were communities pre-
attitude to livestock diseases may be scoical rather than pared 10 commit time and resources to maintaining and,
pro-active. where n('Cessary. replacing tar.gets.
The principles of disease control are idemical whatever Information and awareness are essemial at all levels or
the production system. ·niechallenge in tradit ional S) srems li\'esrock production for disease control. The livesiock owner
lie5 in how to practically apply comro\ measures given the or herder is the first line of defence, as he/she is in daily oon·
lack of resources. information and. sometime!\, the very dif- tact \\ith the animals. 10 Two of the essential elements or dis-
ferem outlook of subsistence livestock owners. Ultimately. ease control. namely early warning and sun·eillance. depend
however. the aim would be 10 promote transformation from strongly upon disea5e recognition and reponing by tll,'llers or
rraditional to commercial systems of production. Given that members oft.he commu r1ity. In spite of the knO\\'l.edge that
1ivest0ck owners depend upon their animals to provide exists among traditional livestock keepers. most are eager to
them with the necessities of life. few or them w<l\tld refuse impro\·e on this knowledge and are highly receptive to lnfor·
the opportunity 10 improve the income or llfl> support gen - maCion. provided it l~ delivered in a form thar ther can ea~ily
erated by keeping livestock. Hisroricall)'· development has understand. The incorporation ofhigh-quallry photographic
depended upon the commerciali7.ation or agriC\llturc. and material showing signs 11nd lesions familiar to them greatly
there Is no doubt that. given the opportunity, sub-Saharan enhances acceptance and learning. Ideally. the information
Africa can achieve the necessary le\•el of disease control for should be transmiucd 10 0\\1\ers by the veterinary service., of
opiimal animal production. Being a latecomer to the devel- the country concerned. who can at the same time identify
oped "·orld. it may e,·en be possible for Afri<:a to avoid some lines of communication for reporting diseases that require
of the problems Lhat are inherent in exC'essive intensifica- government intervention. I lowever. where this is 11ot pos-
tion of animal production. sible, the function may be taken over by prh·ate or non·
Community-based animal health care, although no t a govemmenra! organizations, who may also 5erve as Lhe first
new concept. hos been developed co addre-~s the need for lin.k in the line of conununkmion co rhe veterinary authori-
ace!!$ LO veterin:iry services in n,rnl. frcquent.ly p3$torru. ties. In a disease cmergen<:), participation of livcstocl< O\\q1cn.
communitic~ usually far removed rrom the official ve1erl· in formulating and implementing control acthities is vital for
nary service of coumries.3 It has been format!)' applied in the success of the exercise.6
several African countries for \'arying lengths of rime and Globalization is increa~ingly becoming an impetus for
with rn111ng degrees or success.3 • 8· 12 Obviousl)'. owners change in man)'spheres and lh·cstock produC'tion is unlikely
have practised a degree of prima11· animal health care ever to prove an exception. even in the remotest parts of sub-
since humans have domesticated animab. and there is no Saharan Africa. Thus the present preoccupation of the

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176 scn1os o~t: Aspcc1,; inJ\uen~-Jni: the occurrence oi infectious di.;casc,

developed world with diseas~ such as fi\lD. bovine perceived by the owncri.. Given sufficient incentive through
spongifonn encephalopathr and food safety generallr will increasing market demand and the dwindling land a\'ailabl.e
force any country wishing ro expon 10 developed countries for migration, there is little reason to suppose 1hat Africa
or trade blocks 10 be able to certiiy their livcs1ock and live- would no1 respond. as other regions hal'C. by udop1ingmore
stock products accordingly. An example of the difficul1ie~ sophisticated husbandry sysiems.
that result has been pr.ovided by the events surrounding the Initially. changes in prod11clion systems are mo~c eusily
apparent spread of Rif1 Valley fe,er (RVF) virus from the effected \•,ith the shor1-cyde species. in particular pig-5 and
Horn of Africa to the Arabian Peninsula where. In 2000/ l, pouhry, Currently. large numbers of sca1·enging pigs anll
several hundred people died of R\ 'F in Yemen and 5audi chickens arc an iudi$pen!>aulc pan oflifE.' in villages in many
Arabia. As a consequence. a ban was placed on the tradi- African coun1ries. The sur.il'al of 1hese animals is cmclal ro
lional annual export or million,; of sheep and goar,; 10 1he 1heir owners. to whom they represent hou~ehold food secu·
Arabian Peninsula - panicularly at the time of the Haj - ri1y and. in 1he c,1se of pigs. a mobile bank 10 meet unex-
causing great hardship to the pa~1oralis1s of the Somali en>· pecied ex1)enses. Scavenging systiim~ require no input from
region who rely on li1is trade as their major source of in· the owners and ~erve additional useful purpo~ssuch as en·
come. The imernacionally accepted guidelines on 1he re• vironmental cleaning, but production is generally \'ery low
quiremems for ensuring freedom from RVF contained in 1he and mortality unacceptably high from cau~es ut1rela1ed to
OtE s lntema1ionaJ Animal I lc.>allh Code:?: require updating. disease. such as road nc:c1dcnrs, prcct:11ion, malnuirtnon.
particularly from the African perspective. ,\ddh.ionall)', so and eraclkation because of damaged crops. The productive
far African countries have 1101 managed to bring their tolk--c- potential of pig, is such that relatively small inputs can result
tive resources to bear in addres~ing such problems. II is in sizi:abli' gains. Convincing owners to change from ex1en·
therefore vital chat the African perspective is convc)'ed more sive to more intensh e produc1ion S}~tcm, should therefore
successfully into irnemational fonims such as 1he OIE and be rela1ivel~ easy. panicularll• if local 'champions' can be
the \Vorld Trade Organi7.ation. \,·here 1rnr.lc-relatcd decl· iden1lfied who. l)y precepr and example, can convince other
sions are made. to ensure dun dew!oping countries arc not owner, of the proticabilil\ of improved emerprises.
'yic1imized'. albeit unimemionally l'hc inherent resistance of indigenou~ breeds of dom('S·
Change from traditional ro commercial Uves1ock pro- ttc llves1ock rlisea,es endemic 10 1he region should not be
duc1ion systems seems llllrcahM,c, especially when one under-cs1ima1cd as a means for disease control.12 1' Iuch re-
considers 1he nomadic liv~to1:k systems practised \\idely in search is still n<>eded in 1his field. ltowevcr. upgrading pro-
arid pans of Africa. :-JeverLheless, 1here is no doubt thar In duction ~vs1cms by the wholesale introduction of animals of
some areas change has taken place. at least with respect 10 exotic breeds that are not ~ui1cd to African produ<;tion S)'S·
the marketing of catde. E,,q,anding cities ha,·e increased the terns and arl' highly suscllptible 10 the diseases 1hm are en-
demand for meat by people who cannot keep livestock demic in the, region. is a recipe for disaster. Such animals
themselves. Large numbers of catde, 1;heep and goats from can usually only be successful in artificial sys1ems when!
the Sahel find their way tc, coastal markeLS such as Abidjan, complete prott:Ction from vecrors and diseases, and an ap·
Cotonou and other coastal cities, indicating that their 0\\11· propriate level or nutrition and management can be
ers are 1villing to undertake commercial enterpri$e5. Condi· achieved. Rc1aining and improving the gen('tic resource$
lions under which Livestock are kepi and marketed arc 1har cxis1 in 1he region i5 ,ital f<Jr impr01ing production and
dictated largelv b,· fe"ll,;ibilit\.' and b)' the adYanmges comrol o' IJves1ock diseases in sub-Saharan Africa.

References
t li. • ll.\MROJi:<.\\ r, M \, Iii 8\",1JIRUJ>Dt:\-,
A'IA!':l'U, \\ , \tAUIPll, l- .\ ., i\t>O,f, for PQJ•,•rl)' r.:d1,r,{(,1t mul .qutalnabf,• rurnl lit 1.•IU100</s for Jfrc:;tocJ.
1

J.8., 199a. An outbreak of co11mg1uus bo\lne pkurupncumom.i m .fiumc•r.t Rorm.•: r•uod und A~i1cuiture Ors;,;)J'Ul.ation of the Unllcd
Ngamilcnd dhtr1ct 01 nonh•\,·cs.tt.rn Houwa.n{1. n1t• Ve-rtrtnm, Flt•tor((. '"tion':i..
113.46-18. 5 c..rrrusc.. \\' \., Jll'-'Kll1t. , , .• ,4 6 '.'YAlv\llU\1A, n., .JOCJI, .\1tmut,Ifur
2 .\."\OS. :t00:.1m. . .mnr:01111/ AJ,mut1 /lrolrh (Qdt. runs· om,c F.rtuller,:fo,, nfJ.1tnr11(l l)J>,.Y1~ l,;· Sra,,,,,1,,g !.)111. Homt!"" f-ood and
~ ruu.:nMliurul d-..F\ l:JJLlUOliP At,;'rii:uhuur Ori:;mi.T.,uf:4a or 1Jw Unitcd ~;nit>1h

J r. ,Tu:v. "· • 1.£\'t.A,-n. T., ~001 <.:ommunny pocnfip~nion and she &Jeh\<tf)' 6 _\Jct1Ulll (Jll lftt.• l'r,·pt1ra11'on
GU:Jl1'"C,,. \\'.,\,, ilt)t-01·" l",L.,. & 01&1, 1.U :!UUI.

oh·eterimuy ~~1-v;c1.-s m.Africa.. Ph'l't'n:ur t ,,1~mmry· '\lr,lu·m-.·, .t9, uf .Ymumul ,lmnwl.Db,'O.k·Euu,ttt,mcy· PNplJr,dnm Plmu. Ru,mr Food
95-113. •ndA~riruhu"' Ori..inlr.ttlor> lli 1h1• llnh1-d 'l;;tt/on•

..; C'il'UC1,,;, \\.,\, (COORpj:\.\mAJ, o~tro..~tfl. (i. 00!',;.\Ll)'-O~. ,\.I .. J.\\ff",, ,,••
f drug3,urpt~'t0~-u~1iU'ncc .1nd tm.c:ginR
<,;Otkt,<~. 11 .. :1001 \ 'ett>rtnat)

)lCDLH.MOn, I, U."l.1..\.~U. T. \JOftfll:-, 11 'i,, Pl W\H:-;, .\ , M\\r lf \l \ ,,u, ,1,,1 .. fann1n1-: communltU.•, 1n lhi• )1.tdik,,...,~ l>i~1rh.:t. ~orih Wt~ P'to,1nrt•.
;;,outh .\!Jiu.. ~1mud\'r1 'flll?'iii. Unhte~it,· 1\f P1ct0:"ia.
W..\RP, o ""'\',1'""1 "- zoo1 FQs,--.rsng 1h1• r»IIQ'tlinl<>gut' m t.tlf'fKIT( uf
tm1,rored cminwl ltt•nlth <u11J l'ltf4'fltUJ.t) pl1iJl1r-lwal1h "' m: L"111'' Vhint 8 111/TT"\:f ,. ,:,.1.ZlOL ..... ltM" f a.u & ttf'l-JFfl ,,. o.u •• :mon. Cb;iro1c1c:-t.\llC'S

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 Special factors affocdng the control of !h'Cstock di~ases in sub-Saharan .\frlca !Ti

md P<:fonnnnce of ,illagt animal htnlth worker< and V<!tcrinal\ Africa. Jdttrtto1 u/ 1llcSomll Afritan \ CUJtinnryAs:11,citJiitm, 68. 43'-aJ.
~~Um.ts in ttOrthCJD ~!al:w.1. Joumt:tl of tho South .,Vtietm \'t•Urinnr,•
11 ,ton••~ •· >., IS99, Th• ~pplie~11on or economi<:>m nrum.\l h~alth
,\$socw1fon. 71 160-165.
pcogrammev,,J. prac1kal guicle. s,wmf/k flUd Tt:ehnitnl Rt"1i·i1Juo: Office
9 JOS&S. lf,,\,. QUMEM, ... t.S\1..o\.."iO, T,1.~ MOGC.,\. w & srL.,.t, £ •• 1998, lm,mn1101111I d,~ iipfworin. 18, 30.,.314.
Cammunity,bas,'<I animal ht!•ltll ..,n,ces in so1,t1u,m Sudan: tM
15 .>IUO..RA);Ot. 2001. No, thc,foo, .a.nd mouth ropon: E,·cr)1.rung Tony Blair
experience and the future. J,1; ,tuKAR.,t:11w., .:,.. "Ol'IW<)to. M ,I, IC<b ..
dido t wam ,..,u 10 lr;now about the blll\l<lSI blund« of his prem1Cl'$hlp
PtUf.Ndm,;.~ofzlte IX lltl.1:t,mtlurml 0.mft:tttm~ "J[.~'"',_.imUJu Cl[
Pmnr~ 1:_,,,. :\o,·embor2001 pp 3-30,
lmticuno11to{11opic,1f \ 'Men1101y Medlcirr,. J4-l8S.:p1<•111v<'r 199&
Hllmrt', 7.111'bal,u,.., I. 10;-133, :({ ft05,)"1'JJ\, P-R., 1994. Rlnd1.1:pbl. /n: COl'TJ'..! R. I.A.\', .. fflO!otSO~, C. R. <&·
ru>'T1'1, R.c.. rod,, ln[<o,:1(011; Dt;,,,,,,., ufUw<tQ(k wir/1 ~penal Rf/ertnce
10 ,;nun, ,.,.., 1996. The etree1 oflnfrnmucrn,es en ,ul\ el113nce ar.d
1i,S011rlr,m .-l[rh."ll. (;ape Town· Oxlord Un,vers!ly Pr•&, Sout11.,m Afric•,
momtortng.b}'Sttm.,. Compulit1uil'l• Rt•por!S ,,,, Tt'1mi¢t1I lt,•nL~
prv~ncNI l<' thr lrm:ma,ional Commiul!t or 10R,tgionol ('..qmmf.num,. 17 ~H0'1~' G.R. 19,9, Alternati\'t> for <C)llll()Ulns .tnllll.ll distllM<>
Office lnrtrt1111lonal dn Epr:.oorl.s. /9,o, H3-li3 ;or 236. If Illa Spanah re.ulrlng from int.,ructfon h•n,c,n 11,'t'>tock ~nd \\ildttlc In >ou:he:n
and French \'l!rstons ar,, included I. .\fnc.,. Sot1rll ,tfri,1111 Joum11I o/Sc1tnet•. !15. 71-76
II It.A., 200L0AU/1B.-\R. P.O. Box 30786. :\alrobt. !(enra. Pcl'$Onol
~Q(~. 1$ TltUH>HUD. "" 1995- Vti,,(UIIJI)' F.11id,•111la/~·. 2nd ,'lin O>iord, u~
communtcation. Blocl:w•ll Scf<ntiRc.
u MARG~. w,r,., 19118. Strategy !or llve.t~k d<~'C!Gpm~nt in Mrica. /11· 19 T0)1A, H , uu,oun.. a,, <Et.;::'\'.,\, M .. ats-n, "·· )tt>UTO!\, "·· U)U/'.i\, A,,. UUi,
MUl:AAATiltWA. s. 4'011\\'0LO, ,..,. (ed•).
Proe,..,dinl!J ofthr IX ,,.. 11199.ApplJ.."'1 Vc1vri1111,;· Epid,•111U1/ogy. AP.EMA'. \lal,on•Alfor,
/11t,r,1nrio11nl Conf•rMerofrlss«/111/i,11 of lnst11utfo1u o[Tropica/ ~ \'A:-. 1,Ut MUt\'.'f o~, ~w"x~ c.E • non ti\, C.J,, 2001, Use or et.h.nowttrina.rY
V~t('FfriaQ·,\1t."dic/11c. ,..,_1S Srp1tm1Hr J9!Jll llamrr, Zlmbalm'f', 1. .il-58, rntdicinnl pl:im, In catde by ~t"\\'an:t·)l)eakinJt people in the .Mt\dilrr.\\'E
13 !i,t-\SIIU. P.J .. S0XAX0t. A.'- VA.~ A\'ERBelo.~. w.• 19i97. Tick ,:onuot by ~rea ol lhe Xatth West Provln~ of South Aftl.ca.Jo11mnl of1heSo111h
smrul·scale C;'lrtle ra,mel'$ In the eeotrnl E3St~n Cl!J><' 11,0,•inei!. South ,\frlaJn \',•t,•rirrn,;• AJJocfar/011. 72. !89-196.

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 9
The control of infectious diseases
of livestock: Making appropriate
decisions in different epidemiological
and socioeconomic conditions

13 D PE RRY, JJ :VICDER M OTT AND T F RANDOLP H

Introduction circumstances that would affect the nature of a decision

made today. These Include changed political and socioeco-
This book describes the m any in feclious diseases Lbat affect nomic circumstances. different discase-comrol instlrutions
livestock in sub-Saharan Africa. and indeed in marl) other and authorities. and difl'erenc decision criteria for allocating
regions of 1l1e world. and discusse~ in deta il the technical as- public and private funds and other resources to infectious
pects of their diagnosis, treatment and control/eradication. disease control. to name but a few.
In a n environment with unl imited resources. and with the Vv"hy wou ld we today expect the answer to be 'probably
technical capacity to undertake tbe appropriace measures. not'? Much has changed in the intervening years since the
many of the diseases descri bed here could be brought u ndet' initiarion of ECF control in southern Africa in 1904. l11e
control ore,•en eradicated. rndeed. in the pasc, a few of them magnitude of the threat presented by even the most virnlen1
have been eradicated, but usually at enormous cost. Re- of i11Iectious diseases is different. and is probably much
markably, in many cases, these costs are not available in the smaller now due to our increased understanding o.f the risks
records, indicating lhaL they ma) not have been of prime and impacts of different infections, and a greater invemory
importance in the decisi<ln to eradicate. Take for example of measures at our disposal 10 control chem. le is doubtful
the eradication of East Coast fever (ECP} from southern Af. that !:CF would threaten the very existence of the entire ag-
rica. In his derailed acr.oun1 of 1he Pradk~11ion prngrnmme ricuhural sec1orrotlap1i. i i clid :11 rhP heginningof the pre,i-
that spam1ed the years l 901 to l960. l..awrence1H estimated, ous century fot sectler agriculcural developmeo1. Ar the rime.
from what he called 'fragmentary daca·. that a total or 1.4 the region wa$ recovering from the devastation caused by
million cattle died as a resulc of the disease, and an addi - the rinderpesLpandemic, and in an era when the world was
tional 100 000 were slaughtered However. he noted thac preoccupied by the perils of infectious di$eases, Somh Af-
daia on the costs of control were ·non-existent'. One cannot rica was also struggling to recover from rhe ravages of t he
help asking the que..~tion: would such an intell$ive control Anglo- Boer W11r (1899 10 1902).
and eradication programme, involving compulsory weekly Funhem1ore. it is difficult 10 imllgine the determined ap-
dipping of all cattle. s tr ict m ovement contro ls. quarantine or plication of a technical solution being successfully imple-
atrecced herds. compulsory examination of spleen smears mented today given the political and organizational realiries
from all animals that were slaughtered or died, and slaugh- of much of the developing world Part of this doubt is related
i,er of in!c,,cted animals '>ith comperumtion, all sustuined to the more complitate<I nawre of decisions on resource allo-
over decades. be carried our if ECF \\'ere to breakout in cer- cation and how (and bv whom) these decisions are made
tain sou thern African countries today1 coday. IL appears that an exrraordinary degree of authoricy
LI is in the answering of this question that one confroncs was gh en 10 the highly successful group of scientists acti\'e in
the stark interaction between che technical feasibility and investigaring the causes and concrol of ECF, and this out-
the socioeconomic feasibility of infectious disease conuol/ weighed many of the bigger economic questions. There ha,·e
eradication. There are several importan t differences in been subscamial changes in the seat of authority for disease
178

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 The conuol or infectious diseases or lll·c,.ock I i9

comrol in vim.ially all comers of the \VOrld, from a very cen- which was often in geographically disrinct areas known by
trali7.ed, powerful and autocratic government involvement 10 names such as 'tribal trust lands' or "African reseJ"\·es·. which
a decentralized and ofien impoverished public sector. and have evolved into the so-called 'craditional', 'communal' or
this change has accelerated in most countrie., of the African 'smallholder sectors'. depending on the country and the no-
cominem in the last few years. The change has been compli· menclarure u~ed. These low-producing. often subsistence·
cated by a deterioration in the veterinary lnli-astructures of oriemated systems. relied ,·cry much on the s1a1e for tl1e
many African co1mrrles. and thus the feasibiiily of delivering provision of\"etcrinary sef\ices. which ln general were pro·
even highly technically efficacious control measures Is in- vided a1 no cost 10 farmers. This was done for many reasons,
creasingly questionable. In addition, several decision· both political and technical. A principal concern, 1101 (l!Wa}·s
support tools (such as cost-benefit analysis. for ex.ample) 10 stated publicly, was the belief that infectious diseases could
aid in decision-making are now available and widely used. spread from 1hc peasant sector into 1he more commercially
l;iut did not exist in the early part of the twentieth century. oricmated livestock enterprises. but the responsibility of
In summary, decisions regarding di~ease control have governments for the welfare of the poorer sectors of socict)
changed radically due to a combination of improved know· was clearly also an imponant driving force (see Table 9. I J.
ledge about infectious diseases and of the need 10 take imo Both of these systems have evolved substantially, fom1ing a
accoum a whole range of socioeconomic considera1ions much 1vider set or production systems in which socio-
that were not previously considered. economics play a critical role in the choice and efficacy of ln·
This chapter evaluates how dedsion-making for the con· te1Yentions to control infectious diseases, and as to who pays,
u-ol of infectious diseases can be improved at all le\'Cls (e.g. and who benefits. Overall, the commercial sectors have shrunk
farm. community. country. region) by the consideration of in size considerably, although less so in sou them Africa than in
both epidemiological and socioeconomic issues. We re\~ew other pans of the continem. ln most cases, state si1pport in the
what rhe impacts of diseases and thelr control are, how form of loans and subsidized veterinaryseT\~ces has v2nished .
these can be quamlfled. how such information can be used In manr cases their effie!encyof production has improved. de-
in the choice of disease control intervenrions. and ho\\' such spite I.he economic and political pressures on them.
inten·entions can be best implemented under different sets The changes in the· peasant" sec1or have been much more
of circums1.ances. profound, and it is here that socioeconomic issues usually
outweigh technical issues in 1he op1imol control ofinfect ious
diseases. Firstly. there has been recognition of1he un.fque na-
Livestock production and use in Africa
iure or cenain !ivcscock-centred systems in Africa cb!lJ'acter-
An important perspective on the socioeconomic i:,sues af. ized by pastoralism. These include tho~e of the :Vlaasai.
feeling infectious disease comrol can be gained through an Rendile. Sarnburu, Horan and ocher:; in eastemAfrica. the Tu-
undersranding of the concept of different production sys- areg and Fulani of West Africa, and the Herero of Namibia.
tems in wh ich diseases have different impacts. These im- ,tmong 01hers. ·n1ese groups are characterized by their ex-
pacts vary in terms of che production losses and control treme scarcil) of financial resource.~, but by a high priori tyro
coses incurred. and whether the control costs are borne by disease control for their ·walking bank accounts·. 1bey are
prh·ate or publk coffers. During the colonial e ra. and for a highly demanding of veterinary interventions. in 1ern1s of the
variable period afcer depend ing on the country and region. 'priority" of the dlsease it addresses and the appropriacencss
there were possibly two major production srstems (al- or the solution to theirciroum~l3nces, bu1 if these arc correct.
though such 1erminology was 1101 applied to these) in much the}' are higher adopters of reclmologies to lmpro\·e animal
of Africa. The first of lhcse was the ·commercial' sector, health than other smallholder groups In ,\liica.
comprising different scales of beef. dairy. sheep and poultry Secondly has been the evolution of market•oriemated
production on medium-sized 10 large farms with fairly high smallholder systems chat are commercial. often intensive in
levels of inputs Cin terms of feed. veterinar)' drugs and vac- nature, but very small·~cale. These include the smallholder
cines. and genetically improved breeds), in which many of dairy system of the Kenran central highlands. the ·emer-
the financial aspects of disease control were the responsibil- gent ' small-bcale commercial farmers of South t\frita and
ity of the individual farmer, as currently occurs in Europe. Zambia, che 'small-scale commercial" sector of Zimbabwe.
for example. There was some fi nancial support provided by and the peri-urban dairies ofAddis,\baba, Cairo. and Dakar.
the state to farmers. such as loans for the construction of Some of these have achieved significant success in 1erm~ of
dips, access to vaccines and remedies at governmem-subsi - their producti\~tr. In 1he Kenran context. the smallholder
di7.ed prices. and public veterinary sef\ices that were gener- dairy sector now supplies somewhere in 1he region of90 per
ally s1ate•financed and -run. and delivered at no or a highly cem of 1he milk consumed in the country, 1 15 a dramack
subsidi7.ed cost to farmers. Socioeconomic issues in this change from the dominarion of large commercial dairies
sector very m uch boiled down to the cost efficacy of differ- some 30 years ago. In these sectors there has been a chang•
ent approaches to successful infectious disease control. ing panem oi disea,e priorities. and a tendency for risk
The second production sysrem was I.he peasant sector. avoidance management practices to develop. Again u~ing

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t80 sccno, o~ ,\spects inOuencing the occurrence offnrecllous diseas~

the example of the smallholder dairy system of Kenya. the secror \'e,erinary ser.ices, and the fragile socioeconomic role
use of stall-feeding (known locally as zero-grazing) has dra- of cattle in the livescock component of the agriculmral sector.
matically reduced the risk of rick-borne diseases. with most These are very broad production system groupings. At-
of the fodder being cut and carried to the confined animals. 1empts have been made to develop more structured classifi-
chus avoiding heavy tlck infestation derived from commu- cations of livestock production that further disaggregate
nal grazing. This bas brought greater anenrion ln this pro- and that can be used for anal~'lical purposes and for priority
duction system to the so-called infectious diseases of setting. In 1996, Ser~ and Steinfeld:!23 produced the first glo-
lmensification, such as mastitis. bal U\"estock production classi.tication syscern associated
The other major peasam production system. \\~dely \,;th a detailed data set. For this reason. it has been widely
prevalenc in southern Mica, is not orientated 10 the lil'es1ock used, and served for example as the basis for the lmema-
product market. but ro the crop market This sector. charac- tionaJ Livestock Research lnstituce's (lLRI) recent priority-
teristic ofthe 'traditional sector' in Zambia and the ·commu- setting exercise. 203• 238 Sere and S1einfeld's classification is
nal lands' of Zimbabwe, is primarilr a staple crop (mainly based on the main food source for livesroc.1<, and on the
maize) system in which livestock play many roles. The use of agro-ecology oft he region. They used the FoodandAgricuJ-
oxen for tilling the land ls probably one of the most important ture Organization's (FAO) agro-ecologlcal zoning classifica-
functions of cattle in this system. followed by the social {or tion and produced detailed country tables \,ith
networking) value of animals. The sale of canle into the beef disaggregated data by area, human population. livestock
sector usually comes in t hircl place, and formal off- takes from numbers, and livestock outputs for each livestock produc-
this sector have always been considered remarkably low (in tion sysrem category. Thornton and co-worker.s2l; have
the region of3 to 5 per cem: see for example Perry and associ- since refined and updated the Sere and Steinfeld classifica-
ates).18~ As a result. 1his system has suffered considerably tion system and database. and plotted a global discrlbution
from infectious diseases with the dramatic decline in public of the di[erem producrfon systems.

Table 9.1 Exampl6 of general livestock prooucuon system tategor,es curren1J1 in u,e in southern Afr•ca and 1~eir anima popula:,on numbers

Tne general groupings used for reporting of livestock popula1,on numbers ,n many soijtltern Afncan countnes still fall in the broao catego11es of
11adit,onal (communal. ssclor familiar. or Swazi National Lands) and commerc,al. ln these, 11111 'vadiuo.~al' setters dominate in tarms of livestocic
I\Umbets. except in South Africa

COUNTRY CATTlE (GENERM.J DAIRY GOATS SHEEP

TRADITIONAl COMMER"...t.At TRAD. SMAll·SCAI.E COMM ! TRt..0 COMM TRAD COMkl.

Angola 291 213 59546 0 0 0 1685600° 34400
Boiswana 1562 200 258500 0 0 0 1soe 400 28300 233300 16800
Malawi 579 700 23500 li 600 0 4000 1583400 14 100 98000 t 800
Mozambique 291 213 59546 0 0 1400 SIS 525· i9 874'
South Africa 4 090 791 6153720 0 0 1 433 .!65 4 485953 1433547 3162 06S 23 188 ~S9
Swarlland 491428 142 367 3997 0 t 187 428415' 30887'
Tanzania i5374 il9 58351 0 187 687 24645 10682 434 12343 3J8A 95i ao:1
Zambia 1 700000 97000D2 0 20000 SOooo2 s.:o 25i 71 8()61 27 038 50212~

Note. 1 Sheep and gcats reoorted togettler

2 Also includes 'semi-commercial secto(
Sources References :.-·s. 28

In Z1mbal>•t.1e the livestock populat1ons of 1~ smalll\01cer secioi are 1eported ,n three

categories, =munal, resenlement and small-scale tt1mmerdal
SPECIES LIVESiOCK StCTOR
LAA GE-SCALE COMMUNAL f\:SITTLEMENi SMAll·SCAlE
COMMERCIAL COMMERCIAL
Cante 1679652 3 6896'\0 468056 231 416
Goats 77667 2 582463 188 263 61477
Sheep 136565 45577~ 282~7 19 588
Pigs i3S 216 123005 8826 11674
Donkeys 1'-!613 323550 2sns 8378
Horses 90'7 67 22 1t

Source. Oeoartment of Veterinary SeMces, Zlmaaowe. 200!

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 The control of infectious dls~ases of li\'es1ock 181

In this pro_ducrion syslem classification, four main pro- • l.GH - Livestock only. rangeland-based,
duction categories are identified: landless systems (typically humid/sub-humid
found in peri-urbansenings). livesrock/rangeland-based sys- • LGT - Livesrock only. rangeland-based.
tems (areas wilh minimal cropping, often corresponding to highland/temperate
pastoral systems). mixed rninfed systems (mosll~· rainfed • MIA - ;11lixed, irrigated. arid/semi-arid
cropping combined with lh·escock. i.e. agro-pas1ora! sys- • Mil I - :Vlixed. irriga ted, humid/sub-humid
tems), and mixed irrigated systems (significant proponion of • MIT - ;11lixed, irriga1ed. hJghland/ 1empera1e
cropping in which irrigation is used and Is interspersed \,ilh • i\lRA- .Mlxed, ralnfod, arid/semi-arid
livescock). All but the landless systems were further disaggre- • MRH -),li.xed. rninfed. hu.mid /sub-humid
gated by agro-ecological potential. as defined by the length of • MRT - l\llxed, rainfod. highland/temperate
growing period. Three different agro-ecological zones are • LL - Landless (per!-urban)
used: highland/ lemperate. arid /semi-arid and humid/sub-
humid. In summary. the following IO li\'eslock systems ha,·e The geographical <;overages for each livesrock produccion
been defined and mapped across the globe: system ha"e been mappe~ 7 and rhe dis1dbution of sys-
• LGA- Livestock only, rangeland-based. arid/semi-arid 1ems for Africa are shown in figure 9. 1.

..

Systems 1n 2000
l!Vl:stc>:<. range. artd;se1111-<mc
lw.es,cc,· ran~e. humiCl/sub-hum,d
l-ves,oc< range, highlafld11emperate
Mix.a. ·ainfeo. arid/semi-arid
D Mixed, reinfed. humid/sulrh~mi:!
m Mixed. raittll!ll. h,yhlan~/le:noeia1~
- U!!Jan
O Otner
LJ NoOaia

Figure 9.1 The dis:ticution ot livestock product on svs1ems mAfrica (from Thomtoo et al.•20011.31)

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182 ,,er.ox o, · Aspects influencing the occurrenct! ofinfectious di:.enses

In each of these sy~tems, disease-comrol priorities arc Livestock are panicularly im portant to the poor. It has been
differem. and the efficacy and viability of imervenrlons also estimated that livestock form a component oft he livelihoods of
differ. Optimal success will be achieved by serious c;-onsider- iO per cent or t11e world's poor. 12-1 1\11imal diseases are an ev-
aticm of the socioeconom ic Issues Infl uencing. indeed ery-day occurrence to these people, as animals of the poor are
favouring. optimai delivery, adoption and impact in the parlicularly vulnerable co disease (due to many reasons. u1-
perlicular target production system. cluding ki~k of knowledge about their management :llld COil·
trol. and lack of access to- and resources for- animal health
and produccion inputs and services). Funhem10re. poor fam1-
The role of livestock in African societies
er~ usually have few animals, so the loss of an individual ani-
u,·estock pla~ many dlftering roles in agricuh ure and the live- mal has proponionally greater significance. The~· also have fe\,
lihoods of vircuall~ all sectors of society in Africa (see Table reserves on which co sun~ve during lean rimes. liveswck being
9.2). They are important sources. of animal protein in the one of I.hem. which t11e~ can use tor rccoiery.
form of meat. milk and eggs, their hides and skins comribute u, estock have not always been the flavour of the mo·
to clothing people. the fat from chelJ' bones contributes to the mem. Not so many years ago. much publicity was given in the
manuf-<1ctu!e of soap. their work contribu tes to ploughing media of the developed world to the woe~ of consuming
maize fields and moving product<. to and from markets. and meat, and the negative em~ronmental and animal welfare
their manure fertilizes the soil used for crops. They also play Impacts of keeping large numbers of pigs and pouhry under
an important role in social networking in man~•societies. and intensive management. While these view~ might have been
as a ·walking bank accoum ·. Livestock and their products rele\'aru to the conditions in many dc,·elopcd nations. they
comribme co the incomes. health and livelihoods oflivesmck totally misrepresented the conditions and demands of the
keepers, traders in ltvestockand their products. labourers on poor in the developing world. In 1he conswuer societies of the
farms and in abattoirs. and of COU!se to the great majority or developed world, per capiln consumption of meat and m.ai1~
the human population who consume livestock. products in other livestock products is predicted 10 decline.49 ln these
their diet. The cxpon of livestock and their produets gener- high-income societies. a wide variety of readily a,'8.ilable food
ates valuable foreign exchange. so contributing to national products. low levels of malnutrition. and Increasing levels of
economies. The heal ch of li\'esmck thus has a crucial inOu- cardiovascular disease. diaberes and ocher complications of
cncc on the healt11 and well-being of Africa's peoples. Ol'erindulgence have led to a negative ,1ew of animal pro-

Table 9.2 Oomes11c l1vas1ock species kept in Africa and the,r contribu1ion to national ecoromics and household assets (from Perry ill al. 2002' 901

SPED.ES CONTRIBUTION TO HOUSEHOLD ASSETS

flNANCIAL SOCIAL PHYSICAL NATURAL HUMAN

Cante Saies of milk. meat. Netwwklng mec)lanism Draught pO\\eI for ccop Manure f01 maintaining Household con$umption
lli<!es. animals. draught Social status nditator culriv.mon soil fertility of milk. meat
power services. D1aught powe1 for
transpon uanspon
Savings 1nstrumem
Camels Sates of milk. meat. NalWorking mechanism Oraugin po~~111 for Houssholj consumi,t,on
hides. animals. Social sta!IJS ino1ca1or ttansport cl milk. ireat
uanspon services
Sa,ings ms11uir.an1
Sa'es of animals. Draught po,\l!r f0t trop Manwe lor maintaining 0 1ov1sior. of housenold
draught seMces, cultrvat,on so!I feniht~ ·.~ater supplie~
tr.~spon lesp. waierl OralJl,lhl jl()we~ for
transport te,1) .·merl
Goaisanc Sa:es of milk. meat. Natwork,ng mec0$nism Manure for maintaining Househol;l consllm!)'t1cn
sl'.eeo ~,oes. aoimals Social status ,f\il1ca1or soil rend,tV (Ji md,c. rreat
Savings ,nsirument
Pigs Sa,es of meat. animals Manure IOI maintaining fio~sehOI~ tonsurnB!ion
Savings msuument SOIi feniltty o: meat
Sales oi eggs. meai. Networking mechanism Manure for maintaining Housanold consum,nion
i:r.\1 soil fertility of eggs Meat

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 The control o( infecrious diseases of livesmck 183

tein. These have been coupled with concern over the envi- humans. and epidemiology. among other criteria. ln terms
ronmental th reatS posed by the need 10 dispose of large of their impact, a nu mber of facrors are important in deter-
amounts of animal effiuem from large-scale and imensive mining the scale of impact. These a re:
animal production units. • the number of species affected:
Ironically. these t\\'O factors are reversed in much of the • tl1e proportion of animals affected;
developing world. Malnutrition is rife in many regions, and • the~ize of the geographic.al area affected orunderthreat:
livestock p roducts, panicula rlr meat and milk, pro~ide ru1 • the ability of the disease to spread:
important means to overcome this by providing protein. • the losses associated with the disease in both animals
micronut ri ems and vitamins. The demand for meat and and humans; and
milk is pred icted to grow by 2,8 and 3.3 per cem, respec- • the availability, affordability and .efficacy of measures
tively. per year. dwarfing the grovnh rares of 0,(, and 0,2 per 10 control it.
cem prndicted in the developed world.~9 Furthermore.
livestock are a powerful mea ns of enhancing the purt"has- We J)ro1>ose a classification of infectious diseases based on
lng power of the poor through the sale of livesmck prod- the main characteristic:; of their transmission and on their
ucts. generating income that can be used for tile purchase differem impacts. This is presented in Table 9.4. This classi·
of food. education and health care. When it comes to the fication contains the follo1,ing categories:
ennronment, livestock are a most valuable asset. pr0\1d· 1 J:pWeml<' diseases. main!~• corresponding 10 the list A
ing the essential fertilizer for crop production, often un- diseases of the World Organization for Animal Health
available or unaffordable in any Olher form. or Office International des Epizooties (01£). These
The realization of the starkly differem roles of livestock diseases are considered 10 be an international threat
in the developed and developing worlds, and the predic- and are usually directly transmined micro-parasites.
tions as to how these differences are likely 10 continue and for \\'hlch the risks of epidemics accompanied by high
intensify, has reversed the negative attitudes to supporting production losses are the main impacts.
agricultural development that involve livestock. 2 Diseases from the OiE list B that are directly rransmi.t-
On a global basis. precise estimates of the numbers of ted. fn general these occur sporadically or endem ically.
poor llvi:'stock keepers, traders, labourers and consumers by They ha\'e important impacts on livestock production.
region and system, and the types of livestock tl\at contribute and on lost producr!on opportun ities.
10 their livelihoods. have not been rnade. Thus, it is necessary 3 Diseases from the OIE list B that ace indirectly trans mil·
10 rely on crude aggregate estimates from summary tcd (mainly vector-borne}. In general these occur spo-
repons.111ese do provide a picture of rhe relative imponance radically or endemically. They also have important
of livestock to poor people. The Livestock in Development impacts on livestock production and lost production
{LID) 123 has developed global estimates oi numbers of poor opportunities.
livestock keepers: these are presented in Table 9.3. -1- Farm-level endemic diseases- termed ·other diseases'
in the OlE classification. Their main impacts are pro,
duccion effoccs on the farm. but these have an aggregate
The impact of infectious diseases
influence on national productivity.
and of their control
5 Zoonotic diseases or infecrlons in llvescock in which a
Disease classification based on disease impact, majorimpact is che risk of human diseases, this element
transmission and control of risk being important in control decisions (e.g. brucel-
Diseases can be classified in a varietyoi ways. based on their lo&is, tuberculosis. Tryµm1osomt1 rhodesiense sleeping
aetiology. mode of transmission. impacts on animals and sickness).

Table 9.3 Numbers(in AGRO-ECOLOGICALZONE CATEGORYOFPOOR UVESTOCK·Kffl'ERS

millions) of poor llvesiocl<
,eepers by hvestO(k. EXTENSIVE POOR RAINFED lANOLESS
J)loducuon system GRAZIERS MIXED ll\lESTOC!<
tlrom livestock in FARMERS KEEPERS
Oevelopment. 1SS912;,
Arid n• ssm1-ar«f 63 2·3 155·

Temperate (includ1og tro:i1cal highlands! 72 85

Humid. su!>-humid and sull-uopical 89

Toial 135 387 l56

• Predominately in irrigated agricultural areas. bUi also in other densely populated regions

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184 steno" o,c: Aspects influencing the occurrence of infectious diseases

Table 9.4 Class1ficat1on of infectious dis~ses based on 1he,11ransm1ss1on. types of 1mpa1:;, and level oi imoa:1

CATE GORY OF INFECTIOUS EXAMPLES OF MAJOR IMPACTS LEVEL OF IMPACTS JMP.<\CT INDICATORS
DISEASE (s!!!! te}(tl OISEAStS

Epidemic, 0,rec11y Rinderpes,. foo1-and· Dms 10 minimile epidemic lmemat1ona1, reg,ooat. loc ce~ce or disease
transmit.ea. mouth dis~ase, African :is~ nauonal. !ix.ii a·ea. Coo!rcl t~sts
,mernatioral imponance swine ;eiar classical rrcduct,on losses 1r. farme,
!OIE hstAl S\":rrle ~e-,.e.r st:steP1Jble an11na1s ,f
ep,aemic
!les1iic1,ons 10 rnar;cets and
1nterna1,onal trade

2 Endemlc/spor.i~1c. Haemorr.ag•c Proouct,on los-ses NaMna,, lacal area. Disease ouurre11tB

directly uansm111ed, sept1caemca iniecuous Comrcl costs iarmer Pm1uc;,on losses
farm and national bo1 n~m ncirache1us Aes:11cuons 10 ma1ke1,ng and Cor.:ro 1 :os?s
importance fCIE <Sl Bl !ISRJ, cermataphi1os1s aaGe

3 Endern1e1s;:orarfo:. Babes1os S roWOfll)SIS. Froduclion tosses NatllJllal, local a1ea, DiSe;i$e 01:cur(enee
indirectly iransm,tted, theilerian. Afritan Produai()I) oppor:~n,t,es fos: fa!lner V~tor aa~ncarn:e
farm ana raiio~a' a~1ma 1rvpa10S-Omos,s iies:r1ct1ons to ma·~e;,ng and P,aouc:,on losses
importance (DIE ':S; Bl :.race Control cos:s
Corvo! COSIS
l Endern1c.1sporadic, Mastms FrO<iuct•on tosses Far1Tl1l! rna,10nal. total O,sease occurrence
directly transmitted, ISrap,'tylacaccus au,eusJ. Produwon opportun,t:es los, area less ,m00r:an1J Produc:,or. losses
mainly farm 1m;l0r:ance pneumon a Control costs Ccn:·ol costs
(DIE other d,saases) lpasieural!osis). foot ro,

5 Zoonotic oiseases BruceiloscS. mpacts on human hea:1n Nauona1, l0ta1 area. Disease occurrence -
11ansm1tted d11ec:1v or tubercutos,s. ProduchQn losses larme: a01ma1s and huma1ts
1ndirectly irom an,mals T thrxtesiense Contiol costs f'<oduct:on tosses in anirr.als
rn hum~n~ slAP.p,ng s,ckne~ !lnrnPn ni di5i)~sa 10AlVs)'
m h.;;:,ar,s
Control costs

6 food-borne mfett1ons E coh 0157 infecl!oos. Impacts 011 human heal,h local area. nat,onal Buroei,. of disease (DALYsr
salmortel osis. Casis o! food qualil)' control in humans
campylol:acumosis systems tAln:rol costs

DALY~ 01sa0ilitV Ad1usred life Year a measure of the cost 01 human disease

6 Food-borne infecllons and imoxications. such as Sal· Infrastructure and other features of the ~ocioeconomic
monellosis.. and Escherichin coli 0157 infections. Such milieu in differem set1ings. The impacts of endemic dis·
infections are increasingly a problem of intensi\'e live- eases are main!> felt at farm level. while broader economic
stock production and marketing systems in which there Impacts can occur with epidemic diseases that restrict
are significam opportunltles for cross contamination marketing and trade in livestock and livestock products.
or. for example. carcasses and milk supplies. The occutrence c,f epidemic diseases affects borh poor and
richer lh·eswck producers by marginalizing them from
Multiple impacts of animal diseases and thelr control higher-price liveMock markets ond restricting their capac·
LiYestock play an important role in many agricultural sys- iry for 111lue-added trade. 13.' The overall benefits of the
tems and thus animal diseases that impalr their function control or epidemic diseases are great 1811 and In rich coun-
have mltldp!e impacts at a1,imal. farm. ecological zone. na- t-rie, where resources are a1'll.ilable for their eradication.
tional, regional and international le\'els. Decisions on rhi, option is invariablr pursued. Ho1,·e1·er, numerous re·
disease t"ontrol depend on a good unden.randing of th1ise cem examples highlight how the maintenance of disease-
multiple Impacts. Focusing only on direct lo~esof oven dis- free status, when the disease sri!! occurs elsewhere, is risky
ease can lead 10 a poor appreciation of the relative impor· (e.g. foot-and-mouth disease (f'MD introductions world-
tance of different diseases and of what should be done, or wide, and classical swine fever reoccurrence in the ;\<clher-
not done, to control them. landsJ. The specific impacts of epidemic diseas~s and their
The major impacts of different classes of infectious dis· control include both direct disease effects and potemial
eases are greatly influenced by the production system, trnde benefits. f'or the laner, there are usuall}' differemial

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 The comrol of infeclious diseases of livestock 185

benefi1s and costs 10 different lh-escock secrnrs. 135• t$$ they limit economically important herd management deci-
In recent years. a wider perspective on disease impacts sions such as animal selection 011d optimal longevity. The in·
has been adop1ed 1hal considers bo1h tJ1e direct effecls, as teractiotts between disease. nutrition and genetic selection are
well asa numberofnon-tivesLOckfindirecl) effecrs. An impor- important and emphasize 1he necessity to conrrol the effects oi
tani consideration is to differentiate impacts associated \\ith key epidemic and endemic diseases before enhanced nutrition
!he occurrence or cllsease from those associated \\1th the and genetic progr.immes can make an Impact. Ukewfse. sub·
avoidance of disease risk. These are illusmned in Figure 9.2. sramial productMty and economic gains \\ill nm necessarily
The framework in 1he figure is particularly useful for con- be achieved bv disease control alone.
sidering the more profound impac1s. beyond direc1 produc-
tivity effects. that livestock diseases have on African Risk management - comrol costs and lost potential Con·
production system$. TI1ese include the impact~ of disease siderable costs are often incurred in controlling animal dis·
on control costs. the constraims on lh·estock management eases and investments in control measures paid for may not
imposed by diseaSl' risk, including limilations on species always be effective. This is panicularly tnie for smallholder
and breed choices. as well as impacts on the production of farmers, who often lack information and have limited diag-
agricultural and other products, natural resource use, and nostic support to make appropriate disease comrol and
human welfare. The followi ng sections briefly describe che crea1mem decisions. 132 The overall impacr of c:oncrol mea-
major types of impac1. sures may be constrained by non-compliance of a significant
proportion of the community, emphasi2:ing the need for regu-
Dir ect effects on 1.ivestock producth'ity The direct elfuc~ latory \·eterinary services supponed by legis!a1ion and incen·
of animal diseases on llves1ock productivity have been com· tives 10 comply. Art important principle. emphasized by
prehensively described by Morris153 and are illusrrated in several amhorstM. 188 is that it is nol the total losses associa1ed
Figure 9.3. These range from reduced feed !make and changes witl1 dL~ases lhat are important. but rather lhe benefits rela-
in digestion and metabolism 10 increased morbidity and mor· ti1•e to costS 1hat can be obtained from differem con!rol op·
rall!y. and decreased reproduc1ion, weight gain and milk pro- tions. An important loss associated \\1th animal diseases ls the
duction. They influence product quality at animal level and cos! of nor investing in production inputs and adopting less
their aggregate effects influence herd productlvi[), In addirion, profitable livestock management strategies ro a\·oid disease

~ ANIMAL DISEASE ------...

~ - - - OVE.Ri DISEASE DISEASE RISK

Livestock pl'-Oductivlty 1Mar1cet disrnption Risk management

j
• production losses I . access • S!)l;l;ies and breed choice
• treatment costs • pncerisk • maf'la9emem practices
• preventive control costs

I
'
Other income activities
• crop production (manure. I
Livestock prod~c,ivity
'lost potential' I
01aughl)
• fuel, transpon

Household iocome
levels and asset
accumulation r
I
1Natural resources
Human welfare I.Ji'
• itlness. mortality [2Xl0noses • land 1Jse
,._-~ and food-borne diseases) • settJement and migration
• food security and quality • ecosystem sustainability

Figure 9.2 Diseasa ,mpacts {adapted b•; Pe1r, 111 al.. 2002150 irom Swa11ow. 2000232)

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186 ;oc,,.,,- o,:.: Aspec1s influencing the occurrence of infeclious diseases

Available filed not

ALTERED FEED fully utl1ized by
INTAKE animals-consumed
by decomposeis

O,ges!ion '
Al.TEAED ffED
DIGESTIBILITY
Nuui1ive-vatue oi
reed not fully used,
,emainder excre,ed

'
Metabolism Altered nutneni utillzauon
Mmerals and
Altered
respiratory
'
Body material extre1ed
in fae_ces and urine to
'
Consumed bY
decomposers or
Protein Energy efficiency abnormal extent used for fuel
other nutrients

Consequential T
production
eifec1s in
affected
r--'---,
Ptemarure
death
Changed value
of anir:ials
I '
Reduced
bodvweight
'
j Reduced yield
I
and/or ' Reduced
,quality oi milk. eggs.JI capacity for
I '
Altered production
of dung (used fuel)
'
Body material
divWd for use of
wool. etc work disease agent
amma!s

' ,,,, I
Heid
producuon Reduced prodUCli\"e
''
IReduced fertility and I
' '
Lower precision in
I
Increased yield o!
effects I life of animals I fecundny
recognition of animals of
superior genetic ment
disease agent

meets on herd
maintenance and
' ' '
Allered replacement pattern and reduced capacity for
genetic imµrovemem
improvement

Figure 9.3 The ways '"which diseases may affect the produc:,ve value of animals (redrawn from Morris & Marsh, 1994,s.:,

risk. 231 Such strategies may include a reduction in livestock panosomosis in this laner system are highlighted in Table 9.5.
numbers (for example in tsetse-in[es1ed versus non-infested The in legration ofllvestock into mixed farming systems is now
areas in sub-Saharan ,\frica11 1}. variations in grazing prac- 1videly considert>d 10 be an essentlal componem for sustain-
tices 10 avoid tsetse chaUenge.2 ~~ and llrnitarlons in breed able agriculrural developmem.144
choices (such as the necessary use of trypanotolerant breeds
of cattle and small ruminants in West Africa}.2Z• Ecological impacls Another key category of livestock dis-
ease impacts is on the utilization of narural resources and
Impacts on crops and other farm enterprises Given the ecological impacls. Li1•es1ock have been considered harmful
key role or livestock in mixed fam1ing systems, there are nu- to the enl'ironment and implicated in a host of environmen-
merous effects of animal diseases on other agricultural enter- tal sins including desertification, deforestation, global
prises beyond livestock themselves. These include impacts on warming and pollmion. Thus, rhe control of livestock .dis-
crop production through decreased nutrient cycling and eases, panicularly large-scale disease control efforts, have
draught power, and loss of other products such as fuel and been the subject o( much debate and environmental con·
transpon. Intensive smallholder farming systems in many re· cems have hacl a negative effect on investment in livestock
gions of the developing world have integrated livestock imo all disease control. Thi~ has beetn especially true of tsetse fly
other agricultural practices rosuch an extent 1hatlivesrock dis- and rrypanosornosis control in Africa. However. recent evi-
eases. when they occur, will have multiple agricultural produc· dence suggests that the environmental impacts of disease
tion impacts. Good examples of such crop-livesrocksysrems in control efforts are not invariabl}' negative, bur can be also
Africa include the smallholder dairy production system in che positi1·e or neu1ral, depending largely on how people choose
East Afric;.in highlands 130 and the mixed crop-livestock sys- 10 manage their livestock in response to reduced disease risk
rems in sub-humid Wesc Africa. 130 111e multipleirnpaccs of try· °''
(see Reid i>r lll.2 for a fuller discussion related to 1se1se fly

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 The control of infectious diseases of livestock l87

and rrypanosomosis control and its e!fect on land use. sero,·ars. Camplyollacrer spp. and emerotoxigenic colifom1s.
settlement and migration. ecological habitat and biodiver- II is also likely that public health will be nm
her compromis~
sity). Considering such inter-linkages is crucial in arid and by increasing levels of dn1g residue.~ and antibiotic resistance.
semi-arid environmems, where livestock are usually the key For low-income consumers. this will be a serious problem, as
means for utilizing available natural resources and where they will confront many of the multiple antibiotic-resL~tant
lhc risk of a11l111al discast:s (among other ris.ks) compllcatt:~ infections occurring i11 We~tern counuiei. but 11~thout the
rationale utilization of a,-ailable narnral resources. This has benefit of ne\\'er generation antibiotic treatments.
important implications for poor livestock keepers. who rely
almost exclusively on common-propeny namral resources. Impacts of disease on market access Outbreaks of infec-
The~e common-property resource~ are declining in most tious diseases in an area or country may result in local
areas of Africa. and/or international market disruptions as the movements
of animals and livestock products are restricted. Two impor-
Human welfare and human disease Animal diseases haw talll examples in Africa in recent years have been the trade
multiple, im portant and measurable effects on human wel- restriction on livesrock mo\'ements from rhe Horn of Africa
fare. Many of these have been described in considerable de- to the Arabian Peninsula associated with outbreaks of Rift
tail by Schwabe,220 particularly the importance of animal Valley fever and the restriction of meat ex-pons from
products in human nutrition. the social benefits of li\'estock. southern Africa associared with F:'110. The impact of trade
and the impact of food-borne infections and zoonotic dis· restrictions on livestock rrade in the Horn of Africa has been
cases. Livestock products have been shown to have consid- particularly devastating and is highlighted below.
erable impacts on improving child nutrition. 113 and animal In Somaliland alone. the ban assoc1ared wirh Rift Valley
disease control has also been shown to benefh child nutri· fever prevented 1he export of tll'O million head of sheep and
tion and welfare.44 • sr This is panicularly so in pastoralist goats, valued at USS 100 mi!lion between February 1998and
communities, in which 75 per cent or more of general and May 1999 (Figure 9.4). Because of this, local livestock prices
child nmrition is from milk and ocher livestock products.3 ; decreased by 30 per cem, alt.110ugh these lower prices did
Traditional zoonotic diseases, such as brucellosi$, bo·
vine ruberculosis, hydalid disease and anthra.~. for example.
have impacts on both animal and human health. \\'hile Table 9.5 Mult,p!e 1mpac,s of tf\panosomos1s m c1c;i,li\<estot, systemS
many ofche zoono1k; diseases are not among the top impact
tma:nly from S\•,allow, 2000""3.l)
huma1l diseases globall}•159 - where the impact is mea-
Direct ,mpac1s on livestock proaJC'l<Vlty
sured by the Disability Adjusted Life Years (DALYs) lost. they
- retiucec calv10g 1a1es, l-· 2%i tolecan,. 11-20% s<Jscep:iblel
are very important among livestock keepers and workers - ,r,:•eased calf mortaltt, tO- 10% 1oleiant. 10-20% SUitept,blel
handling livestock producrs. A recent studr in Mongolia has - small •uminants Oamll'~91k,dd1ng rates dec,easEd 4-28% /
shown cost:benefit rarios of approxlmarcly l 0: l for the con· 37%)
trol of brucellosis in livestock and 2: l for the direct healrh - aacreased milk production (cow: 10-2611: to18l'ant.
land-area· - 83%)
coses such as hospiral care and drugs. 212 In addition, h wa.,
-(!weased animal offta~e !hew &-31%: land-area - 97'iol
estimated that one year of human disability {DALY) could be - d,ug use (SSA-USS 35 mill,onJ
avoided for USS 34. This study highlights the importance
and benefits of policy changes to enhance inrer-sectoral as- Impacts on hvemck nsk manageme~t
-aecreased cattle numcers (and- 14%; sub-humid - 27%.
sessment a nd action for controlling zoonotic diseases.
humid - 77%)
The intensification of livestock systems in many areas -grazing changes t 1 ,;e
of the developing world'19 ls expected co Increase the trans-
mission and impacts of food·bome infections and imoxi· lmpaC\s on agncuitural and other p!Oducts
cation~. However. because of poor surveillance and - decreased draught e!fic1ericy !40%)
- increased crop produi:t,on (;25-45% per unit lane; +14C-143%
diagnoscic facilities. their impacts on consumers, labour-
oe• •Jnit labour)
ers and traders of lh·estock products in many Afrkan and -a,mease in agricu,wra1 gross domestic pr()duct (GDPl ;n alfec:ea
other developing countries are likely to be poorly under· countries Hrom &-i0•,1
stood and ineffectively controlled. Currently. low-income
consumers face grearer risk from food-borne infections Effects on naiu,a1 re~ource use
- change rn migrat10Nseitlement patreros (variable elfec:s)
such JI$ anthrax and hyd:uid disease, bec:iuse they often
-lim ted to moderate changes in biod,vers,tv assoc,ated with
buy livestock produces in parallel markets in which meat :se;se 1,,· control
inspection is non-existent. In addition to these classic dis·
eases. new zoonotic risks are li kely to develop, particularly lmpac1s on human wellare
in the area or food-borne infections in the more industrial- -loss of ir.tome and assets 1e!a1ed to rmpacts above
-lr·,estcci( reservoir of sleepmg sickness. Eastern Southern amt
ized production and processing system, of developing coun·
Wes! Africa Jlim,ted 1mponance ,n Central Afr;ca,
tries. niese food-borne infections include Salmo11ella

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188 ~1.CT10, o,L: ,\spects influencing the occurrence of infoctiouij diseases

not persist. Unfonunatt:ly. resolving ll\e removal of this ban generation and a.sse1 acquisit ion. especially for the poor.
\\~II not be easy. since they usually have limited access to cash to pay for
Sanitary and phyio-sanirary CSPS) requiremems ofmn animal health services.35, ~s
have an even greater cffec1 on market ae1:ess in local mar- In addition to impro,ing the welfare of livestock keepers.
kets. In many African countries. sanitary regulations for live- enhancing livestock production through the control of ani·
stock products han: been adopted from those pm In place ma! di.seases can have multiple bencfi!S to consumers. trad·
for industrialized systems In developed countries. These eIS and labourers. This is panicularly uue if disease control
may or may not be reasonable or appropriate, depending on and other livestock developmem benefitS are focused on
local conditions. In Table 9.6. the health risks associated poor livestock. keepers such as smallholder mixed farmers
\\ith the marketing of raw and pasteurized milk in Kenya are and pastorallsts. Enhanced livestock production on small-
e:,;plored in tenns of protection of public health versus mar- holder fanns tends to be labour-intensive. using both surplus
ket access and income-generating acti\·ilies. family labour and to a lesser extem labour from non-family

Impacts on household income and asset accumulaUon

. members. ;9 1mpro,·ed smallholder dairy production ha$ also
been shown 10 increase labour opportunities for smail-scale
The benefits of lh·estock as a regular source of 'income', milk traders and 10 lower transaction cosrs so that milk prices
both for cash and barter. have been detailed in 10 poor consumers are considerably lower (Table 9.6). rm
numerous studies. 1:?;s J\ number ol observers (cited In
LID. 19991~ have shown I.hat poor farmers (both those with The importance of animal diseases to the poor in
small land si7.e or tho~e who are landless) are increasingly sub-Saharan Africa
rel>'ing on livestock as their main income source. In addition Because of the impac1 tl1a1 animal diseases can have on poor
to income, livestock are often rhe maln way in which poor communities. disease control can be an important tool for
farmers can acquire real assets, providing a safety factor improving poverty alleviation and enhancing social cqulry.
when difficulties strike. The asset acquisition patlm·ay usu- This topic ha.s been investigated in some deta-U in a repon
ally begins ,~ith poultry. followed by small ruminams commissioned by the Luter-Agency Group of Donors Sup·
and/or pigs. \\~lh larger s10ck such as equids. cattle and porting Re"'eareh on lfrestock Production and Health. and
water buffalo (Bub<//u$ l.mbali,I heing acquired at later sponsored by the Depanmem for lmernational De\·elop-
srages. Animal diseases are a key constraint to both income ment (DFIO) of the Go,·emmem of the United Kingdom. 190

- -- --~
large-scale livestock
------>
Pen Sudan
Livestock taken by
boat to the Gulf
expons originating States
from Kon!ofar and
Oharlur

.,..)
(
/"'

---
• 3erbe0a
Sudan ( \.. Somalia
_.,,_,•
!..
·,.\ __],
Ethiopia

___/ .
/ ./
•

' ,<' • ......_ .,,,,

. .,_r _____ _.. .,_ ·--- .../""'7
). I
\ I
.
( 'Uganda • Kenya J
. ( '
r...r.-·---. ~ D Coumries included In ban
, ._\
'-' <..
Tanzania ' · , D Malo areas affected by ban
Figure 9.4 Main I,ves-..ock produt:ior, arfras ,n the Hom of Africa affected by the Aift Valle/ i8\18r ex~r: tan !source: FEWS NET 2000"'1

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 Tht comrol oflnfectious dlsea...es oflivestotk 189

Table 9.6 Assess,ng anc' manag,ny

mn,.t,orne heallh ris.~s for ;r.,e benefit of In Kenya. aop,o~imately 90 pa: cent oi mil~ ,s produced ov s!l',allholder fanr.ers M~ch o: this ,s
tonS11mers 1n Kenya (from vmore e; al coosurr.ed lotally l:lut inc1ea:smg amounts are being collec.ea. transponect and sold In 10w11s anc
"' ciues by smal!· and large-scale :,aders.
2000""1
Omore ano cc:leagues 113 assesseo a variety of nSks associatec w,ih d1f:eren1 milk mar(etmg
systems. O•ierall. llcth raw and paS1e~r,ze:I milk rare:Y met thlllt atteptiro standards for ~cterial
counts Ninetv per cant of milk reaching consumers was not pasteurized but 95 pe• cen, of It was
boiled hafo:e consumption. Worryingly 5 to 15 pei ceni ol milk sampled had ani1bacterial residues.
however. Satmo11e//aand E co/10,57 were only rare!y detecteo Zooooses lbn;cellosrs and
oov,ne,soorce t~berculosisl were not ae:tected in milk iro'll srnallnoloer farms ba: ani,!Jod1~s to
b•ucellosis in milk indicate that :here •s SGme rlst that increases as milk iHo!lec,ed and bulked
from many so.irees

'n,s studv nigr. 1gms :he trade-offs :t>;;; will need to oe corsioered. in we;gning ooth pubiic hea1:h
and eevnom,c !le'l'ei1ts lo ~rodllCer$, traders and coMumers Two mdk marketing sys.ems are
eme1g1nQ - a fol'IT'al market 'or h1g~·Quality mil~ and m,!k ;:roducis that ,viii !l!l!Ul'e the highest
milk QuahtY s:andards and an informa1 mil~ market ,nvolv,ng small•scale prooucers and traders wllo
sell unoas1eu1ized milk at a thirc less cost to consum1!rs. The informal secror prov1ces employment
to thousands of people The au;~urs recommend measures to ,mprove me quaht; and safety 01
milk from the informal sector (advice to c011Sumers on bOihng milk. training arid imp:Ol/ed handling
containers 'oc small-scale traders! and maintain economi0 bereftts for poo: 'armers. tradeis and
consumers

:Vlany of the points made thus far on the muluple impacts cines and other disease comrol tools, will need to be de\'eloped
of animal disease are particularly relevant to the poor, who are based on newsciemific methods and approaches.
often at the greatest risk of suffer:illg from the deleterious It is generally acknowledged that poultry are very impor-
effects of animal and zoonotic diseases and who have the least mm to the poorest ofthe poor. The major disease constraim
capacity to control disease or cope with disease risk, Here. we to poultry keeping throughout the developing world is
briefly outline the main pathways in whfch the control of ani- Newcastle disease.228 However. greater benefits from New-
mal disease can benefit poor people and illustrate this \,ith an castle disease control could be enjoyed if other inputs such
example of the control of Xewcastle disease. Readers are as the control of oLher important diseases e.g. Gumboro.
referred to Perry er al. 190 for a fuller accotmt of tl1e Impact of fowl pox. internal parasitesJ plus impro\'ed housmg and
animal diseases on the poor. feeding were also a~-ailable as a package. Essentially. some
The control of livestock diseases is likely to improve the benefits in securing poultry assets ,,ill be achieved by
livelihoods (see Carne).32 and DFID, 4~ for further accounts providing thermostable :-.:ewcastle disease vaccine alone
ofiivelihoods Issues) of the poorin three main ways: but greater intensification and income generation benefits
• Secure the current assets (human, financial. social) of could be obtained if input packages were delivered. These
poor people who keep livestock. cot!.Sumc livestock prod· are e:1.'l)andcd upon in Table 9.7.
ucts, market livestock and livestock products, a.nd work as
wage labourers \\ith livestock, by reducing the risks they
Measuring the impacts of infectious diseases
experience through animal and zoonoric diseases.
and their control
• Enhance the marketing opportunities of the poor by con·
rrolling the diseases that affect the movement and mar- There is a series of data needs required for the e,·aluation of
keting of li,·esrock and animal products, primarily disease impacts, and of the impacts of alternative in terven-
locally. but also regionally and imemacionaUy. tions to prevent or comrol them. These are similar in prin·
• Reduce the constraints experienced by the poor co live- ciple regardless of the scale of assessmem (i.e. whether a1
stock-based pathways out of poverty through intensifl· the level of herd, communit}', production system. agro-eco-
cation (improved productivity and performance logicalzoue. country or region). These fall into the following
efficiency through the use of inputs). broad categories:
• • Size. structure and composition of an!mal population
For many d[seases, control effons targeted at poor livestock under consideration.
keepers wlll simply require the transfer of a\'ailable kno\\·ledge • Characteristics of animal population and its cnviron-
and tools: for other diseases. currem tools and deliverysystems mem. panicularly in cerms of agro-ecology and feeding
\\ill need to be adapted and impro\'ed so that they are appro· managemell!.
pria1e for local conditions, and in particular for optimal uptake • Disease occurrence. in terms of prevalence. incidence.
by the poor. ln some instances, new 1eclmologies, such as \'aC· distribution and dynamics.

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190 >Kr~,~ o,f: A~pt.lCt> innucncing the occurrence or infe<ctious diseases

Table 9.7 Assessing ~ptions fo1 Newcas:ie disease(NOi control bv vaccmat,on ao smallno c!et subsistence poultry system1 oi tile d~e1op111g world

Socio-pol111cal and er.v,ronmental context

- L1m1te<I mpu!/ou,ll\,, svsiems
- Variety of social networks
- Relatively weak public support inst1tu1ior.s for credit. animal health and product,on inputs
- Markets variable but some local demanc

J¼lu1si11on or capita'
- Village pouluy aCJ:e,sible to paor aoo oi:en provide an effectwe starting point for acquisiuon of additional livestock assets (pigs. small ruminants);
livestock asse,s maimain their value in Environmems w1111 weak banking systems and are crucial in umesof crisis (such as heal!h emergencies, e«:.l
- Can be instrumental in ;irov1dmg funds for school !ee1; ihuman .ap,rail:93 enhanced notn:,on - particularly 1mportan; for child davelopment 113
- Depending on scale. mar',e1s for village ;ioultry are relative!·, 'tlbus; (physical capital) but will require enMancement for large·-scate producuon
- lmportal\Ce o' poultry as $9(ial caprta -ab1l1tv 10 fulfil sm:,at obligat,onsm
Eval~auon of opoort111u1,es
- 50% mortality rate main reason for lowouiouL 50% of mortalities due to 1nfecuoos c,seases; NOmost 1rr.pcrtant and widespread m!ecuous disease
of pauluy 1711 !Annex ' 2 ln Reference·~)
Potential op1,ons
A) Oi~tribution of thermostable ND vaccine in feed (S.E. Asia) anci by eye drop (Mo,ambiqi.ev19
Bl Enhanced 1nput-outpu1 systems includr~ NOvaceination·21 ; \'BCC,nation. micrc-uedrt 111proved breeos and feeds
Ass&ssmem of opt10ns
Opuon A' Apore;,riate vaccination s1ra1eg1:s 10 secure assets
- success depends on development of ap0top1ia1e and very inexp1ms1ve vaccine p1oduc!icn and clis11ibuu011 systems\being deveio~ed and assessed by
ACIAR and partne!SJ
- limited bene.lits m terms of aqu,s1tron ol capital can be caotured ov ooor peoole in a •,-ar,2ty of comexts
- economic betl8fits 1ccsttlenef1t r.mo 1~ 1 '!:-<) rmoonant flUl hrr ten due to cons1tain1s from other hm1t,r.g t~puts and ma:ket i)Otenual dependr~ Oil
market access
Option B. Vactinanim and other inputs 10 <mprove the intensTcation oi rural poultry keeping
- greater success possible ii more lntegia:ed support systems dimloped (housing, feel1ing. etc) but this is ~i an easilydeli\<Sred opuon evarywhl!re
1successfullv appl~C mBangladesh710 out needs assessment elsewhere)
- has greatest pctent,al for mtensilicauon and helping to reduce poverty b111 requires e~hanced markets ior both input delivecv and outpu~

• Effects of disease (o r ofimer\'entions) on producti~ity (or Us e of intuitive response

other ke~, indicators). :-Ian}' impact assessments use intuitive valuations of some
• Value of producti\'ity losses of disease/ incremental gain or all of the parameters listed. This might sound very
from Intervention. non-scientific and obvious. but is the rule rather than the ex-
• Value of control costs of disease/ incremental savings ception at the fann level regardless of the produaion system.
rrom intervention. Even at the national level, where resources are inadequate and
• Impacts of zoonotic diseases {or of interventions) on the when impact assessmems are demru,dcd wi1hin n short time.
human population. this technique is common. As a general rule, when made hr
• Environmental impacts of disease (or ofimerventions). fam1e1'$. this tends 10 be based on a combination of personal
experience and financial commonsense, when made b)
This series combines demographic. epidemiological. pro- economists it tends to be on the basis of sound economic cri-
ducth~ty and socioeconomic data. illustrating the !ncreas· teria regardless of technical considerations, and when made br
ing t,end towards a mulri-disciplinary approach t0 disease veterinarians il tends to be on the basis of technical consider-
and disease conuol imp(lct assessments. Clearly, a-• a gen- ations regardless of economics!
eral principle, the broader the range of data included, and
the better the quality of the data, the better the resulting de- Passive su1veillance systems
cision. However, ~he collection and processing of dare also The o,tandard source of quantitative data on livestock popu-
have a cost in tenns of money, time and communication, lations and infectious disease occurrence In most countries
\\Illich muS1. be taken into consideration. a m! may have a sig· is still from national passive reporting syMems. 111 1.hese.
nificant effect on the quality and \'alue of an~· impact assess, livestock population sizes by adminiscra1ive boundary (dis-
ment s1udy. trict, province) and cases of a given disease observed by the
For each of these categories of dalil. there is a set of veterinary depanmem are compiled in periodic reports. The
techniques that can be used for data collection and assem- advantage ofusingsuch data in an impact assessment is that
bl~. and thes e are listed and discussed in general terms they are a\-ailable. and generali}'at no cost 10 the user (but of
below. course often at a s ignificant cost to the state in their

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 The control of Infectious disea~es or livestock 191

compilation and publication). However, there are numer- procedures, enhanced l.>y the information and communica-
ous disadvantages. which usually far ounveigh the advan- tions explosion, and the Internet. There has been a greater
tages. The data are often long out of date, with some annual focus on priority diseases or disease complexes. on greater
reports of veterinary depanments in Africa published 10 privatization of the data collection and analysis, which then
years late. Funhcrmore. there is a tremendous variation in become a marketable product. as well on the greater use of
the method of reporting used. As far as livestock population active surveillance techniques.
size., are concerned, there are only a fe\\· countries that carry The decision as to whether or not 10 use official reports of
om regular livestock censuses. In many cases, populations disease occurrence based on a passive reporting system wilt
seem to mysteriously increase annually by a standard index. depend on a deeper knowledge of the quality of the data.
There is also a large variation in the categories of livestock and on whether standard reporting procedures are being
quantified, some 1aking species. some dividing wi1hin spe· followed. This can vary dramatically from place to place.
cies (such as beef and dairy), and very few quantifying on 1he and from year t0 )•ear. depending on the emhusiasm and
basis of production system. . diligence of individuals. and of course on the resources
The continuation by most national veterinary authorities available for such a service.
In Afriea to report li\'estock populatiOtlS on the basis of spe- ln passive surveillance. consideration is rarely given to
cies rather than production system severely hampers im- reporting tl1e producthir.y effects of a disease (or a control
pact assessments. We have. for example. eiq>erienced this intervention). producti\'h.y lossesfincrememal gains from
recently in studies of the impact of F1'1D and its control in an intervention, or control costS/incremental savings from
rhe countries of south-east Asia. Tiw pig populations of an lntervenrion.
many countries in the region are reponed as a single cmity.
Hidden within this are at least two contrasting production Ac1ive surveillance systems
$}'Stems, in which both the infection dynamics of P.11D and As a result oi an expert consultation. FA060 developed a
the economic impact of its control are entirely cllfferem. 188 frameworkforanimal health information systems in devel-
:.tedium- and large-scale pig production present in Thai· oping counrrie~ that could serve the varying needs these
land. the Philippines and to a lesser extent Vie1nam is simi- countries face. A key component of this report was a recog-
lar to commercial production enterprises in many pans of nition that the passi\'e disease infonnacion systems that
the world. with intensh·e hygiene measures, regular preven- have been in place for so many decades are often of littJe
tive FMD \'accination, and little in the way of FMD occur- value, frequently being our oitouch and highly inaccurate.
rence. In contrast. tl1e vast village and 'backyard' sectors are Strategic active $Urveillanc;e. and strategic sample surveys
highly vulnerable with little vaccination. hygiene and move- and studies with focus, were considered co be much more
ment control being applied, and as a consequence, frequent useful. It has been encouraging to see this approach
ourbreal;s of the disease occur. adopted in some regions; for example, the southern Afri·
As far as disease reporting in passive surveillance sysrems is can countries recently held a workshop in Pretoria on de-
concerned. there 1s much variation from country to COUllll)' . mand -driven animal health information systems in
and several different indices are used. For example. some use collaboration with the OIE. FAO and ILRl.
number of cases, while others use number of outbreak$. ln Rabies surveillance in Ken}'a pro,'ides a good illustra-
both instances. there is often a mhm1re of clinical and labora- tive example of the problems of passive collection of data
tory records conrributing to this index, which may go undiffer- and the potential opportt.1ruries for strategic active d.ato
entiated. This is particularly important with infections in collection. !'or many years. passive data collection for ra-
which the mere presence of an organism does not necessarily bies was considered to pro~·ide .a reasonable indicatiori of
mean it has anything 10 do with the mortality. di~ease or rabies OC'currence in Kenya, albeit an underestimated one,
production loss under investigation. i\lere identification of particularly in non-endemic rabies areas. Potemial rabies
haemo- or gastrointesrinal parasites, ror example. is often exposures were reported ro veterinary offices for follow-up
tak.en as diagnostic when their presence may have been merely and any humans considered e.'Cposed were referred to gov-
coincidental. ernment medical officers for free/subsidized post-expo-
Generally. passively derived systems have been at their sure treatment. However, overtime, the laner form of post-
best when not applied nationally, but rather strategically exposure treatment was withdrawn, without alternative
and demand-driven for specific diseases. The overall de- suppliers being established. and many people gradually
dim,,-0( passive!) derived disease dam has probably been stopped reporting cases as there was no Incentive to do so.
less in developed countries and in the more commercial In order to beuer assess the rabies situation \\ith a view to
livestock enterprises of southern Africa In these areas, there improving control. Kitala et al. 106 established an ac1ive sur-
has been a significant shift in data collection techniques. veillance system using stratified random sampling for
through greater focus and more use of active methods. human and animal rabies. link.ed co a network of private-
The$e changes have been supported by improved technolo- sector vaccine suppllers. :\ctive surveillance through com-
gies ior animal identification and more effective trace-back municr rabies workers uncovered 40 times more rabies

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192 si;cno., o,,: Aspects influencing the occurrence or infectious diseases

cases than the passh·esurvelllance S}'Stem. as well as provid· ing the occurrence of something (of a disease. for example).
ing data on dog demographics and transmission parameters whereas snidics invol\'e the comparison of the occurrence
for the development of models ro assess rabies transmission of the evenc of interest in different groups of animals, usually
and the potential impact of control measures. 107 Depending undertaken 10 elucidate causal associations. The major
on resource availability and priorities. countries could con- characteristics and useful feamres of different survey and
sider mi.xes of passh·e surveillance (If It can be linked 10 srudy types are outlined In Table 9.8.
animal or human health service provision) and active sur- Obser\'ational srudies are srructured comparisons oi
veillance from sires chosen using probab ility-based sam- the rls~ or rates (depending on the smdy) of disease (or of
pling methods. other parameters indicatJl'e of disease, such as infection,
or decreased productivh-y) between different subsets of the
Observational studies animal population being srudied. Successful studie~ de-
There is a large number of different types of observational pend on the accurate identification of I) the presence of
sun·eys and smdies that can be use~ 10 gain more precise disease (or infection) in the study group. and 2) the pres-
data o n livestock populations. their characteristics. infec- ence or absence of attributes that may be determinants of
tion and disease occurrence, and disease causality. Below the disease (or infection. or deviation from expected per-
is outlined the general characteristics of such studies. but formance). Thus in its simplest form. this results in rhe
ior more detailed discl,lssion of their relative merit~. and generation or a 2 x 2 contingency table. as illustrated in
design and analysis issues. consult the n umerous texts on Table9.9.
the subject.143. 1so. 201. 221. a;. 2.39. 200
Cross-sectional studies In the process of beuer under-
Surveys The 100! used to gather further information rrom standing diseases, their causes, their impacts and methods
lhe field on a particular subject is often termed a 'survey'. to optimally control them, a sensible starting place is to de-
Thrusfield,239 a stickler for detail. differentiated between scribe disease pre\·alence and what factors or attributes
surveys and studies as follows: He defines sun·eys as count- may be associated with iL For common diseases, cross-

Table 9.8 A. sum'llary ot the main lea,ures of-surveys, observa1:onal studies and thnica' tnals In vetennary epidemiology

RETROSPECilVE PROSPECTIVE
TYPE SUP.VEY C•lOSS-SECTIONAl. CASE-CONTROL LONGITUDINA.l COHORT STUDY CLINICAL TRIAL
STUDY STUDY G'ilSERVA TtON
SiUDY
Disease or Prevalence Prevalence Prevalence Prevalence and lnc,dence Incidence
1nfec1ion 1sometimes ,ncidence
frequency umt inc,dence)
measured
Comparison 9.•oup No~e (counting Nor,affected Selected controls. Usually non. Selected animals Ma:thad animals
cases) animals ,n the matched \·11tr not exposed to exposed :o
s1udy ·cases' tor fat:ors potennat causal infe:tlon or
(e.g. breed. agel determinam disease. bot not 10
treatment
Pnmaiyda!a Sum. mean. Re!auve nsk. Odas rauo. Depends on Relanve rist Relative rotk
ana)ysis procedu•e med•an attributable r,s~ amibutable r.s~ ~rame1e~$) lleir.g amibutabie tis~ atul!J\itable risk
nonuored
Ease of conrluct Very easy ana Easy and chsao Modera1etv easv Hare and Hard and Hard ano
cl\aap and cheap expenswe expensive expensive
level of causal LO",\t Low Low to mo(ferate High High High
proof

.
Deorea
. of
invesiigator
\Jom1 Very 101'1 Low Mode.ate Moderate Hlgh

control
Rele\eance tO the Moderate High but poor for Mode1a1e and High and good ior High and good fc: High and goOll for
neld situauon impact measures poor for ,mpac~ 111pact if s,~dy impact if study impac, if swdy
measures POpulation is populauon 1s p011Jlation 1s
repni-sematr,e representat,11e representa11ve

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 The C1>mrol of infocuous diseases or li\'csiock 193

sectional studies are the logical. quickest and easiest first have been acquired at different times than the disease it-
step. For rare diseases, when disease records are a1-ailablc. self. and 1hus causal relationships can be difficult to estab-
case-control Sludies 10 compare cases with different non- lish. However, causal relar1onships are somewhat easier to
case groups are commonly employed. After preliminary in- establish for attributes that are relatively unchanging and
formation on a disease has been collected. more powerful characteristic of the production system in which the ani-
but costly study designs, such as cohor1 studies ()r clinical mals are found. On balance, the priocipel benefits of cross·
trials. need to be employed to assess lhe impact of risk fac- sectional studies are their ability to estimate prevalence of
tors on the incidence of lhe disease and to assess the impact disease (which may be a rough guide 10 itS impact) and
of control factors on direct and indirect losses associated how prevalence cases are distributed by potennal risk
11ith the disease. groups.
Thus, the commonest st0dr used in the field in sub· Ca~e-control studies are also relatively easy and cheap
Saharan Africa has been the cross-sectional. !"his has been 10 perform and have been used as a first step in trying to
the starting point for almost all effort$ to study infectious assess disease impact and factors associated with disease
disease occurrence in the field. It can be used to establish occurrence. The initial step is to identify ·cases· of the dis-
the prevalence of a given disease or infection. and the asso- ease wilhm a study population. Strict diagnostic guidelines
ciation 1\ilh different natural (such as l4lro-ecological) or must be established to define the case; the va lue of this
human-made variables (such as mamigemen t or leeding). 1ype ot study hinges on the quality of the diagnostic crite·
It has been quite extensively used to study the epidemiol- ria. particularly with respect to their specificity. The same
ogy of ECf over the last decade or so. 47 • , 5 • 136 1\t the com· guidelines are then used lo select non-cases. or controls.
mencement of a cross-sectional study, the onl}' number from the same population. Dependins on the rarity of the
that is known is the total animal population under study disease. multiple controls per case may be selected. Case-
(or tl1e number of the random or otherwise selected comrol designs were initially responsible for linking
sample of 1hat popularionJ. The number wi th and witl1om smoking to lung cancer in humans, 53 and have been used
the disease, with and without the anrioutes under invesci- in the veterinary field to study a vJriety of conditions in-
gation that might be associated with 1he disease, and any eluding feline urolichiasis, ~ 3 ~spiratory disease outbreaks
quantities 10 be estimated in the study, are unknown. lm- in cattle200 and an acwe and highly fatal imes1inal syn-
ponantl}'· ii will usually not be possible to differentiate be- drome in horses. 18" While often very usef1.JI, case-control
tween ol.d and new eases (of disease or incidents of s tudies are susceptible to multiple biases. which has led
infection in the case of an tibody prevalence being taken as them to be dismissed by many. 61 Some main drawbacks in·
the indicator of disease) in this type of sn1dy. The cross· elude trying to exm1polate results from a specific control
sectional study therefore measures point prevalence. The populaclon (based on hospital or slaughterhouse records)
attributes being srudied as influencing disease risk may and changes In 1he case population after their derection

Table 9.9 Basic data analysis ior assessing tne association between a t1•10-categoiy disease 11s< factor and disease or infection ocrurrence ind tferent
types oi observational studies

DISEASE (OR INFECTION! STATUS

P'lESENT ID+l ASSFNilD-1 iOTAl
RISK PRESENT (F+) a b a-b
HCTOR ABSENT (F-l C d c-d
STATUS TOTAl a~c N=a - ti-c+d

TYPE OF STUDY SEQUENCE OF RISK FACTOR ANO OfSEASl: STATUS DEFINITION CAlCULATION o: RISK MEASURE
Cmss-secuonal F+ (}... °ID•/F+l IRelatwe osk from
F- ().. '(0+/F-) p,evalence datal
Information on risk factors and disease status collected at ooe time
Case-comrol F-.1 F- ~ 0- :jO+/f+! / PIO-/f+I (Oddsratio)
0- "IC+/F-l / PIO...iF-1
• Disease cases ano controls chosen and retrospectively checked for
risk' factor stat11s
Cohon f./ o.,o. 'tD+/F+I IRelauve osll from
F- '(0+/f-) 1ncide11ce datal
Disease-free populatioos wuh and w1tt:out the r,sk factor are
followed to determine disease incidence

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194 """'"" c.".: Aspcc:1s influenr,ng the.> c,ccurrcnc1• ofinfcctious <hst!O~<">

e.g. dairy farmers with antibiotic residue violation<. who benefits can be otm1in1Jd u~ingspC1cific .ira1egies to control
subsequently are better infomwd of sound antibiotic ad- the disea$e, rn 1 Clinical and lield trials pro\ide the optimal
ministration prac1kesJ. These bia.es can be reduced to method ior assessing the impact of comrol s1rn1egies on
~omc cxwnt, ior e;,;ample by im·c~cigating multiple control di5ease incid<mce. Ii\ eswck production. reduction in con-
groups (oflcn a control group in which the disease has bel.'n trol cost, and orher direct nnd indirect disea~e impacts (see
definitively excluded ui)d a contfol group from thc general obO\'C), Tlw key ad".imai::c of 1hcse triab i~ t.hot th..,, arc con-
population). ducted in real populations or interest. either indh'idual ani-
mals. !arms or disease control arens. The more the stud)
Longitudinal studies Both rro~s-sectional and case-con- groups re~ernble the diseased populations ol' .merest. the
trol studies pro~ide a ·snapshot' or disease in a population. greater the ahiliry 10 e.,1rapolnte re~ults \,idely. Another use-
pro\'iding useiul spatial data but lacking in temporal infor- ful feature is that new control strategic"' can be compared to
mation. lm·estigatlon or what ,ausaJ factors precede infoc- the current s1andard strategy. thus m·oiding I.he difficult situ-
lion or disease incidence and thl' sequence of disease and ation of tl)ing to force farmers to apply nn control in the face
control impacts are best invl>stigated in longitudinal ~tutl- of disea., e risk. Finally. because rhcirdesign requires the ran-
ies, in which study populations arc monitored m•cr time. dom allocation of animals or forms into trenrmem j!l'Ottps.
There are three main 11ariams oflongiludinnl stud) dt'$lgn. clinical or field trials are extremely po\,·erful, pro\'iding for
The first does not 1necessanl~, involve a compansoll stausucal compansons tO be made without the worry of con-
group, and !~ simply the monitoring of pantcutar disease founding. which is a cons1ant Jlroblcm in obscrvarional stttd·
fea1urcs O\•er time. This may he ,imply to derermin<.> the ies. l's~<:mi111ly. clinical trials have a very ba~ic design. A :,tud)
dynamics or a panicular infectious dise(lse under denned population. representing the reforem·c populallon of inter-
field conditions over time. or to determine 1he relationship est, i, assembled and randomly nlloc-ated into dilferent treat-
between Jeveil, or infection challen~e and ho~t respon~e (in ment groups-. depending on the treatment or control
terms ofimmuue response. morbidity. mortality and other comparL<ons co be made. Trcatmems are made according to
productivity parameters). t\n impnrwnt Ccature Is the abil a standard protocol and, when possible. some degree of
ity to mea~ure infection and disease incidence. and dis- blinding. In whkh thost• who a»sess the disease and
ease-specific morbid.it)· and mortalicy. all crucial components productivi~ outcome,- :1rc unaware of the trc-atment suuus.
of impact assessm1:1n1 studies. llowever. longitudinal stud· l·o!Jo\,ing treatmem, standard follow-up protocols a.re
ies are e;-;pensil'e to set up and arc logi~tfc-ally demanding. itllplcmented to record lo~~-w-l'ollow-up and clinical and
particularly ir the smdy size is large. In additjon because protlucuon parameters or interest. Hecause individuals or
of their expense and complexit). Lhey are usually c~rrie·d herd, an: rnndoml) as,;igned to trearmem group,. the ~1atis-
out under very defined local condition,. and il may tical comparison 10 a~~c,~ difference\ between group~ j,; usu,
not be nppropriate 10 extrapolate the result~ co oth(·r ally ~u-a,ghtforwnrd unless there has been differemial loss-10-
circumstances. follow,up. Funhtor details on cliniC'.t.1 and fit.lid trials can be
Two other ~ludy type~ using a longitudinal '11"\lnure. fount! m,tandard ve1erinary cpidemio!oID· texts, a.- well as in
beginning with a disease-free population and monitoring :--ordhuiz<:n er tJ[. 1t,.j and :O.!einert. MB
rhe incidence of disease over rime. are cohort studies Can
ob,ervation i;mdy lype) and clinical 1riab (an experimental Important considerations In conducting epidemiological
studv type:. In cohort studies. the starting point is identiry- studies in the field :,tandard surve, and srudv designs in-
ing a group of animals exposed 10 a pankular su~pec1 , ariably need to be adapted to field conditions. The cha!·
causal determinant. and then the Identification of a lenge b to maintain the essential fcaturh of ob~Cl'\'31ional
matched group of animals not exposed. Hoth groups are or experimental de!-igns that "~II lead to ,·atid, meaningful
monitored for the developmem of di~ease. This design or and lnterprerable re~ults. In Table 9. 10 a checklist of key ge-
study is commonly used in human medicine with regis- neric ~teps that need to be followed when planning a sul'\·ey
tered members of 1he population \\ith certain character- or study ;s pro,1dcd.
istics (e.g. located in a high-ri~k radiation orea, or with Proper ~ampling Is crucial tCl d<Jsigning a study that pro-
diabetes) 10 monitor their subsequent health compared 10 1'ides maximum information for minimum cost and to ob-
groups withour lhose c.:haracterisuc.,. In veterinarv medi· taining 1•alicl results Given Its importance. there are
cine, this design is primarily u!tecl where detailed herd surprisingly few comprehensive textbooks on sa,npling that
IJ10nitoring schemes are in plac-c. One example of a cohnrt can be read b)' non-mathematician~. F.xception, Include
smdy used to assess the incidence of ECF in differcm risk Cochran H Levy and Lemeshow. 121 and \'ates.258 However.
cohorts or female calves was conducted by Git au er <11. 71 1h1:re are a number of papers. repons and books that de·
scribe useful methods for Important sampling issues in ve1-
Clinical and field t-rials In assessin!( the impan of dis- erinary field smdil!l,..i~. 10 t:, 117• H '.I Tite imponam principle
eases and their comrol, the most important aspect is ncH in all sampling is that every animal ha~ an equal or measur-
what the total losses associated with a disease arc> but what able probab ility of ~electl<>n.

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 ll1e control of !n!cctlou1<diseascs of lkcs1ock 195

As part of the different field ,tudies discussed above, and different caulc type~. and it maI be more effective
1he following practical 11ucsll<>ns usually need 10 be 10 strat.ify rhe di,;trict on the basis of these variables. Stan-
addressed: dard statistical tcns41 and epidemiology texts 1~3 describe
• estimation of 1he prc,·alcnce or incidence of infection or the sampling option~ available under different d rcum-
disease; scanc~
• esllrmulon uf tlw tlbtril>u tion uf rbk factor.., lrnplemen· !·or lnfl'cllous tll~ease;,. il b usual that the rlskl. uf lllstm,e
tation of control methods and other factors a5sociated for animals in a herd, agro-ecological zone or other group-
wilh disease: ing are 1101 independent. \\'hilc non-independence r~quires
• comparison of disease rates. survi\'al, mortaliry and pro- a modification of standard sample size fom1ulae (un In·
ductivity cffect5 for different risk groups or under differ· crease in sample- size if the risk in a group is positively corre-
em control ~trategies: and lated), and l~ thus considered an extreme nuisance by most.
• detect ion or disease presence. ic acwally provides key Information on the \"ariability of dis-
ease risk and transmission. ! lighly contagious diseases. for
for all of these questions, the sample size wi ll depend on e.xample. tend to infec, almo51 all animals of the susceptible
che level of precision desired for the statiscical estimace of species on a fann. The clustering patterns of other diseases
interest. 1.he hJ,·el of confidence required that che estimate can pro\ide infonnation on whether ,,ithin- or bem een-
of imerest lies within a given bound. the independence of herd risk factors are more important or whether factors
sampling units from each other. and the performance associated \\ith ecological areas are most influential (as
(sensitivicy and specificity) of tests used to measure happens with many vec1or·bome diseases. 118 Methods to
disease. For 5Tnall (finite) ru1imal populario1ts, che sample account for disease clus1ering. both in time and space. have
sl1.e may be reduced as a function of the total population been an acth·e research area in recenc years and there are a
size. t ' 3 Standard sample silie calculations a.ssume sampling varlet) of techniques that can be used, ranging from simple
from an infinite 10 000) populacion. Olien m complex sur- to complex.
veys orsrudies with multiple risk factor$, it is difficult. a priori,
to estimate 1he size of important comparison groups. In the
Table 9.10 Steps 1n planning a i,uia sU1Vey or stucv
end. as with mosc things in lifo. practic.'11 and logistical
queslions will play an imporcant role in arriving at a final Clearly state me OBJECTIVES of the siudy.
sample size. - wflal parameters are tc b'3 estimated'
- what is the U~IIT OF CONCERN (animal. iarm. a~al?
/\ staning point for csthnatlng sample silies in cross-sec-
tional studies is to estimate the likelv pre\·alcnce of the dis- 2 Nha1 s me TARGET POPULA'ION?
1

ea~e or infection in the swdy population. Well. you might - all animals
say, if we knew that. we wouldn't be doing the study! The - an age. sex or 01h~r s~bgrmip
Iariest sample sizes are requi red when the estima ted prev- 3 Can a SAMPllNG FRAll'.E ll!STJ o! :ne TARGEi PO?JLAnON be
alence i;. around 50 per cent. and the smallest when the es- oota,ne.i?
,t YES. collect,:
timated prevalence is very high or very low. There is a reason - ,fNO
ior this apparently paradox.ical situation. The estlmation of - wllat lists are a~a11ab'e7
prevalence requires an estimation of the proportion of pos· - does this Its: ,ep,esent the TARGEi POFULATION7
ltive 10 negative animals. rather than the ab~olute 1mmber of - ls tliete a listmll ot :he ur.ITS OF CONCER!I! or large1 UN'TS
positives. TI1us a larger number is required to demonstrate a ;1om whicll l1sis o' p11marv sample units Cc" be obtainea a:
!Ol'. er costs?
50:50 relationship than one of 90:lO (or 10:90). Thus. if no
estimate of prevalence can be obtained, tl1e sample size 4 To mee1 your ob1echves. wl-at DATA and BIOLOGICAL SAMPLES ~ee.
to be collec1ed?
needed 10 de1em1ine a prevalence of 50 per cem will define
- State tl1e data neet!S i!Xp:tt1tly
1.he upper limit. - Consider the logis:ic difficulues
Depending on che distribution of disease. simple
5 Sele•· a SAMPLE SIZE t<ms1s1ent w,th your ObJECl•l<?S
random sampling proc-edures may not be very efficient and - Wtiat PRECISION (e g w,t111n 5%) 1s required
more complex sampling methodologies. such as stratified, How CONFIDENT 195"· 99%1 do vou ,._,,sn io be 'Ni1hycur
cluster and muhl·~lage sampling. may greatly decrease eslimate7
costs. increase sampling eflicien~1 and provide estimates Can •1ou ao1us1 you· sample size beeause ,M target
of dii>ease occurrence at 1':11-ying lcH.!ls of interest. for p~p1Jiatton IS FINITE'
ke responsesoi mot'.,dual unns ma r.ero or other cluster
example. a well-designed smdy to determine the pre,;alence ,ncel)elldent or Nrre'ated w,th 1esoonses oi o:he, 1nd"1v1duais
of an infeccious disease m a district may be of liule value n that herd or cluste:7
in dl!tennining the impar1. or in deciding on an inter· 5 W;;:~ oa: a detailed \VORKPlAN
,·cntion. ff there are significant differe11ces ";1hin the
7 PRETEST your survey or stuo~ ~saumems (questtonnaiTes. diagnos:,c
districL These differences may depend on factors such as
tools. etc.I
agro-ecological zone. management and reeding practices

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196 lllcnu~ t,st.: Aspec1s influendng the occurrence ofinfeclious diseases

Asimple and usually adequate procedure to applrif dis- /\tore common are Impact and scenario development stud-
ease risk varies between herds is 10 inflate ,he sample size ies chat informally evaluate ai1d combine data from previ-
estimate by a factor of I + (m - I )p, where m is lhc median ous studies. 183 In both sets of circumstances, an obiectf,e
herd size and p is the intra-herd-correlation coefficient evaluation of the e,idence presented b} different studies is
(which measures the non-independence! of cluster mates). an important step in impact assessment.
A practical example of the use of this method in a study of
anribody prevalence to tick-borne pathogens in three dif· Models
ferent agro-ecologlcal zones of coastal Kenya can be found While evidence based on empirical field studies is critical to
in the publication of Deem er al."7 More complex methods assessing the impact of a disease and its control. the wide
for two (such as animal and he,rd) and multiple (such as range Offield situations, control scenarios and orher unique
animal. herd, community. ecological area) levels of poten- disease circumstances mean that all the empirical impact
tial correlation have been described and applied to a vari· data required are unlikely to be available. Because of die
eiy of veterinary and medical situations. 122• 126· 127 12ll \\~de \'ar[ety of disease scenarios possible, models have been
Another important disease samphng application is to as- used ro build a plausible framework that can be evaluated
sess the presence of disease (or vectors) in large-scale control and validated with empirical field data. and if reasonable.
and eradication campaig11s. 123 Cameron and Baldock29• 30 used to make impact predictions under different disease
have developed very useful methods for assessing the prob- and control scenarios. A wide selection of models has been
ability of disease presence using tests of va11ing sensiti\iry used for different purposes to assess livescock populations.
and specificity. These programmes are available and easy to disease C}rnamics or combinations thereof. Based on their
use.~ malhematical formulation, models are either:
,o\s noted in Table 9.10. rhe mosl imponam and firsnask • deterministic - in which a system of equations with an
in developing any plan for a field survey or study is to care- equilibrium solution i.s solved:
fully define the study objectives and from these identify • stochastic - in which random probability elements are
important derails such as the parameters to be measured, i.1wol\'ed that can lead to exci nctio11 or outbreaks at various
unit of concern and target population. Invariably. good points; or
studies rest1lt from appropriate and well-defined questions • simulation - in which computational algorithms are
being asked and poor smrlies result when the questions to used to incorporate a wide variety of influences on differ-
be addre~sed by the study are not correctly formulated. ent outcomes or states.

Evidence f~om the published and unpublished literature Livestock population dynamic-s models have been used to
ln assessing the impact of diseases and their control. mucl1 predict herd growth and productivity for different systems
of the impacr information requi'red may be available from in Africa. One of the earliest models was developed by
previously conducted studies. Unfortunately, study validity Texas A & M Univer.sity215 · 216 and subsequently applied 10
and quaHry vary greatly. Some studies can be relied on to beef cattle production in Botswana. 96 The model ac-
provide good quality impact information While others are counted for the interactive effects of genotype, breeding
best left unread. Deciding on study quality is a damning task system. nutri1ion. reproduction, growth. milk production
for the uninitiated; however, \\-ith common sense and some and nutrient intake.Animal performance was simulated by
basic guidelines on how to review papers. most readers can various sub-rou1lnes, the most important or which were
become very discriminating as to the value of most pub- animal growth, cow fenility and mortality. The model was
lished studies. ln Table 9.11 at\ outline of I.he criteria that used to predict the productivity of beef production systems
need ro be considered in evaluating obsenrarional studies. ln Botswana. t\lthough it permitted a very detailed analy-
and in Table 9.12 an omline for the evaluation of cllnkal or sis, it was correspondingly very demanding in its data re-
field trials, are given. These checklists have been developed quJremems. The model has not been used to assess the
and refined O\·er many years \,ith a number of colleagues at effects of disease on productl\'ity.
the University of California, Davis. USA, and the University i\ herd s.imuJation model was de,·eloped by lLCA to pro-
of Guelph. Canada. vide the user with an array of potential policy options. such
Given that most information is imperfect, persons em- as for weaning. breeding. milking. buying and selling live-
barking on a large-scale review of evidence on the impacts stock. so c.ha1 herd performance could be evaluated under
of a disease and its control will probably wish to consider different producllon reglmes. 1111 A funher model at ILCA,
information from as many sources as possible, but to originalli• designed 10 assess inteiventions of fodder cultiva-
weight the information sources by the strength of I.he evi- tion and feed supplementation in Mali and Niger, has been
dence provided. Meta-analysis of multiple smdy outcomes developedz.16 and subsequently modified to assess animal
has been widely applied in human medicine to S}llthe!li7.c health interventions. specifically the economic impacts of
the results of multiple studies.50• 213 There are only a few chemoprophylaxls to control trypanosomosis in the Coast
examples of formal meta-analyses in veterinary srudies. 24 Province of Kenya.100

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 llwcomrol nf infectious disease. or lin•,1ock I 9i

Other approaches to herd d>,,amics modelling have general!~ incorporate some aspect of socioeconomic data
been taken 20 · 144 and models of the herd strocmre and dy- collection, and chis becomes increasingly imponam in
namics of can.le in the tradi1lonal sector of Zambia have smallholder production systems wherii li\'es1ock activities
been developcd. 1-14 These models were used to assess the ef- are less commerciali1.ed. While the emphasis has been on
fects of val)rjng cah1ng rates. and calf and adult mortality on epidemiological merhods. !.here also exist a number of de-
herd structure and dynarnics. sign Issues and me1hodologies specific co socioeconom1c
For in£ectious diseases. mathematical models assessing data collection o.nd a1,aiysis. Refer to Poate aJld Daplyn 19~
transmission a11d infectiou dynamics have pron•d ,·cl)· for an oven·iew.
useful in assesfoingand predicting the impact of changes in Two pracucal poims are worth stressing. First, deciding
key risk factors and the impact of vaccination and other the sample size for socioeconomic data is more likely to be
control strategies. The mai11 concepts and major applica· based on a rule of thumb conditioned b} logistkal con-
tions for irlfectious diseiise modelling have been set out by strnints rather than on the standard formula. For epidemio-
Anderson and t-.la\'' .
. in their authoritative text on che sub-
Ject. The essential feature or mathematical transmission
logical studies. the focus i, often on a ker parameter of
interest, \\'luch drives the sample size. In socioeconomic
models Is quite simple, particularly for microparasites (e.g.
,-iruses and bacteria). Transmission persists if each case
Tebte 9.11 0.;tt;r,e for me e,a ' ,a,,011 c' obsel\a,,or.~I $Wd1es
can produce one or more secondary cases. The rate (or
ratio) at which primary case~ become ~eeondary case~ h Were the ob,ect,1es of the study clear!, stated? Yi?/N
known as the reproductive rate of the disease, measured as Was this a elate soooping/prelim,na:y s:udy? \Relates :o Yi?/N
eitherthe effective reproductive rate {R) or the basic repro- mul1,o!e comiarisons and in1erpre1auon of resultsl
Was the stutlygioup llkelv to !:e ,eo:esentative 01 a la:ge, Yf!/N
dvctlve rate CRq ), and transmission persists if 1hi:- rate is 11arge;, DOOulatton? (OnlvtOlltemed with extrapolat,oM)
greater than one. The basic reproductive ratio represents r'/lle o· s,e:ly7 CaSl!·tor.trol JI. Cohon I Crcss-sec,,cnal I).
the reproductive potential of an infection while the effec- Lo~;tod1nal I l. Hyb11d ( 1
tive reproductive rate. intluenced by ~everal factors. is less
GROUP FOAMAT,ON
than or equal to the basic rate. If by circumstance or con- Ware exc,us,0~ 1,~cius,on crner,a well speciiie.ci? Y;'I/N'
trol effort the reproductive rate falh below one, then trans- Was sample size considered a {)flar/1 Y{//N
r.1lsslon ceases. The basic reproductive race can be Was the F- I0-1 <;•oup well defined? Y,'1/N"
estimated in a number of ways, including: Was the F-· (0-1 g·oup !tom tile same source? Y,'/N
• as a function of the number or density of susceptibl~: 5 Was ,~e :. 10·1 g·oop well def,ne.11 Yi'.IJN'
• as a function of the birth rate. average age at infection fOLLOW-UP/P!ACEBACK
and the duration of maternal immunity~· or Was :he fcllow-~p nrace!lac,l ~erforrr,ed
• directly from outbreak data in closed populations when al compt:1e~,., Ytl/N"
bl b ,nd :or· !D+J status? Y,"1/N
secondary cases due 10 a primal) initial case can be de-
Was response bias w1th1n 1ole:aole i,mits? Y,"//N"
termined.42
OUTCOME and ANAl YSES
:\ variety of Influences on di~ease tran~mission can b41 c~ti· Was th outcome D-(F+I sraius Y11/N"
well uetmed'
mated via ll11 • Of key interest for control programmes is the
deflneo ,r,tepenoenu, 01 F• 10-1 siatus? Yi'I/N"
critical proponion of the population 10 be immuni,ed If tesiing a speci11c t1ypo:~e,:s, were canfoUnders ce•,ne.: v,"?11'
Ip,= I - {llll0 )).' R0 can also be e\'aluated for other comrol apno1r?
options. Was confounder control auequate7 Y,1/N"
Mathematical modelling approaches have been applied to Were tl11lre adeq Jate efforts to control/explain poss1llle Y,'7/N
a number ofinfectious diseases. Including brucellosis,11<> ruber- b1ases1
Were approp:;a:e analyses done? Y/'.1/N
culosis.'!lf.. 2" 1 sheep blowfly infestation.'" F:v!D. 12~- t.S, lll!l. ~
2:,· rinderpest, 102 211 rabies,r, 111.,12. 1'19.24, theileriosis..o. 1,s. 1.,
Were ana 1•1Ses done !apparently! correctlv? vntN·
Wat powe· ,<el~ l<i be adequate? vnfN
heanwater. 11>11· 1ij9 fasciollosls, 71 • :-3. I;"? ommagiosis. 72 17" and Ir useii. was '!IU1bple testing managet1 exola1ned •:.. ell'
1
Y(I/N
uypa.nosomosis.jjl). 117 ,ia. ir.o.2011 IS ther~ ev,ce~ce ol a dose••es~anse re!ationship? Y,"1/N
ln general. mathemarical models have been very useiul PRESENTATION and CONCLUSIONS
tools for assessing many infectious di•ca5es, 111th most dis· Were tile Methoils presented ,:ea1lyi Y(!JN
ease 1ransmission and impact or mujo1 control smllegilll> Were tl18 Resu/ls aoo Analysis l)lesented clearly? Y1?/N
reasonably well predicted by relative!} simple models.' Was the Discuss-on summarized clearly? Y;?IN
Are the authors conclusions supported by the data1 Y,'7/N"
Key socioeconomic information required Was the s:~av ctes,gn appropriate to araswer the Y,1/W
Object,vesl
In most cases. socioeconomic factors will be a critical com· r~e rn:>fe .,. and "N"s'. 1/1e .ass ·,alid the stlllt;
ponent for understanding tl1e impact of disease and its con- • inc cat;;; aserious prob'er.: rt r.o'.
uol. The epidemiological investigations described abo,·e

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198 ,ocr.,,~ ..,,,: Aspect• influ~ncing the occummac, oi Infectious di~(>(l~

Table 9.12 Ouuire fo• :Le evaluation~; erini~I :iials process has been highlighted by l.ee er al. 116 Since man~ of
che typ~ of data collected are standard {e.g. fanner charac·
OBJECTIVES ANO EXPER1MENTAl POPU1.AT10N
v,,,t\ teristics. livestock management practices, and expcndi111res
Weie the ob1ec; \ as o' ~·.e trial clear•v s,;,teo'?
Was the trial grc.;p ld,ely robe represllnlati.a of a largai on healih inputs). it is important therefore to review and draw
ttargetl pooulwon? iOnly contemeil w•:": awapolauon.J Y7,N upon existing proven questionnaire strategie, from standard
socioeconomic ~urveys and borrow as needed rather than
GROUP FOAMATION
Was mal s,ze fi.e ·r.·1 cons,aered a pr,orii trying to reinveni rhc wheel and repeat past mistakes.
Y/11N
Were e~clus,011/intlus:on cmeria well sped,ed1 Y('l1r,i:· Allowing adequate time ior pre-testing questionnaires and
Wha1 was the stated metnod oi alloca::ng thorougr training of enumerators are borh essential for
·.reatments - - - ensuring quality data.
Is tllere evidence to.at formal randomira:ion was actually
used? Y(I/N'
Measurement of other indirect impacts of disease and
Were the 11ea1ment and procedure!sl wall :!ef,ned' Y/1/N
What was the ~ompar,son groupls)? 1uck anal their c:onrrol ,\Ji noted a bove. animal disease and its
other treatmems - - - control generate a wide ra nge o r' other outcomes beyond the
other leve!s oi same treatme~:--- ob\ious direc1 impacts on li\·estock and human health. Both
no treatment--- productivity gains and expenditures on control have impli-
FOLLOW-UP PEP.100 catlotis for farm and national-level income. Secondarr
Was adequa;e follow-up performed. economic ourcomes such as these a1c discussed in the
al complete!;" follo\,in,: section on C'conomic approaches since their
bl blind to treat!llent s.aws7 ·rt' N e\"alu:ition often wrvcs as the basis for decision-making
Were !Ila managers a:!eq11o1e!y blinoeo about :,ea?.ment1 Y/1J'N abouc control. Within thi.> framework of such analyses. inno-
Was compliance v.,thin adequate lim,,s? VflW
\'Btivc approache,- often must be dC\·eloped to address
OUTCOME ANALYSES ~pecific indirect Impact, \\11cn evaluating a potencial
Was the outcome{sl well dehned' Vfl/N" \'atcine again~! bovine 1rypanosomos!s. for example.
Assessed indep9nden;Jyof treatmen· stau.s' Y(IJN'
Kristjaason el al.· 11 a11cmptcd to value the lost production
'vi/as the outcomeislapprocrrate lor the staled ob1ect1ves' Y/l/N
Were there aoeq,,ate e!forts to controltexpra,n possible Y/l/N po1en1ial-cat1le production a1.1ivities not undertaken due
biases1 10 the di~asl' - that could be recovNed by the introtluc1io11
'Nere approoriate aoal·,ses done lie. consis1ant within of a \·accinc. They e,timated this value by assu ming rhat
design}? cattle densitiC$ in t~eise area~ would rise to equal d1ose
Were analyses aone 1a0paren1ly) corrwly1 V/7/W found In non-tsetse areas. and in this manner predicted the
Wts power li~-el1to !!e adequate for main outcometsJ? V/7/N
ls there evidence o! a dose-response relationship? Y/11N added value of production.
Indirect impacts of epidemic diseases. in panicular,
PRESENTATION .:.NO CONCLUSIONS may be much more se\·Crt' than the direct impaccs. Effec-
Nere the Meihodspresented c!earl1·' Y/'!iN
y,1,t.; tiYc control of dhC'aoei. that restrict international trade can
Weie the Resu//s a1d Ana/1-sis p,esemed ciear,y1
Was :he D1scuss1on summarized ctear11·' vn1N offer access 10 new. higher-1"Ult1e export markets for
Aie the author s conr.lusions supoorted b1• the d.ta? V(//W !11•es1ock and their products. generating added \'alue and
Was the ttial design app,opnate to ansv,e· tne Obiect,ves' Y/?1N' emplO) mcnt not only within the livestock sector, but also
The more·?· ano 'Ns". the less l'llhO l/le :nal. rhrough rht• rt'sr of rhl' nMion:il Pc-onomy duP 10 rnulriplier
• lno,cat1!S a ser,oos problem if ·no·
effects. The substanrial impacts that an outbreak oi F\10
would create in :\trstralia in terms of lo~s of exports and
secondary effects beyond the tive1>toc!: sector have been
studie.~. there Is usually no single oarameter of interest. but dcrno1is1rated.fo11 The 2001 outbreak of F;'\lD 111 the UK
rather a n umber of potentially innuential factors. The rule of highlight~ the magnitudes of these impacts, including
rhumb. therefore. is to rry lO have sufficient numbers of re\'enue$ lost to tourism. Mullins t'I al. 156 report a similar.
units o f a nalysis (e.g. farm ers, households and vlllage~l for more modest. effect of rsctse comrol on tourism in
each major subgroup by which the sample has been strati- Botswana. though no anempt is m ade to value the impact.
fied, to permit ~tatl~tical comparisons across groups e.g. In each case. the measuremem of indirect impacts will
t-tes1s). !>1)cial scientislS tend 10 be comforrable. therefore, require the development of appropriate economic models
with a sample si7.e of:I::; to 30 units of analysis per subgroup. adapti,d to the ~pedfic impact, once the impact has been
Second. r<1searchers often u nderestima te the care that is proper!) identified
needed during the design and implementation of insm1-
me111s (questionnaires) for socioeconom ic surveys. Ques- Geographical information systems
tions must be appropriate!~ fonnulated and asked if they arc fhe repor1i11g and anal)~ib of disease occurrence in a spa-
to solicit effectively tlle information sought by the researcher. tial comexr can be a useful tool in impact assessment. For
The error that can be im roduced simply in die tnm,;lation years, large map~ peppered with coloured pins denoting

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 The ,omrol ofl11kctlou• di$ease< of ll\·<'-tol'k 199

che sources of reports of diseases and rnc1ors have adorned imponam fn lho differemiaLion of dairy. pastoral. mi~cd agro·
1he offices of veterinary departmenis 1hroughout •\frica pastoral and ranching sys1em$. for e)(amp!e.
With the advent of computer power. 1hese have dc,·eloped Gi\'cn the fnadequacl£><1 of available da1a on the distrlbu-
rapidly in10 geographical information systems (GIS). Wha1 1ions of host populations. some attempts have been made IO
do 1hey do that the pins did not? model their distribution~ U$ing ClS. and mo~1 no1ahie is the
Geographical Information sy,;tems have computerfwd pro- work of \\'Im ,•r al.:;~ These authors used a combination or
cedures to allow input. storage. manipulation. anaJy,;is and available data. including 1hose ru.-semblcd b~· ILRI. and mod-
displar of ~palial dam.~-o-They ha,·e been applied to the repon- elling techniques (Figure 9.6
lng and srudy of man~' infectious diseases, parli<:ularl~ the Similar to data on the distribution of host species, al"ltil-
vector-borne infections in .'\frica.'11 \\'e do not intend to cover able spacial data on animal diseases in :\lrica an: in the
this huge area in any detail in this chapter. but will surnmam.e categories of recorded obser\'ations and model predicLions.
lhe main fea1ures of this useful mol. Gi\'en the inadequ.icie$ of many dlsease·reporling systems.
Spatial data require that a location be assigned IO the there is no continemal distribution or 1he main diseases
recording of a disease event, in the form of lati1ude/longi- based on recorded ob~enations. f-lowe1 l!'r th<-')t' do exist for
1ude for example. This then allows this piece of data to be se\'eral coun rics in the condnem. :\ notable example is
nggregated wi1h other data describin~ lh!! e1wironmcn1 in Zimbabwe. 1,hkh developed a GIS capacin in ch!! Depart·
which that event occurred. Common!~·. the fir51 step b to mem of \'etcrinary ScJ'\ice~ fn the late I 980~. Thh unit
express the disease occurrence in relation to administrn· produce, distribution maps of disease outbreaks and of
rive boundary data (such as province. district). and this is interveniion programmes (such as vaccination campaigns
how diseases are generally reported. However. while thi~ against :'\ewca~tli' di~ea~e) at regular imen·ab, which are
serves to locate where a disease lucident has occurred. it used to keep scakeholdcrs Informed. but alsu 11s feedback
does lirtle co e\'aluate disease risk and impact on produc- co veterinary staff to encourage bencr disease reponing
ti\ity and other parameters. Therefore. the nexi set of data (Figure 9. 7).
commonly included in any spatial reporting of disease Is Considerable progress has been made using (;JS for pre·
u~ually agro-ecological zone. kind use CUlegory. \'egetation dieting the distribu1ions of disease ,·occors. '.\losl notable in
and communications infrastnJciure tsuch as roads:i. Re· this field is work on the distributions of ticks and 1se1se flies.
markably. in most African countries, these types of data are For tho latter, pioneering work was done b} Rogers and col-
now much easier to acquire. generally at quite high levels leagues at Oxford.2 0!1 This has recen!l)' been followed up by
of resolution. than data on animal dbcases or on the host. McDermott et (I/.. "11 ,_.ho u~ed models of predicted climate
they affect. II ts also possible to access, through 1hc Inter· change on the length of the growing period m predict how
nee and other sources, high-4uali1y satellite-derived da1a the distribution of tsetse speC'ies might change over 1.he next
on climate and vegetation cover. 50 ye= t Figure 9.8).
In many African coumries. GlS units ha\!e been e~tab· The use 01 GIS hal< also ex1e-nded to mapping disease risk.
lished within the canogrnphy dcpanmems ofgovernmem Considerable effort was devoced to predicling the current
ministrle:. and planning dcpartmen1s that have assembled and potcnrlal dis1rih11tion of ECf and it~ vector. the brown
much of 1hc natural and infrastructUral feature$ of lhcir ear tick Rhipi<:efJll(l/us appendic11lm11s). In ,\Irica 1hrough
countries. However, there arc relatively few thm have es- 1J1e use of climate matching models. ' 20· 1g6 1"" but more re-
tabli,hE'd the ~=e degree of capacity in the li\lestock pro-
duction or animal disease sectors. On a contlnemnl scale.
cently the focus has been on predicting risk of the disease
through the Incorporation of a wider set of \'ariable~ includ-
the lmernational li\'estock Research lns1imte (fl.RI) de\'Cl· ing r.be satellire-dcri\'cd vegctadon indices. This has been
oped a map of cattle densi1y for 1he continent of Africa done both ata country level !in Zimbabwe1 5 " and at a conti-
(Figure 9.5). In doing lhfs, II.RI used the most recently nental ll!\'el'~ (f'i{,'Ures 9.9 and 9.10).
available data at t.h e highest resolution from publbhcd
sources. The result combines ,ornc dcnsitie~ that are at Combination of models, observational and
secondary admlnistrarive boundary level, and possibly 1O ell.'J)erimental studies
years old, with new data at a ,·cry high le\·el of resolution tn the abo,·e sections an array of tools chat can be used to
(such as Zimbabwe, with caulc data down co the dip tank determin" tbe impact~ of diseases and of method, LO con-
level. revised and published annually). rrol them have been described. More likel~ than not. no
:'\Qt onl)' are there difficulties \\ith the quality of dilta on single t0ol will suffice in pro\'iding the an,wcr. As a rc~-ult,
number;; and distribution in space for a given spc<:i1$, bill should resour<.'C~ be a,·ailable, a combination of 100b can
when it comes to impaet assessment. these lil'f!Stock dlslribu- be used. and the resuh, normally r1>nder more 1han the
lions do not differentiate between the different production sum of the component tools included. Epidemiological
systems in which any one species is used. and ~ the tools ha,·e played a particularly imponanr role in ~tudying
different impacts that animal diseases and their control will the d}11am1cs and impac1 of tick-borne diseasc~. 182 A
have. In the case of the cattle example. this will be particularly practice we have advocated in several i1we~tigations has

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200 ,cc:no, 11,r, Aspec1s i.Mluendng 1he oc<:urrence of infec1ious diseases

... ,, '
..

.- ""~
# _ _ _ _ ... , . . . . , ,

.
.
,,.....
... ...._ ... .... _
,'

,. -- ... ,- .. l

~,
~ - ....
.

- -- . ~ \

\,
'

0
1-5
5-10
10-20
20-50
5o-100
> 100
Nodaia
Water body
Natio11al par6

Figure 9.5 The d1stribu11on of cattle density mAfrica (source: Kruska et al. 1995111)

Major rivers
C3ttle per sq, km,
0
<10
10-24
25-49
= 50-99
a >HlO

Figure 9.6 The preificted density of cattle ilrom Wint et JI. 2000';01

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 The control of iniectious diseases of lin•stock 20 I

./,. --·.r..y
.
Provmt:,a! centras

D*
0
FMO 0,101eai:s
FMO.lree area
,utcpe 5 2S'OSID2

Sobendle 28/08/1)1
;
( '.r~-.
\ ··--;r,

D Proposed expon zt)(lB to RS A I

•
D
D
Wlldhie conSe!\'aocies
Buffer zone ,._ . .,
Surveillance rcn£: '\ .....- -••
D G3me parks -.
D FM0-Quarantlne11 areas ~
·1
••

OistntlS
\.,.
""\
.·-·.....
.
.. ...._,_.. '·

Figure 9. 7 An example o' Output flom the GIS unit at the Oepar1111ent of Veterinary Se'Vrces. l,11bab\~e In this case 1he loca1ion of foo: ,and-moutll
drsease outbrea~s ftam August 2001 to October 2002, in thecon:ext o• the CXJuntry's disc..ase control zoning. Is 1llustrated2$l

been a combination of cross-sectional and longitudinal Infections. ~tajor gaps in lmowledge. and in parameter
(usually cohon) studies. The cross-sectional is carried out values. \,·ere identified. and these formed the basis for
initially. stratifying the population by key variables 10 deter- the objectives of a serit>s oi experimental and obsen•ational
mine major differences in prevalence \\1thin a parent popu- studies. E.-;perlmentnl studies were used to investigate
lation of animals. farms or households. followed by a principles and. in some cases. quantify infection processes
longinidinal study to determine incidence and impact in the under experimental conditions. and obsen·ational stud!~.
contrasting prevalence populations. Such was the design of principally longitudinal studies in clearly defined areas,
two sets of smdies on the impact of ECF in cwo difl'erem re- were used to determine parameter values under different
gions of Kenya: the cenrral highlands75 • 76• '17. 78 and the ecological and fanning systems condition~ in Zim-
coastal region. 1sa 13~- 110 Both of these series of studies l)abwe.191. 192• t 93 • 19·1• 1~" Concurreml>·· surveys were set up
ended with robust estimaces ofinfection prevalence, disease to characrerize the differem livestock production S}'Stems
incidence. case-morbidity and case-fatality in the different in which hearrwater occurs, and their economics. 38 JS
agro-ecological zones, cattle types and feeding manage- Armed with the results from this wide range of studie~. It
ment regimens pre\·alent in the study areas. The studies in was back to the models. The first rask was rouse the models
the central highlands also derived impacts of infecrlons on to predict hearrwater incidence and mortality under the
daily weight gains. :'\either series progressed into e,·alua- variety of production systems in the country. based on the
lions of economic impacts. outputs of the cross-sectional and longitudinal studies.
In another series of studies. carried out on the impact of The second tasl:. was 10 use these estimates 10 calculate the
hearrwater and Its control in Zimbabwe. two additional economic Impact of the disease, and of different control
components were added ro rhis framework ror impact scenarios contemplated by the Depamnem of Veteri nary
assessment. The first was a mathematical model of the in Services. i:1;
fection dynamics of rhe disease, which served as a starting These economic impact studies were subsequemly ex-
point for the entire study (Figure 9.1 IJ. 168 Using initially a panded co include the other countries of the Somhern :\fri-
conceptual model framework, the major parameters and can Development Community (SADC) region. 151 although
pathways in the mainrenance of the infection were pro- wtth the large region and multitude of production systems
posed. and supplemented where possible by parameter invol\'ed. considerable reliance was placed in this smdy on
values in an a1tempt at understanding the dynamics of secondary data. Finally. ft was back to the models again to

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202 ,, , 111>, '" · Aspects influencing 1hcoccurrcnce of infectious dlses•~s

evaluate how new ,<acdncs under development might work cn,ironmcnlal pen.pcctivc. Then an overall assessment of
under 1he differem control scenarios comemplated. 169 both risk and the unc.:,rtainty of the risk-assessment proce,,,, is
These example~ tllusrrate tlw need for a good conceptual made b} combining the fik('fihood and con~ucnce analysc~
framework behind impact assessment studies. rhe impor- with judgernems on !lw c.;uality of data a,·ailable. Once a risk
tam role of good epidemiological data in economic impact assessmem ha~becn ~-onductcd, it~ re,;uhsneed tobecommu-
asst-i;smem~. and lhc advantages of combining Im·e~tlgatiw nkmlfil IO and discu~ed \\ ilh all slakeholdt:1» ~o Lhal it ca11 be
tools for ma'(imum effect. incorporated into rn-k-managemen l practices. ,\s di.o;ease con-
trol and food saft!t} become incrca.~ingly more complicated.
Risk assessment formal and iniomtal risk assessmem and management" ill be
The introduction. spread and control or diseases aU de- an imponant feaiure of decision-making In animal disease
pend on a sequcoce of biological evems 1ha1 are uncenain. COll!TO).
Rules regulating tmdc in animalsrutd animal products (e.g.
the sanital') and phyt.o-sanitar~ iS?Sl regulations of 1he Measuring the impact of zoonotic diseases
World Trade Organizaticm (\·VTOJ/011:. and decisions re· on the human population
garding what disease control and food safety strategies Within human hcnlrh. 1he disability adjusted life year
should be adopted all depend on assessing and managing (DAI.VJ is an instrument recently developed to alto,, the
disease risks. Hawn.I Analysis and Critical Control Point relative burden impo<ed on humans 10 he cteter.nined. rhis
(llACCPJ methods used in indU$lrial food safe!) is one ex- was used as a common health measure in the first Global
ample of risk analysis methods that have been widely Burden ofDbease swd~ commbsionecl b~ the World Health
adopted. Research and development in risk analysis, both Organization (\\1il) 1"" that documemed the human health
technical and policy issues ha\•e greatly expanded in status. worldwide. for the year 1990. Ji bas subsequent.I)
recent ycars.90. 137 1a\.1,i;, 2 i9. 2;9 been u~ed in re,i~ed global disea.e burden estimatcs mo,t
A risk-assessmem process for pownrial hazards (such as recentl) in lhe WHO World I lcalth Report :woo. in which the
disease lnrroduction) and methods to conuol them needs to health si.uus for l999 is described.:!.>"'.!
be considered by \'ariou~ categories of decision-make~ in ani- ,\5 well a~ being u.~ed as a means of ranking the ove.rall bur-
mal heaflh. This should be a formal process but ma) be ei!l1er den of indiVidual disi::ase, !lie DALY is also particularly useful as
qualitative or quantir:icivc. A kC)' i11itial step ls to identify and an omcome measure in economic evaluations of inren·en-
characterize 1he potential ha;r,ard both in terms of its cause.~ tion~. including l-O~t-effectivcness analysis. 60• nr Consem,us
and co~equcnces. The lik<;>lihood or a ha1.ard (or hazards) oc- cost-effoctivcnes~ ratios or US $150 per O,\l.Y and US S23 J:>er
curring is then cstimmed. The impact or consequences of haz- DALY a\'ert<·d are agr~-<:d lhrcsholds for di1;ease comrol inter-
ards then need to be a-se..ssed from a biological. economic and , ~mions to h<' con~idered ·anraeti\le· or 'highly atrracti\·e·.

D Correct a:isence D Ne change· allSent

D Correct presence D No change presem
• False presence a Present to absent
• false absence • Ahenl to preseN

Figure 9.8 Pre1,cted diswbuuon of the Gloss1mi morsitarsgroup of tsetse flies compared to known d $tl•Dut on llefll. and preaic: or.so: how tlus r,,10!":
oeaffected bv climate tha~ge 10 the vear2050 !from McDerm:m er al. 2000' 3 l -

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 lh<' conuol ofinfocnous diseases of Uvesiock 203

Zo~as Est,mated 11ur:ibe·

o1anima s I OOOl

1an1
Jr.creasing ris,
of disease based
0
1
2
H.J 4 253' /t 03S~
2 028 12 371
on probab hty of 3 1 OS9 2 SJ~
vector presence 6.06/" !F,i
4
5 1 i45/ZGSS
6 • 41 702 /3 3-l5
~ Coun:ries in :ne e~e,n,c region
b Countries po1ent·i , a: ns,:· Vector
suuabil1:y llut abSence of dlSease

Figure 9.9 The predicted d,str.buoon c· ris~ of East Coast fever 1te~m lvhnJauw e1 e/. 2001.15 u~puo1:shed data.152 and based on Cwnmmg, 1999:'l

Ou:ilreak
probab11t1y
• <0.05
• 0.05-0.1
W 0.1--0.15
• G.15-0.2
• 0.2-0.25
•

-- 0.25-0.3
0.3-0A
0.4-0.5

-• 0.5-0,6
0.6-o.7
0.7-0.8
>O.B

Figure 9.10 To.e predicied a:striOu!IOn of risk of rheilenos,s n Zimtabwe lfrom Ouchaieau er el 1997;,sl

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 /
.,,,----'-----------
, ____ ,,. .,
•I 8. T. qA. qN ) - •• - •• • • - TICK
\

.. .·· DYNAMICS

,, '
.'

Innate Maternal
proteciion
p

Figure 9.11 A model of tM infection dynamics of Ehrllch,arummanrium infection Ifrom O'Cailagllan e1 a/., 1998'..;,

respec1ively, in a de,·elopingcountry setting. and used co ad- elits. which gives a lower weight to health benefiis that occur
vocate onesrrategy o,·er another.2 s 1 further into the fuwre. ha~ been criticized as inequitable4
The DALY is a measure of healrh outcome that incorpo- and has auractod considerable contro\'ersy and debate. Age
races both premature death and morbidity/disability. The \\'eighting of 01\l.Ys is also highlr controversial and should
DAl.Ys caused b~· a given disease consist of the year~ of life be avoided as no consensus on the use of age \\'eights hns yet
lost CYLI.s) and the years of life lived with disability (YLDsl. emerged. and the impact ofrhe.ige weighting func1ion used
In quamif)1 ng the cost-effectiveness or an intervenrlon. in the Global Burden of Disea,,e has been shown to be
death$ averted due to the implementation of the Interven- cotmter-intu!tive:11. i;s
tion are convened to YI.Ls using the age-specific life expect· The use or a gene ric measure such as the DAL\. rather
ancies found from appropriate life tablel>. In comparing the than intermediate rnea~ure~ of benefit$ such as cases
DALY burden across different settings a common life table is pre\'ented or cases cured. allows for comparison of pre-
used {e.g. for the global blUden or disease sLUdies a Japanese ventive and treu1ment interYemions. Moreover, as DALYs
life table with a life expecmncy at birth of 82.5 years for incorporate both m()rtality and morbidity/disabilit,. it is
females and 80 years for males). For cost-efl'eclivcness possible to make comparisons wirh other intern>t1tions that
Sludies. comparing different interventions in one seuing. a improYe the quality of lire. as well as with those that ~ave
local life table should be used. 159 livt:s. Whilst several limitation& or DALYs have been identi·
The years of life lived \\ith disability arc calculated based fled in recent critiques:· 21 58· 1;;;, 24 ~ they do have several
on the durar.ion of disabiliry/morbidily. and disabiliry imponam advantages. particularly in cos1-e!TectiYeness
weights given 10 each condition. The weights ranged between analysis,ut :md their use facilitates comparlson 1,ith the
O and l, with O representi11g full health and I representing results of other recent cost-elfectl\·en1JSS studies in develop·
death. The weights for each condi1ion used in the Global Bur- ing coumries.1rri
den of Disea.se study159 were derived using the person trade- Official global ~timate<- exist for 27 infectious diseases of
off method developed at an intemarional meeting held In humans. with the most recent ,·alues calculated for 1999:152 as
Geneva in 1995. For the ,-asr majoricy of zoonotic diseases no sl10\\11 in Table 9.13. Using a strict definition of documented
such consensus of disability weights exists. natural transmission bet\\leen animals and humans, 20 of the
The YLLs and YLDs are summed to give the DALYs score. 2i dlseas~ are classed as 7.00noses.l.3t> HowC\-er. from a con-
The DALYs may be discounted to account for time prefer- trol perspective. targeting che animal reservoir is unlikely ro
ence and the risk premium. The discounting of health ben- have a ~lgnificant impact on the burden of human disense in

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 The com roi or infecuous diseases ofllvcs10ck 205

13 of these 20 strictly defined zoonoses (Table 9.13). as the sertificatlon. deforestation and global warming. The conuol of
amhroponotic transmission cycle (from human to human) li\'estock diseases. particularlr large-5cale disease control ef-
is relatively much more important in these than any forts. have been the subject of debate from an em,jronmemal
zoonotic cycle. For the remaining seven zoonotic diseases impact perspective. The greatest environmental concerns
- trvpanosomosis.7·1• 2011 schistosomosis.235 leishmani- have been in the large-scale control of iserse fly and u11>anoso-
106
osis.11' Chagas disease.&l Japane.,;e R encephalfris. 11 ~ mosis in Africa. both the indirect efl'E!<'.rs on habitat and wild-
hool.."<vorm 196 and hepatitis (caused by hepatitis E ,'iros)226 life 174 and the direct effects of chemicals used £or tsetse fly
-there if> e,~dcnce suggesting that the animal transmission control on non-target rJrganisnlS {including people). water
cycle is important and that \'eterinary incervemions could quality and nmrient cycling.161
ha,·ea profound effect on the global 01\LY burden of the dis- A number of studies have assessed direc1 impacts of
ease. However. it is debatable whether any of these diseases chrunical~.s4• .;s. ID'J In general. newer insectkides and rhe
falls in the most Important category of zoonoses. in which methods by which the}' are applied are considered to be less
the animal host is the sole maintenance reservoir of human persistent and have no long-rem,, irre\•ersible effects on non-
di~ease. and where blocking the tran~mis$ion between ani- target species.221 Howe\·er. risks always exist for personnel
mals and humans will eliminare the disease In the human handling chemicals. Rola and Pingaliun provide a frame\,•ork
popuJation. The conclusion is that DAJ.Ys are yer ro be cal- and examples for assessing the economic Implications of
culated for many of the Important zoonotic diseases of im- such ri$k in a related application for crop pesLicides. Another
pact in ,\ frica. potential impact is the development of resistance to pesli·
cldes or drugs. such as in the c:ase or rrypanocldes In sub-
Measuring environmental impacts of disease Saharan Africa.69 Fleischer and Waibel 63 describe an
and its control approach based on natural resource economics for assessing
Livestock ha\'C been considered harmful 10 the environment the social "alue of resistance for crop protection in Germany:
and implicated in a host of emironmemal sins including de- the approach has yet to be applied to the field of animal

Table 9.13 Burden of mfe1:tir:ius d,mses m disability-adjusted life year$ !DALYs) by cause in WHO reg tons. esiimates ;or ·99S. The zoonotlt sratus o!
each disease ,s 10dicated (from WHO 2000'~1)

CAUSE GLOBAL AFRICA THE EASTERN EUROPE SOUTH-EAST WESTERN ZOONOTIC

Ai\o!ERICAs MEOITER· ASIA PACIFIC
RANEAN
HIV/AIDS 898'$ 74A49 28'2 2172 72S 88S6 70·
_:, No
Oiarmoeal diseases
Ma:ana
72063
44998
24321
36838
25-;4
76
10123
2774
i 13i
2
30017
3071
3955
2235
I Yes
Yes
i ullertulos1s 33287 8721 l 11~ 226ll 1258 14101 5832 Yes
Measles 29838 17463 31 3293 HS 8308 59' Yes
Tetanus i2020 303S ;50 1821 i31 5302 566 Yes
Pe!'.USSiS 10905 4937 '89 1415 ·97 363~ &33 No
Meningitis 9824 3620 722 1472 ~33 2557 1005 Yes
Chlam·1d,a 7969 2233 779 250 581 397!1 l.i5 Yes
Syoti1hs 5081 3189 i 13 648 5 2067 59 No
Gonorrhoea 5686 2425 313 ·01 125 675 '"
~:> No
lymphatic ii;arios1s 4918 1834 8 11 0 2788 278 Yes
Hepatitis· 2790 1172 212 109 57 706 535 Yes
T:ypat'Osomos,s • 20,:8 1991 I 56 0 0 0 Yes
leisnmaniosis· 1983 256 50 210 0 1467 0 Yes
Schlstosomosis • 1932 1637 i33 98 15 i8 30 Yes
Pohcmyah.is 1725 279 152 19 1 1095 178 No
Hookworm· 1899 406 173 93 0 862 156 Yes
Traclloma
Onchocerc0$ls
Japanese em:!!pnahns·
'23S
i085
10L6
,~
434

0
0
2
Q
237
0
0
0
62
218
0
505
0
No
Yes
Yes
I 0 828
Chaga; disease· 676 0 677 0 0 0 0 Yes
Ascarosis 505 202 ,:3 43 0 186 31 Yes
Triclluros,s 48' 198 155 3 0 120 5 Yes
leprosy 476 79 60 10 0 318 9 Yes
Dengue 465 24 0 i 0 440 0 Yes
Orphtlleria t51 24 2 11 1 110 2 No

. Disease in which the zoonot1c com~nent ,s considered s,.~1fican, compared to anthrooono,ic traism,ss1on

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health \'a.nous indirect environmental imparu of H'Ctor due to inadequacies in data. It i5 nece..sary 10 articulate and
comrol of tsetse Oy on wildlife, biod iversity and land usc quantify accurately the components and coses of the differ-
have been studied and found not to be lnvariabh nega- cm im1:rvendon strategies under e,·aluation. and further-
254
tive:!!la. ?a,, but ro depend on ecological circu;stancc~ more. what outcomes the} \\ill produce. In both
and people's rnspon,e to reduced disease ri~k. Approachc·s epidemiological and economic term~. ln,ariably, there are
ha,·e yet 10 be adapted from natu ral resource economics for data gaps aJ,d key assumption~ have 10 be made but as
,-aluing these type, of impaets. long as the~ are rea$<>nable. and are well articulated in the
analysis. they need not be an impediment to developing
qualiiy decision-sup port aids.
Identifyin g and deciding between
Secondary dara arc often available from a broad range of
intervention options
~ources, based on the types of information systems or stud·
In this S1!Ction "e focus on tools and methods ior deciding ies described in the preceding section. with considerable
hetween different control strategi~. for diseases judged 10 variation;. in accuracy and precbion. \\1wn lhe appropriate
be of high priority. 1\ much more extensive review of the secondary data are lacking, primal') data collec1ion may be
relevant issues and methods. and from which much of the necessary co properly capture the specific cpidcmiologit-al
material in this section is drawn. can be found in the Au- and socioeconomic consequcr1ce\ c,f the different control
gust 1999 OIE Re1111e Sci<mtijiq11e et Teclmique.181 The 1e_~t- options under considerarion.
book on animal health economics by Di}khui:r.en and Ir is often relatively strnighlforward 10 measure and
~lorrisnis the most up·to-date source for details on how to quanti~ the dlrect impact:, of disease control in ttlrms of
apply many of the standard analytical meihods. the market value of changes in production losses and con·
tr()! coses. ldentlf}'i ng and valuing additional indirecl or
Steps in evaluating disease contTol interventions secondary Impacts may pre~cnt much more of a chall1mge.
Four major steps are general!~ involved In determining the For example, indirect, longer term impacts of lmpro,·ed
most appropria te. technically feasible and economically reproducti\'l" performance may require use ofa herd simu·
,ound conrrol or inteT\encion .strateg). These are: ~cenario la1!on model and a set of as,umptions about ho\\ herd
development; qu;rntlfication of the intervention options structures and off- take rates are expected to e,,ohr O\'Cr
and their cffecis in epidemiological and economic terms: time. Ofi'-sile impacts. such asrhose on the environment or
de,·elopment of clear decision criteria; and e.t post evalua- on other ports of the marketing chain and beyond (e.g.
tion of the results of the decision- making process. market prices. employment. human health). may require
au,~iliary anal)•ses that draw on the economic literature.
Scenario development In ~cenario development the Still others may be 'intangibles· that are much more diffi-
framework for decision.making is constructed by listing the cult lO \'alue. bUt clear!)' need to be recognized. flow. for
intervention options that could be considered and the differ- example. can cbe value of a control option better targeted
ent consequences of such options are de.scribed. The sce· to the poor be quantified compared 10 one targeted to
narios might be based on different geographical expansions commercial producers?
of a disease (such as that described by Mukhebi era/.. l 999. 15• ·n1e challenge i~ 10 assemble information irom disparate
when considering the spread of heartwater in Zimbabwe). sources in a meaningful decision-making framework. com-
different technologies used for disease co111rol (such 8.\o that of bining both epidemiological and econom ic information.
Minjauw et Cll.. 2000151 when comparing acaricide-based and
vact'ine-based strategies for heartwater control in soluhem Development of clear decision criteria This is an area that
Africa), a combination of the two (such as in Lhe ~!ukhebi er isslill poorly developed. The commonest criteria used are just
al.. 1999 1;;; example cited ). or differem lengths of time ror a biological. based on efficacy. Does rhe imervemion success·
disease eradication programme 10 succeed (such as thal or fully comrol the disease? However. as has been repeatedly
Perry <!r al., 1999. 1M when considering future !'~ID eradica· demonstrated In empirical studies, what makes sense techni·
cion in Thailand) . cally (the technical optimum) only on occasion makes sense
There are many important features of scenario de\'elOp· economically. The decision criterion is lherefore usually fl!·
ment. but the most important is that lhcy be realistic. This re- framed in cenns of: does the ,-alue of the impacts of di~ease
quires chat they not only take inlo co11siderarion national control jusi.liy the effort (cost in\'ested? Or. if various option~
policies and aspiration,. but also chat chey are based on the are being considered. which yields the b~t value forthtefiort
best a,-ailable information on di,ease occurrence. interve111lo11 or re-,,;ources (cost) ln\'esred? Translating d isease control in·
effects. logistical considerations and resource availabilitY. tervemions and their impacts i.1110 dollars and cents also pro-
,·ides rht benefit of a <:ommon unit of measurement that can
Quantification of the intervention options and their sen·e as the ba,;is for comparing apples and oranges. e.g. con-
elTecrs This s tep In the process Is often the smmbling trol that impro\•es reproducci,·e performance versus another
block to high-quali~ and credible decision·support aids that improves weight gain in an animal.

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 The comrol of infectiotL< diseases 01 livcs1ock 207

If the l'!conomic criterion is incleed 1he one adopted for special consideration, for if eradication is to be a viable
decision-making. then se,·erul differem measures of eco- scenario, (·cnain crtwria must be met.';2 lhcse are summa-
nomic worth may be considered. The most popu lar include rized as follows:
1he benefi1:cos1 ra1io. the net presem value and 1he internal • effenive tools for breaking transmission are available:
rate ofrc1um-all measures from financial in, es1men1 analy- • epidemiological and population ecological feature~
sis. .-\s noted above. however. these quamira1ive measures of che disease make it amenable 10 eradication !o.g.
will oftt'n need 10 be qualified or supplemented by addi- clinical case~ and/or infections are easily de1ec1able.
tional crllerla for impacts more difficult 10 quamifJ or, aJue, the siw of the area affected i~ 1101 too large and has a
such as the degree 10 which benefits are distributed equil· relatively stable population):
ably among differe111 segment~ of society. • the di.e~e is of-.uflicient ,ocioeconomic priority:
Also affecting the criteria is the level at which the deci- • there is compelling evidence that the cost 10 benefits
sion is to be made. Decision-makers work at all the different ot eradication omweigh those of comrol: and
levels - farm . community. local auth?rity, national. re· • adequate managerial, oper;uional and financial
gional and international. Although ccnainly not the rule. as resources are available.
the decision moves from che farm to the national levl'L the
criteria generally rhangt from financial to economic. Finan- With thei,e criteria, the feasibility. advantages and risks of
cial analysis refers 10 costs and returns associated \\ith cash eradication \·N~u~ control can be carel"ully 1walua1ed be·
transactions at nominal market prices clireccl~ incurred by fom a decision 10 embark on an eradication programme is
the decision-maker. where~ economic anal)'sis broadens made. The most powerful advantage of achie,1ng eradica-
the scope to include non-cash costs and retumi valued at lion is that there may be no subsequent disease control
!.heir 'crue· prices (e.g. \\ichour distortions introduced by costS. depending on whether eradication b global or local:
ta..~es and subsidies). and evaluates the \•alue of contml for in the lattl'r cast!, co~ts cominuc to be- incurred to ,\Void
society n~ a whole rather than thC/ individual user. l'inanclal reintroduction of the disease. However. if eradication
analrsts is therefore appropriate when trying to a~ess fails. con11nu111g dlsea~c cnmrol costs remain and pre-
whether the control intervention will be reusible from the in· ,'ious ,ira<.licati,m programme costs (often substantial) are
dividual u~er·~ perspecth·e - L·an farmers or other, handle sunk and do 1101 accru<' the expected bencfitS. ln addition,
ics demands on their resources and cash now, and bit suffi- eradication programmes ma: di,·en 1unding [rom other
ciently anracrive in terms of finand;,J benefits co merit 1heir more general public or animal health programmes of
panicipatlon? Economic analysis, on the other hand. 11mm; henefit. 81
10 be used when the decision-maker is tmemstcd in the pub-
lic good. cspecinllr if public re,ources are used to support an Evaluating other control strategies
intervention. If eradicacion i~ not being considNed. the nc.xt set of deci-
The choice of criteria. the per$pec1ive of the decision - sions revoJ,es around how best 10 livewit.h the disease. Thi,;
maker. and the particular context oi the disease under con- mean~ whether to treat, to comrol, or to prel'ent (which usu-
sideration rhen inform 1he ,pccific anal)'lical approach or al!) infers 1he 11$e of vaccines. but can also be lhrough
cechniquc co be used to evaluate the disease control chemoproph\'laxis and the use of genetic resistance) . .-\1 the
intervemion. farm level this decision will be influenced mainly by the
type of infec:tic)us. disease. 1he impact the di$!.'as.e is ha, ingor
Ex p osr evaluation of the decision-making process Ir is is likely to have. 1hc (global) avnilabililyof diffcrem technical
clearlr imponam that scenarios de\ eloped. data used. analy- oprions. the local availability of those options. their prob-
:;e,,conduculcl. and symhcs('!: prepared. are appropriate ford1e ability of i;uc;cess. 1hc impact uf ~urces:.. the relaci \'e co~rs of
decision-maker in question. in which case rhc adoption of the different intervention options. the availability of resources
results ~hould approach 100 p<!r cent. However, they rarely do. in the comexc of other consLmims and obligations. and any
One way to avoid the pitfalls of developing d<1clsion aid:: for social or regulator) responsibilities of the farmer to follow
farmers or national stra1eb'Y development that arc not used is certain procedUies with a given disease. At the national
to leam from each analy~i.s carried out. and conduct an et fJ0$1 level, deci~ions will be influenced b) thl' stune factors, but
impact a.~~ts,ment of the CYaiuation. This is rarely don!.'. bLll is rhe assembly oi data and the evaluation of options \\ill be
an area that needs developing if veterinary epidemiology and infinirely more complex.
econo.)llic:s are to ha\'e greater Impact on the quality of deci·
sion-making in the furure.
Approaches for financial and econontic analysis
The special case of eradication A wide ran~eofanalyticnl techniques has brcn successfully
In scenario development the most common framl:!work adapted from the economics litt:raiure for application to de·
compares alternative control opLions. However. occasion- els.ion-making for animal disease control. Before briefly
ally an cradfcatlon ~cenario is included. and this requirt'.s summari1.ing some of the most commonly used. it is useful

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208 "' 111" o~,, A>P<'~ts influencing the occurrence or inieaio~ diseases

to note three features of these applications that distingu ish ing larger-scale control interventions. often invol\~ng public
the field of animal health economics: resources.
• The nature of livestock production, particularly for Comparing costs and benefits is the basis underlying the
larger animals. Livestock are often a production as;e1 rnajorl[) of financial and ec<>llomic analyses applied co an-
and at the same time. the product. Cont rol affects both, imal disease control. Most may be broadly termed as ·cost-
and its benefits may need to be valued from both benefit analysis' in spirit, though we must be careful 1oa1·oid
perspectives. This requires taking into account the confusion since this term is al.so the name of one of the spe-
productive lifespan of the animal. and the multiple cific techniques in this class.
production cycles associated with off-take of livestock
products. For this reason. economic analysis musl often Enterprise budgeting and gross margin analysis
Integrate some type of herd dynamics model that can The~e are standard tools for iarm management in which a
describe how 1he production assets and off-take evolve budget is dc\'eloped for one or more fo·estock production
over llme. activities. The budget details all inputS and their costs.
•
• The nature ofinfecllous d.isease. RL~k of disease and the together with the putp u1s and revenues, so tha t the net
impact of control often are not limited to the individual profit can be calcu lated. Enterprise budgets include both
animal; they may depend as well upon the epidemiologi- variable and fixed costs. whereas gross margin analysis
cal dynamics within the host population. Again, eco- only considers ,·ariable costs. For disease-control deci-
nomic analysis must try to capture this dimension.. and sion-ma ldng. d isease control is treated as an integral
as a result, there have been a number of anemp1s to inte- part of the production process. and so budgets are csti·
grace epidemiological modelling with economic tech- mated for production actl\ities based on different control
niques. This also means tha t economic costs and oprions. Okcllo-Onen e1 a/., 17 1 for example, treat three
benefits of disease control may extend well beyond the cattle herds under different tick control regimes (no con-
individ ual animal. herd. or farm. trol. oncc-a-momh dipping. mice-a-week dipping) as
• The multiple roles of livestock in smallholder forming different production acthities and estimate gross margins
S)'SCems. Economist:. like simple production processes for each. :-tore typically, these approaches could be
where an addedjnput results in an easily idenrlfinble added expected to be used in cases where the disease cpntrol
output. in a commercial li\'estock woduction system. in- inter,ention affects use of uther inputs or the quaUty of
puts and outputs are u,uallywell defined and quantifiable, outputs, 1hu$ fundamentally changing the produ.c tion pro-
though complicated to some extent by 1he preceding two ce~s. such as comparing the u~e ofrrypano-tolerant breeds
points. In smallholder farming systems. as described in. of cattle versus regular chemotherapy with trypano-
preceding sections, livestock serve a number of different ~usceptible breeds.
purposes. many unrelated to commercial sale. ils a result,
the benefits of disease control may need to be ~-alued Panial budget analysis
tl1rough their impact on draught power and improved crop Th is is an abbreviated version of gross marii n analysis.
production. or through other intangible impacts such as Rather than estimate a complete budget for a production ac-
enhanced social starus. Similarly, farmers ha,\'e multiple tiloity, it considers o nly those costs and revenues that change
objectives. so their decision-making may not be a function due to the control intervention to detennine if. and to what
of the cash cos rs and benefits alone. which may Violate tl1e degree. incremental costs associated with the imervemion
underlying assumption of many of the standard analytical (cost of the intervention plus any returns foregone due to
methods. the control) are rewarded by incremental gains (added om-
pm value plus any coStS savings). The a11aiysi~ is usually lim-
Each of these features has considerably complicated the eco- ited to a single production cyde or time period. and
nomic assessment of disease control. and the collaboration provides absolute (net return) and relative (benefi t-cost
between veterinarians. economists, and Uvestock produc- ratio) indicat0rs of investment profitability. With its mode~t
tionists co address these challenges has led co the develop- data requirements. it is one of the easiest and most practical
ment of a specialized field of ani mal health economics.52 approaches. Pegram er al. 177 used ti1is approach to compare
In Table 9.14 the principal analytical approaches used the value of intensive lick control versus no lick conrrol in
for decision-making based on economic criteria are sum- traditionally grazed cattle herds in Zambia. Gummow and
marized. The first set rerers to farm-level decision-making, ,'llfapham 82 provided a well-organized example of partial
and are used ei-rher 10 advise the farmer which control budget analysis enhanced wilh a smthaslic d imension ap-
option will be most profitable co use l)y comparing cos rs and plied to experimemal trial data from South Africa for evalu-
benefits. or 10 u nderstand and predicr what level of control ating a vaccine against Pas1eure//a lwemol)'tica in cattle:
the farmer is like ly to adopt b>' evalua ting the trade-offs in while Kamau e1 flL 10" reported a similar analysis to e,'aluate
terms of overall fam1 resource allocation. The second set of cypermechrin pour-ons to control trypanosomosis in
approaches is better suited for the decision-maker manag- Kenya

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 The Nntrol of infectlou, di,ease, of 1h esiock 209

Table 9.14 Memods for me 1inanc1a1 a!!d economtc analysis c: o,sease and ns control

FARMLML

METHOD OBJECTIVE fEATURES

Gress margin/enterprise Evaluate financial viability by c.cmpanng all costs and - Su,ted to cemrne:c,at produc1.on act,vities wnh
b\lcf9ets revenues for a given producuon ac11v1ty 1ru:orporatiog a rna,~et·'<alued inouts ano outputs
sl)et,!ic a,snse comrol suatsg•1 - fa·!s !O 111tOfllOrate imiiac:s t-evonc tne specific
i:toaut11on actl~11Y
Parua o~cgets Eva!ua1e f,naM,al or economic ,,a~,1 :y oy compa11ng - Smg:e 11me period
1rn::emental costs 1ncor1e:i ano ·e,,-enues genera1ed bv - leas, oemaoding in aaia ceQwem,;n1s
adopting a spec,flc disease control 1mer,,,emion - Muit1p!e ume periods
- Cos: and revenue flows d1scoumed to account for urr.~
Cost-beoeh: analysis !CBAI
value of money
- lndicaiors· NPV. BCR, IAA
~-
Decision analysis Compare the eipec1ed ~et •eturr.s from a process oi - Su,teo' tor analysis of res;lQ1'1se ro ep,o'emic disease
seouential aec1s,on-makmg ar.c proca~11it-:-we19hted cccurrence tnai can cs express.~ .s rsSk probao1hl\', er
0!.ttef'r'eS a,~ase conttol hav,ng m11lt1p:e :x,:.ent1a1 numimes
- M,mics farmer dec,s,on-m.king
- =ails 10 .nto1pora1e secoodaiy ::iµac.s
Optimt2ation models M~1m,1e an O!l1ecrl\la funct,on (ofte~ profnsl, subject to - Usaful for estimating opuma level of disease coot101,
resourca and other co1istraims re,;ardie-ss of wheiher OJl\t1ol 1s 'lumpy or cont1nuo11S
Yanable
- Eg.. linear orogramming, c,ram,c orogramm1ng,
etonometric agricultural r,~"sehOI~ model
Simu1auo:1 models Use a systems modal to ~e·!orm ' ,•Alat-il' scena1io - f·1JJ•callvbased on btoli.g•ca, mocels le.sp., herd
ana!vsis 1,)'!)am,cs modal. li~esioc, ·cr::o mooell
f!EGl()NAUNATIONAL LEVEL
METHOD OBJECTIVE FEATURES
Cost·ber.efn analysis !CBAl Evatuata financial or economic viability by comparing - Mu :,pie time pericds
,~ciemental costs incurred and ·e",enues generated by - Cost a~d reva11u,; flows d1scoun1ec to aet0unt for :ime
aaoptmg a Slletific disease con;rol m,erventioo value ol money
- Indicators NPV, BCR, IRR
- Can b~ extellded :o ·nclude a :.1:ie ,ange of ,mpacts 1
l.".ey can be ass,gned c rr: ar'(e: value
- E,tens,on oi CSA 10 capture impact on market pr,cas anc
Econom,c surptws
d;saibut,onal 1mpatts between producers and
consumers
- f(P:callv used for eval~ating returns 10 research
,n·. estments
Oetis,o~ analysis Ccmpa,e :he exp&ted net returns trom a process of - Su1teo for artaly$1S of resi:oiss; tc ep,demic disease
seQ~entia1 dec;s1on·makmg an~ probab,htv·we,ghted ott11rrence that can be :Koress~ as ris< p,ol:ab,litV. o·
outccmes o,suse com,01 having mu,:,pie ootent,al outcomes
- M,m,cs \elerinary serv,ce ~ecis,on-ma~ing
- ealls to incorporate secoooar1 Impacts
Opt1mizat10n models Max,mne an obJec11ve function foften profitsl. subjec1 10 - Useful for est1mat10g oot,m3 ie-,er of o,sease conttol,
resource and other consmimts regardless of whe1her ccnnol is ·•umDY or co<11inuous
va:iab!e
- Eg li~ear programmtnQ. 01 ~am,c p10g:amm1ng
S1muta11on models a'ld Use a svstems model iO µe:iorm \o,haH:· scer.ano
---- -Can m:egrate a ·iarie:, o! ;:rc:iuction and
system$ ana1V$•S ana!vs:s i)OS:-producuon systel'IIS

Cost-benefit analysis (CBA} of money refers co the idea that S1 today is wonh more than
Thi~ is essencially partial budgeL analysis extended over a St a year from now, recognizing that today"s SI ca11 be In-
longer period of time. Incremental costs and gains are> vested and expected 10 earn interest (In a risk-free sa,'in~
identified for each rime period. and discounting is lmro- account. for instance), )'ielding SI .05 next year. In addiuon
duced 10 capture the time 1•alue of money. The time votu.- to profi1abillry indicators of absolute return~ (Net PreMint

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210 ,, , no, o,t; Aspects Innuencing the occurren('e of infectious disea..~,

Value, the sum of discounted scream of net rerurnsJ and Optim ization models
relati\'e returns (Benefit-Cost Ratio). an Internal Rate of Re· These models use informanon on a household', or firm 's
tum can be estimated that expresses the a\·erage annual in- resource base, market prices, and input-output relation-
terest nne yielded by the investment. This approach ships for its production activities co solve for the resource
is appropriate when the costs or benefi ts extend beyond the allocation and mix of production activities that 1,ill maxi-
current time period. Gittinger,' 9 an authority on mi1.c some defined ol>jecth·e. typically faTm or firm profits.
CBA. offered an extensive explanation of the method a11d One class or models is based on mathrunatlcaJ program-
numerous examples. ming techniques {e.g. linear programming, dynamic pro-
Cost-benefit analysis tends 10 be used mainly for evalu- gramming). Building a mathematical programming model
ating larger-scale interventions. Penne and l)"Hal.'Se, 1711 for is quite demanding in terms of daca and research sophiS1i·
example. conducted a CBA oivarious F.CF control options in carion For this reason. such models arc used primarily for
Zambia. Benefits are estimated using a 30-year simulation regional or national analyses in \,·hich models arc c!e\·el-
model of traditionall~ managed cattle herds. and the results oped for representative farms or herds; they are not practi·
indicate that from an economic perspective. immuni1.ation cal tools for livestock producers 1hemsel\ es. Researchers at
makes more sense than chemotherapy or tick conU'Ol. \\"ageningen University in I.he :S: etherlands ha\·e de\·eloped
Stem131 applied this approach to evaluate a public vaccina- a number of such models for incensivc commercial sys-
lion campaign for pcsre des petits ruminants in Niger. and tems 10 advi~e on optimal timing for disease-related cull-
demonstrated the high economic return yielded by the in- ing and stock replacement actions. 9~· 101 In a panicuJarly
tervention. Similarly, Perry e1 nl. IUl1 evaluated the remms co innO\'ative policy study looking at farrnerincenti\'e~ to par-
F!\10 eradlcntion in Thailand. and found that such a pro- ridpate In \'arious components of bovine tuberculosis con-
gramme would be justified if it led 10 l.'\-port trade 10 high- trol in :S-ew Zealand, Bicknell er ai.2: demonstrated how a
\-alue markets elsewhere. Considering the case of a country mathematical epidemiological model can be cransformcd
with export trade already established. Beremsen er a/.26 into a d}'narnlc bio-economic optimitation model. Appli·
used CB,\ combined with simulation model:, to evaluate dif- cation~ for developing counrry sitUarions are much less
ferent control strategies ifF!\10 were to be reintroduced into common. ,\11 exception is a study br Elall et 11/.60 in which a
the J\:etherlands. dynamic programming model was fomrnlated 10 e\-aluatc,
oprimal ECF co.mrol strategies cank dipping, immuni1.a-
Econ omic.surplus models tion) in communal herds in Malawi over·a five-year period.
l11ese are an extension ofCBA in which the impact ofinier- The other main class of optlmi7.arion models is based
ventlon on s11pplyand demand in markets forlivestoc:kand on economccric approaches. but these hove been used
live$tock products is captured. This can be important: if a little 10 date in the field of animal hcaJch economics.
given intervention has broad impact and results in avoiding Chilonda and van I luylenbroeck.40 however, suggest how
mortality. the supply of meat \\'Ill increase. lowering the rel- an agricultural household model 225 might be adapted to
evant equilibrium market prices and therefore the esti- examine ,mallholclt•r farmer management beha\ iour tor
mated value of rhe benefits from the intervention. Changing live~tock health.
prices and amount::. marketed also translate Into changes in
consumer and producer surplus, and these can be quaml- Simulation m od els
fied in the model 10 measure the welfare gains and losses to These models have been used extensively in animal health
cuch group. KristJanson e1 al.' 11 used thb ap1>roach to economics, but arc usually limited to biological processe:,
e\'aluate the potential Impact of a vacdne ag-.-1insl trypano- and applied to measure the impact of different control sce-
somosis. and predicte.d that due 10 lower market prices, con- narios as an auxiliary component within an economic a1,a-
sumers \,~II reap 55 per ccm of the estimated USS 1,3 billion lytical technique, such a~ CBA or optimization modelling. As
in net benefits generated by the vaccine. versu$ 45 per ce111 described above. analyst~ often include some type of herd
for producers, who gain from lower production rosts. The population d~11amics simulation model in order to capture
impacc or the recently completed Pan-African Rlnderpest benefit, of disease control over time, and near!) all of the
Campaign was similarly anaiysed.2H $tudies noted in the preceding paragraphs contain such a
model. In some cases. tl1ough. simulation models have
Decis ion analysis directly integrated financial or economic parameters as
This is particularly useful for disease control imervencions well. Such model~ are often packaged as high!)' speciaJized
that involve diseases with variable risk of occurring. managememsofrwarefor intensive commercial production
multiple possible outcomes. or a sequence of actions. This systems. Examples include PorkCHOP. 51 PigOR.•\Cl.E and
technique estimates the expected net returns for a set of DairyORACLE.112 and Dairy;'\l>\J,.92 Similar models arc rare
actions (decisions) and outcomes by conditioning for smallholder livestock production systems in de\'eloping
e;1ch associated cost and return by their probability of coumrie$, which can be anributed ro a large degree 10 their
occurring. l01vcrcommercial orientation and the difficulties of model-

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 The control of ,nrccliousdiseases orlivestock 21 l

ling the multlple roles of livestock in such ~ysiems. Excep· ing the robusmess of the quantitath·e results. This may be
tions are Xyangito er nl. t6!. and ~yangito 111 til.. 167 which accomplished either b} estimating results for a wide range
model che complete farm household econom>' rarher than of potential scenarios- ra ther than limiting the analysis ro
the livestock production system alone to evaluate the finan- only one or cwo - or by changing values of individual kei
cial and economic impacts of immuni.7,.aiion agalnst ECF on parameters that umd to drive the analysis. Stochastic mod-
smallholder dairy farms in Kenya. els build this type ofsensitivity analysis cl irectly inro the ana-
While certainly not exhaustive, this brief review oi eco· l)'tical model. and researchers ha\·e found the spreadsheet-
nomic anal~'tkal techniques and examples from the animal based tools of@RISKTM (Palisade Corporation) particularly
health economics literature should give the reader an ap- useful for this purpose since irallows the analyst 10 simulta-
preciation for the broad range of slruations and trpes of de- neously represent multiple key parameters by different
clsion-maklng to which economic analysi5 can be applied. types of probability dislributions. These rypes of sensiti~il)
Many relevant economic tools have yet to be applied or ap- analyses can be very powerful in estimating notional confi-
propriately adapted to animal disease ~ontrol probiems. dence inrer\'als for quantitative results.
particularly in the developing world. A numher of opportu- Sensitivity analyst~ alone, however, doe~ not ensure re-
nities and some or the challenges that remain 10 make eco- liable results. Even if the data arc accurate, the conclusions
nomic assessments more meaningful for effective decision- from the economic analysis may be (Jawed if the scope of
making are discussed in Perry and Randolph. 1e:1 1he analysis is coo narrow. Unless all potential impacis of
disease control - both in terms of costs and benefits -
Estimating the cos t of a disease have been captured, then the anali1ical re.suits will be
i'-01 mentioned above are a number of studie$ that have es- inacc;urate. It is essential chat the analysis be a& compre-
timated the economic cost of a given disease, usually at na- hensive as possible to consider and quantify where pos-
tiorwJ or regional level. E.'Zamples include Mukhebi er a/. 156 sible the full range or impac1s. Studies are usually good at
ror ECF in eastern and southern Africa. Freyre w al.67 for fdemifying and measuring the obvious direc1 impacts, but
1oxopla,;mosis in Uruguar. and '.'igategize e1 al. l(il for O\'ine nre often less ambitious about addressing chose that are
faciolosb in Ethiopia. As caucioned by Perry and Ran- indire('t or secondary. Thus. while senshi\il)' analysis can
dolph.183 and earlier by '.V!clnerney, 133 • 13•1 such esrimates compensate for cenain data inadequacies. attention also
may pro\ide an indication or the relative importance of a needs to be paid to the comprehensiveness of the analysis
disease, but \\ill be misleading for decision-making abom of impacts.
investing in its ccmrrol unless this preliminary analysis is
then supplemented by information and analysis of the costs
How can co ntrol options be best in1plemented?
and realizable benefirs oftl1e control itself. The point here is
that the economic cost or the disease 11evcr equals the po· Once the impacts of an infectious dise-ase and its potential
1ential benefit ro comrol. even in the case of eradication (e.g. control have been evaluated and the optimal con trol strat-
doc$ not capture changes in comrol costs post-eradication). egydecided, what remains isto implement the control mea-
and in any case, rhe C<lS1 of com rolling disease also needs to sure,;. Despire improving knowledge of disease dynamics
be factored mto the analysis. Estimates or the economic cost and the availabilit) of better conuol technologies. the
of a disease should be viewed as only a preliminary step to· rt->cord of di.sea~e comrol ha~ often been disappointing. This
wards a proper assessment of a control imervemion. section explores a range ofissues that need to be considered
for effective di.sea.,e con 1rol.
Reliability of economic analysis
Economic criteria should be critical 10 responsible decision- The his torical context
making abom disease control. But with what degree of con- In rhe introduction 10 this chapter. reference was made to
fidence can we trust the results of economic analysis? How the early efforts to eradicate ECFin southern Africa. At that
reliable are they? Schepers and Dijkhui1,en? 18 describe. for 1ime, how disease control should be implemenred was
example. the sometimes contradictory conclusions about clear. The prioriry was to address the threa1 of panicularlr
mastitis comrol from different published studies. There are contagious diseases and their devastating impact by Im -
cwo key issues to collsider in answering these concerns. posing authoritarian measures on all livestock keepers for
First. the quality of lhe anal~'tical results is obviously the public good. The need to co111rol the major epidemic
largely. a function of th,;: quality of the data used, with key diseases was in fact the primary impetus forthe creation of
epidemiological paramernrs plai~ng a paniculatl} impor- state veierinary services in Africa.99 For these diseases. the
tant role. 'Garbage in, garbage out' is the oft-quoted adage. technical solmion proposed by the veterinary ser,;ices
Sufficiently high-q uality data an, often not available. how- probably coincided fair!}' well \,ith what was economical!}'
ever. and the needed primary data collection Is not likely co justified.
be feasible due either to resource or time constraints. \\"ell- As the threat of epidemic disease was brought increas-
designed sensitivity anali•sis therefore is critical to e\'Bluat· ingly under control, public veterinary services continued

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'.! 12 ,.,.,.,," n,r- A.~pens influencing rhe occurrenc,:• of infenious disease,

to ,•xpand their bureaucracies by extending their mandatt':- rily through commt,>rcial channels, although recommenda-
'° prowcting commercial ii,•estock production rrom the re- tions on their appropriate U$C may be dis~eminilted through
inirodurtion of epidemic diseases or the imroduction of public extension semces. The end-user is the livestock
ne\\ exmic disea~e~. and b~ beginni ng 10 addre~ 1.oonotic keeper. and which liveMock keepers are ~,oncemed b) a
and endemic disease,-. With the sh ift towards endem ic d is- given disease comrol technotom will depettd not only on
casei. disuase comm! became less a question of ensuring disease incidence. but also on Lhe l)pe of livestock produc-
compliance to Imposed control intervention<- and lnc-reas- tion system ~ well as their perceptions or the dise~e and
ingiy one of encouraging farmer-adoption of comrol teth- the utillc~ of c-omrol. The institutional environment is also
nolug,cs. The combination of centrist siat.epolicies a11d the an lmegral component of a delivery system by conditioning
,hunag!l ol trained human resources meant, howel'er, that the infrastructure and rules by which lnput5 and senice.:.
dbl.'.i,e comrol deci~ion-makin.g and impleriwncation con- cnn be provided.
tinued to be conccnmned wtchin state veterinary servic:cs. ror a specific disease comroi action. the appropriate
Mtt:n discour~ging the de,·elop rpen t of private $ector composition of the generic delivery system described in Fig-
pruvi,Jon. In thb co111ex1. successful implementation of ure 9.12 will depend on the nature of both the disease and
dis1:asl' comrol was usually measured in rerms of cost the control suat<'ID':
1:ft1:1:1h•en~>ss. • The n ature of 1h e disease In Table 9.4 disease~ are
·1 he public fiscal crise, experienced b) African countries classified inco ~ix categoriel, according to the type and
beginning in Lhc 1970s ll'd to pressure on scate veLC'rinary ~cope of impacts they engender. The rwo key charaner-
»enices 10 reduce coM-,. and activities. This coincided wich istic~ distingui~hing these categories are: Cl) romagious-
gro\\~ng acceptance ot the pnrodigm that the private ,ector ncs~ - 10 what degree the disease is limited 10 the
can pro, ide many !)'J)CS of dii.case control more cOlciently affected animal or herd or is likely u> spread ro other ani -
and elfe1:tivel) chan the public bector.~·1 -\; a re,uh, efforts mals ,md beyond: and (2) whether human~ are affected.
wc,re encouraged to privati1.e iin•stock health ~enices and These characteristics have Important implications for
sh1fc 1i1e responsibiliry ior muc-h ofrhe decision-makmgnnd deciding who has responsibilicy for deciding the control
co,1 for disease control w lhc;,tock keepers thcmselvei;.2111 re;.ponsc and its implementation.
Sud1 l'ffort~ hn\·e been underwken to va rying degr~·es and • The nature or the control s trategy Severul different
\\ 1Lh \':<ryi ng level,- of commitment in most. If not all. African comm! opiions mav be available for controlling a dis-
countries. ease. each having its particular rcchnical requirements
The gencrai consen,u, i~ that mt.her than Improve the and uconomic characteristics. As an example of a purely
suuation. the redurtion oi state scnices and privatization technical characLeristic. the thermostabiliLy of a vaccine
hil\e in,te-dd rcdu,ed capaciry to control diseases dfec:tivcly detennines whether or not the vaccine requires delh·ery
in ·\frica ' 31'· 211 These trends have prompted research and structurl!d around a rigorous cold t·hain. Technical prop-
de\'elopmcnt efforts m b(•ncr unch:rsrand how to improve errie5 ot a comrot opLion may also condicion its eco-
thc dc':<ign and performance of delivery sy,,cem~ for imple- nomic characteristics. From the livestock keeper's
1ne11Llng disease control. perspec,ive. ino.:enti1e~ for using a control option meant
co protecc his/her individual animal from a disease may
Dell\·ery sysLems be obviou~. bu t iris less ~o for a control option that tar·
:\ 'deli wry ,yswm· b n<>w ,·ommonly accepted a, a g,merlt gins prot(·ction at the population le\'el. Whether a control
1cnn fo1 descrihmg ho\\ distase comrol is implemented, strnrnin· i~ primarily curative rather than prcvenci\·e is
whether it im olvcs a ont:·time public-managed di;,ease- abo importan t.
cr.1dka1ion campaign or che ~u~tainahle provbion of \'e1cri-
nar1 dntgs for managing endemic diseases. In Figure 9.12 Due 10 the various technical and economic- characteristics
th.: basic compon,mc, uf a delivery system are prt'>emed. of a disease nnd control ,trategy. rhere is a range of ap·
Dhcasc control in\'olve~ two main inpui,. - veterinary proache, used, as illustrated by the dotted arrows in Figure
products and knowledge - tha1 are deli\'ered or adminis- 9.12. \t one extreme, state \'eterinary services may try LO
t<·reJ by livestock health ~t:n·ice,;. 11w suppl~ and db1rlbu- impo~i! control. ~y. 10 limit an outbreak or a highh coma-
tion of th<.' inputs. a~ \Wll a~ the provision ofwnices. may be giou~ di<:ease (a public good) Ll1at requires ll\·C1itock keep-
the n,,ponsibility of \'Orious combinations oi accors in rhc ers co respect a quaramino order. In this case. successful
· public. prl\'ate. ar civic ~ectors. ,\ <:ampaign to erndican:: a n di~ease control is associated ,\ith ensuring compliance. -\t
t•pidemic disease may be pn.:domi11amly public sector. with the other extreme. a livestock ke<!per's willingness to pay a
vaccine. manufactured bya parastatal agency. being admi n- privare veterinarian IO care for an an imal is an example in
isu,red free of charge by government veterinarian, based on which succe~srul disease control depends on the lh·estock
~un·eiUance information and guidelines is,ued by publicly kcep<!r's demand rather than compliance. Many disease
funded research mstirmes. Amhehnintics, on the other control interventions represent a mix somewhere between
hand. arc more likely to be produced and distributed prima· these two extreme~.

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 lnt cnmrol of lnfectiou, di<;>.t<~-,. of 11,ce.~tnck 21 :1

lmprovi.ng delivery Thormon.2 '- for example, estimated thai diaRnosis am!
l'roductiviry remains well below potential in the majority of consequently control) of salmonellosi~ after iti in1rnd t1C'•
livest0ck produc1ion systems in the developing world. and tion into '\:cw Zealand would b(• delayed br eight month<. if
disease continues m be a critical constraint despite the the comempora~· animal health surveillance acti,ities
a\ ailability, in many cases, of appropria te comrol LcchncJlo- were reduced. Effons 10 improve animal health lnfurm a•
gies. Inadequate health management Is often attributed to tion , whether 1hrough more e!Tecth·e informati<>n ~y~te ms
poor delivery ol control technolo~ies. I\ number of research and nem'orks:1'•· ~ 9• 170 im1>rovcd cliagnoscic 1ools such as
and development efforts have been devoted 10 idenrl~;ng user-fricndh pen-\ide rest~. pr solkiting informa1ion di-
opponunitie~ for impro,ing delivery s>·stems and efrecci\·e rectly from ramwrs31 can therefore be l'Xpectcd to con1rlb·
U$e of disease control technologies. Many of these efforts ute 10 more effec-th•e implernenca1ion of disease control
target specific component;; of delivery :;ys1ems as pre~cntcd strategics.
in Figure 9.12. Better Information is needed 1101 only 10 know \\ hen 10
implement conirol. but alsn how 10 implement ii properh.
Better products Certain characteristics of a control tt'ch- Kenyan farmer~. for example. report that the lack of
nology can constrain its dcli\'ery and adoption. The classic primed information or advice on using amhelmimics con-
example is that of veterinary products Lhai require a cold tri butes to under- or o,•crdosing, and ocher inappropriate
chain, i.e. that must be kepr refrigerated at low tempera- prnctices. including the use of ~ul>~tandard or counterfeil
tures to remain viable. I..ack of infra~tructure. especiall)• Jn product~.i;o Hoth the private and publi, ~ector, are in-
areas where livestock keepers are widely dispersed o r \'olved in gencraclng and prmiding information using a \'3·
mobile. has m;1de delivery of such produces problematic in riery oi media, often through ad\'ertising or ~:<tension
much oi Africa. Ont1 way to overcome these constraints is ~ervice~. and targeu:,d at both ~l!nice provider~ and 1he
10 adap1 veterinary products accordingly. i\n important farmers themselws. Research b no" needed to explore
theme has been the de,•elopmem of thermo~iable pro- "uch information channel~ and how rhtir efft'ctivenes~ can
ducts, panicularly vaccines such as 1ho~e again$! rindN- be impro\·ed.
pest.1~1 Ne>wcaiulc disease.2. zr.. 230 and ECI'. 1"11 ,\ nother has
been simplifying techniques or atl mi nistralion to make Better services initially. impro\ing thl' sen·i<;e c<nnponcm
control more accessible or improve co\'erage, such as coat- of disease conirol was largely conccntrate<l on upgrading
ing chicken feed with thermostable Newcastle disease human resources tJ,rough training and. more speclflcally,
vaccines or administering them orally using a ~imple eye- graduating adequate numbers of \'!'terinarians. :'\lore re-
dropper.25 In the case of crypanosomosis control, the cemly, Kleeman and others have begun examining the op-
development of the Sterile Insect Technique, though very ponunitles for improving the quail[}· 01 scnicei. through
e:(pensi,·c to implement, avoiru. need for farmer irl\'Oll'e- better management, and to this end Kleeman 108 propo~cs a
ment and many of the environmental drawbacks of alter- number of tC'chnique~ as a toCllk.it for analring senice<
native \'CC'lor conrrol options.3 and their efficiency.
Veterinary produc1s can also he impr(1,·ed In term'> uf l'ro,'iding adequate liYestock heahh ,cnic;cs in areas
their economic charncterisrics, especially their price. \f. where li\'CStock produc1ion is nm market-oriented. nr pro,
fordabil!ty can be enhanced by more efficient production ducrion units are relatively rew. di~persed or mobilc - c\pe-
and disrribution thut lower cost either through technical or cially among paswralists - has pre~entcd a particular
instilutional improvemems. For products chat provide challenge. Such art•a:; orwn do not offor ,uflki<-m li11anC'.ial
slgnlticant benrfits beyond che farm where the} are U$ed, incentive< co support prh·atc vetcritrn.T\' practice, anti in the
public ,ubsidie~ can directly reduce the nominal co,t past depended large!) on ~latc-pro,ided sen'iccs. As state
borne by end-u~er~ and thereb> encourage up,ake. 37 wterinary sel'\ices have declined o,·er rhe past three Lie·
Increasi ng the effectiveness, safety, or case of application cades. a number of research-development efforts hnve t"x-
of veterinary products also hll!i the effett of reducing perimented with community-based :,,ystems that could
economic risk hy minimizing productivity losses provide ba~k care in thc ahscnce oi formal ,eterinary ~cr-
and wasted investment \\'hen the products do not work co vices.33· " 4 The.sc schemes typkally Involve the training of ;i
satisfaction. local community member .is a Communi!.) -Basetl Animal
Health \\'orkcr to provide limited parave1erina~ Sl.'1·,·i<:e,
Better information As alread}' noted in previous sec· paid for by 1hc community. 1'his approach is belnj: 1estetl for
Lions, epidemiological information is critical !(Jr c\·aluating example to provide ,·etcrinal)' care to backyard poultry
when and how much disease control is appropriatc. 189 p roducers.~
lmpro,ing the availability and quality of epidemiological
information enables delivery systems to target their intcr- Under standing demand i\s implementing dbeasc cnn ,
\'Cntions better and hence become more effective ,tnd rrol shifl~ from the public 10 the pri\'a lt! sec101 , deliver) , y, .
efficient in their provision of control. Sanson and rems need increasingly 10 1akt' Into accout\l the pa rticular

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214 ste1io, o": Aspec~ influencing the occurrence oflnfcctious diseases

Figure 9.12 Ageneric del,ve,y system

Institutional Environment

PROOUCTS
INFORMATION
ProduCtlOfl
Ois11ibu11cn

SERVICES

COMPLIANCE
(PUBUC
GOODS) -
,.••
•••
••••• DEMAND
lf'lllVATE
GOODS!

••
LIVESTOCK KEEPERS
Ptoduction system
Rislc perceptions

requiremenrs of livestock keepers and their production sys- ery for disease control. it has become evident that better
tems. TI1is is important because failure to properly evaluate techniques are needed to measure \·V fP. One or ches~ tech-
farmer demand rfsk5 undermining the financial \~ability niques draws upon \\'TP methodologies used in marl:et
and longer-term susrninabilit)• of li\'estock health services. rcse~rch and contingent valuation adapted to llvestock
Animal health information systems pro,ide only one com- health senices. 1 TI1is technique has been used to e\'aluate
ponent of farmer demand for control: disease incidence and the willingness of li1•escock keepers in Ken~-a ro contribute
prevalence. How livestock keepers respond to disease risk their labour and money to maintaining tsetse traps 10 con-
\,ill al$O depend on the role that livestock play in their trol animal nypanosomosls.56 Dara collection and econo-
household economy. their perceptions and understanding metric analyses ror other factors such as farmer and
of both the disease and the control strategy. their resource community-level characteristics that influence farmer
le\'els. and the relative costs and benefits of using control. demand have also been reponed. 1 In a similar analysis.
Evaluating costs and benefit~ 10 support decision-making Swallo1\l' e1 a!. 233 evaluated demand £or pour-011s for trypa-
have been discussed above. Various approaches are being nosomosis control in Ethiopia. Another teclutique adapted
de\'eloped and u$ed to assess the other V'Mious components from marketing research is conjoint analysis used to assess
of demand. consumer preferences regarding specific attributes of a
One approach is to solicit needs directly from Livestock disease control strategy.202 Each technique is likeI) only to
keepers using parrlcipatory techniques. Many of the tool~ capture one dimension of farmer demand, so se,·ecal may
developed to date including diagramming, mapping, scor- be needed in combination,
ing. ranking, and interviewing techniques. often used in
conjunction with establishing community-based livestock Better institutional environment The institutional emi-
lieahh services ha,·e been summarized by Catley. 36 ronment refers to policies that dictate or influence which
The other principal approach is based on demand actors are involved in disease control, the roles tbey play,
analysis techniques from the field of economics. Se\'eral of and the rules th<W work under. As noted. the current insti·
these techniques and rheir applicarion to livestock health tutional environment governing veterinary delivery sys-
services in India, including assessing farmers· willingness um1s in del'eloplng countries has been conditioned ro a
to pay (WTP) for specific services and products have been large extent by external macroeconomic trends over the
descrfbed. 1 As pressure has mounted to ensure cost recov- past three decades (especially public fiscal stabilization

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 The C<lntrol of i11fec1lou, df<ea~e,. (If livt,1ock 215

programmes) that have reduced capacity of srate vete- been pub!ished.91 ll is not clear )'et what impacL this re-
rinary services a nd encouraged privatization and decentra- search has had on public veterinary policy.
lizatio1, of livestock health services. These changes have oc-
curred at a variable pace across Africa, and ha\·c created Assessing perfom1ance
considerable confusion regarding public and private sector A general approach for assessing the <>verall performance
roles in disease c-Ontrol and the appmpriate arcompanying of deliv.-ry syMems for disease control is s1ill lacklng. Re-
adjusimems in the regulatory environment. The result has searchers at Reading University in the UK are in the initial
been considerable research devoted 10 developing a frame· stages of developing an approach ba~ed on the following
work for e\·alualing and improving the inS1ilutional environ- four criteria: 136
ment for delivery s~·stenl$. • accessibility, referring to the \,illlngness of pro\iders to
The evolving framework draws upon economic theory. par- supply disease control and time taken to find a .senice
ticularly the New lnstitutional Economics. ro characterize the provider or product when needed:
underlying ec.onomic characteristics and incenti\'es inherent • affordability. the actual cost as well as ability and ,,illing-
in disease control products and senices. Central 10 this type of ncss oflivcstock keepers to pa}~
analysis are the concepts of e.\'lemalities and public goods. An • acceptabili~. the form and manner in whiC'h lhe product
ex'temali()' occur.; when one person's~ of goods or a ~ervice or sef\iCe is presented; and
generates impacts - either negative or positive - on other • sustainability. both financial and inslirulional.
people.1! these impacts are not properly captured in the mar-
ket cost of the good or sef\Tice. then the consumer is likely to These criteria capture a number of the elements discussed
use either roo little or too much from a societal welfare per- in the preceding paragraphs. The challenge now remains
spective, leading to what economiSts term as market failure. 10 idenrify indicators that C'an be used to evaluate these
Marker failure occurs when markelS are unable to prO\'ide cer- criceria.
tain goods or services in the desired quantity or quali~·. Public Research at the lLRl Is currently exploring r.wo additional
institutions have evolved primarily in response to 5uch market techniques for assessing deli\'ery sy:.tems.162 The first draws
failures, and ro mke corrective action by pro,iding public upon the Sr.ructure-Conduct-l'erformance paradigm from
goods. For example. smaller li\·estock keepers are likely to 'un- Rgricultural markets researd1. \\'hich compares delivery sys-
der-im·est' in vaccination against a contagious disease. and ,o tems to \\hat might be expected in a perfeetly competitive
public veterinary services may be justified in undertaking a market, especially in tem,s of the nominal cost of the final
vaccination campaign to ensure adequate population cover- product. Using this technique, delivery systems are exam-
age. External ities related to disease conU'ol are generally a ined for evidence. for example. of monopoly power. barriers
function of the comagiousncss of the disease and ilS zoonotic to entry, or inflated marketing margins that would likely
impacis. In I.he absence of e.~tema!ities, economic theory co mribute to inefficiency.
dicmu:s that markets (i.e. the private sector) are more efficient The seq,nd technique, ba~ed on another theme from the
and cost-effective in providing disease comrol products and !\cw Institutional Economics called Transaction Co~Ls Analy·
services than the public sector. sis. shifts the focus to non-price factors that may con tribute to
Evaluating the degree 10 which a given disease control poor deli\'ery. Asymmetric information between different ac-
action is associated 1,ith externalities thereby provides the tors ill a deliverysy-stcm, for (!);ample. gives rise to a number of
basis for judging wherher it is best left to the prhc1te or pub- potential transaction costs. 1\ livestock keeper has difficulty
lic sector to deliver. T\\'O criteria are often used: excludabll- judging whether a ve1erinari1,1n is treating his/her animal cor-
iry and rivalry (see definitions In Figure 9.13). .As shown in rnctl)', and therefore faces an added risk-and added cost-of
Figure 9.13, veterinary products and sef\'ices can lhen be losing the animal plus the value of the treaunent. An effecth·e
classified as public, toll. common pool. and private goods. delivery system is one that mi11imi1..es such transaction costs to
each ha\·ing implications for the appropriate actor 10 imple- the extent possible.
ment, or ar least finance, the disease control action.94 A gov-
ernment supporting a ,·accination campaign. for example. Institutionalizing impact assessment: the role of
may find it more efficient to contract services from private epidemiology and economics
veterinarfans ra ther than setting up and managing its own The effective and optimal use of epidemiology and econom-
vaccination teams. ics in impact assessment. national disease control planning
The manner in which this framework could be applied to and implementation depends very much on how tl1e use oi
\'t'teri nary services was demonstrated in Kenya118 and then lhe discipline is 'operationalized'. There has been a fairly
in Africa more generally. 119 It was also developed as a meth- \,ide range of modalities tried in different countries. some of
odology for classil)~ng individual livestock services. z43 In which have been high!)• successful and have endured. some
addition, an excellent oveniew of progress to date. includ- have struggled. and some have been quue transient. As epi·
ing a number of examples of how thi& approach is being demiologists and economists. it is imponant to evaluate
used to address specific issues and sugges1 solutions. has what has conuibuted to these differem outcomes. and learn

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216 ..,,,-,°'' ''": .~pect~ inlluenclng 1he ncclmence orinfectiou, diwa<c~

Figure 9.13 Two key economit EXCLUDABIUlY'

ccaratte·,suc:s of disease control
lOW
(source riolden. i 9999'! I HiGH
PUBLIC GOODS TOLLGOOOS
- Ep)demic er zoonot,c d1seasa - Vaa:ine producuon
control (including suNelllance. D1agnost1t sel\ltC8S
movement control, quarantine Veterinaiy clinics
servici!sl - Dips
- Some extension
- Some research
Control of food-borne diseases
- Drug quali(Y control

COMMON POOL GOODS PRIVATE GOODS

I- Tsetse comrof on rommunal Endemic disease prevention
cl; land using ttaps, targets or and rontrol
aerial sp1a11119 Sales of drugs arid vaccines
- Some extension
- &>me research

&c!udabihiy the ab1lliy 01 .he user to exclude OLiers who ri,we not paid from
oenef1ting fio.m the use(s ,nvesiment in a gocu o; servica
R,val:y conSlimot,on o' a good or seMce b\ on; use· preclutles its use bi• otMr
users

from the succt:Sl>e~ and failure~ in the adoption of our disci- been quire dilfert>nt, and much more modest. There ls defi-
pline in I he pnst. nitely a raised aw,1rcncss. and the value of epidemiology
Possibly the greatest succes,, in 1hc field oiepidemiology units as advisers to directors of veterinary services has been
and economics has been in r~search. Duringthe lasrtwodcc· recognized by man) donors who have supported training
ades there has been a ~teady improvement in the focus. rel- programmes. As a result, there has been a significant de\'el·
evance and scieniific quality of publications in the field of opmem orltuman capacity. The major obstacles to a more
ve1erinary epidemiology and economics. This has been par- active adoption or the epidemiological and economic tools
ticularly important in the development, testing. ,·a!1datio11 developed have been a lack of polilical understanding of the
and application of a variety of methodologies. \\'hich have potential returns from their use, and tack of resources to
very much consolidated the accep\ance of veterinary epide- allow the discipline t() be effectively institutionalized and
miology and cconomi~ a;, a /Jon(I fide discipline. It isimpor1- practi~ed by its new disciples.
am to acknowledge the pivotal role that has been played in ~\bat are the uecessary conditions for the elfecti,·e adop ·
thi$ process by the journal Prel'e111i11e \'eterlnary• ,\fedici11e. tion and use of the research pro<lucts of veterinary epidemi-
This progress ha5 been made by individuals and groups in olog} and economics. from journals sucih as Preve11til,e
many countries of the world. Duri11g 1998. for example. the Veterinary \Jedivi11e. among manv others? First, there ha,·e
distribution of manuscript submissions to this journal was to be trained epidemiologists and animal health economfats
roughly one-third from Europe. on<:-third from the /\meri- in a given count!)'. In many developing countries, such
cas. and one-third from the rest of the world. H 9 Bur what people. If the) exist, do not exceed a handful at most. There
impact has this scientific success had beyond the research is probably a highly skewed geographical distribution of
communily1 In many coumries of the developed world. it trained epidemiologis!S, depending in many cases on the
had a significant impact. There is evidence ,hat It has con- previous existence of technical co-operation projects in
tributed, for example. 10 the development of several na- which capacity building in the field of veterinary epidemiol-
tional disease smveillance programmes, such as the ogy wa~ a priority. ~ext, there need ro be positions ear-
Xationa!Animal Health Mo11irnringSys1em (;-.;AH'.\lSJ in the marked for cpi<lemi1JlogiMs and economists in animal
USA. and t.o the teasing out of health and management con- health research and development instimtions. In most de-
straints at the fann le\'el 1ha1 ha5 probably had major lm- veloping countries these would be unlikely ro exceed one. if
P.!!Cts on national livestock productivity, particularly' in the any. Third!) i~ thi' condition that there must be adequate
more intensive dairy. feedlot beef. poultry and pig produc- funding for this person 10 operate effeclively, including ac-
tion systems. It has also had an important effect on policy cess 10 journals. c-mnil and Internet. Public sector funding
and strategy development for national disease control. for for veterinary epidemiology and economics has had to com-
example in the work of Dijkhuizen and his colleagues in the pete with more operational aspects of "ernrlnary senices.
;-.;etherlands on F:0.10 and cla~ical swine fever. in most and during the ~1n.1ctt1ral adjustmt!nt proce,s 11nder1aken by
coumrie~ of the developing world, however, the impact has man)' countrirs, they ha\e suffered badly. J\s far as access to

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 The comrol nf infectious diseases oflivcstock 21 i

elec1ronic media is concerned. we were all made very aware appear to be increasing in their impact; the;;e include
of what ,,.was iabelled al a recem G8 summit in Okinawa as groups in Thailand, Colombia and Chile.
the 'digital divide'. It is not only the high cost of. and thus In all of the developing.a.swell as much of the developed.
poor access TO, computers. but also the inadequate telecom- world. units or groups have tended to be kd by epidemiolo·
munications infras1ruc1urcs. and the high cos1 or access to gists. who have been more comfonab!e developing the tech-
them. The botrom line is that the adoption and impact of nical dimensions. As a result. there has been a reluctance on
$c!entific developments in our field lea\'eS much to be de- the pan of both research and de1·elopment groups 10 ern-
sired in regions where this impact Is needed most. rhis iS plo)' economists, so ~everely limiting potential impact of the
particularly the case in Africa. discipline. The \'eterinary Epidemiology and Economics Re·
On the developmental side in which epidemiology and search Unit at ReadingUniversit} in the UK is one of the few
economics func1ion as a component ofa na1iona.l \'eterinary academic institutions 1hat truly incorporates both disc!·
ser.ice. various modalities have been used to intt'.grate the plines. Public sector funding of staff positions is in shon
di,cipline into national disease-control bodies. These in- supply, and economis1s are considered a lower priority
clude insdtulionalizing it In departments of veterina~ ser- human resource 1hot cannot be afforded.
\'ices or in veterinary research instituccs (or both). and the Internacional organizations also ha\'e epidemiology ca-
development and use of university dcpanmems. particu- pacity 1hat may play a role in nmional animal disease con·
larly those with a mandarory ·extension' runc1ion. There are 1ro!. The F.AO has a small epidemiology group within its
a few 'classics' around the world that may sen-e as role mod- animal health service in the Rome headquarters, and ind1-
els for some se1s of circumstances (but not all). and the vidual epidemiologists and animal health economists in its
Vcrcrinary Epidemiology Depanmen t at the Veterinary many field projects. In addition. the Ole has an epidcmi-
laboratories Agency (formerly 1he Central Vererinary labo- ologis1 at the headquarters in Paris. Somewhat surprisingly
ratory) In Weybridge. UK is cenainly one of those national for such prominem international organizations. neither
epidemiology units that has endured and gained an Interna- has an effecth'e integration of epic.lemiology and econom-
tional reputation for focus and fi.mctionali ty. In 1he develop- ics. nor do they employ agricuhural economists within
ing world. epidemiology and economics have been the their animal health srnrr groupings. The ILRI. an lmerna-
·navour or che moment' in many countries, but while there rional Agricultural Research Centre of 1he Consuhali\'C
are several success s1ories. epidemiology units in many de- Group on International Agricultural Research, has an epi-
veloping countries have generaJI~ suffered from a lack of demiology and economics group, the mandate ol"which is
sustainabilit)'. ·mus, in Africa. where epidemiology has been cmirely research. t\1 a regional level. some epidemiologr
recognized as playing an imponom role. given the history of and economics capacity exists in the African Union Inter-
a population. ra1her than indi~'idual animal. approach 10 African Bureau ior Animal Resources, and this capacity is
disease control. national units have been set up in several currently being enhanced \.\'ilh suppon from the European
coumries such as Zambia. Botswana. Nigeria. Zimbabwe, Union.
Tanzania, Uganda and Kenya, but 1heir functionality and Dis;ippointingly. most of these in1ematlonal and re-
impact ha,·e ebbed and flowed due to their heavy reliance gional units are driven . in 1he 1radltional disease-comrol
on donor agencie.~. whose support has varied over Lhe years. manner, by 1echmcal issues, and the small economics com-
However. on the brlgh1 side, there ha~ been a considerable ponents. ifthe}•exis1 a1 all. are more ofan add-on to provide
dcvelo):>mem of human capacily during this cfme, which the necessary figures to justify programmes. Here is a real
bodes well for the future. In ocher parts of rhe developing oppom111it}' for mor<? integration 10 provide berter st1ppon
world there are some examples of new national indigenous 10 strat~gk decision-making.
units. supported by funds from their own governments. that

References
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218 ~r=, osr: Aspt•c1s influencing lhe occurren<:e or infectious diseases

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~!.!inland Volume- I ond II. S1.1,i>tics Un,1, Mmlscry of Agrkull\lrc "'1d lhc S;maai region o(Somllllland- Pa.,tornl lk1"1/a11m<lll .Vrrwork Pap,..,.
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Ep1d<-mlologi· Llnll. Ocp;tJ1mcm of Anlm•I Prndu<iion and He.,l,h, /,r: GOODM 1· \. a.-TltHU>flliU>. , .., ... (Cd'!o;, s«u:y for l'tlerltull)
Gowmmcm of Zambia. Lu,a~a. Epfdaniol~•and P1,11•w1tlJ·~ ,\.!rdtd,11•. Pnx<t"11,1g1 ofa ,\!~tin& hel,I 01
13 A,c,;,.., 199;, R•lnt6rtoAnnu•l. 1999- Dlrc~o 'Jorlonul d• !'KU~na. Unl,mitJ•C,,11,g,•, Clim<". 9-11 \p,.;J 199; pp. z.u,.
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Storistks-and ~fanogeml'nt lrlformatlon. 1'atlonal OoP"rtm<Ol of
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36 CATLD,A.• 1999• \/c.,,Juxls
. tJn 11:,• mo;·t' a r1.wie11• "I1\•rerinar,· tJ$rf'S bf
1~ \'\;0-.:,..1998. Nntion:aJ Li\~:od: Dh·eJopmfnt MlJ.\u,rP!An . .Xarional
pnn1clpntor:,·opproac/m ll1:d n:tt/:ods f«u.smg 011 eX'/Nf(i~nc<.1 m
u,•c,u,ck o,~·•lopm~nt ~laster Pion Ta;k Fo,ce. Dtpomncm ofAnlm•l
Heahh and I,:du.my. Mini my or Agrft.ulturc ,ind IITigotion. C.o,·cmrnen1 dryl.and A/rial, London..; Jnu.•mat1onal Iru.l\ttHI!" for Em,ironm.(':nt and
1>1:,-dopmon1.
oD131l>l\'i. UJ0111,~,·e. pp. 71
:r.' C.\TLO' . ., .. 1~99. n,~ lt~rd. mJtintt -CJuldte,, tmd liL'trslot:k tu th,• lfotn of
10 .,,;os •• 2000. 7Jmbabw~ U\'l'Stock l'opulonon Dn,:,. Compu1or iilC$.
Ooporunen, o!Vecorinacy Sef\1ctt. llnmr~ Nrl,;a. l.ondnn: ~,c 1hc ChJJdlffl.
l';" J\SHJ.O'RO, R.W.. 1.998, Tht. h,t~hmaniti,&. Jn~ VALMt'ft, ._ M.., LORD ~OUl~lW k
38 (UA\!IIOJ,;O, T., ,1v..:nr.:tlJ, ,\,W., o·CALL\GH.\;\", r.., . . l'LT}R. T P.• )o.:RUSM, H..l ••
,1tow c;.,. , '1AU4.~. ~.M." rtiur.·. o.o.. 1999. The CQnrrol oihl·.anwoter
il-d,._;, /.00111114·.s: &'alogu:nl. C(Jnlt'ill Prartict c,nd Pul,1/c'
j,b.;1'10:S, U,1.t1 .•
H,alrl, C,,nrrol. Oxford: Oxford Un,,er,;iry Press. pp.sz;-4,14, on l~c sc alt commtrcinl und smullhoWcr fanns in 7..imbobwt
Pr11:.'1mi:.·n h•rcrt,w')' MtYiicii1t -39, J91-210
t8 eu.c:ox, P~), &-MltCno:-...\l.O, o.w.. 198o. To control rnhit-r.: ,-accinate (oxc,...
39 CRA'f!:10\;.(), t,. Mtnl'reBI..\,\\ 0°CAU.AGltAS, C,f., Pf:UM, T,f., ).~U.S~'\. IU,..
NewSdvmist, Si. b40-&;5,
\Jt,_OLJ:\, (i,p_ ~JAH.'\:-., S.M. f. PIJUtV ~.8,. \9'9!:f. lit:i'lrt\\'tlttr and 1he
19 s'1.rr:.·wll'C1<, 1 "STMI. , . 2001 Unpubll.shed resc~t<h repon. ILRl. ~onomn:s of li\'l\',tock produc11on on lorsi: ~eai~c<>nnm:rci61 und
N'airobl. sn,ollh<1ldtr farms III Zimb3bwc.R,•:·r,~ (>'1)/,•1·agH1 du.\1~,cin,
20 IW'TJ'ST. 11.. 19&8.Herd :sntl flock produc1i,1r1· a1,.~ssm<•n, u"ng irand3rtl WJ.trlnoircdn P/l)'j Twpii:nu,,. S2 3l3-323.
o!Trake and tlw demoirum. .iJ,'ri<ulrurtll S_VJwm. 28. 6.,-;s. Jo e111tosn.\. P." \'.\S 1111,u."'.BkOlO:. y,.\., 2001 ..\ <:onccptuaJ fnam(1,or~
2.1 11.i\ilt.'\DRli.<lTt J,J .. uc>x:-.Eux, 1. Ii'. \'AS i>IB ~MS. f.J .. 1~9tt DALY!. lhc lor the economic ~naly"s of f.tctor, lnllw,nctng d,>cision•rmiklni of
31,..,·w~lgh1s on bal~n~. Bu/lr1/11 ofr/:f Worf(/ /{,rrlrlr Orga11tsarfo11. 1-1. smtll-~lt liUlne-r.s in ,1n1mal hcahh man:Sf:<'ment. 0/ESrJMrifir
·139-13. Ttthnical llec111a. 20, 6$'i...70Q,
~• n.,a:<ER. c, ~ <;RFRS. A., ,,96 Opemng ,he debate on n.\l.\'s. H,nlrlr /lr,1/cy ,1 COCliRA~. ,, ••. , 19:7 !i<m1p/111g1ecl111/1/u1•s. 3rd cdi1loll. -.:uw\'or~, lohn
a11rl Plmmill(I, 11. 179-183. W11l'y.
23 B.\5"rrr. T,1.• 1993 uind u.,~confl!ctsinpa>1or3I d•,·•lcv:n,mt In ~,2 c:ou.,tA.,. 1•.f.. 1. 1)\-r, c.., 1~6. lmmunll..it!Dn t'.O\'tra&c requiu~d to
nonhi:rn COted•h.om.•, /11: "\.SSm. T.J. *" c:Mu:o.rMr·r. D.f., ttd.,s.:.1..tmd ,,, prc,·tni nucbr<ok> or dog r.ibie,. \'ara11, l4, 1115-186.
African,\g1ann11S>>tt111x. ~l•dlson. Wi><:<>nsin. U$~:Unl\et1-!cy of --43 C:U)!~Jl!':G. t.i.!!I •• 1g99. nie e,·oluc:omrry ecoh~l orAfrican tick..,
\\'lKons;n Pres...- pp. tll-15 ,. Unpubllsh,'<I n Phil : n,•.<is. llni,·e"ity of Oxford IIK.
u IIEC.lri:l"TT, :r,., :997. An appro:u:h 10 me1;1.;t-nnl)~h. po,t,poutum GnRH .,ncJ .1-,; CUAAY J.I. JIU5'S.·A~Ot0flf", l,l..~. Pf.MY. ll.O. ~ \111>..,n.,m , ;\.\\', 199(;, A
lhu «i))roductlw pcrf<.>rmlll1ct o! d•II)' co1,~. 1!11itt,mialogft ti Snru• f'nlmewori< /or lh• analySis of gentle, imru-househokl dynomlc, and
Anlmalo, 31-32:IJ,03. l-3 livescocl: dis,.,..· con1rol. \\1th rxamplo, !rom Uo,in Cib'hu OiW!ct.
,s aui.1,G., 2001, Compari,on oftl1uditTe1<:n1 \dcdn« avaUable(o,ih~ Kon\'.a.Jo11111a/t,fl/11mm1 &ology. ?,. 161 189
cc,mol of '/11\,·ra,lle di;ease in ,111ago ch,ckcn<, /11: ,IJ)f.l><, ft.G. ~ ,s OE ""-" c. 19g.,. De1·elopmen1 ,uppoq uml U,·estock ,emces. lit
<PRAOBROw, P.n .. u:d,,. S·\DC planning \\"Orl~hop ,m ~l:\\'('M°tlc diw.ue wu...so~ R.:: . r11u1, ~-..- \tAO., ~. led~\ u,~stt>rk °'11't.·lr;/Jn:1m Srratt!gi.·f
<ontrol !n ,111ag<' eh!ck,•n,. ~tdlngsof1111 /11r,rnatlo1:11/ 11'orl.JJ:op. for Low /11<Qmf Co11111rl1!$. '.\airobr, FAOdl.RI. pp. Z!-2,
Mnpr<to, Mo:ambiqu,•, 6-9 .\lnn:/1 .:?COO. Ai.UR Prq«<d/11~'! !l:o, 103.
.J6 OE.U :fUW, p,:,.;,, M<'Jll.Jl.\tOTr. ,.,.;., U.Rtltf. ~.... H., 1990, :-.ionltorlng c,f
pp.56-00.
u,~1ock hcol1h and ptoductlun tn ,ub·S.1',~r.111 Africa. Pm.,mlw
26 HtAtt.,"TS~. P.11.M,. om:-HU11'f~. A.A.. j,,,0Sfi:A.M, ,\,J., t992, A d~namtc modtl •,,•r~rlt11>1)'!>1"1klim, ZS.195-212
for~c»l•bcneat •n•l)'Si• or foot•llnd·"10Ulh di sea,, conuol illl!tcg,t.->,
41 DEil.':, ,,L, Pf.k~Y. R,U,. ly\J'L'l>L J,M.,. MCDI.RMOl'f. 1,1,, '1AHA.\. S.M.,
Pt,i,i111i1·r l'erqrlnnl)· .lfrdld11~. 12, 239-243
UU.00. ".u .. \S()ft7,.UltA. , ,i,_ \IUffl""f, \.f J. nD\vt.A'\0,,C.J .. 1993,.
r e,a,,;.u. i:.P.. \\1u,~. ,.,•• ,ww,n. R.f.. 1!)9<J. Publlcpolicyand pri,-:ur Variauor.. tn pn,\'alrncc r:uc, of ,:ck-borne dl•••se> in Zobu taltlr b)
incentives ror li•e<wck djseasc ronrral.A1.,uo/l,111 /011r1U1/ of.-lJ!rlr-11/wr; ugro=lc,g!cal mne· irnpllcation, for Ea,,t Co:1>1 fe\'Cr lmmun1sau011.
nnd R=u= &:onomy, 32, 50Hi2 l. Pm..m1f1\'l·'rr,'tl11ar;• ,\ft.'tl/<:111, 16. 1;1.18~
28 BOKCLI.A. 1.G., 199;. Beci. She~p :.nd Goo1> Mnrkellng Re,·iew. 1996197 ~ OliP\fU"~1£:0.'T f01l ll\,"lllHNA't10'-AL DE\U.0$':'\U!.~ J (u.-m), lia., :.t()OO.
Planning und Marketing Dep3r1mo111, MnrkNing 0•\'l>IQplnont Bureau, Siutalnab!e ll\'tllhood guidance <hec;5 /i,.,1/11CIO(UiiJdfi1('(01•.11kJ,
Mini.try orAgrk1rhuni und Co-opomlves. Dar"' Saloam. pp. 25.
*'
.~,g OEtGA.D(t, ( . RO~Vff.\.~l'. ~, .. STE1SJfl.D. H . ruui . .s. COURtlOlS C,
29 C.\M.atON. A,R,"' 1.,LDO<:J.:. ,.c.. 199&"'. A new probabiliry fc:mnul.s for 199~ Uw,1ock to 2020: th,· next food 1e,olu1lun Food. Atriculr:,r,•nmf
filll"\'f.'}'~ co ~ubsfanllat<' frctf'dom from dJ.sc•nst-. Puwntih· \ "tt1."rinan• rlr• Eni-1to11m,m l)tsn&to11 Paper :!t Woshlnglon D.C., IFPRI ILRI
,\1ttUtint.34~ 1-ti. FAO.
30 G\.\ttROS'. ~\..A..• BAU>O(S:, ,.c .. 1.9981\. l\\'O·Stagc .samphng in.:tm"\"C}"$ lO so Dlo:I'Rj!X, ~" aEAJJS', 1. , •• 1992. M£';.a,a.naJy.4-i\!St3te of thtl' sc:11mc.c.
wbstanti,t,ite freedom ftam disease. Pr&11tm1ioo \'ttarutat)' ,Hrdic.inc. 34. F.pidtm10/0K)'R•vim,,,_ l•l, 1;;.i....160.
19-30. l-1 OlJKJiUft.L..... i\,A .. NOMOW, ,, ;. ,o,,. y:orou.:;. n.~. ,ssa. J>ori.>(;liOP (($tT
3J c,s~o~. R,!<.t. ii kOt ft,1'., 1962... Lit'i!Stock D1,.c,:(W• ,!;11r1't!)·1; ,.;, Fteid .\'!1mw1l mon:wl. Un1,·cntty of Minnno[.2.: AJnlcuhurnl F...xu:nslon Sorv1cr.
for l'a,erlnnr/ans, C;anbemu .\usrralmn Govo111n,cm PublishingScm~e. .;, l>IIIOWl1t., •., .... •· '10RRI>, 11.S .. l9.S':'.M1mal Health Econom1~
~"2 c,1t~EY. o., 1998. Sustamob~ nual lil•dlJuxxJJ:: H'1uJt conrribrmon can u,• Pnnc.pio,. 40d App!ie,tio1is. Pr,>1 Cr,ut«mmn Fmmd1,1i~n In ~,>h'rirwl)·
mak,l London: Dep.mment for lntcmatlonol lle\i,lopmom Sctene>?. Un1,c1'1lt) ofSydney:Sydne; S0u1h,

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 The control or infectious dlsca5c, ofll,·estock 219

:,3 oou.. A. 0r 1t1LL.-\,B., 1.96-i Monali1y in rdouon ,osmoking. Ten yt:tf',, s1Mrso~. 0.1.11 . •odsJ. l'.oon<1,,"1: 8/o/o;:i,...,.1, Cltniea/ Pra(tlrealltl P11bl"
obs<r\'~::onsofBritlsh doc,or; Bm/$11 .l/trlica/Jo11rrwl. 1. 1399-HlO. HM/th co,,rrol, Oxlt>rd; O,.iord Unlve(Slty Pre>~. pp.501-51:!.

ilJ 1>01m1wwF. •·•· • ·TIN<:t,E. c.c.o.. ·~~.:. Dl)tm the Tropic,: 7111· fm111t<t ~ OrT.\U, Y."-.. J'ft:J{~\. tU.>., M1l~OL p,t .. \tCDll~IOTt.J1f., )tURZ.\ffL\,.i..P, &
or. 1\'i/dl/fe In ~imboh«-cof Gro1111d Spmyf11;;forT,.,$<' Fly Crmtrol. vou~G~ .\,5., 199;. The- pn.•\.iJcnc-e or ti(k..bome fnfcctlon,i: In smaUhokfor
Chatham UK: Natu.rul ReMlUree> ln1i,tltulC. daat') furn1> In ~luronG'a DiWiCL !:en)'!\: u <ro>s•,ec,,on~! >1Ud),
i.i DVCH.\1'1:.\U, L.. J:RU~~ R,L ,e. Pt'Mt4\·. 1.:,.. IS9?, R1.1ducing"' ,pi>tlal Pr,1,·e,1111.,,e Vi•:c.rinary M<!1llcm'-'· 30. ~ur;·
dornl>M<' 10 I<> rfl'#cll,~ d,mcnslonalltY for lngl,iit tCg.<e$>iOn an:.11..1,oi ,o c;rr.tu. i...x. P.ilK\'
11.0.,. '\fcuauon. ., .. 1999. Thl' fnc:iclcmre. calf
lh·ebtock dis.ease dl:,.t:ribution da11\.. Prtt'lntii-eVtteri,mry .\tMidnt., 32. monallt!' and c.tlf morbldlay dut 10 Tire/1,ria 11<mv1 mt'ee1ions In
10;"-?i8. smallho!dt!rdi1i11· farnl) in Mu.run-g)1 Dist:ict. ir{ea~,1. PM-enti:~
'.16 lCHF'-~Ot. r,_:;,, '(\".'AU.OW. O.'-!, ._,~\1.\M\, 0,\\'.1,C.URft\, J,J,. 199•,
Vt1rrl1imy .\1fdicm,. 39. 6s-r,.
\\'Ulingnc" «> c<>ntribucc labor Md momi• to tSCts.>comrol:.>pplkat1on 1'; ClTAU, t..~, \ICD1-R.M'11T, I.I .. l,.ATLXDF, p,:., ()t.Au.AGtlA~, ~.f., nl'O\\~
o( comlni;_cnr vnluatlon ln Busta DtMric:. ~anya. t\-'orld ~: ...topn1,..-n:, 25, 11.~. • ,uu>v. n.r,.. 2000. l)!ITrn-ntes Jn thceptdomiology or thollorlosi>
ZS9-2Sl. on ,mnJlholci,,r dn!ry farms in contras,ing •gro·l:colog,cal ond gra1.lng
:,7 csoo1J1. ,.111., 1999. The rolo or ,h, publk ,,ctor in the de,:olopm~nt and m11ta of hlghfand ~•nya. Epllltmlotog; nmt /11f,~rlu11, 12<1, :12;;.3~.
implcm<!.n:atlon or ruti~ h~Jth polld.eoi Prew,1itft'lf! Vowruuuy 78 CITAV, <:.k .. \!O)E!lltMO'M'. ,., •• ,",1CD£.ll.\~OTT 8-.. ~ 1'} JtkY. n.n .• :tOOt Tht'
\/tdici11r• .;O, 101-ll~. impact of l'lre/lcrio /J(IT1., ~tfl'tlion and othor facto!'< on e.\lf me.m d;uh·
l8 TI',\,:\S, o.a II ut•Ftt..EY.~.F.. 1,,9:,., The appUcatlon of economtr ~\o1lun.11on "'"lghtgauu in <mullhold,-r dairy form< In Muranf• Olstrkt. Kenr•-
Pr..'t·t11J/h't• \'iwrlml/1! J,flVli((fti>. St. 149-!60.
<t><hniqucs rn •he h""1tl1 soomr. th~ st~te of the on. Jo11mnl of
lnrernnlfnnnl Dwelop,mmt. -. 50)..524 79 Qri·n,= R.P. 19tsz. F.c,111011111: .-\na/J~ss af.-lgric11/wml PrOJI/CtS.
Bollimo,~. MD; John, llopkin~ Untver,;~· Pr•s..
~ u·ucr.s. ,. w. .,. lti:OE!lit1i.~·. J.tL, 1:98r Thr C'Cf.alogJcal impac, 01 in6C'Ctlcsdt•.s tn
conm·crfon ro ~hf." control of b'et\C!' Or,~ In Africa.: s:i. TC\'fc-...•• In: ""'""'J.OMO. Uo cul~'"="'· P. & MIUs, A.. 1.999, Co~1-e!T("Cd,c.nc-~~ o( maJ.ana
UO(ft'),-\.\~, ,.
~ .• (~d.). /111~//(ML-d T"11.<¥ f~1· Comrol: ,\/eth()(/s tllrtl Strm'l{fu,- control in ,ub,iiahnran ~frlca. l.1mce1. 3.5-1.178-385.
Pr,m..,din~, oftheCF.C lrtteriu1uo,u1/ Sy111JJ(l,lu11>1lsµro. 4.--6 March 19Bo. 81 COOClMi\.'>:,c., COi.US.\'\', p. £> \UI.I.S, .,•• 2000.. &Ollom/( A11t1l}·Sf$ o/Ma/arltl
Roacrd,m, A.A, llalkema, pp. .;~s6. Control i11 ,11b-Sal,am11 A[rfct, S1r.11•gk R•,••rch ~ertu,, C;Jobal Forum
fio "'"· 1994 Repon of expcn ronsultouon on 1he need fonn!olltlluon rnr Heal;h !le;eJ.rCh. \\~iO.
)Y:')(cms to ,trtn5:,'th\'n \·c-tennary .scrrn:cs an dewtoping.<"Ountn("l; 82 Gt.JMMO\\'~ 8, ,. \11\Pll.\.\l, t•.11 .• ~ooo.A SlOC:h.s.<tk
. pan:t.il-bud{!Ct rut.llysh,
Rome: AC'.A,81,. FAO. of an e,tpcrimcmal Ptut~u11.·lla J:wmolytlca fet-dlot vu.rcmc 1.rial.
61 PTl'STl'J:<, c>.a.. 1985- CJtntcal cpld,mtologi, tlu,,rr,hlA'Ctur< ofr/infral Pr.•t\!11tlt't' \'~1tri1;n17 .\lrdlrint. 43.. 29-42.
rNMrrh Phllado!phin. WB Saundct>. 83 G0Jt1'U.1t,,. H..a;,. CO:lttll.'. J.l,., OCIJ<E. M.C-1..AUIUCI tJ..A. M.A.. CtltJIJ.1< 1rs

Gt Jlh.,~t-.M. 1. !~82. Clob& ,:rndication or ,im-.:illpox. Rt1 ftwo/ lnfertihlt •

1 '-E4URA. LL.. 1988, ln.llucnre of humans and do1nestlc ruumal.. on 1hc
r>,,.,,_, ~- 9)~922. houU'hold ptC\';'l]~ncc e>f fr)'J)tmosoma cru:i in frllltomft fnfrMnn,
po1,ul,nlons In nonhwest A~~ndna Amerlrt1n )1J11t111d o/Tropfrol
63 FU!ISC.Hf.N, r.. ~ \\1.,\JREt. R,, ;.~01. £l<l'n1L'J11\0fecon.omk res-lM,lnCC
Mcdkine 0111/ fi}'~o/cn,·. 58. 748-58.
innrnag,'l!ltnt >1ra1egi~-wmc cmpirlCI\I evld•ncxe r,om t'll$e s1udft< In
German)'. Eco11om/tso/lw.<is:011uC<m{rn111tt• lio/11 b;• .~emurr,,, for,,., 8.J HAU. ,\.1 .. 199r Edi~ori,.I: F.rndica11or. or rnruml1 Tmlffcal M,.,hrinennd
f1111,r;, ,\irlle Hou~. Vuglnl•. s \ptil 2001. lnti!f/lnn11nnl Htalrh, 2, ·I 13~ 14

6.1 Fl'.\•'- ~n ,ooo. Hom of .\rnca Food Socuri•l Upda<<: Octobo: 20. 2000 H.~,. & f:U.i,\~lt. R-.W•• 1998. ModtlHng lhf' t'('Onumk,. of
BS MAU, D.C l,,;,.lbliM

l:ttp't/u·11...·w.feu;:t.1U!.tl. unlm"1 h<alth c1mm,I pTog1-.rr1> u;jr,gdyn~mic progr.un11d11g.

.~grkulnual Sysrt!ms-. SG. 12.S-14.i
65 FRfl(CALC. '.!001. D<>wnloadabl~ irom lmp:lfepiJJJi'b.mnss.,y.ar.111:1
86 IL\8Tl;,M,\tittA..,: t.M .. JUIPJ,,\~NA\, n... !lfl:.\t.\S'Z,. H 1-. t,, THI-I), t It., •98'3 An
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cpidcmlolo!11c Sy•unm 4Mlysl< model ior .-\fncan 11)'1'•no1omia,i$.
&6 FNt'iCJt. s. p,, \\';\U. n. .& MORGAS", x.t...199-~ . A shnulation mocfo1 of sheep Prciwu/i-, \J,,,cri,wry.~k,1fcltw. I, 125-136.
blo\\1ly strike c.1used b;- l.i1cilia s.·rlMta (Oip,cra: calliphor!da,,J Kcn1,1
ii!:ni11.nri.n11, 18. 379-381,
67 l!Atlrt\S.A.!UA:\!. t _,i.. flUPJ,,\NS£1A.. R.•• RJ(\t\,~S. H.fi. '1 TIIU:0:, J,11., 1983.
Epidemic ur.d ~ndl•inie th;,racwdstks of U),panosoml3\is in c~ttlr.A
6; fRn'Rl. , .. 1)0-:'if"-01 1•• r.u.r.();\, , .. C\>TU.LS. o •• COKML\, o. k C.UAR~"tO. A.. &imul.uion model. Pn!t1~ndttt Vtrtr.int1.r;· .\!i,dldnt. l, 13i- J,JS.
199r. The- mcu.k-nce and l->conom:c: s.1gnifc:111cc of ovi,ne ;oxopt:1.s:mo~ ln
88 H..\.l\TL'\L\!UAM, t,M,, ,-uPPA~~fJI. n. 'R1£)L~XS', H,P, 6:"l'lta~. J.H .. t983C.
Uruguay \'~terinnry PtlNM11<1I~·. 73. 13-15.
E\illuation o:t ryp::ino-.omiais control n.ircnud\'t!'I u:,;lng ..ut cpl.d~miologic
68 G:\R.S~- ~.o. t,,· uc::i,:,. M.l\,. 1ggs. An cva:Ju,u(cm o! u.lttmBtivc conttol stmu!atlon model t:1 Nwmfl.v Vffttri,uuy .\Jtditint. 1. t-l7-lS6
~uaa:gte, for loot ond mouth dlse.ue ln \u,;tralla: a regional approach
&,.a1c;c,"!o, tu:.. 1996. Oc-s,gn and cn'iUUiltion or a ,-ctcrim,ry
89 ti.A.Kl. ,t.:i.,.
Pre:,~nt111t... Vi1ttrlmrf)· J\fc.Yiidnc. 23. 9-32.
information s~-s,e.m (or ~nm1bfa. Prif.-tntlt-e 'i t'1m·n(ll)• Me((irlue. 26
69 OIi.RT<., .. HOL\IE.S. P.u .. 1998. Drug managcn,0111 and paroslle 23!l-:UI .
re,l«anc:e in Lrypanosomes in Africa P.v\T '(,c/mlc11/ a,ul :kl•nrljic
90 UATI<AWA''• ,.c.. 19113. kisk anal)~is and meat hsg,~M- /J.I.F.. Sc/en11/k
Serl~ I Rome: Food •ndAgncultu,oOrganlurlun.
a1td Tecimknl RtM11<t, 12. l:l:65-1:?90.
;o GETTl~O\·. G, i, IJ'l'llO><, \\'., 1989. Tho dymunio of Ea>! Co.1st (e\'<!l': A
91 HAl. ~..... Jl~\1)0WH. $,f,. i/. .tt(,ti.Elb, D.).. 200(). Re.mot(\ .,eniln,g n..nd
mod•lling per,pecth'e for tho it1tcgr.i1lon of kno,,1C<lgc. PomsitoloJ:io·
g,'Ogmphkal lnfnnmulon •)':<tCJns in cpfdmuolOji). Mtvinc,t; 111
Todny. :;, 66-73.
ParJui1olog1·. \'ol 4• I.Clndon. .-\cadcm!c Pr~•
n <.>'mNln. G.,. ,,,o:u.. ,. s.. 1~,9. ,\ mornx fom11~~(l<>n of th• life ()'cl!• of
92 UA'ff.S, D,P.. Pft.Jf.TLlt. n.u.
tv \H)RRIS, R.~•• 1998. Produc1fon :u-id
li,-tt lluko. Pr"""edi11gs oftire lril/1 .~aw,my, 7911, 15$-167. r<productl,·e relpoMc ,o us&of Dlllr)MA.\/; A management lnformat!on
/,l rurr-ttSRY G .• l1At8D>.'.1.. S.: •• .\IUtOUR. J. ~ 8(S-ITU.~u.s1nm. c..1919-. ,, S)'Sttm for 'lew /-<mnnd do.it')' herds. /ounurt of fJniry Sctu11~. 81,
pc~dinfon 1nodel rot bovine ostcrtog2ub,.. 77,e Vetcriun.t)' Rt'C-Ott/, 105. 2362-2.368.
5i-S9.
93 HEFr:t:R.St\.S, c.;.
1,; \U~TURfU.l-. f .. QUOO, The dCli\'el"\' or Vt-corfmu)' services

73 Gtuso,. T.1...19~. The 'M1' ,;ysrem for foro:casiing the prm·alence of to the poor P<'<'limfn•t')· finding; from K~n)'J. Re1xm o/JJ/11) 11rojecr
fasciolirts-is /n: c1asos. T,£., tectJ. l\t"tulirrnnd P11rtcsiti~ .rlnimal v,.,t.¥".Sb. £1,3.i!!. Re>.dlng. UK: \'EF.RU.
l'.'brld .\1t•ltt>rt>logic.nl Oragan~1io11. TL~hn!rol Nore .Vo. !59. Gc:ne\-a_ 9.: HOLDts • .).., 199:9. The.- economicso, the debv~rrof \'et.crin:iry smius.. In:
pp 3-S. rER«v. n.o 1c<!.,. Tlr~ Eai1:~m,cs ojA11t11111r OOlllUC<>11trol 0/E
Sdenllfi,:am: Ttthnir-nl Ri'ili,w, 18. ..2~.

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220 <>:1:-no~ <l'L: A~pec1s influ~ncing the occurrence of infectious diseases

,5 noulu;:,. t..u.r,.. 1995, Economk optim17.adon or d1.:Ci!ltons \\1th Tc;,p1;c1 tu In.: lnr\!gmtcd CA.'ngm1,1l1lr lnformarum :.-.iyµ~~ms lh~fulJnr,1';mtaltuuJlt'
dw') cow health mall.\g('m~.,,,. l'hl> rhc<is, Woi;cnlngen .\gncultmal ,\frrrmgemrr11 Qf,\htmnl R,wmrcn i11 -~fnca. Procc,'<llng orthc Afric• Cii$
Unh·cDil)', l995 meeting. ·\bidJ;10. CiHc d'l\oirc, ~tnrch ~'i. iJ'\Jl'lAX!9SI.? Unned
96 UU(H-S·Jt)l\t-..S, , ....... cu.1..., R.R. 1, Fl~l 1'0.'1, ,t.M,, l!J:'.i.Alt A.Ut"#f>h'nt oftiH·
~;Jtion~, Crem·,o1,
FJt11/fc"1/u11 o/lk>r•/11, Eiruq,l/u,o, /11 l:11i;l1111dm11/ Wal"'. Study :S.u. 19, UJ L\nL\M. ,,. 199-r. I hunt'" 11utri1Ion tn 1hr dt.'\·~Jt1J.>1ng worlcb, r'.·\O Ft)()(/
lJnkcNt)' of Rc,1dlng tmd \'t11ri1/m1 l 1,11N1 29. RQinc:- F..\O.
n;- 11u~~A.,U'1014t, 11. '-' et1ruh·, ,.J., 1992. lmpnc1 or lmprn,·t.'"d lh:C'\tot:~ 114 uwRt- ,(.t 1 , 1992. I lb-tot) or OOvint' th1.-ilcrio~i~ in ,nu1hem Afnca:. /It·
di--.. control on hou,ohold dirt Jnd 1wlr.1e: a studi· In U.-in t ,1,hu ,01t\'.\t... R..\J .. P!RRl. h,U. ¥t \6USG•.\_.;.,. nit' EJJidrmiolo~· ufnmJ[('rfOJi.J
Omrf<:1. l:tny.1./1.RAD r«1111/tnl 11,·J)Ort >:u 2 '.':;u,obl. ~enyu In .'l{rlra h,m!on; \t:,tdcmlt l're,.-<. Pl'- l•'.19.
ln1crno1lon;1l Wbor.unr;· for \nlm•ll Ui ....r-1.,,•., us u ..u:. t.. •99H, \f<hquiLO•bomcarboviNse.s. /1,: ,.AL\11.Jl. ~4M. UlHUSOUl~\'
g3 11c_,, 1978. ~lathcm.at!C".il ~toddling of U,·lht<>CX Produt:tton ~}~tern~ "'<fMP50'.'\, n,111 .• 1cd,) .ttNJn~ !S!o!o,rtcol. Clm:rt,T Prm·1ran11d Pui!/r(
Apphrntlon ofrhe Tcxa< A& M Univor<ily l.leef Caul• Producuon \ludcl lfrn/1/1 Ct>lltfflf, o,1ord: Oxfonl Unlwrslr,· Pr.,s. pp.•IOI -115.
I<) Bot<\...-ana. lLCt\ S)'lit('lm, S1mlr \!'Q,). t\ddl<1: Ab:ib:1 lnt.:maciPnul 116 U£., J ., .1...,1 .. ,1oru:. S.J... (U 11\\'.\,, n..s.. l99S Prub1Cm/\ UQIL...1:mng a
1

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99 ILCA. 197g. TO\va.rd.~ an ttonomic ~>M~Mtu:nt of ,·cterirr."t~· in pub u, 1\.187 IS,!.
tropital Africa lLC\ Workin~ Oocum,·nt I , Addls.Ab,1ba. lnmn,.11innal II~ utc.11. t.11 .,. .. ,., U.IOi., ,:,c., 1919. ,>:nt!~:iCnl li1utl~·mf1t!c>gy In \·C(<·rimtlj·
Liw.iock Cemre for .\!rlt'~ ill.<.:.~. 5c/,nc,. 1.,,ndom <:lmrl"' Griffin and Company I td
100 nn. , .• CJlt'}L.\. ~. n·u:rtikE.~ G.D \1,. DUIU;;IS. J•• l.J~\Ji:, S.C.A .• MAf'.IIL, llti 1F.Ox.\lu,. n,,._, 1~.li7 Thc ,,,ppl) ot· \·cumn~· ,arvk~,; >-:t:n}aJI k$.;oM,
J.ll,H., )1,\1 ()O, SJ-t •• '1UX£l\Df. F •• '.\(;l).\ , f-~'.\f , RARJf:\ .\. f-'1- fflOll'l, W ~ Journnl n/ ·',8rlrulwml Atln,{1:btm1i011 muJ 1txuosit,,n, l6. ~l!hZ.'{~.
fk\lL. r.c.~1.• 1988. Economic a.,petb of c-,u1fo produ«:tjan und of
ny uo.,· U(O., v ~ ,~. suucturJ.l 1tform ofthr ,·e1~rim,~· prnre,!Joo in
chemoprophyl:00.S for c<mttol or 1ryp;1nosomiash in •,ilfa~c Ea.,t Ahitan
Zebu caulc in Kcn}'l\. ln: Lfl"•i:w,·k l'tt>tfoe1io11 i,, T$nsi• ,J..lr."tti-d..t,,.,,.,. hf
4
ACrtt'a :ind thL' /1\t.m lo~tHuhonul F.conomk<. Ot•!•!Jlbpmc.•m a.»d Chang~.
24~2.?i-26i.
Afrltn. Prorti'dln&' ofc.t Akc.·ting oftlzt, 1.V'rleo,, Tf)'/)Qllotoft·ram Ll1'\•&1ocJ.,·
,·~cwork. Nairobi. Nllirobl: fmcrnatlon.iJ Livestock Ce-nm.' for Afnc:i QHt.l 120 l[$,(,\l.11). I',, , ..u,urr l'l>:Wtt. JU.. , !\'OJW.\l... lt.,\.I.. KtlSllUtT, K•• 1)()1..\,\'. r1 .,
lmcmation:it L.ibo-mtbt} fur Re'li'ittt;h 011 Animal 01$N1,c" ll.tl\01. CIU)Zl. ti, \\'.\LKUI. I b .. In'.\ IN. A.U. ~ l'ft.JI\', n.o.. l!l{IO. Gl•.ogr-Jphlt.QJ
pp. 3(;()-376. lnformMlun ,.y-s1t"mr. rnr ~tud~ing tht.• tpfdt1mu)tng;- ur <'i'llClc dl.sr~s~
cauit!d b\' Tht•ih•1Jn JNm'/1. 1'1~r \"t:1,.•rillnry· flri"htd, Ilit 2,r;s...263
101 J.~L\'t~·Gu. ,.\v.. 1993. Uyn.un1c lt\'e"toc~ modcUin~ tor on.farm d,:t'Non
,upi,on. PhO Thr.si!I. \\".11:L-nlnw.<.•n ,\j;.ricultun,J Unh·t,r,.i~. Tht.• i-u: LU'\', p_,,., ., u .\1f~tow,s.• 1991. S<.tmp/tng ofJ)()Jmlmlo11S; '1h'tlwd,Hmd
;\l'tlicrlands nppJirmitJtu. '\Cw Yo1k; Itthn \\'ih:y.

102 tAMr~~A.D. • fCOS"trrtJ'I. , ,. ,9@9. ,\ti cp1d1.'l111nlogic:-.iJ modf1 of 122 u..,,<. K•• ,· a 11 "t.A, :,..1.•• 19flJ. Rt:&rl'.s&ion Qni:,ly,J .. (or ,1,rrtl.1ted dMn
Mdetpes1. I Ot!scrlpnuu uf1hc mud~I Tm111m/ 1\11i11tt<I /let1lt/l(llld Amm11( Hc.•1•14"turo/Pul1/k llt'<1lch, 1-t, .;'3-f;.P
Prrx/11<1/011, i I, 59-68. L23 1,1 1..-1.<rocx ' ' OfVUA)P\lf'l''liT IUD), 1~9. L11-.•stor·~ un Pm't'ft)'·Fuc·,mvl
103 JAMl!'OS D,T, • .MOSLE'i'.. \\,ti., Ml.i\SU\\t, A.ft. & l\ODAOltL.._, f.L, 199!. DfSt.'U.¾' L>et't'lopmrm. l.h•c..,wc.•k in J)r\.tloprnrn1: <:n·wlccmt.•, ~m~t,-t't, UK.
Contrr,I Prioriries In D1.•1...-lnpl11,: Cotmtri(I.J.. Nt!\\' Yori:: Ptth!i,hed tor th1•
1:1.: ".tU:.\UU\'. 1.U. \UlAQI N. ' - ~(;\\'. f,t SUSDQU1M, WU ~ :O.UUJ R. \\,"°l.,
World Bnnk b}" OlCford Un!,,•"lt}' Pro~, O<fortl.
l97!f.. A Sttuf~· Oj rfo~ Potvmiul lmpdet oj Foot and Mouth /)/Jt•M· In JJu
100 1c,u,STU:<, J., l\l90. Hool ah and producti\ity ohiltag• pouluy in U11itMl S:m,.,,·. Wa.,hin~mn O.C.: Uni1L"d S-tatc1o Gt>\'<'frlm«mt Printing
S<>urhcaiLASi:t. Economic imp:tet of tk\'C.luping techni<1U1.'S to ,ttninate. Qine,:
b"d> omUy ~lnsr i'l<n'<:.»tk di,e.,,.,·. :IC/AR l'l<>rktn,: Pap._•, .\"tJ• .11.
l:!5 \l~Dt.Jl~ton r., .. 1000. E,•Jdrnct- r•rcd fot ~M.shlb.hln~ the.- :1bsen,c of
r..tnberrn. Albtr:tlla.
lS('t.si' .i.nd tr)'f'.:U)<).',,l'\1lH».ho .:i~oclr.uet! \\ith t.wl!i<' ttr;)tJlc-a.tion
souso·u. l,\t,. \\'.\('fllAA,
IOi J,..\'>t\U, S.\\'., OMU~VB... , , .. )r,U\,\GU. , •• \1,\-:C".U..,\, .P,. programmt,,. /11; rm,.xc.1'Jt. lt.• Ced t .4.rtininl 1'')'JH1II0$0111U~t.c.: D1<JR,r1JJI~
P. • '1Wll.tTl, o.. ::s.x,e Finuncia1 "naly~is.of animal rt)'J)t11,011omio"1., nnd f:ptth•mwl~·- \'l<•nna:. lnu.•mntion.al :\:omtr f.ni:rsy ·\Rl'ncy.
comro1 uNrig cyperm.ethrJn pour•on in Kcm"3. Prrt•t111tit1r t c!tU111111')' pp.156-1&1.
M.xlir111t1• .i.i. 231 ...:uo.
~26 \tl.OI .A.'-1011, ,.,. • ~:uuJ..a,;t:x. ,·.11., 199-;. A r~vkw of l'' litthod'\ to adjust !or
106 KJTAL,. l',\1., MCl)utMOn. J.J •• KYVU ,,.~ ~(;i\1110'.\I.\ J,\1,. :.."OtlO. ctu~rnr-f'ttccr... In l!XJ!lrm;nnry cplt.h.•mioloW(:'11 "'"wdu.t\o :anlr'nal or
C:ommunlty·bMCd acm·c wl'\·t·1U1mcc for rnbtesin \lachi\ko, Ot~u·ict. popul~uon, Pli't't'11tl11t• \'tttrl,utl)· \ft·dldn,•. 18. l:t,.\-17"J.
K~nya l't~t,•111£1~ ltw·nmn,· \lr,llc:,,,. ·M, ;"J-85.
127 ~lC:OLl"'~()rl 1,1, ~,:uu~t.S. \' H.4, "\,tlOU-.:KI ,t).\. 199..: ~lUd\• i;ft... ;gn rutd
107 l.CT.\L\, T' .• ) 1ct>ER-'10TT, ,., .• ).."YUi r ,,,':\,. C",\TIIU:01.\ l,',1.. l'fRR'l, n n. N ano.t:uc mi.:thod> f~,r dtna ciplltc1td from d~ttr)I,. f)( ,mlm:lJ,., Pri'I.Yntfl ~
w.,,.o,u,i. , .. ,001 Dog «ology .,nd den,ogrnphy ,ntormauon to \ritrrinnry .,1c'tllt:uU.'. 18, 17J-l9l.
,;.uppon 1he- pl;1nnin~ of rnb1~~ con1rol m M.1cMli.~DJ.)mct. t\~n)·i'\- jktt,
128 ,1r.0LR.,:crrr f 1J'c.,1 L\firl.\:,r., ,.. .;.: Ji.\.DOJURA,. ,1 .. 191)7. f.,,,,•m.mtts in
Trrmlcn, ,8, Zl 7-230.
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106 KU.lN.V~. <,., t999, S.•r,'i(t: ,\lmutgl•11u•111 In J.i1¥1i<kJ; l>m'i't,,pmrut.· di.wases. 1:,1u/r'miol,mie ,..1Stmu ,\mnml.t, 31-32: 1:t2:S. t.-3.
(.'onrep<.< and Ef,m,111.,. 111abodcn: l>eut><he C'.escllsthafl ftlr
IJ!I ,1cm1ff\f()TI ,., •• Jr:.,\l)()Jllt\A, \I., o·cw.\t:H:\~1 C ,I, I( !'<UCIIO,JU, \t.'.\!,. 199-;.. \
Technlschc Zusammennrbvh :(;I~, C.mbl I cump.,ri>un of diffcr,•111 mu,Jc~ ror ~<e,....;.lng ,·;matiou, in th1.1
1® ~ati,1..\,. Ht .• aus. r. ~ tAXM s. \\, t9SO, Tht em1tt1n.memaJ tm11~c1 of ~ro•prcvalenc~ of lnf1,.."Ctiou\ bu\1nc-rhinotr'1Ch\."'lt1' b)• (am,. 41t3 o11td
L\cl\,e ro11tmt op-."'tutfOJh ,\ rc[lOn on rn~·nt knowlfflse- F.10.-lnlncal dtstnc, tn ~cn~.1 l'~"'llTll't... \'Nm,:a,y .,ttdl(mr. 32• .?19-l.14
Prorluc1nm Ht..vtltb P<lp1•r 7. FA(). Romu, pp. n
)JO ,,cotRMOM ••••• AA,'\JX)t,•11. T.,. '1 ST\.U.. ,.J .. Ht(J~ TIU.' l.'conomf~ of
no Jrrio, "-''DIW.\S, P.A.~ "'rn nso,., taaa
<:mrtr Htf'd DJ•11nn1ic.-s•..\n
'-1 .. op1lm.il tu:.ll1h Jnd produc1n1~· In 1om.1Uholc.fe, li\·t\tot"k: ..r-1wm, ln
lnt,og,•rnnd Stoc/ttJjftC ,\IQ04'J for !:t'fthmrins Prmlu.t:titm."'Jt.-r111itnw: dt:\'ttlQping coumrie, /11: Pt:lUt\, \.l.o., !l-d.. n,,.
f.-rn11nml1,:1, ofAnlmnl
I/.C1 IIJJSJYUl'II Report Vo 2. \t!dls \b•b.c l11ternnrional u,•.,.,tn<l tNnre Clis<~<U'CmrtflJI <>.I.I'. :ir/Mtrjicnn,1 T,,·hmml Jt,wrw. Ii!, :1\\9-419.
for Afrfco. 1.31 \tf.nlft\1Ul'I.J.J .. ~Hl!oilJA~:-v,, I'.'.\!,. lt,'ltU!ot,,:A. ,u.., t\f,U,, IC."'·· tlOli1'....;os.
Ill ~Ml1.l 1AX~O"S', 11...)t., ,w.\UUW. h . \1,. kO\\'LA-'\'O..,., G.J .. ~'TI\lr,~\. 1\,1,, >.,. l ,P.. C:01.L.\lt\X, f'G,, JO','f).. l",G, t;, mOk\.TfJ'\, v.~.. i.!.flnl, ~lfC~b of dlm,ltr.
m LUU\,. r.s.. ,9i)9. Mc:,~'>u.tintnhi:t0:o-t.~ofAfri(nn animal human populalfun ;mJ ~<tdO·«onomk eh.1nµi" cm IS('l'i.C·tran.,mnk'<J
ttyp3.no-.omin,i,, 1he J>l~lentinl ben4:fihof contrnl and tt•rums 10 I.J;'JldOu"1mfa,;• ttl 2050. /11: wrn. a. 4, •ue,., , l<'<M WurLf Cln.<S
rc~orth. Anr1r11/J1,rn/ ~,.,.,,,.,. 59. ,9--911. Pmtuttt':i. Vn! I 11wA{rfrrm l~1>nn~«1mr,. Bo~mn: Klu\\l!J pp. ~38

U.2 ~U~KA. A.I.. , PER!i<"i, "·"·· ,n1n. R.<- 199,l, H1.'\'CJ\t prosrL....Sin dw lj:! 'I.\CIIIL\. \:,. t,.hU~.. ,u ., \\'.\-'"'i'.\SGU, ~\\ '1t:UI .k\10! 1. f.J•• \',;UJJUW~,' f
dM·~lftpinent nf d~itlnn ""uppon ~Y"ilt·m.; rnr lmprai\ ~ .:s.nirrtnl ht•alth. J. M,\lrt,U\:, 1.. .200Q. l't'fl.\epti<U'1J ot (,m.lc- b\\,1ll'r-. in tfif> corurot 0£

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 The connol of infecuous diseases orlivestock 221

Afrlcw1 bovine lT)lli:mo,;omlo} ..1ti., i'mrt't6'1'11n,:s of 1111~ \'tntl: lm,•a1(ninn<1l 1Sll ,1r.\J \\1\\, ,.- .. I L')\\'(Jlffl1. n., \\1il.>, c; .. '\$\\\UR.\• .=> , "ltU~li.,\, P., ,~uhl~',QS,

:iyn1WS'lt1m "" ~·,rnma,:. Enfdrmil)io;yand Economco. 7-t l August t ... x:t-nRa1,.., a.. ~1 1:a.,1 c.oa-., ft"l.cr ln )ub-~1hamn.Afnca r<l-«('r.
WOO 11rcokenridsc. Colur~do. lnh.•rnrmo11.1.J u,,l'..S<H.:l.. ft~t-aich 1n,1ituc1::, 'lUll '.\airoh1, ~cn~,.4.-
13;1 ,1c1~Mt.._t,. J.J 198ft. Th"• C't'onamli: o1n,1J~"'ii1 <If h\'l tCJCli..dhez.)\.•: the.•
1. . 1..
'~' ,,m,11.1 .... u..,, \~ij~ Th,• appltl'JUQn<'l econo-min.in nnhnal httalth
<L,•\clC\fllng l"ramL""Mff.... It:' \\1UJIU IUj,, r .. \(it.IR. J.f' ,._ JtU\•.\,S, "·"·· p:or;ramme~; ~ pMC1icaJ ~ulde ht. roun e..n .• !t'd. n:e 1;·co,1<Jmla of
it!'d~;. f>roawflng, uJthe Sth /mtm,11101111/ Sympu;:um un t itt.·mmry ,ln111111/ t>J;,,'1<' Gmttol. 0./ f .S. m:lifir n,u/ Ti<'/mtml //,,,.,,~. 18.
i:p!deu:.ofogy and F-:t,mr,mir-,1,, ;\t.'ttt t '•~t,•fl11<rrfn S<m:dUmt•i<a, 'WS-311
311ppkrn,·1w1m, ff...l, 03-7.;, 1:;.1 1-:nnMs... ''- , ... ~?.\Jt.1;.11, \\·,r,.11-t•,~. The rtl,uton.."hip lwt\\'tt•n fnfectiun!,,,.
1,-.: ,-,os[Jt'\"t.\·, 1,. ;~fl. Old ttonomlc~ for n~w 11mbll!Jll\•lil'l"trtd dt'-e.1.\\.', dtw...t ~ 1tnd tht-lr tronmnk 1.•llt..an~ In; Pt1trw. ILu." 11A,..,_L~. I,\'"· •:d!L
/o;,rna/ of,tgdcul/11111/ F,'()r.om/(1. 4- a1s-aa, ~lrJ\.lellin~ \'«tor Homt: .lnd 01hur r.ir..l,ttir D1~w'-. 1•ttX'.t\1 tllng1 of,,
l3S ~:a.too. ,. & u~Ut. , .. ~DOD. Snc-io•C<"Onomit imp.trh nf fte,·dnm fmn, 11crl;s/101101;w1i.:,-d by 11.111111 ,ollnb<JrdlltJ•1 u-/111 F.10.;!3.27 :>.o,·cmber
lh.t,i.totk dl,.c~sr ond MCpon promo1Jon in dcvcfoplng tOW'ltrlc<. 1~9.2. Xot1n,h1. !\4.•nyn. fmttrm1ti111111l /,e1lt01t1:t1,;· l« Rcttartll In ·\nimnl
I.Ji t'S10t"('. J"ftJrntl.llimt tlntl P1Jliq.• l>JR•tiJ..t.(nil. P.tpN \:o, 3. Rt.Ur.tr. F.\0. /]/Je,i.w,, '/L/1.W,. J>p. 192-~13.

t36 ,tntoo. \ .- \'.ttSMQHl ... t .• '260:t. 0e1,,.tf.\'O! :1n1n,:d h.:Wth c:irc ro 1hc ts,", \.IOrtkl~ Jt" , •-"fil''II 111, f,\\ ,,ut:-., }I,\\ '-''«,' flll..L, ~ ,i l'\l•V,()'.'I.,

poor. ft re,.i~w. /,r. [tERa\· A.O. lt\,,\:OOLPJI, T .f_, \'101Nt.MO-rr ,.,. ,0,1-..•'h
. 11:.k. ,t.A .. 1001. Pf4.•,d!rt1,~ s.p;uial modclUng oi alttm;lli\·t.> ronrrol str1ut1;tes

/m,,'t!Sring 111 ·lUflitll/ ff('u/11, fb.'Si.Vlrtb 10 All1..••ril1tl•

/or 11(0-RS:lQ'.'.•, 11,-K••
for 1h\" rotU ..i1nd•n\ov1h ~hCi'l\<" (1tldcmtc in Gr~I fJriti1in, Tiw
POt1'!tl)'• X:llrobl: ln1t•rn.ational IJ\'t<S.mck .R&•ard1 JnMittJrt· 1ff .RI,.
l1•/,•rln11,y/!,.,111/, I 19. 137-H•I,

13; -.1cxAU, w.u•. ,99&. 1\ g_encrdl fouueworb. iilu••tr;JUng UJI app:o.ach w lS6 ~rn.,;m:nr. u, .. Prmt\. rt.o. 4' um~h:.\ n.t_ 199:: El'l,umah:J L'<'t'II\Untlr~ ot
th,•ik.•rto .. 1,c.·untrol In AfriCJ /Jrt't'l'J1l11'r l'cN1'/m,,,• .\11·.dlc:int,. Jl. ~ -
quanmatl\·e n\icmbial rooc;f tn(c,~· r~k o.<-wssmrm. Journal o{F,H.KI
Pror«1io11, 61. 12 l!>-1228. l!'.i' MU~}ll(II. A.'i\,. f:IU\f(t());.Q, 1 o·c.\l.UQII--'\~. f" J .. t'i;TI-J.l. t.r•• J.11U'i~\, IU...

138 ~IAl.bO, ~.II .. TU0R$11, \\'., tW:JtJ. ~. !\'.QU\fl, 11,. )U.)\Vl..\i'e.t)•.,. G,J1 _., t'flll\\", lt.1),.
,1>1>1J\ r..1 .. M-\Jt\X.~..,,.~n1un· 1u1. 1~ \n,t.,~cn.'SmtntrJfthi.,
2001.. !)c-coprcvult•nti.•:. of ,~.,t·tron,.mlttc,d inf1X'1lons of ,malJ,hofdrr i.'1.:<U\oruic: impac1 Qf Ju,<ltcw.iwr rr.ouvlrin rummamium inf(:Cli<.m> .i.nd
1~s control in t..tmb:.b,n•- l'rcn"'mi1o;• v,,1.,•rinar,· Metficin.~. 39. ti'J-..189.
daif} c:;mfc In co.ulal 1'cnyil. Jlrr1,rntli~., Vrti•rit1<Tf)' Mf",ficinf', :i:?. 1-lU.
I~ \11)11 "'.',· (,.,. \I L .. Uf>l', H , ~ .. un,u,
f' , ~ot \ "' ,,, , ,, & f>ftll JI M()S ... \{U1 °:'!U.
139 ).\AU)(), ~.u ... U(IWl.M.;US. 1•• 1•• I lfUAf'l \\' (,I n,~,w# c.,. & rUIR"\', •• o .. !t(l'OI
A!Qngiludiru,J s111dy ol d,,• .,.. tnoidcno,• ond c,., fotalll) rM,• nn , .. 199.;. lhc t'lr«·I ofl.M"~ Uy :md ~c~· ,·orurof on mun-irn in 1lrc
,1n:aJlholdt:rfann.s fn co;.:r-tal h<m~a />rt!WIIIH't' ''tri!rinao \lrdla,,;,.52., r>kt1vang1> Ochil of Ror)wana Pubhrati(III no. I l~. f>tor~Jflll,..".f t;/thr
:Ntlt m..:t·:Jng 11/tlt(' l11t('tnlUttJm1l ~nr111i.m: C:Qimtil/fJt l'r,11m111.,umut.si..,
17-2:9.
R,·.k,m:h mid C/Jmrt,l a.W..TRQ. ~1.ipUID~ pp. !"J.Ji~ ..,562.
t.;O \~AtOO. "',ti,. ~(;tl.\U, P,, \UK>C".0, ~. WllLt\,Nl:., ~. tHUHP> W, Jt()\\1.A'' IJ~,
IS!l \IUJOV.\ . C.f .\. • LOT'r:1:. h.O., lfl'lb. (',lohf11 Burrf.,•n af /:JISt'4tM. IJo~1on:
c;,J., "'""'· 11.0.. :001 . IJonufication of• 1Jr~t1 l>Uflulatlun< for
inununl~uon 118.sin:,.t Fw~t Co1ht fcw:r 10 ran.. raJ Kcnyil. /1r1•1·if1tW't' H:,r,.,,,rd Unwcr5hy l'tt-&\
\ ·r1..,r1nary Mrtlk/Jw, 5:2 31-41. Ilk> \111,0ltl,I ,. i\,, \H>N/.\fU\• ., a. '\l·~I , \ •• 1 ~ \ubunil \'Ut't.-ln('"I- (01 1h1..•
I.a \\.AJUMJL i.t.... .U...\llW,\J. U.~1.1.). 10¥A.\:.6, J .lO\b'\5. S . .a.. _.,_r..L,·~\OtJ A. l':),l. (0Utro1 of n:t•llf!riil Ptlfl'(1 Jn· ~tQfV,.\kt.\, .. WIU.IM1~t)S,.,.... ,e!d" ·, l.J\'t."
I,.

Communh)·-b.,scd vlk<"dnauun \\·Ith tht'rmus.t.tbh.• \'f.l-M n 11l-;t(ft1ph.."tl v,,cdn....., for nk'flt'rin p,mvr: n~µ1oymcnt rn E.1;-.?1:m, Cen1rnl and
-rinderpe$t ,•3cC"ln<' ..n1L".rrmwt&X1. Ott .'1·11y11 \:'tJu'fiuurlm,. 18, ~o:-.,:;og Sn~chcrn \frico, PttK·,..-dln/1< ofnn HO, OAl/·1/JAR,mtl 11111 l\i>rks/lop
hd1l ,,r l,1tunnt/rJnr,J t h,·ttc>ck Rrsvnrclr t,:-sutu1.1". tO-t~ ~tMch l ~
LU '.'itJ\Jl<;II, \'f,l., t«J37, Pt.iccii:.:d computtr modcb !or ~ie:SSiOAproducthit) \,nrob1, Kt:n),1 1pr», 125-130
h1 pl Ap(oduetlon 11,ystcm.), /11: 111nusf.-JCLU. M.,,.. (t•tU. t11«t"l'tlings 1Jj tht!
Socwr;)i>r \ ·,1li.'riMI)' F.11UlrmiQlo;o· ttnd PrtJ'W1ntlrl\• i\lt•tfil:iri.•. Un1,·1:r--i1~ 161 ~,i.n.,,.. F.1wu,nm11•m(I/ \lt.mllorlnl,! H,md/}l()(Jk/t>r i~·t6tC.'ontrol
1Y'f::i,

of Eclinb\u):h. pp. 9S-106 01••ra111w, fdlwd by ch,• Scl<nliff< f.n·,ironn1<01dl \lonhonn~ Group
l'."tl~Jc; • Regional Tx:L~ ..anr.J rr)'p,mu,-omlJ.sl, Control Prot::r\ltnfr.C.
l'"t.'ll!rina-o· Ep1tlrmrt1f()K,':
1.;J -..i,;,101" • .S,\....... Ml:.U•.\,ti, ir \\i.IJ.hUtG. P. 1987,
Hnraro, 1.lmboln,·•·
Pri.mi.pT,-,, <md ,\ll.'tlu'Jfis. Ames-: ro,\·a :-ttUh! Uni\'CP.iity Prl?',..".
1fi2 '1)U,c.'u I.\\",, M.\'WKH NI, r.l,. coni:u:. <,,. ,mt.Utt:, M,C...a. Pl!Klt.\, n..o.
1µ ,tAnHlWM.\!\'.,n-.,\', & PlR:R.Y. e..n.• ,g.95, Mra~uring the bc-11efl1~of d,)('a:,.c 2000. An tcunc1mlc :\.;.,;e+,.,m<ml uf t·urt1.•ru dt.•1h.cr,,· (l3thwny~ for tJ1t
conuot tht' relollom,bip ben"ecn herJ ,,rucrufc and herd pro.dJ1C't1\:1~·~
Trnp:cnl -\umml fft·lllth amJ PrcHfuc-ti011. 17 JB-31
cnntro! of n<}i;-bon1e d!$l~3~'""' In ~'-·n~.1 /tr .... ,u.,."··
,1.0 .. \tfHUJ ,, r,:,. ,.
MUt IH~\UU. 11 '.nh- PttX\..'\fflln," of tltf' \ lmh Symrm>llllt, oft''"
0

n;; ~1n1><;0H. $.G 1984. 1>11,~· d<"~lopmcm aml Jnwrmtl n,QCl.t•dn~ ,n l111,•n1nti«mur 'S"nt:i1.·l)·j(1r \'.·wn,wry· l:r,UJ,•rni'uln~·11ml Fco1wmta. 6-11
,ub· <aharnn Mnc.\! l'ecfonnan~. policies >nd up1lun,. Ii, .~1url; P,,,1/ry A1ii;u>1 iooo. llr<'<~i•11rtdg~. Colorudo. C',0111p,1Ct <IISJ._
Un/1 l\'miir.;;P111H!r \'a. .ii..uld!, ,\hah•, E1hlop1H: Jnlcr"'1tlon.J u 1.-.. 111A1111,, c.. l~Jl. Fmanci.al lo,...<e• au«<! b\
l6l ~(o\lH,tl.J.. l' It.-. ftl ... ,
uw,tuc\.C~nm, fo1 Africn (ILC:\J, o,1n~ ftl,c10lla>1> ln cl,e 1Jh10plan hlj!hl,u1J, TtN/llrt.l .~11/tr.,tl /1,'IJ/tll
1-:t; ,1wu,. cu-., ,99.1 Th,: 1ran11mh,,.fon d~-nnmtO> orn,~,,mu /Hin"· 1,,:· mul P1{J(iut1to11. ?ri. 1S!i-tt;I.
~todelllllg ,,'(>tOt·bom• and other parasitf~dl...,.,c~ lntrmatfor.'11 t(;..i ~ORntlUIZ•Y f.fl.T..\L l•MA!'l:~t~.\, X .. NOHifk. II. I, .._U l I .• tl}93. F!dd
L.abomto:y for Rt~a1ch on Animal 01i.t1u~ fll.ki\01, Sai;ub1 , trfa.1.md error 1~1'1t1.·t.Y1mp, of tla1.• lntt•r,uufo,mf ~·mi1u1.r with t~'urkJhupJ
?P· 105-117. o,t th~ /)('~o~ 0.J11tlut·1 and Jn1~rpti::atio11 offlt1 ld lrilll.J.. &,-;:~,J Dal.
U'." '1WU\'~ G.J., rl1tR\·. tt.U. • \()U'.'..(,•., ....... l!t'}..1, Ptc-J1mim,~· .,n;\I~ sj .. nf thL' /11l• .\,•Jlh.,fm11ls.. ;1;"-28.·1/Jftl 199.1 \'.'.t~cnin~c.m; f.pji.h•c:.on
trantmi~aon d'.i,rthlml~ of th.thJr.rin""L~ in e1ntem ,\Crfc.i... Pr:rlllll<JloS:.,,'\'., t65 ~on:1t.,us:,;.. s..• r..,Wll'JF, C,. 7,WtFI lkJ\,;(., \!.II. e, MU,I>, C~t L995, The
106. 251-2&1, li.\c:c:r i.:om·t.•p1~~pcciticat1011 ot t"tilvri.l u>tin~ quanut:niu~ tlM..
t:.8 M£1~urr. c..L. 1986, c:J;nft11l Ir.al.~-il1.'Sign. tumlurt and 11ntt(~•,.t,. Oxford: .h.w,.,m,•nr. Fr,ml ,',hnul1folt1':_t·, 12. 81 90
Oxford Unl..-t1-rsity f't\.'$.... 10& :\'\'.\"-{0110, 11,0... RJtU.\ElUSOX- I,\\" \IU');.lll8J, \ W, \l\l'.'1.0\•, I>$ ZJY.)H.l, r.,
1i;9 ,,1aom.1.M:N. '- :ao01.>. f.lk\i~ puhlfc'1uons, \ms(crdarn.. f>t•r~rnal s.v,,...:1 ,. 1. ., l'lnJt\ n.u ... 1991 \\'b,:.h: iJ.Tm cconomk ::mal)'>Js of E.a.\t
oommunlcacit,n, C11aM (!:\'Ct inununv:.sJJ<ut !1.ll\llegic, Hl ttJlifl DJ\tria. Rc•nrn. Ptet.-.:n:it'<
1sn \Hu.JG-',, r.1. , .i. ,, u~~. x.n._ 1~88. ·\ modl?I or bet..,e; triU\.....mhh.-d ;nlmal Vtt.-rintuy .\/t'ffi(i1tt. 2.l. 21~:!l.1.
ttypan050mia,j,,, Ptir,wtolog;, 96. 211-239. tti; sv,:-,;crrn, H o .. AU:HAA&Jj():-,; ,m>:tUlU. \.\,, 71,1,,n., ,.., ~ \M).l ,. ,. 4,.
f.W,

15t 'tl'-'.J,\UW. a .. K11U~"-" R.. ODE.PO,.\ .• R.\.\:OOU-11. T., ,10nu,·.~un..J, f'UI""· e.o .. 1~•J. Who!t- form "mnd:tiuQn ;inul~':'l~ ij( rt:on,uml' nnpacb
'"'L\'- \ A 1•1:11",·· n.n.. 2000. f;(onoinlc impt1ct .t!',~K:imtnt t1f (.tJwtlria n! F:i~, Coast w,·cr immuni::ci.1,1ion s.inttCK(es on mtx,:d c;rop•livJ.~to('k
ru1111111m1/11m lnfo<.ilim ~nd IL< concrol In so111horn Alnca. Pro.,wl/11~~ of fnmb in ;_:,m~..i.. -¾:m.'11/ruml ~)'"!ilf!ms, s 1, J..;:._:
1/111. Vfmh lntuntnttount !t)'lflJ)<J#um rm \°<ttt.'rltt(Jt)' l;"pUlrmlCllfl!t, ,w,I th.It (')'f,,\I L'\CII\'.\.. C'.J. \tllH I:\,,..... "' 1"1 .... T.I·• .,. l'l·UJ(\. n.1>., liJ98. 1\nWy·,i~ of
&<1M11tlc:<. i-11 /\ugu>I 2Cl00. BM:l.cnddgef.0. UM. the cpidC'rnlo1og)· of hcar1wac<>r (('ou.rlrw o,mirra,11ilm1 infl'<'llun> In •l

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 '.>2
2- ,,rno~ o,:c Aspec1s influencing the occurrence or infecriou, dlsease,

u:in,mls,iion d~nron!;;s ,nodol, Paro.i,o/ogy, 117.~9-&I. 188 PERI\\', 8,P•• t,;.\1.PaWtDH, w.. COUi)t.\.~. ,.c.~ HOMT, H.S., MCl)EltMOli ,.,••
169 r)CAU_\GUAS. c ..J.. ~!EDLEY. C.F.. PCT£R,. 1.F., '.\tAJtA'.\'. $.\1."- l't;Sllt\'. e:.n.•
IIA"OOIJ'll T.P. b <.:t.lil!>t>ll, 1..1,, 1999, The C<;OllOmic (ll1pR_Cl of

1999. The effect oh-acc!oation on the tmnsmls$Jon dvnaml<> or

fc,o1.and•mou1h di<ta..., and h, control in .;ou1hea~1 A>,ln! • pnellm1rnu,
heanwater (OJwdrln rumfnautlum tnfocdon). Pr~vt:tJ;tft,'t \ 'e:eri1ttuv a~tment " iih.pecinl reference to 111allo11d.1n, PffRIIV, fl,D•• (t~.f. n,~
.\ftdlci1I<', ,12, I 7-39. • F.ro11omlrs ofAnimal Dl.,e<U• C,,ntro/. 0/E S<lom11'{ic and Tuch11/ra/
R•1,.,,., 18, ~ill-!95.
170 OGVS"OWt. <,...,.,·•• OLUOJ:\JS" .S.B, ic UUMUO~). G.t>.. :gag, fhe d(Wctlopmtnt
and effitleneyof tlle ru,imal healt'h tnforma1tons,'!<tem in-'•igerlo. ,ag PrR11,-. a .. >ict1En>1crrr. 1." M~1>0U>11. r •. 2001. Con cpld~mlology and
Phtr't"Jltit~ Vi·t~ritUJ.I)' ,\/ffllcinC, i. 121-135. «onoml~ m•~ca meaningful oontrlbulion to narionn! ~nim;u-discM•
control1 Pn,1y111/,., Vt1eri11ttl')' Ml!dirine, .,a,231-260.
17J O~'U.0•0:-0t..~. J,, MUJt:Hl•B,. />.,W.. n,l,;,-\HII\WA, 'F-M,, ll,'IJSlS.I, G, ROD!', t..
fl.Cl~OXtif\, .B•• P-Orul:\', B.J). & OPUDA·.-\SU)O• .t.. 1998. Finar:d.!t.f anah.~I~ of
sgo PliR::t~Y. l\.D.. Ri\~UULPU, T .f•., )!ct)f::R,.,t01T, J.J .. s.o:,;ts, t;:.R. .Q TIIOl\~"l()~.
dipping stra1egi"s fonnd1genotJS <tmlc under ranch conditions.in r.,., lMI:. hwc.iins in animal h(al1h te<earch tq ~11¢Vfnte po, ony.
Uiand.t. Prt!vt111h•e ra1erin.a1)' ,\f1.'llicuu,. 33, 241-250 ln1i'm~tlonlll Uv.,..ockR~a,ch !n.uw,e (ILIU), Xalrobi, l:•nra
l9l N:i'Efl. T.r.. N'.KRY. 6.U.. O~C:AI.L.\GKA:X, t,J .. "2f0tf'\', G.f'.1 SHUllftA. w..
1:"2 oW:JU:.s:,K,\W. C-B.. 1900. The apprQacb to foreca.sti"ng the incidtnce-of
~lAOZl~I.\. \\',, BUIUlltll:E.. .\1.t. ~ :,.t\ll-\X. ~.J.1.. t998. The d isnibUli(Jn o(
!sscioll:isis o,·erEng!and and Wales. )951H96'L,tgric11/wra/
.11eu:<1rOifJ!;Y. 3. 35-5-1. herutw~ter in the hlglweld of 7.imba.b\<e, 1980-J 977. 011d,r,r,µoor,
/oumnl o/Veterl11nl')' Rerenrell. 65. 177-18;.
1;,s OMOIUi. ,.o.. )td>muton. l,I,. Sl'.\.\t.. ~.•.Aftl.Mf: $,M .. $;A',;(:'mn:.. ti.~•
nu~... >..A .. 2000.Anal)~b of public health risks from con~umptlon or ·~ rrr{e. T.F.. PlR~'(. Jl.O•. o· f'.\1,1..,\GltAN, (;..), Y.t1)U.Y, (;..F.. Sttt)Mt\~ w.,
Mi\021\U. w.. BUJUtlOG'I!, M,t, ttMAll.\~. ~.M., 1998,. l>bitrlbutio:}Ofthc
informalli• marketed mill: ,n sub-Sah.uan Afric.•n coumrics. "'°'''tl/11gs
oftlrr Sfor/1 lmcmailonal S;mposlmn o,, Vett1rinary Ep1rfemlolog;,·and \'tttorsof heartwnter. J\mblyomma l:tlua1wn nnd Amblj'Ommct
&onomfcs, llrc,;kenndgc, Colorado. 7-11 Aui:ust 2000. iw.,ittgarum M.c.ari: lxodiduol in Zimbab\,~. F..rperfoumrnl &App!fl!il
.tc,uolo~ . 22 1-16.
1,, <>RMERoo, ,v.n., 1990, Africa "ith 31)0 \\ithuu1 i,111,tt. Jn$1!('t Sdr11reand
1.93 P'l.-71'.k, T.f . 4.~DERSOX, f.C., fl"UtUtln<.i!, \t.f., flJ,-~kY', 8.0. & M,\K..\.~, S,M ..
luApJ)licnri/)11, II. 455-lt;I
19.99,. Susceptibility and carriers1a1ui o!impnla. snbit ;tnd L~ssel)c for
crtrlcal f(!Vit!'W
115 r,\.\L\lM. M., OJ!lr.,ID.\M. H,. HA\\'I((.".: Lal: :,)\'JU I\J, f>., l9'98, \
Cnt1'l/ril, n1111Ji:antiwn infection lhe;1n-wa1<r1.Jo1m1al o[Pnraslrolog;·.
of prforlt)" «nlng In 1he health <eetor: th~ m~1hocfok1~· of th• 1993 8.5. 3] -3.;.
l'l'orld Devclopm•t11 Re1>on. HIIDltlr Ptilfcya11!1 l'l1111nh>g. 13, 13-JI.
19., PST'fft, T.r. BRYS()!'(, :q.. Pl!Jm\, 8.0,, o·CAU.,on.,s. C'"-J,. MffOUY G,f..
ir6 P>.TO!'.\ G. & Cffl1~8Y, c., ;98,3. fhe conU'<)l ofn pnrJ.Sillc nematode
populntioo In sheep rcprt'SCntt"d b~·a dfscoo,e timc- Metwork \\.ith
\fl,A~)lW.O., HOKA~. I.(.., HURSUP<ie. ~i.,,
f..,'\tAUI\.V, ~.M .. 1999. Cot4'(/rit1
rummonrillm 1ofocdon in an Afrirnn :'\;nurc res.N,·e, 'f1te \.'r.t.eri,utry
;1ochnstic inpurs. Prl}(,'l/din~ 0{1/lr lrffll >11:<1,itm;-. 83B. 267-280. R,•cord. 145. JQ-l-30.'.
117 P.f.G1:t.,UI, Jt.C., JAME.Ii. A.D,, OOSTFJ{Wtl~; C.t.1 .M,0 KIU.Oft,S'i Jq .• ~tC.H£. J..
1~5 Pl<T'E.K. 1".f-.. l'tfl.J\'r, U..t>.. c)°t;.\U;\(ilii\..'\'. C.J .. MWUi.\', G.F-•• )ll.AMBO, G..
G.HIROTI't, )f., l'CKU..-Z., CHfZ\Vl:A. U.G.D .. MW:\SC.. E.T. "CJIIZJIUKA. F,, 1991~
8.hRBET, A,F.41\:AHA.~ , S..M,, 19.99. Pre\-u?ence o( Cowdna rumfnan:ium
Studies on the ecooornicsof tic.ks in Zambi!I. T:x~rl,mmtnl and ,.;pplicd lnfecuon 111 Amblyomma lrebra,im, tfcl:$ trom he3m,'3ter endemic
Jlmfo/ogv, 12. 9-26.
ur,-a, of 7Jmboh1w. EpidJ!11110/og:,a11d /11/erno11. 123. 309-316,
1;8 PE~-:<t:. I:. & o'11A£SJ:, 1..0., 1999, Thclltmosls com.rol modollh,g
ro,;pori~nces fro:n Southern Pro,ir.ce, 7.ambia), Trov1eol .\lrdirine,n1d
196 rmm H. o.u.. =•~.
M,W.,. \!ORRIS. •-s., 199-4 POSS POP: A gwg~phknl
,imul•lion :nudd ur l>t;vlnc tuberculo~l> lnfe~1iun in a "ildlli<
J111rmar/n,1a/ /frall/1 , 4, A.~.\70.
po11ula1lon. 771c Kc11y1111 Ver<rtnnr/a11, 18. 313-315.
17"8 l 1l.ftMt, , t\, kMAOS~x. ,, .• 2002. Uu:~·roture revfewot dlse.ueconll'OJ
197 PO.,tt. c..o. &. DAPL\'N'. f•.F., 1993, Data fo: .\gt'arian l)evelaprmmt.
delivery. adoption and iml"'cl Ismail hol<ler poulu;? /n, PUlllY, n.o., Combrldge: C:.mbrldgc Uniwr<ity Pre,;,
R..\NDOU•H, T.F., )tCDe:RMOTT, 1.,.• ~(l'?\T:S. a;.n.. "ntOR:<\""lOX, P.J:•• 1m'e$lmg
In ·tnimal fleahh R(';fearcli tontlt:iliatePor'frtv. ~3iro0i. Ke.m-a: 198 Pl«>\1e, r .• 1998 . .loonot!c hookworm inftttlons tan~1·1os1~mn,e.1.1n:
lnterno1i1mal Uvc,mx:k RCS<."1l«h lns111u1c (IL.RI). • VAU!E.R~ £.R .. u.,,u, snut...ssY & s1.MPSO..'i, o.t.u .. 1eds}. Zor,,,ow: Btologfcal.
Clinlc<>I Prm:rir~11111/ Pu/1/lt: Hll'Q/111 Omm,f. O<ford: Oxford Uni~-crsll\·
180 .,,._.,., n.o.. 19811. The d..ign and use of supportive "Pl<kmiologirul Pr.<;. pp.80~22. •
<iudlc,. hr. LESSARO, "· & •u~,. •.n., (eds\, nie lnt~rlgprlon 0/1)/sc,1,r
Outbrtttknwd /mp(lirill PrO<l11t'IMry. ~~""""'J'
Cl/111,:; of\·ur1/i 199 J'O:'\O.. 8.J\- tt"Ol.ili.ltOv1.s. Wt .. 11,u~1 II.ft.~ \l)IGT. u.,.... 198.a, USi!I'$
Gu/ile 10 1/te 0111ur/tJ 1/nbi.,, .\loil,t. O,uor:10, Que.,"" lJ0£,ers!1;-.
A111.rici1. Philudolph,n: \\".B. Soundtrs. Pl'· ~7-108,
200 PJ\CTUURO, o.r._ tt>WAtcOs.s. • ,\10111.••Mll.\. s.r., 1983, Casc-contto1.srudlc.s
181 P•n•v. s,o. (ed.), 1999. lhe/Jco11omic-so/A11/11111/ D/,,.,,µO,mrol, OIE
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In cattlf~ Pmt:i!l!tllngs o/1ll~Sm:ietyfor \'etcrinnry 6pfd~mlologyn11d
182 P~Ak1, e.o "ro111<c. ~.s .. 199;. The pa$t and ruiuro rolts or cpldem!ology Pm'>'mit~.Wo't/irinv, &ll11l,11f/llr. pp. 13l~l38.
and economics in tho contr()l of tlok•bc1mc di<c11$c> of111•cs1o(k in
\fricn: the cn.;e ofthe.ilt:rio,is. Ptrl..J1!1ltiW V1t1ritttlf)' ,\/t.'li/Clnt!, 2~. 201 l'\m $.~.II .. SUAW. +\,P,.M •. \\OOUS, .\.I,, nu.n. i... &.. JA\fGS, A.I).. 19\'ii,

107-120. 1',terlno;ry· Epidemimo;ry nnd /:l:1Jnomi<S i11 ,{frita. AddisAb.'lba:

ln1enmional IJ,-cstocl< Ccnrr~for Africa.
183 Pruv. a.o." MASOCII.PII. r.F., 1999. lmprovfug 1h1• •-mcnt or1he
economle lmpnc1ofpam,hlc dlWll5e$ in 11roduc1!o11 •nimaJs. \',t,riunry 203. ftA..'\'"OOU'H. T. 1,.NOU~c·u. L., ;:qoo. Gender and u11J1sac1ion co~c ;J,
Parasl10/11gy. 8-1, 1'13-IG(i. conjom1 anal)~i, of cboic• oflivcs,ock heal1b $Cl'\'icc among
;mallho!der dany farme,s an Kenr•. /i((tl:grountl J)ll{}l!rfora fXJ1ltr
184 P.E.fUt,. JI_D,, ,\IWA."MU~lO, n.•• S<:Mt.l.S. lf..f... £ltJtcn, l. &?.AMA...~. M,Ft., ;98..; . .,\
pri:>lntt'dat 1/11.• Twe11cy-founl1 Ct),,gr1!.csof1JJ~ lmetnQfionttl,¾sotiatio11
siudy of health ond productiv!t)· or rrndltiomill)' managed eaule In of,lgrl(Jdwral E</o11omfsrs. llerlln. 18-24 M)\U11 2000. 1''11Irobl:
Zamb!n. p,,.•1ltlmfllt V,\1,,rfnf1ry Mrdit:lm.-, 2. 633,.65:t
lntetnallonal U""stock Rt.,,eatch ln,lltuw.
t85 PtMY. ts.D., PALM GR, J.E., TIU)\.ftt, H,t•.. OJRCU, J.(t. \IOKJU!f. 1>,. tiHKIC.H. ~1, ~<I
w.~.. ()l)l a.o. A,S,, TIIOK?\"'TOS.
~OJ A,\~001.Ptl. 1'.>' .. ):'ftft,TJA~~()N, 1',M .. O\t,\MQ.
• n1J..,u1~~- \'., 1.986. A·case-.con1rol '-tudy or Po1omi"!:c horse rcve:.
(ramework (or pnonry
P,K., Jl!IP. 8.S.. ROl\.lN"SON. T • ..f< 1':\'A.'\, J,G,. lOOJ. A
Pr1t·,,m1il~ Vt•11,ril>tt?' i\1£111-cbw. 4. 69--82~ scmng in lnrcma1ional liwstod: ...,.,..,,,ch Rl'$/:atth Eval11a,ion. 10,
186 Pll.JUl\", 8..0,. UiSS.MlO. P.. XOll\",U.. R.1\.1., J;tfNJ)hll1', ~ . & ~U$1{..\. A., 1990. 142-160,
Oima1e, <<,getalion tr.nd the dfmibu1lon of R/11µ/c,11/la/111 20..; 1u1m. R.~. 1997. Jmpnc1s uf conrrulhng crypcutosomosi.S on fand-u$e a.i\d
appc11dlr11/111r"' In J\friro. f>Mmi1olog,• 1«1<1y. 6. t 00-10-1. lh<:l!O\.l.[Onoumt: ~ate of ourlmuwledgi..'" t1nd ru1urc dsr~c:uon~.
187 1°1.i.~\·. U,O,, A~VS.AA, M., U',$&.\.«1>, I'.. '\:Qlt\'AL, 11,/\,1, ~ XU~OER.1 • J,. •• 199,, Pr«Mltugs aft/Ii rw..my-fou11l1 .\/<t'1i11gofrite lmcn111r1011alSrlr111,flr
Esum~ling the distribution and nbund;u,rc of Hl1iplc~pl11;,l1"' Co:mril for Try-p11110,omias1.< R,m:r<i> anll C,,11rrol. ~ lapu10,
app.,1diru/a111sin Africa. Pre,·,mii'C V.ti'riJ1n,yMedici11,, 11. 261-2f,8, Mcw.ambique, 29 Sejlicmber-3 October 199;,

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 The control of infectiou, dlseases of livestock 22J

!!05 AUit. Sit~. nu~t:.A. 91.L. \IUTIIUS. , •• TAll' A. \\OTIOX. "'·· wn ..0"- t
a. bnd o....,,,ond for 810,>dinjl 51.Gtl;. f'AOAnm:nl Ht1tlrll nr:11 Prvd11c11on
\IUIMU. w.. ,ooo. i..,nd ,ao and land coH·r d1·namlC'S 111 n:,pon,-· 10 Pa1wrGm.11on1e, f.\().
cll.m~"' In dfm;Uic. bfologlc.J end «i<klpolhlc:al force«· thee•-<' ui 225 Sl!\"GH , .• SQU!Jtt, L &:~1R.\U~S, 1.• 1986.Agrkulturttl H1JUS(f/1<Jld ,\lodt·ls.
,outh\,t<ttrn Ethlupio. Lt11,ds,,.,,..
&'Olag,·, 15 330-$.'\. l¾llimorc: John> H~ptlns Unl,er<il)' Pr•s<.
206 nr:mu~c..11.n, k, k nM1P:., c.,-., 1qg9~ h tht linn1eMi<' dos; 1Cltflis/ltmlll.nrl;r.)
U6 S)r.,0~1mu.. s.1 .. 1$!,~. f:ictt>~ in the spread ofhapatirls E. ltwr.rt. 3s.J •
.1 r~·oirhost of Amerh:an atUinC'Ous lci<Jhmani:n·hr1\ crhfcn1 rcvtc\,' I0,19-10.';0.
or curtcn1 evfdcn<e. ,im"'rca,, /o:mu,/ of7'roµktr/ .1t.vllr/11, ,11ul 11~-giei:,.
2:r; ~"t1n1. M.u .• 1991. 1/nerinoryClillital Epfd('miology:.r\ Ptubl1·m·Or#mr<d
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.o\pprQ<iclt. S1onchum; Butu:n\·unh•Hl';ncmunn.
207 nosr,~os. r .• ~oon. Sp,i:ti-a!srnti-.Ucs :,nd g<'O)?tl1phir:11 lnfomtlldon
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R.O<lLRs.. 0,1.• teds). Rl•motc~t>r..srng m:d C';ro1:tm11hlcul /11/om:unr.,u
D~elopmcnt <>;ttlon, for Farrill)' Poultty. lntrQl/ucrory /¥1/ter-Fim
$.v1rctm.1 h: Epltf«;m/olfll:O·, .·\d,tt1n(n in P,ut11ltolo,:y. \'ol . .;;~ London.: /SFPDIFAO fkcrronl< Coufero11« on Flimll>· Po11/1ry. CiUlil'F.. FF., ,«I,!.
\~~_.mlc Prt•,s. J>p. al-124. Rome: FAO,

208 ,,oc-,11!il, n.J .. t983. ,\ gcne-rnJ modf'f fonhc Afrtc.in U)'J)Q.no~omJ.~cs. :!:...""9 ~PRAntsRO,,.', P.tt .• 1q93,.~c:,..,·ta'-tlc di~a.~ In ,i.11.aie chicken-;. In~ Poultry:.
Part1Mtnlo1t1. 9i. 19:l-212. Sc!J!nc.• Rtw1r.w .5. pp. .>i-96.

zosm.lGWt~ D.f. 2000. Sawltites, space. timeand.Vr.ccan u·ypo~O)omc;,. /m ~JO~k..u,axow. i-.a 4,, t:OPL\..'o:u, ' ·'"'·· 1~f'.16~Ptoductlo.n or thcrmosrablr
UA\', S.I RI\S:DQL.PH, S,L& ROG(.R1. 0.1 .. {l-ds), Rtmo,~... S<-nslug and
~4.•,,t·a1llt.• dl-.<."1l'ltt \firw,.: In dc,•1.•loplng coum:rfes.. P~l·t•111it~ \"ttt<'r,,u,ry•
(kog,o.11hiral lnfomuu;cm Syuems in E1ii1J~mi<Jlugy. Ad1'{u:a,,t ht Medltim•. 29, 1S;-ls!I,
Parasiloloio•. Vol. ,1; Lt111don: Acad.omic Pross. f>Jl, 129-171 231 Sl'L\I, c.. l9~,..~n ~ononlic .u1al1·,lsol lhe ;,=ention n! p<=sic d~"< pcm,
210 Rot.,. A.c." ri~GALJ, P.L, 1993, r~tteide-., nu, producd\')t}, .and formeis· ruminam~ in XiSl'ffa11 {(o.1ts. Prt1tt•ntln~ Vi·t,•rinm:-,- Ak.""(/itlm..1. 16. l-11-150.
heahh; :.sn ~c.:onomic ~ment. ~filn.lla; 1nrcrn:i1 t<HlaJ Rite R<:~"1uch !l,32 :,wAuow u.M., 2000, lm11m:ts of Tr:,•11<mor<>m1a..<is cm ,Vri,"rr ,\,¢,:ru/W"\
.tn..)1in1te. l'MT I rt'ltn:cal and Xitnnfic Series :>.o. 2. Rome: l'AO.
w R0$Srn>t. P.JL & JMIE~, A.D, ,989."11 tpld~mlolog!cal model of ~3 'l\\'.,u..c,w. ll" ,nn..\11', v•. ,., U-..AJr....~.G..4... ,995,. Po1en1r-.al dunulnd fo.r ,1
rin.dl.'rp~t.?. Sin1ul~1tiun~oftl~ bi:h;niourof rrndcr').,e,jt ,1ru.,, l:\ m!x«l puhlk·pm'llW •nlnml h~i11tlLlnpuic <''alu,ttiun or., p1tut•o11
poputada0$.. Troplcrtl Ammat Htnlth and Prrxl11crfr.1n. ?I, f'\9--84. 1n..~cc.k1dt- (or cone rolling bel~..__tranmhted U)'J)rul05om~ls In
212 RCTT11. ,. •z.t.'<~ u<;. , .. :.tOOL &onom!t a.rul1) '\l~ of bruc.rllnM-. cormoA in £thiop;.n. Pri'i--c·mlu \ .,.,.,~,1,u,f'J· Mrdicm-~. .?4. 165-2i5.
~longolla·lmpro,emc111 orhuman hraflh 1h1ough Ilic 1ctcrinJ" <•ctor 23,, "TUUII, L:-r. '\)Al"''· 0.\\,. 'o.\U.:Ht.81. ,\.\\'., RA.'\'UOLNS. T,F... lgtJ9, f:.('(>namk
-lh• c.1-0 of anlmol bmtt'IJG<i; con1rol
Basha Swi,- Tro1•1cal lnsihute. lm~<t ~,.,~,meni ul rfnderpc,1 oon1rol ln.\lrica. /rr; PlRR>-. 11.u. 1ed,1.
::ttJ !.fOTl-l\t..\:'I:, J.-J. ~ CJl•L.1·'1t.A.I\I'), ~.• 1898. ,\ [()(fem Epftlrmfo(Pf:O'. ?nd rdn.
ni~ ftQnomic, of ~,1un11/ D/$i.'<1JeCbnrrol Olf Sd.nri{ltoml T,,·hnl<al
Philodolphio: Upplncon-Rn,·cn. R,wi<w. I 3. .;.s.a-4 ':'7
;t1i; S:\tlOIJ> A.. 2000. Scntin8,•up: ccirical f'1(10rs ln lcadurship. -:tB.S 1.,,it>H, ,,,.r,.. 1998. SchL\10-:.omos"is. bi: P.\l~rr..n.. s.R•. 11lnu sovt.,m· ~
manaJ?.emcm. humun r~urcc dcv·clopment ond ln.s1itmh>n buildiog tn """"'~· 0,1.10, ru,1. ;,,,,,..,,$1!$;
R/tJlugirol. C/111/ml Practlrt-nnd Pulillc
i;o;ni: from pflot pro1<c1 to large seak implomenmnon: the Bl~~C fle<llrh C~mrol. o,l'ord: OKlord Unl,·•"'''I' Pn;s, pp,717-730.
pouluy model in 8anR,fndct.h. /n: t'KJLBLnt.. r." 11n1 ft,\t~. 1•.u. {ed$~. :!,36 TAVlOR, LIi .• L\Tlt\\l, .... M. I. WOUlJtOU~,...L ~t:r.J,, 2001. Ri>kfoC'IOrs fur
t>oult()·Ostt J'aol fn POt'cft)' liradfuuio,t nud Prvmo1Jo11 c/QJmJ~r human dl$c?MC r1ne-rszcn<'u. l 1hiffn411l1ft.nl Trcm14~-rio11 ofth~ Roynl
lil1<1olif). Proc«~l/r,gs ofo Workshop 111./d o,, 22-26 .\111((/t 1999. <ti Tune. s«iet)O (Sdrl«< B, 810/ogia,/ sc1,,1cr,;/. 336. 9!13-889.
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11t1ss1:-i;t;u. , I.NISTI,\N','°;\:· P,'.\1,. ACID. It,'"··
~ts ~,arm_ 1.0. 4,1. t.\Rn,·R101rr. 1.c... 19:9. A t,tencral c:.tu1c pmducuon Ank.,o..... onfJtQ•., "~OlG\\,\.. r 2.002 • .\topping potvrt)•ttnd lit lt.!t()('/.:
.-,,cm, modrl. l.,1i'Tfrulrurol Syn~ms. 4, 217~227 itt tlt,:da't,.1/opmg u'Orld~ f1,1cmatlonal Ll\"'~stockRcse:uch Jnsrnoit
'ILRI • :\nlrobl. Konya.
,z16 ·M~'DLf\"). 1.0. itCAnn, R10111. r,c.• 1979. Agen-oral an1le ptcxl\Jct1on
>)-,.tom, model fl. l'roceed,n~ u>ed !or-imulonngomo,al ~rfonnot1c~. !f.38 nmHSTO:\ i",K. M~OOr.Pll, 1,f , KAl'ir,.,~"'S,(),, Jl,'.\l.. OMA.MU, w :s:.. Q[)t:lttJ•
.,tg,iculwm/ ~y.<1rm,1. 4, 211$-309. \.~.,..,.,.A-,, J.G .. :moo. Vnooty 05s:,-ssmen1 ior thC""-lntC'fmuiunof
Liw,tock ft,(>.)l'U.rch tn:o.HIOIC 2000 {Q 2010. n.RI lmpatl A$Jt'Simr11, Si/fl~
::?17 "'"~'.',(. R..L "THO{(Nl'OS. R', ~ •• 1-997, .\
modcllin~ npprc),IC'.h to lhl'
ci11•m11\cu1Hm o(tltc b,·n~!icsof • notlon~l ,mwl!Juntt pri,gn11nmv. AO. 6, '{3irobi.
Prtwruf,~ \14.•h"'rhmry,\lcdldne, 30. 3t-r:. 239 r1tRu~t:1nn. M tg9_,;. \'"t'terim111· E.J)fdrmiQ/oyy 2.nd edn. Oxford;
218 )(Ht;Pl'I\S. J..\. "'01JxuuttU:'I,:, i\-'- 1991. The economh,.-s of ,nai;1io.; "'.nd Black~«u Science
mJ...,ti(f~ control in d:ilry canl~ ::1 mu-cal amdp,l\ nr c-,tlmuh.•" pubti11ht"d J,J .• mnuun, n,1 UOtT. ,1 .. 1:0UTOU. l , ,. ....,~\. ,, .. 1~91,
~ .O T'();.M, r,.,, fl(.:-.n
,1nc:c :9-;-o, Prt.•tvntfl•1• \ ~:crmary .\f«Jicim•. 10, 213-224. Gio.qmn, ,frµid,w1/olo,ilt•1wlm11t,•. \la, ion>•,\lfon: Edition, du J•uJn:
219 ~t'.JUU1l0M~ \'I\,\.\ U:\. T.\\',,6,.DE IL\.\.'\.. C f9~S.. rrrndsJn tbr
Vl:l.l:rlni1trc.
or,:311izatlon ond iln11ncing of li\'ostock ond 311frn:U heal,n '""'let>< 241 TRn,·1ctu.,. \\:,J. &. \~Ul.fba~. 11-.~t .• 1903- ~tod~llinR bO\iO~ 1ubercuJ0~1s
Put ..•nrlt,.. \'t-1NJ,wr,1 ,\frdicuw. Z.i, 22.5-240. In badger< In; TIIR!l>flloJ..11, "·'··· 1...S.1. />roettdlrr1, of r/u, S<><•t<'f';/t>t
220 SCJIW"-Dl, t.'.\',. 1984, Vt1trmnry Mtdit,1UJ aml flt mum Ht"<1llll, 3rd cdrt. Vtt,rltrar:,o F.pld,mloloi:-;· (fn(I t'r,•1•,mii·t ,\/«ltrint. Edinburgh,
Bakimore USA: wmr:ims ~nd \Vllkin:s. Unhtt,lt)' of F.dinhuri;h. pp. 7~.

~2J ..,.I{\\ \Dt.. <·.w.. ~Jc,.iA~:-. n.1,." FH.\ ...-n, ~t~. tt,~ Epu.lemiQ!~· m
:u2 uw.nt. \. ~ J.\Q:so~. l, 1 199s. ·nu.· World Sank ..ind ln1~nu:itiono.l hu:1lth
\"ermna,;• Pmai<r, Phlll!doiphta: Lea and fcbfgar. polfc:y~ a ,:rhlcal r~1ew .Joumol o[lntt·rnn(io,inl D. ~1~/op11wnt.
. i.
52S-5-ll.
2:2~ S<:n:~u;tc fS\,K()~ \ff~'TA.L MO~J1'0JU'G anoaP. 199), lmp:u:1 of uu-rinl
,pra11r1g411d odour·b•h,-d mtge~ on OCO.')'Stt•n.•. ll<1,or1 /!llt;-/9!JO. 143 U\ ti',U, tM... JT:»Ut, G. lC' 1)1; IIA.\S, c..1992,. TI1r bnt,mc..: bCIWt"C'fl publlr

.\lanitoringoftktsr control r,JNratfons fn Zombifl. cmcl /.imbab:t,r, EC :lnd pri\'3tc. ~L~c1orat:1lvitles ln the dt.~h'l'ry of h\?Stock service., 1/(JrM 4

Bank llfa'CIIJ$(()n Pn/k'T lti.'I. \V3Sh1'¼,'lUl1 0.C.

OeleptiQn, Harar~. %imhab\\e.
223 ~rJ11r.. r ",n1,11- u•. 1t., 1"0. \\'orld U\'t:-ttt>rl. pnxluL11on \.~""itNn!-. 244 u...:oP. 199;. Hun1.;sn llL\\IClupmrsu Repon. l99; r~ford; Oxfortl
atmmt '-lilt Uh i~c.Ul"To und
trends ~40.~llmal PrtNlt1rtln1t mul J.l;tttlrl: U Oi\·~ilr Pr~,;:
Paµ,r .\'o 127. Rome: Fnod Md .,gncuhu"· Organi,;auon ot 1he Unl!•d -=:.:is \'OIGl U,'R.. TT;'\~£, Jt.H, 4'Bk01·5.U()\'i,:-, t.H•• 196.;~A spad:i1 \JWUl:mon
'.\.-;ulun~ model fnr rabl.., conrrul Irr. s.,co~. r.1 .. ,cd.t. Pnp11/a1'Q11 /)yr1nmf,~of
22.1 \lL-\W, A.P,\t, • Hf>\"ft..t'.H .. 193;. T.rypanotolernnl c:aulrand li\·Nocl.:
Rabin Ir, \\'lldllf,. !.ondon: r\Clldctnlo Prn>. pp. 311-349.
dc<clopmc'1ll u1 We,;, nnd C.-nmtl A!rica. \'qi I. lmt•rn,lU<>nal i.uppli 246 \'O'- ):.\UFN,\.'\"', .... Mt:1'-"Tikl .. ,. 4. tm, P.. 1~90. /LCtl Blo-L'tonoruf,: 11,,f'd

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224 -i,,nc" o.~o:: Aspects influencing the occurrence of Infectious <lisense.

.\/,:,;/~/for rhr .\flrm,:tmrp111,·r, Addi,;Ababa: lnu,mutlCltlal u,.-.1,x,k Ea>Joglc2l

25-4 \\"U,.,os. c.1 •• ftt.11>, R,-~•• '11"A'\l0' ,.L ,. Vl!RR\, 8.D .. 199i•
Ct111r.,. for Mrir:a con'l!cqu~ncc .. of rontr<1Hing 1he lWt-.e ~· ln M>uth\..~tern Ethio-plJ~
2,47 \'Oht. I)., 1996. Qttant#ldtil ~ Ri~k Attfll)'Si~· A Guitlt' I() .\Jomt• Ctrrll> e/foct, ,,rt~nd•us,, on bird 6p{-cie1> dl\'e1'i[!', C<11,sumulon Rlnlog;, 11.
Simulation 1\.l0tlt'li11J;. Chtthe)1er, UK: John \\'lh..'}' and S<:mt '43.S--1·;;

r .. 1994,Sp;ui:al
248 W\OIUt. T 1•• )UI.LJC:.,\S, 1•.'.\t 11,\\\"U~GS. f",t,:i;S()W. \\' '2.S..-=. w,,.,., \\
~l: ~t.t sna-Kc;n. J, ,.. RC}Ct,:R,t,, n .. ::oou. u,-C\,otk Ol.Stribudon.

f:t(tOt,. in the 3S$C!'SSmcmt o: u·rp.inoM»m~is ch~lh:ng~. ;,r: PE-Rk'\', a.o. & Production ~nd Dl, c~>c,· to\\ard, • Clob;tl UvestockAtla•. Con,uJ,..,,f,
1u~-,L, r.L, (t(h:J, ,\tudr.lling Voccor~bo,rn~tmd Pnro.sitit OisMU.s, ,rpon. food Jnd ,\~iculturc, 01J1•nlza11on m 1hc Unitc..J Kauon> 1FAO:,
:,;,urobi: II.RAO and l'.~o. pp 115-IJO. Rome.
2.;9 w,\1.Th"Lh·Totw~. o."' ,1u.-w1.:~. ~. \.., 19'9! Che-mica I Tt:",fdu~ in food." of
JS{, WtMJUlf:U,l"l• \t,t., U.'\\"J)O;\, O,T ,, PL\l<St)S \. fl lrl.'JtCJU:O.(t .•• ,j.. 19°fl6, F~tlUl\:
of,..lr.cma11on to pre\•tnl fooc-.:md-mouth dbcase 1:j,;demlologJroruf
anim.il origin: O\'C'T\1("\v and ri~ as')CS.smem. PM,cmuv \ t·rlrinnt)'
/11/r:ct/Q11. IJ6, 363-.171.
.\l,,vtlrln~. 20. 161-1 i8.
~7 woouuH1>1. M,l • t->C>t{.\LUSO:>i. t... ;."\lUQ, Thc~fenteofcomroUing
:tSO \\,\.:\l"AS.CU. "·"'·· U.\lX, A,)... AUGUTl. ,., •.: .• xca-.:-\'I, ,. ,, .. '\IUG,\.\HU, J,\I,.
dt~c~outbrca\:5,,. Vr,llm•,.iJO. 31.5-SJG.
19!\b, Antht.\lrnfnch: ~ba1nc~r timongA,t ,hcep itnd goa~ In Ktny.i.
Pm.r.!mlt't' Vi!tt'r/m;?· .\h.'dlc,,n,. 21;, U.=t-2.90. 2.;ll I \I,., ... ·~81 """''''"'& M1•1/lud., fl•( CA!IISll;es m,d S11ru,ys. -llh edn,
2S1 wuo. 1!196. Jm'f!'Sllng fn J/(!ll/llr Rdmm.:h tmd Dtrt't.'lopmt!lll: R~pc,rt of the l..1.mdo1t· Churl~ c:ri!Un.
Ad I loc Comm Inet on Hl'llhh R~••0tch R~fotlni; tP fumr. lnt•f\'l:ttilon ~;<3 z-rvt1w.. c. Ml'l\N, r-s. & MtlPl'A\1'.1t,11 .. 2001. tnu.•m:.1IPmtl rrudc.anilttal
Opt/ans, Gl'llcv~: 'l'OR/Gon/qO, I, heal1h and ,·c1mnary c"ldcnilotom, ehallcn~c< ruul 01>J>0m11111k'S.
~?,2 WHC,. ~00. World H('o]tlt Repon 2(1(/(), ~n.,, .... World I h,alth Prr1..,•,111w v,.,ci,'llfmy.\tt."tlld,r.1. 48. 2o1-:rn
Oqt:anl7..4tftt11. 26011\flf.\U\\1' Ot.PAR1,1r.~Ttlfl \'[Tl;MIS-.\JIIY ...t'll\ IU ,, 20UI. fno, . . ;ind,;\tC'JUlh

-m ,-.1r IfBERG, r, .. 1975, A c~M.""c-omrol stud)" of some fundamental llisrmOutb1rak ,\ul,'U'1200J . \tap do\\nloot!~d from
dererm1nam3 in the, C'JndemiolQ~· cif 1hc feline, urQlogy ~yncfrom1.• . lm1•·J1c~·u·u· 11Jrlaw11lrnt- ro.W~l\'ll/md.htm on 19 Xu,·ember 2001 ,
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 10
Infectious diseases of animals
in sub-Saharan Africa:
The wildlife/livestock interface

R G BE~GIS, .RA KOCK, G R T,HOMSO:-.: t\i':O R O B!GALKE

Introduction The pres~nce or disease~ tharoccur in both domesric and

wild animals and which may be transmined from one to the
The critical need for rural de\'elopment to $\tppon thP rnp- other constitute5 one of t11e constraints on land-use options
1dl\· grrming human population In sub-Saharan Africa is in- in sub-Saharan Africa. Although the interface resuhs in
creasing the pressure on ,,ildllfe in the region. Po,·eny and some disease transml~sion and economic loss. it should be
lack of food sccurtry arc roor c..iuses of the massiw de$tnic- appreciated that \\ith the mas:;lve decline in available habi·
1ion of habitats and animal popularions.:1!1· 58 • ·,9 '" Wilder- tats for wild herbivores in most regions of Africa, the extent
ness areas arc being encroached upon and arc often of rhis Interface is ac:wully shrinking on tl1e macro-scale, yet
perceived by rural communitlc, .1s legitimate source~ of intensifying in other region~. In addition. amongst the \\ide
natural resources from \\'hich they are unreasonably pre- variet} 01 wild aniodactylids in Africa only o few species are
cluded access for livestock gra;1.ing. collection of firewood, epidemiologic,11ly imponant with regards to serious eco-
thatching gra-,, and other produces. and suh,istence hum- nomic lh·estock dise~e:.. The,;e are African buO"alu (Syncerus
ing. On thu other hand. the po1en1lal or the rich ,dldlire heri- cafferJ, \\'aTthog !Ph,icod1oer11s africmzusJ. bushp!g (Po1amo-
tage of the region for sustainable exploitation and 1hereron1 d111<'T11S sp.). greater ku<lu (Tragelaplms ~irapsitero$), bush-
economic development h olwious. It is also important for buck (Tmg8uphus scri11ws), and ,,ildebeest (C<>1111oclia<'res
sub-Saharan Africa 10 preserve the diversity ofil:.' fauna and ~pp.), which hm·e all berm linked to one or more of the follow-
flora because nowhere el~e rm Earth ~ there a comparable ing imponant disc·n:,cs: rinderpest RP). foot-and-mouth dis-
resourct. Ith. difficult 10 reconcile these e~semiall~· conflict- ease rF~ID). African swine fever iASFi. malignant catarrhal
ing lmpermi\·es. fever ()!CF), East Const fever (ECF). bovine petechial fe\'er.
The grO\\ th in the li\'e:-tock population of Africa i, no longer and uypanosomosis. rhese major infectiOU$ diseases of live-
keeping parn with human populaticm increase. and Iii estock ~tock and wildlife are comprehensively r>rescnted elsewhere
productivit) is not improving in most regions to compensate in this book.
for the increasing need~.% "I here are ;t num\>er of reasons for With this background. the objectil'c of this chapter is to
this '<!atic livc,,mck economy the most important being land discuss the keywildlifo-related issues rhat impact on disease
degradation. 1o;i declining livestock production and faltering persistence and control. These include free-ranging \\'ildlife
dise~e control senices.'ill There is ,ilso a decline in \\ildlife as disease rcsen·oirs, alien diseases introduced with domes-
populations, even within many prote<-red areas. with O\'Cr 60 tic animals. pnstoralisrn and transhumance. the use of fenc-
per cent of all such areas now being relativclr depleted of ruue- ing. and compliance with animal health rcquiremems.
lop!!and large bO\'ids:18Thi!lresoureewill 110 longerpro,ide the which are necessary to gain access to international markets
supplementary (ru1d free} food enjoyed by past generations. and trade. In addition, certain current trends in wildlife
The concomitant increasing need for land on the one hand, and management, such as the de\"clopmem or gnmc conservan-
greater awareness of the lntrin~1c and economic: value of \\ild- cies and trai1,fromier parks. and tl1e increasi ngly popular
life re,ourccs and sa,1U1nah bindinirsity on the other. result in introduction and reloc-.ition oi ,dldlife to pril'ately owned
the issues at the interface between livestock and wiidlifo be- game reserves. which may expand or imensifv the interface,
coming increasingly acme and politici..:ed. arc also of concern and are discussed.
225

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226 "Ct"" <>'r: A.<pects inlhicJl<:lng 1hc Pl"C11rrence of inle<'liuus dis~~;l',

Historical perspective lo wildlife and their endemic diseases and par3liites. over
many cemuriei;. 17 19• 111 Tick resistance, local tl')'J)Bnotoler-
Sub-Saharan Africa ha:. a larger number and d!\'erslty of in- ance and parual adaptation to certain \iral, rickettslal and
digenous free-ra11ging \\1ld mammals 1han any other com· protoioal di~ease~ were probably the re~ult of natural selec-
parable land mass. Thi~ is espedail)• tme for ruminants. As a tion pressures on the livt'!\totk of these early migrants.
possibl.e conscqucnce. an equally impressi\'e array of pa1ho- It was during the colonization of :\frica by Eurepean na-
genic micro-organi~ms and macro-parasi1es. as well as ar- tion, during the past three-arid-a-half centuries 1ha1 the
thropod vectors of certain micro-organisms. ha\·e co- introduction of imported lh·e,tock breeds•- ,o: 105
evolved with their hosts for millennia. which h&culm!nawd unmasked the pres-ence of many \frican endemic diseases
in evolutionary adapta1ion and tolcrnnce beiwcen hos1 and lhnt were ~ilently cycling in the free-ranging 1\ildlife. These
pathogen or parasite. and the <levelopmem of endemic sta- distase< cau5ed significant morbidit) and mortallcy in the
bility. Infection is usually 'silem· in the dt.:finitlvc \\ildlife poorly adapted. immunologically nai\'e European breeds.
host;,, which serve ll$ reservoir~ of specific pa1hogens. lnfoc- and had a ~ignificant inOuence on the day-10-day lives and
•
tion of \,ildlife species other than the definilive hosts may acthities of the ~e11lers. due to their deleterious effects nn
resul! in disease. which Is genemlly mild. lnicction rates draughc transport and production .u1imals. rurthermore.
mar be high, but clinical expre,~ion of disease b generall}' effort~ 10 ·upgrade' the indigenous tribal cattle by cros:;
cryptic In these 'incidental hosts'. breeding also probably had negatrve effects on their
The evolution of the wildllfo lliveswck interface in sub· disease resisiance.
Saharan Africa i& inseparably linked with the more recent Since the emergence of embl')·onic government vcteri·
history of Homo sapilms on rhl~ c.ontinent. The best nary scn~ces ln Aftica. some ten decades ago. epidemic dis-
examples of the recent history or human migration and eases ha1·e received Lhe most a1tention. and were the most
interaction with li\'eStock are tho~e \\'hich iook place in feared becau;,e of their de\·a.iating cffec1s. Disease control
southern Africa: some of these are u<ed here to illustrate cmphosis was accorded tn domestic ll\·estock. which were
certain major factors that have played or continue to pla) a so imporiam for transport. production and trade in the de-
role a1 this inierface. veloping coloniel>. One faclar. which has coi\lribuwd signifi-
The San (Bushmen) were 1he first modem humans to oc- cant!}· 10 the persistence or infe<·tious dise,1ses, is the
cupy southern Africa. They were humer-garherers. 19• 9 7 who cominued dependence 011 pastoralism a:; a livelihood for
left u:- ample evidence of the distribution and di\'er,,it~ of the large number, or the African people,. For example, over 50
residem and migrarnry wiltllif11 in tl1elr numeruu~ rock per cent of the Somali community arc involved In. or rely on,
paintings and engravings. These relatlvel) nl1mndic and pastoral c~ule and small srock for their livclihood.58 · 5' 1The
sparsely distributed inhabitams of the sub-continern culti- large-scale movement of animal population!. across borders
vated no crops and possessed no livestock.; · and V,'NC' in some regions, in search of pasture64 and associa1ed ttade.
therefore unaffected by the anima l disease agcms that were has boen the single mosl important factor in the spread of
cycling in the wildlife etwlronment. epidemics such a$ HP. peste de~ petit-, ruminant> (PPR).
The next Incursion of humans imo the sub-cominem conragious bO\ine pleuropneumonla (CBPPJ, and FMD.
were the Kh<>ikhoi !HommcotsJ. whu are now regarded as The imponanc:e of international trade on live~tock eco·
being closel)• related to the San. 19• ~7 They migrated from the nomics and the impacl of discast' on this trade must be
central interior of Africa t0 1he grazing lands of the coastal mt)ntioncd. The more develop<'d nations. where much of
regions of the Western and Eastern C.ape provincl'S ofSomh the 1radc is cemrcd and a high commercial value for livc-
A[tica. in particular. They were pastoralist,. herding fat· s1ock exi5t>, are prlmarll) concerned with food safel") and
tailed sheep. caule and posi,ibly also goats. and were prob- hygiene, whereas food sticuriry is che major interest of devel·
ably 1he first people to bring livestock illlO ~ouihemAfrica. 15 oping nations..c, 1 his difference has resulted in a greater em-
It is possible that their migratOI') rome, which wa~ mainly phasis being placed on disease eradication and control in
south-westerly, was influenced by 'belts' of tsetse flies and some regions. and imcmational regulauons and rcquiN·
hence lrypanosomosisY~ and they must have experienced mems to ~upport these aims have been implemented. As a
the presence of 01hcr serious livestock disea~e!.. ,uch 11s consequence the developing countrie~ are frequently se-
theileriosisand heartwater. \\ hich were prevalent in the cru;t verely penalized in economic terms ihrough their inabilll")·
of the sub-cominem. This was later followed by southwu.rd 10 trade in these more lucrative markets.''" In e:;Scnce. al-
migration~ of different t!thnic, but primarily Bantu, group, though thc~e poorer nation~ are high!~· dependent on lh·e·
and C\idence of their presence in southern \frka goe, back ilOCk for human survival. their li\·cstock is not pro\'iding a
some 2 000 years.~ 5- 66 Ther had caule of the Sanga (::S-guni) means of escaping from the po\·er{}· trap. Thi~ issue has be-
type as well a, sheep and goals. There is. however. evidence come more prominenl with 1he recent experiences in dt'\-Cl-
of a degree of resistance to certain endemic diseases and oped coumrie) of the effoc1s of epidemics of foreign animal
parasites in caule. sheep. goats and dogs that ha\'C an his- diseases such as l'MD and ASf on their livestock industries.
toric associalion wilh .\rrica. resulting from their exposuru Nt•cent developments. howe\'llr. including some a.~sociated

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 lnfoclfou, disea~esof animul~ in ,-ub·S3hanin ,\fritn:The\\ildlife'llve5ux;l< mtcrracc 22i

with lhe wildlife/livestock interface in Europe and :"ionh wich wildlife is typified by the pastoral sotietics in certain
r\mer!ca. may result in more Oe:1.ibilhy In international trade t\frican savannah ecosystem~.
rules.~ For example, with regards co bovine luberculosis,
transmissible spongifom1 encephalopa thies.65· ,G 8 1D. and Diseas.e, livestock, wildlife and ruraJ subsistence
da$sical S\\ine fever. 1he current comrol policies are being Wildlife-main tained diseases that infect livestock From
hotly debated. an ln1emarional trade poim of view. F\ID and ASF are prob·
ably the most imporiam 1,~ldlife-maimained diseases. and
present che greatest constraint to rural agricultural develop·
The wildlife/livestock disease interface
ment in sub-Saharan ·\frica.
Many disease-causing agents 1ha1 cross•infect 1111dlife and In the case of FMD, the pivocal role playe<l by the African
livestock have been identified on most continents. 1-1 Some buffalo as a S)'lvalic maintenance host was idemified in the
of the more important of these in suu-Saharan Africa. where late 1960s39 -11 and possible maimenance mechanisms of
range and habitats arc shared or where close contact is infection ha,:e been discussed.~ 1 !14. 101 Strict management
possible. are listed in Table 10. l. This inrerface ma) be lin· of the disease around national parks is necessary in :some
ear. as along a dividing fence•line: patchy. reflecting habha1 coumries. a good example of 1,·hich is the Kruger >:atlonal
preference of a disease host on·ector. or diffuse where range Park in South Africa. This Park is fenced and bu.Jfer zones
and resources are shared. The sympauic sharing of range have been creat{'d along its South Afri<.-an btirdC!rs: these

Table 10.1
Diseasas of DISEASE AND CAUSATNE AGENT TRAOIT!DNAt 011 ENDEMIC HDSTISI MAIN EPIDEMIC HOSTISJ
impommce at the foot-and-mouth disease Caltle and Africa~ butfalo Mcst c'o¥en·hooied hvesrock and
wildlife/Jivestock ~L'1DVIIUS ,,,..,ito ,fe
inii!rlace
il'nderpm Caitle. wild n;mirt.ar.:s7 Cat:le ano 'llOSl. wild Afr:can
Mor!>,7111 rus rum;nams and su,ds
Air,can swine lever Tampans IDm1thOaoros1
As~·virus Wild African s~•ds
Can,ne d:stempsr Domestk: dogs Domestic dogs and certain wtl~
Mow,//ivirus carnivores
4f11can hoise s!cl:ness Horses. mules and don~e.,·s
Oro1v,rus
B:ueioague Cenain wild •u-:::na~:s a~d cattle? CarJe steep and goats
OtDNilr.JS
R,h Va'ie~ lever Aedine mosau110es Sheeo. goats, cattle anti certe,n wiJo
Phlebt!virus rumman.s
fl.lahgoani catarrhal lever Blue and blac~ wlldebel!St sheep Ca.He cervids and cerrain capfrte
Alcefiiphire herpesvlrvs I wild !:ovids
(),, ntl lrerpesv11us 2

fl>.l~•es Numero~s w,ld ca ,o. fel1d ar.d

0
Mu,1,p.e
Lvm VI/US I v,vernd sornes ci0:r11m,c dogs
ttea"tvvaier Cenam v111d :u,n,•ams. cheromans Ca,:le. shee;:. goa!sand ~ma w,10
Ehrr,ch,e (Cowa11a/ rumm;;mwm and gallinaceocs b111is ruminants
Bovine petech1al fever Bushboci< Cattle
Cv,oecetes ondili
T,vpanosomosis Wild bovids. sJ1cs. eleµhani anc! Canle h~es. pigs, sheep, g:oa:s
lli!(l(IUS TIYtlilMSOIT'-8 SfJP black 1hmoc;eros ancdoas
'.:orr,dor direase Afncan buffalo Cattle
A!:1tan buttalo-t1erivad
Tne1fer111 paNa
Amnrax Mult,ple
Bae11/us anthrac,s
Bovine tuberculosis Caule Many w,!d ·uminams, su1ds. felil!s
M;wbac1elium bovis and !)limates
Bnn:ellos,s Cattle. sheeo, gca:s. o,gs a"d Africa• buf'aio. eland and
Vanous $.p<!ttes ana b1otypes of manne mammals ,,.a1ertuc~.
Brucel/e sp;1

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228 ,t,·r.>, '"r. Aspects mfluc·ncing Lhe occurrence oi in(ee1ious diseases

lOgcLher with vaccination of caule and regular 1'1\ID sur- ory'.Y·,. ,able antelope (llippozmg,1$ 11iger) and impala
veillance and testing of animals in the buffer 2one~ have (Aepyc:ercu melamµus) are, ior example, the preferred hosts
been reta1ively effective in comainlng the disease in Lhe only ror all three stages in 1he life cycle or brown ear ticks Rhipi·
endemically in fee red area in that counfl)'. cepha/11.~ 11ppe11dic1//a111s and R ::ambezle11si.~).71 Buffalo, eland
\\'llh regards toASF, the elegant ~111dies by Plowrigh1 and and sable anwlope ha\'c also been shown to be iong-1em1
co-workersff2-a,<; elucida1ed the maintenance host and \'irus carriers or several rl1eilerit1 spp. parasi!es. some of which can
amplifkation role of argusid rick,, (Omirl1odoros ~pp.I in cause dii.case In llvei.tock or in certain wlldlife hoSts. The
Eas1 Africa. The role played by 1,ild sulds in th!$ regard in biolo~ and epidemiology of Corridor disease caused by
both Eas1 and southern Africa have also been describcd.a.1 . 9!I buffalo-adapted Theiler/a parM in cattle has been \\'CU
The sinia1.ion in West and Central Africa is. howe1·er. differ- studied. lniection with lhis organism, which I~ ·s.iJeni In
ent as there is no e\idence of a \,~idlife re,ervoir. The in· buffalo. causes high morcalhr in cattle. making fanning of
creasing use of vinually free-ranging pigs in subsis1cnce cacde in the pn.•.scnee or infected buffalo and a suitable
agrkulmre in the coastal regions of.West Africa a11d rt'ccnt ,·ector a hw.ar<lou~ underiaking. 7i Cattle arc general!~
outbreaks of ASF experienced there have brought th!~ lssu,· 'dead-end' ho,1s. being unable ro infoct the lick \'ectors.
once more lO Lhe fore. As a funher example. giraffe Giraffe/ tflme/opardali.<' and
However. at the rural. extensive. communal farming black 1Dic'eros birumls) and white Cem101/wri11m .<imum)
level in sub-Saharan Africa, his most frequently Lhe Vl.'CtOI· rhinoceros. African buffalo and warthog are preferred hosts
borne diseases. such as nypanosomosis, hearcwacer and for adult tick,; of the genus .·\mblromma. whereas the
thelleriosis, that are the greatest disease obstacles 10 local nymphs and larl'lte feed on a variety of larg(! and ~mall mam-
livestock-based agrlcullural devclopmem and prosperit}, mall;, bird5 and even rcpr!le~.-·• Several of these hosts. in-
although certain \•Ira! diseases such as ASF. ~tcr and Fl\1 D cluding buffalo. gallinaccous birds and tortoises hal"e been
also play a role in thi5 respect- It i, in this area chat conOicts shown to bl! potemial carrier,, oi E//rlichia (Co,ctlria rumi-
between livc,,1ock-dependem communitie5 and wildlife 11m11/um, ihe cause of heanwater in domestic n11ninams.-~
most often arise. ns the owners of livestock frequemh· un-
dersiand the role chat wlidlife play in the maintenance of Diseases that may impact on wUdlife h I~ appropria1e
these disea~es. An examph.• of this is the role of wildlife in the that those diwa~cs he considered 1vhich may scriou~traffect
maintenance of rsetse flies (G/assi11c1 spp.J and trypano~o- wlldllfe populations direct!}', or may undNmine wildlife
mosi». llie resultant rcs.crkt(!d acc~s oflivestock to infected management efforts. and ultimately dcwnnine the rare of
areas being a major con,traim on agriculmral de..,('!Op- development of the wildlife Industry and consen-.i1ion ini·
mem. -:o Ccinversely, from a conservalion poinc of view, try· 1ia1ive, in Africa. Diseases ,hat conslitme a livestock threat,
pano~omosis is held to be a major factor fe\'Ouring the such as f~ID. t\Sf and theileriosis. an? generallr managed b~
sunival of many of lhe remaining wildlife popula1iom and the impo~ition oh('\'ere restriction$ on wildlife use. translo-
their refuges in Africa. cation~ and cenain management acti\ilies.
rhe important role plis\'ed by wildebeest tC()11nocl11:rr1e.( foot-and-mouth disease containment mt·asures in
$pp.) in the maintenance and sea$Onal shedding of alcela- several African coumries restrict the translocation of wild
phine herpesvin1s 1. a cnuse of MCF in cartle. has been cloven-hoofed ungulates out of endemically infocted areas.
elucida,ed71• 80· 81 In ea~1ern Africa iraru;mbsion i~ mt)$! thus effecLi\"ely isolating such populations imo ·consena-
frequt.>lll during the wildebeest calving period when pas- tion i~Iand<. South /\Inca also re!!l."s on delineated ASF and
toral cattle. grazing on shared gra,,-sland, come into t-omac, Corridor disea~c comrol areas 10 conrnin 1heir spread. and
with wildebeest foetal membranes or young cal,c.-..88 legislation is in place. which limits the 1ranslocniion of their
The economic- impac1 of chis disease on communitil'< b not natural wildlife hosts ou1 of 1hes<: ar;:-a~.
well defined but is probably limited given the pastoraliscs· On the Olht!r hand. 1here are certain diseases. such as RP.
knowledge of this di~case. t\s a prevenrivc measure. anthrax. rabies. bovine tuber<:ulosb and canine distt>mper.
lfv('Stock are kept off pasture shared \,id1 cahing ,,ilde- which mny have sl(lnificant direct lmpa('tS on the popula·
becst in Maasailand in east em Africa uncil la Le morning tion dynamic, of cerlain wildlife species. on various spatial
by which time the sun and time) has rendered infected and temporal sCillles. Some of the agent~ causing 5uch dis·
placentas safe. In southern Africa the increase in wilde- ease~ appear m be historically alien to ~uh-~aharan Africa.
beest populations associmed with game ranching activities and \\'ere probably introduced onlo the i\frican continem
has seen a resurgence or :>. ICF and tl1is is not ah,'ays asso- \\~th th~ imponaLion of dome~1ica1ed animal i;pccfe,.. lndi·
ciated with direct contact with placentas oryounip,ildebeest genous African free-ranging mammals (within simll,Lr
cah·es. The actual mechanism of transmission oiinfoction In taxonomic groupings 10 the traditional domesticated main-
this latter situation has not rel been determined.0 tenance host) have little inherent g~·rwtic reshaance 10 Lhe~<:
\\1th regards co certain African infectious tick-borne dis- foreign agems. Significanl morbidity and monality may
eases. wildlife mar fill Lhe role of lick amplifiers or carriers therefore be encountered m wildlifl:! where C'On1act with
of the causative agents. African buffalo, eland I Taurormgus iniectcd domestic animals occurs.

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 lntcctious dlscases<>f animals ,n sub-::.aha.rnnAfrica: The ";Jdli(e/h\'CS1ocl: lnccrfacc 229

Astriking c~amplc of a disease with a dcl'a,taiing impact population~ In !:iomh Africarn. 1"· ~ and Uganda, 311• io; 108 as
on populations of wildlife Is RP, currently still prc,em in .-\f· well as in a Kafue lcchwe (Kolm.~ /eche kafue11si$) population
rica in southern Somalia and southern Sudan. The ,ims is In zambia. 3 1 1L has ulso hccn diagnosed in buffalo and wilde-
repmed to have hcen imroduced into Eritrea from India by beest in the Serengeti Narional Park ecosystem in Tanza.
the Italian army in 18Si/88. or by a German military cxpedi· nia.61' Buffalo and lcchwc hm·e become m1e maintenance
tion that brough t Infected catde from Aden nnd Bombay m hosts of \his disease. and sporadic ·spill-over' of infection
the Eal>l African co.ist. : 2• M :Vluch has been wrincn113 l>n the has been documentt!d i11 greater kudu. u. 5 1. wz chacma ba·
grave social, political and economic rcpercus,lons or the boon (Papfo 11rsil111s).''1 lion (Prmrhera /col and cheetah
ma,;.5ive Ciltdc mortalities caused by chis disease as it spread (Acino11rx 11100111.~ ,;r. warthog, 10• 108 eland. topi (Dama/is·
progre~sivelywestwards and soutllw.irds. Countless ,,·ild ar- ,·us lmwnis). giraffe.69 leopard (Pamhera pard11s). common
tiodactylid~ also perished, with buffalo. tragclaphid~ (spiral· genet .Generra gene,raJ. spotted hyaena (Crocuta crocwa).
homed antelope). wild suids and \\ildcbeest bemg most bu~hpig ,Pocamorhoerus lari•ams), and honey badger l.\.lel-
severe!} affected, wirh only relic populations ~un•hing in /i11om cap,msis).'2 The long-renn effects of this chronic pro-
liOme :ircas.Y~The decimation of both c11'1tlll and wfldllfc ap· gressive disease on \\~ldlife hosr populations at ~ustalned
parcmly led 10 \he disappearance of tsetse nies (Glossinn high pre1·alence rotes is unknown. bin earl~· indications are
spp.) from cemti11 areas. and may ha,·c cmuributed 10 wild· that it may negatively cLffect population scntcture and dy-
life dls1ribu1 ion anomalies, such as the formation of isolated namics in buffalo!!<, nnd llon. flovine tuberculosis i" prob -
meta·populaLions of species such as sable antelope and nbl} one of the most serious wildlffe health issues currently
greater kudu. 119 confronting con~ervationists and ,·eterinary regulatory offi-
An incercstlng progression with regard$ 10 RP in wildlife cials In South Afri<-a. 7.ambia and Uganda, and possibly
during the twentieth century has been 1he recog11ition of some other sub-Saharan countries..
,·arious strains ,,~1.h apparem differing pathogeniciry and Canine distemper i$ thought to be another foreign ;mi-
host preferences. The s1tains that were most common!} mal disease which entered Africa in the post-colonial era
associared with high mortality in cattle have been relath·el} and has demonstrated the potential to impact severely on
e3$r to eradicace. hm the apparemly milder strains for cat de certain wild carnivore populations. Less robust populmion~
ha,·e persisted. Although mild in caule. the strain current!} or social carnivon•,. such a~ wild dog (Lyu1on picfll$) and
circulating in Ea~t Africa. characterized as µan of the lineage Ethiopian wol\'es {Canis simensis) are especially vulnerable.
2 group.8 can be high I}• pathogenic in \frican buffalo. iesser This disease and rabies are considered t0 be partially re·
kudu ·. Tmgelaph11$ im/J~rbis) and eland, but apparently mild sponsible fQr the demise o'fthe wild dog in the Serengeti eco-
In some orherspecies such as 1,~ldebees1 and wanhog. How- system in Tanzania O\·er 1he pastdecade.2 Canine distemper
ever, ho,t su,ceptibility appears not to be consistent, nor in has also demom,trated an ability to cross the 'specie~ bar-
fact is pathogen icily c<>nstam with an}' one strain. ~pccies or rier' as severe outbrea~s with monallry have bet'n reported
population of animals. \\itb both temporal and spatial ,·aria· in Serengeti lions.117
tion occurring. Erratic and low rates of vir1.1s shedding orcur Certain endemic multi-specie. diseases that occur on
in cattle. ·\nothcr t<>rni)licatlng factor is the 'atypical' ,·irol- most continent« may also ha,·e se\'ere t>ffects on wildlife in
ogy of some strains with antigenic epitopes poorly detected the sub-Saharan region.
by struldatd test systi:ms. which reduces the sensirlvity of Anthrax ombreaks have been documented in wildlife
sun•eilla nr:e prmocob. populations ,,itl1 no live~1ock link in the Et()shu National
The role of wildlife i11 tile persi•tence of HP has become of Park in Namibia" 2 and Kruger '1/acional Park in South :\f.
i11teres1 in the last decade during the final phase~ or eradica- . 11 z; 29. ,, Large-sea Ic out b rca""
n<ia ,.. may cros.H I1c Imer,ace
'
tion of th!! viru,,, \\~th the identification of 5trains that especially where domesticated and non-domesticated spe·
appear to sprec1d efficiently e1-en in panially immune popu- cies share range and resources. as has occurred in the
lations. This may be the key reason for du: JJcrsi,;tcnce of the Luangwa Valley :>:ationai Park in Zambia.2ti nnd Serengeti
strain in East Africa despite massh·e vaccination campaigns ~ational Park. 611 and more recemlyin \Iago :>:ational Park in
on the continem between 1960 and 2000. rree-ranglng wild· Ethiopia, where a die-off of many cattle and more th,m I 500
life and pastoral livestock ca11le espcciall}· in times of lesser kudu \\llS documcnted.37 9 ~
drought. may be n key epiderniologkal factor. \Jost l'\.i· Rabies is another multi-species disease that is endemic
dence. hO\\twer. is that the wildlife is only acting as an indl· in many areas of sub-Saharan Africa, with sporadic epi-
cmoi.and vector of this disease. which is in fact maintained demic cycles. It has been diagnosed in 33 carnivorous spe·
in cat1lc populations. cies and 23 herbivorous specie~. \,ich a regional variation of
Bo\·ine a1bcrculosis caus~d bi· Myt·ol!actt!'riwn bo1,i$ ls dominant epidemiological rolc-players.9 5 ln spite of rllls. by
another foreign animal diSell$e that was most probablr in· for die largest number of rabies cases reported in the devel·
troduced into Africa \\ith imponed dairy and Bos ra11ms- oping world occur in domestic dogs. \\ith 'spiU-m·er· imo
type bcei ca1tle during the colonial era. This disease has now Other domestic li\'es1ock. humans and wildlife. Rabies and
spread to. and become endemic in. several African buffalo canine distemper are thought to be partiall)• responsible for

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230 ,r<:Tio, ..,~.,,: A~pects inlh1er1ch1g 11111 uccurri:nc~ c,f infectinu~ tli'M!.sSl!S

the demise of the wild dog population in the Serengeti eco- tions arc unhelpful. In the Botswana study some fences were
system3• and is considered a major disease threat 10 the successful in the control nf diseases such as l'MD and CBPJ>.
highly endangered Ethiopian wolf. In Africa. endemic rabies and were effective in reducing animal 111ovemem. The im-
(caused by both viverrid and canid biOt)1lCS) is often asso- pact of tl1ese fences on \,ildlife was based on population
ciated with certain communal burrow-dwelling species, 1rc11ds. which did not show a c-011s1s1cm picture across all
such as the yellow mongoose (Cy11ic111.f µe11icil/nt(I) and the $pecies. the implication being that some species benefit and
bat -eared fox (Otocyo11 mega/oris). as well a, jackals 1Ct111fs others do not. In the end 1hc environmental. social and
spp.). Rabies outbreaks in certain host species frequend~ financial costs of sonw fences may ourweigh their benefits
appear to be density dependent. In u unique documented but the decision on this will depend on complei- anal} ses.
outbreak or rabies in kudu. where non-bite transmission including em'ironmental impact assessments. The recom·
probably occurred as a result of salivary conramination of mendaiions of a study bv Scon \\'iison in ~gamiland.
browse, more than 10 000 kudu dit>d over a four-year period Botswana. 91 included the remo\'al or realignment of ~ome
in Xamibia.• Abnormally high population densities of kudu fences and the endorsement of others as effecti\'e and
were cited as being a majorcontribuiing factor. necessary.
There b aho tht< realization amongst e.:ologists that is-
Fencing and wildlife Sever-di of the diseases associated land communities of wildlife are unlikelv 10 ,,J.Ni\'e iman in
with "ildlifc 1ha1 have major epidemic potential and there· the long term.;-3 In recent years. this has driven some con-
fore alfect international trade in live~tock and their prod- ~ervauoaist~ 10 think in the direction of romplex cons.m-a-
ucts. are highly contagious viral lnrections. These discnse~ lion systems. including traMfromier ,m)as invol,ing
may have severe adverse effects 01\ livestock-based export communal. privme and public lands with corridors along
economics. It is particularly in this contc.xt that the \\ildlifel \\'hich wildlife can move. and dilferent land u,;e.s "ithout
livestock interaction is crirical. The most significant or these nece%arily excluding human ac1ivi1ies. 9 Thi~ approach is
diseases are FMD. RI> and :\SF for which fencing has been a intellectually appealing but highly theoretical, and does not
major tool for control by restriction of animal movement. take into account basic humanocemric ,ocial and economic
ff wildlife carriers or vectors of these diseases arc allowed realities. and it certainly has no consideration of disease-
co range freely in contact with Uvesrock. creating a diJfuse control implications. Nc\'erthdess. the impact of fencing on
Interface. then there is little possibility that exporter:. will be mignuory populations, such a, wildebeest in southern Af·
able to develop trade in live animals or animal products be- rica. has prO\ided stark e,idence of the deleteriou~ effects
yond 1ht:iir immediate region. certoin policiescnn have on popula1ions.;n, 59 The debate on
In certa in coumries in southern Africa, where there is a appropriate conservation policies in Africa will go on and It
well-developed livestock production industry. the me1111s of i~ definitely not the intention here to criticize or promote
pre\'Cnting the maintenance and spread ot these diseases any particular approach. but only to highlight animal dis-
in livestock has been based on the separation of wildlife ease aspects that need to be considered, whils1 as~essing
and livestock b) strict land use policies and fencing at the risks and impact of policy change.
Interface. Borswana, for example, has a lucrative trade with Decisions made by governments pertaining 10 livestock
Europe based on an early and dear understanding of the health are affected b)' the present glohal re$lrictions on
epidemiology and control measures necessary fc)r contain· trade as set out In the recommendation~ of the Internacional
ment of P:-10.a• As the economics of wildlife enrerprises Animal Health Code of the Office International des Epi-
haYe proved to be more competitive in marginal, often 7.ooties (01 F.). For certain di~eas<:S, this Code does support
semi-arid areas. rhan caule ranching, there has been a uan· the concept of zonation of infected and non-infected area:,,
sition of la1\d use from lh·estock co multi-specie~ or e.xclu- \\'here appropriate e1oi11rol measures are in place at the in-
sive game management system~.24 The Fencing was already terface. TI1u~. wi1ho111 a fencing polky. commercial lil'e·
la place in many of these areas, and only game-proofing up· stock industries in unrestricted wildlife areas cannot
grades were necc..~ary. Game fences are also popular with de\'elop in the conventional sense.
commercial ,-.'ildlife ranchers due to the value of individual The concept ofspcciescompartmen1al.iza1ion is also cur-
wild animall. and their need to retain them on their proper- rently being discussed al the OJE. This concept accep!S 1ha1
ties. The fencing debate has beconw more comemious ns If an outbreak of a listed diseasl' (all of the OIE list A. and
the ecological impact oi fencing has become beuer appred· some list B diseases) occurs In a tree-ranging wildlife popu·
~ed Pence~ are frequently crltlciied because \\ildiife are lalion. but docs not involve domestic ll\'e.st0cl;. then the re·
precluded from following hls1orically importam migration porting country should 1101 necessaril}' be penalized br
romes, which are netessary for survival in ecologically mar- agricultural trade restrictions. Some ~omhern :\frican coun·
ginal areas \ I here surface water is scarce and grazing a,·all- tries have taken a \'igorous approach to disease control and
ability is seasonal. Detailed studies of this real or perceh•ed \iable ll\'estock industries ha\'e developed
problem are scarce and reports that exisr on the emiron· In eastern Africa the more laiss~-/airea1ti1udc remains
mental effecb of fencing in Afrlca;8 · 'll sho\\' thal g,meralizn- a conMraint 10 commercial livestock development. and

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 lnfoWOlll> di,l'a,e, oi animals ,n ,;ub-Saharan Atrica: Th~ wildlife, lll'es1oc:k inicrfacc 231

governmem, are under increasing pressure 10 ~eel- eco- where Lhcy are the onl> definitive maimenanc:e host for a
nomic solutions to the Increasing poverty. The issues oi specific \!lsease. The role IJf wildlife as re,er.oirs or ,•ectors
land use. disease comro1 and fencing have 1101 bel·n rc- of disease arr:: sometimes misunderstood and ma\' be over-
~Ol\'ed due lO conflicting economic in1eres1~ invol\ing the stated. even among \lt:terinarian,.
e'(panding and lucrative 1ourism induslry. Ennourism. \ lany pilstoral communitie.s are poorly supported by, e1-
based un game vicwiJ1g of free-ranging wildlil'e and envi- eri11ary services. ir at all. so control through vaccinauon o(
ronmental appreciation. certainly provid~ ,ignifkant dome~tic .animah i\ rarely possible. The issue 1$ whether
t!Conomic benefits to a region, but frequently. only to a these free-ranging Hveslock and \\ildlifo populations can be
small propor11on of the community.n Profitablc ecorour- isolated from commercial ranchin1t.to enabk• a sub-sccroror
ism and well-marketed environmental appreciation have the livestock indu~1ry to function and trade. In ea,tem
also made a major conrrlburion ro the maintenance of the t\frlca. for example. without effectl\'e movement control.
world heritage of animal life. and indigenous biodiversity. there is Uule likelihood of FMD ever being controlled tiue-
A, a result there has been reluctance by go,·emmcnts to spcctive of \\ildlife). :-.:evertheless. wildlife doe, present a
tackle the lives10ck problems. \\hich ihey belle\·e would further complicating factor because. even if impmv<.'d vet·
Impact on the wildlife estate, e.g. through fencing. Ironi- erinarycomrol such as vaccination were initiated in pastoral
cally it is lhe very d~ire to keep lhe ~y~1cm~ open that re- livt!,tod.. wildlife could ~till harbour pathogen~ and ,o
sults in lhe easy access by livestock to the grazing rr~ourccs mai11taltt a rcs1:n·oir nfinfection.
in protected a reas, leading not only t0 intensification of the Pa.s1on1!1sts frequently have extensi\'e ethno•botanical
interface between livesrock and wild life, hut also to con- and ethno -veterinaf) knowledge. The)· understand many
flicts bctw<!en humans and animals, inrfuding predation infectious disease proces.~cs and often treat animals suc-
and disease impacts. In practice. "~th rapidly growing cessfully without sophislicated medicines.21 u . 111' Often
human populations. fencing of wildlife-protected area. b simply b~ the abilit)· to move stoc!; rapidly over long db·
ine\·iiable and l>eneficial in many aspect.;. Fences al,o ef- tances, thev avoid contact with dbeased animals at appro-
fectively reduce predation anti depredation on liveslOck priate times. bu1 this cun equally assist in spreading
and crops of adj<Jining communities. which are often of a infection. Thi, lmowlcdge-base also e.,ist~ at the wildlife
subsi$tence nature. Some negative ecol<>glcal effects of livestock imerface, amply demonstrated in the case or
fences may have co be accepted in order 10 ma1main, iable MCF°8 by the response of livestock keepers in avoiding
populations of key African wildlife species. cnule comact with wildebee~t par1icularly during the
cah1ng ~eason.
Transhumance, pastoralism and wildlife 1\uempts to improve the deliwry of veterinary sc~ices to
The key constraint to achie,ing an impro,·ed animal heal ch pas1oralis1s \,ill continue in many countries, and a spin-off
siacu~ In much or Africa that would be acccplablc 10 the may be reduced disease challenge to wildlife. fhis challenge
maJOrity of livestock trading nauons, is the current unre- will, however. remain mherently difficult as delh•cry of ex-
stricted trans-boundary moYement of people, lhe~tock and penise and v.:terinary , 1ac:cin~ and medicine~ is logistically
wildlife."' problematic and economically non-sus1alnablc unless li\'e-
The animal health code de\'elope.d by the OIE." 6 de- stock value~ and nmrk('tS improve. Therefore. diseases such
signed to reduce disease risk and raci litate international as P.,,11), ASr. anc! COPP are likely to persist for rhc fon~:,ee-
trade In livestock and livestock products. has become com- al1le future. It is only after 1.onation of land as prnc1lscd in
plex: and requires a high le,•el of managemem and invesr- southern Africa, whic'h identifies problem populations and
ment in ,·eterinary sen1ces. Although thb code has greatly regions. and the designation of tl1ese a, tli~ease control
enhanced the economic potential of the livestock indusrry. areas. that the resultant Mrict mo\ement control and re-
it has also set rccommend:uions and standards 1,ith which duced contact between ,,ildlifo and li\·estock will impro,·e
many African countries cannot comply. Without the ben· lhc simation. It i. historicall~ intl.'rc,ting chat modem
efits of international trade. the value of the livestock ,ector human society ha~ ~cwn the decline oi the hunter-gatherer
to these national economies has not improved and this has communities and most pmbably wiU see the rrext 1ransition.
deterred the govemmems concerned from maintaininl'( in- which will be the disappearance of true pasroralism."JJ. •"
vestment in the industry,9 ~ In addition. as pastoralists are
often politically marginalize<l.~Qthere has been linlc incen- Wildlife in d us t ry and conservation area s
ti\-e t.o shift resource~in their direction. 1'he three categories of land 11,e in,·oh~ng wildlife are essen-
In re,1ie\odng the ,\ildllfe disease impact it b important tially:
nm to over-emphasize the role of non-domestic arliodacryls • national land set aside for conser\'ation of wildlife.
because, In pastoral systems. there arc frequentl~ adequate habiuns and ecosystem processes:
populations of fret'-ranging lhestock to enable the mainte- • private land on which a variety or\,ildlife-bast?d
nance of these infections in the absence of wildlife. Thus. acti\~tie., arc ccmduc1e<l; and
wildlife population~ are not always a critical iactor. except • communal land with a wildlife-use element.

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232 srrtm'< ro,..-; .\sp~cts influencing the occurrenre of infec1iou< di,-eas,s

Formal nature conserva1ion In Africa is only ~lightly more a, foot ro1. ,·em1ino~e,. coc:cictio,~. and nutrition-related
1han I 00 years old. The ear!) conserva1ion initiatives were problem~. are more pre\'alent. On small game ranches.
\\~lhin national programmes. with land scl aside e~dusi\'ely where 'free-range· wildlife arc often fed supplememmy ha)
for na1ure conservation. In broad per~pccth·e, the fir5t in the d11· ~ea~on and during droughti:. a common problem
50-odd years was n period of consolidation. during which 1hat ls encountered ts relati\'e O\'erstocking. which re,uhs in
recreati<>nul 1ourism was only a small consideration and a the so-called sick habiiar syndrome. This is a multifaccorial
limited income ea mar. The second half of the l\\'entieth cen- ,yndrome related to progressh·e depletion of palatable
tUl)' sa\\' a mpid growth in the tourism indus1ry in formal grasses and browse, hush encroachmem. soil ero~lon, n pro-
conservation areas. \\'hich coincided \\'ith 1he de,·elopmem grnssi\'e build-up of ctto- and endoparasiu:s, and stress and
of the firs1 private lodgc:s and privme nature reserves. ~·luch lick 1oxicosis,related immuno-$uppression. TI1e net re~ult is
of this growth was based on foreign visirnrs aud facilltated a loss of body condition. anaemia. tiC'.k-bite abscesse> and
by improv~ travelling opportunitic~. aud in both formal monalities that may be assucia1ed \\ith nom1ally benign
and pri\'atC sectors. facilitie, and opportunities for touris1s agents. The relative· t1vcr,rocking due to the inabilit> of the
increased sharply tO\\'ards the close of 1he cenniry. :>:ational game to move out of the area. also fal'ollrs the spread of den-
and international demands !or wildlife consen·alion still siry,dependcnt infectious dbease».
continue but as human population growth continues. with .:Vlore e~1cnsive syslcms. !>Itch as national parks, game re-
Increasing hunger for land, conflict arises. :\s a result there ser\'Cs and gam<' consen·,1.ncle~. mar 1~, in size and may be
has been a recent !rend towards communily-based conser- fenced or open. The more narural the ecological system. lhe
vation iniliative~.9 · • 3 Biodh·ersi1y conser\'ation 1,i1h 1he less likely lhal a single dlsease entil) 1,ill be problematic. since
presen·atlon of local lh·ellhoods I i~ an example 01 some of 1he relationships between hosts and indigenous parasi1es
the recent thinking. fhis thinking has e\'Olved as academics should be relativelybalanced.11 In 1.hese systems it is predomi-
believe that previous attempts a1 conserva1ion through nantly rycliral climatic factors and related animal population
national parks and protected area.~ have been ecologically densitles which l<?ad to 11:\'J)ress!on of di•e:tSe. for example dur-
inadequace or unable to contribute siiinificamly to human iniand alierseveredroughis. Undenhesecircumsmnces. \\ill,
development needs. the clumping of large concentrations of herbivores around
In lhe priva1e S!!CtOr. differences in land tenure policies d\lindling regources. and concomitant nutritional and social
becween countries in sub,Saha.ran Africa and the righ1 of s1rlll>-s and immuno-suppression. parasiw burdens may in-
prlnne in.dividuals 10 own land and wild life has fundamen- crease to pathological levels \iral and bacterial dlseru.5 are
cally affeccecl 1he way In \\hich wildlife is valued. In some also frequently associated 1,it11 these Cll\ironmerual stress fac-
southern African countries where private cmnership ofland mrs. e.g. anthrax. epidemic ~lD and possibly RP. 1~· z; llow-
and 1,ildlifc is enabled by lhe coumry's constitution. exploi- ever, truh na1t1ral CCOS}~tems defined as having minimal
tation of wildli fe resources is gaining momentum and be- human lmerforent-el are now rare. aJtd a natural state in re-
coming a major economic acti\'it)'. t\s a result. borders spE.'Cl or disease ecolog\' is probably the exceprfon.23
between enterprises inl'olved in ecotourism and fannland. Looking to the fu1ure, disease problem~ in the 1\ildlife/lh't'·
as well as publicl)' owned 1,ildlife areas and tradirional com- ,tock interface are li.l:elr to increaw and possibly intensil) in
munal areas. are becoming increasingly sharply demar- some regions of .sub-Saharan Africa. as many are related to
cared. more rect·m conscn•aiion dc1 elopment,. such a, the ne,,
Eas1 Africa is quite tlifferem because the land use system communit} ·bll>t'CI 1,11dlife inltia1lves. large-sca!e transloc,11ion
is much less developed. there are fewer fences and con;ider- of l:ey wildlife speciE>.s, enlarging consen.mcies and l13flsfron-
able mixing ofpeople. livcstork and wildlife. The main wild- tfor parks?' Wilh regards to transfrontier consen'l!tion areas
life use is ecotourism, which Is based on a complex of public ITFCA.~l. h i~ 1101 the intention here 10 por1ray these initiati\'es
and. priva1e conservation areas. as well as subsistence and in a negam-e light. From an emiromnemal conservation.
some commercial huming. biodlvers!t}. heterogeneity and ccotourism point of,1e\\, the
In West and Ce111ral Africa the situation is more complex imegrarion of land :icro~s imernational borders as well as the
with, on paperac leasr. a mulcitude of parks as \\'ell as hunting consoiida1ion of sra1e and pri1·ately owned land in joim ,·cn-
rescn·cs. HO\\'ever. non,,ustainable C1'ploitation has resul1ed tures is Jm,tifiable, and may have major economic benefits. The
in a massi\'e decline In numbers of wildlife in these conserva- m<K~age t11at need~ 10 bo: coswered, however. is thal ail in-
lion arn:is. The result is lhat. rnlle~s this trend can be re,·ersed voh·ed parties and role players must approach these inittatil·es
io these' regions of Africa. the remaining1,ildiife will be isolured fully aware and forewarned of the potential animal health im-
in smnll me1.1-populatio11$ wquestcred in resen-es. plication, aud challenge<, that may be expected when increas-
The disi,ase problems assoda1ed ,,1th the va~ing ap- ing the geographic range oi \'arious pathogens. The possible
proaches to wildlife land u,e are also different. In the mo.re implication, for mtinlal dhe~.: c:ornrol are profound and lhe
intensively managed systems. e.g. game farms where ani- potential impact on thu economies of the region through to~s
mals are in pl'ns or small camps. and are fed artificially ofliJ'e-su<eaininglivestockandlortradeshould be calculated. ll
throughout Lhe year, the production-related diseasei.. such is totally prcdktable that ,11thout imcmational bounda11·

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 lnfccuous d !:w.,51,;; of .inimah in ,ub-Sah.iran Mricn: I he \\ildllr~. '"·~rnck imcrf:tce 233

banier fences and with a 'biological bridge" fom1cd by contigu- auendant parasites arc diffit'uh if not impo,~lblc Lo retriew.
ous \1ildlife populations. any infoctiou~ <li~cas(I present in an\' Thus strict quarantine. disease-screening. and ecto- and
one of the participating couimics or areas will C\-emuallv endoparasilc comrol are cssenual requirt:>mcm~ for the
spread throughout the cnrin.> consen,ni<>n area unle:.s con- re~ponsible trnnslocati1>n of \,ildlifu. 1<19 Disease hazard
tainment or control measures are cstablisht'Ci. For example. in identification in founder popuia1iom,, follo\\'Cd by risk a5·
the recently crca1ccl Great Umpopo Transfromier Parl<. which sessmem and risk management s1ra1cgie.,. form the hack-
incorpora1e.~ consen•ation are& In :\to;r.ambique rGa;,,a :S:a- hone ofregulatory disease control for wildlife 1ramlocation:.
tlonal Park). South Africa Kroger "1ational Parkl and Zim- on a national and intemmional level. Endemic disea5es and
babwe (Gonarczhou :s;ational Park' . and \~hich will e\'cmually parasiies in thc ecosystem at the destination oftrnnsloc.ired
constirute one of the largl'st con~ervarion area5 in the world. ,mimalb ma~ :il~o influenro the succe..s of these exercise~.
there are animal disease risks 10 all three regional partlcipams. and ~hould be taken imo considcra1ion.
Once imernatiorral baniers ha\'e been removed and contigu- A current major Issue is hm, 10 enable increasing com·
ous wildlife populntions have become cstabll~hcd. b01ine tu- ml.'rciaJ cxploirntion of wildllre r~l)urce:. while at 1he same
berculosis may spread nonhward, and ~a~twards from the tinw ensuring that the local communilies do 1101 become
infected Kruger National Par!< componem. tSl'tse rties ;Glos- dispo~sesscd. It is :i difficult balancing act 1ha1 Is being tilted
sina morsicans) mai' recolonize the Kruger Park from thenonh. b}' Africa'!. ra1>idly growing human populations. The Camp
rabies may enter this pal'k from the north and nonh-east, new Fire Programme In Zimhabwe07 and rnlatcd community·
topO[}'Pes ofFMD \lirus from Gonare7.hou National Park may based conser\'ation initiatives clsc.•wherc have achieved
emer the other two parks. and caule-adap1ed '/11ell1?11n p<111-r1 some limm:d success in enabling local communities to par-
mairmo,·esoulhwards and westwards. \\iith this scenario, pro- ticipate and ~hare in 1he bencfil'~ of eco1011rism and com-
active disease management srra1ei:ies need to be discussed at mercial hunting. ~evenhelcss. theni is no doubt that more
high level by participating countries and agencies. progress on this from needs to be made if the confiicling de-
There is also a trend to create larger ecologically sustain- mand~ Of \\ildlife con~en•ation. commercial agriculture and
able conservation areas (conservancies) by the pri,-ate deprived rnral communities are lO b<> reconcilcd. 41;
sector. In marginal cattle farming areas, well-nrnnaged ec(>·
tourism based on "tjldlife is potentially more profitable than Disease su rveillance and con tro l at the inte rface
livestock rarming. Farmers in rhese areas are increasinglv I he responsibility for disease surveillance gencrall) rests
abandoning traditional cattle ranching for commercial with the: ,·eterinary cegulatory amhoritie,,of a given country.
\,ildlife-based activities, pooling their resources by creating and the sun eillance 1cd1111ques used include pas~ive rc-
jointly owned reserves enclosed by subs1amial perimeter poning. iarm Inspections. problem in1·es1igmions. nbauoir
fences and with internal fences removed. 1 These consc•ffan- ,un'O) ,. ,erological sun·eys. and dedicated testing for spe-
cies may be \'Cry large. and several in South Africa and Zim- cilk di~ease eradication ,cheme,. Unfonunately. legal
babwe exceed 100 000 hectares..,, This does not. however. frameworks and respon~iblllties (Including financial re-
preclude the keeping of \1ildlife and livestock togetlwr, which ~ponsibilitie~, for wildlife disease sun•eillun,·e. inve1.tiga1ion
is often done. but curren!lyaccepted '\\isdom· has it Lhm pro- and reporting arc nm clearly defined in many ,\frican coun-
ducdve livestock fam1ing and ecotourism centred on \\ildlifo tries and 1ree-ranging paswral c::nule and wildlife do not e-0s-
are incompatible on the same land area 3S a result of the ily lend th<!m$~ive~ tu monimring.
higher risks of disease and for aesthetic reason,. '\O. -:- Given the depressed state of veterinary ~er.~ce~ in many
This semimem is probably true for much oi Ea,a. Central countrie, m .\frica. it is clrfficuh 10 envisnge anr syMematic
and \\"est ,Hrica, a~ well a~ parn, of the c.sstem lo\vland~ of collection of ~u1veil!ance darn from li\'<.>Sto<:k in remote
southern Africa where many or the wildlifc-as~ocimed live- area~. The bt'~t hope for ch<>,e area, is c:ommunity-bac;ed
s10ck diseases are endemic. Thi' den•lopmem ofchese large approaches 10 monitoring animal health. 4- On the wildlife
consen·ancies sometimes conflicts with Lhe polides of de- ~ide. a growing im(•rest b~ wildllfe managers in wildlife dis-
veloping coumries, which place a premium on agricultural ease, and their po1emial impact. b impro\ing information
and Industrial development. and has led m the critici~m that flow ,md should be imcgra1ed in.a conventional epidemio-
\,ildlife is held In higher regard than people.''8 logical reponing :.ystems.
These ecorourism-based enwrprises generally require a The nred to impro1e the comro! o f animal disea$es at the
wide divers!{'.\' of game species and adequau: popularions of interface is a complex evohing issue. Comrol strategics will
them, for easy visitor \iewing and frequently necessitate lncrea~ingly have 10 un;ommod,.ue the conflicting require·
large-scale n'am,location of wildlife. which in itself may have ment~ oi a varlet)' or land u~ers and lr11(1rest groups, and
inherem disease risks. Translocation of any animal Is. in adaptive policies coupled 10 pro-acth·e thinking and appro-
fact. the cranslocation of a ·bio!o_gical p.ickage· consisting of priate pla1\ning are nect!ssary to pre-empt the developmem
the host together with its nrumdant macro- and micro-para· or ~crious animal diseai.c evcmts. 100 lnno\·a1ive possibilities
sites. Furthermore. it is important co realize that once re- should be examined. For instance. iollo\,ing the repeated
leased into a free-ranging system. such a.nima.ls and their failures 10 ck•velop an ell'l•ctive ,\SF rncdne. would it no1 be

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234 ,ec:111" "~" Aspeets influencing !he occurrence of iniec1ious diseases

more advanrageous and appropriate for de,·eloping coun- Uganda, and lechwe in Zambia are examples.
trie$ that do not export pork, 10 embark on a programme of in general, control or diseases oflimited epidemic potential
selection and breeding of pigs that arc gene1ically resistant is best addressed at the local lcYel. "ilh the assistance of veteri-
10 ASF in order to adcires~ this disease problem in subsis- narian& and wildlife expe1ts \,1th detailed knoi.vledge of epide-
tence rural communities? Another innovat Ive stramm• is to miological and etological detem1inants. For infections
select or breed wildlife that are free of cenain diseases. and capable of causing 1rans-bounda1y epidemics, comrol needs
10 use them for stocking or translocatlon exercises. The to be co-ordinated at both national and intemadonnl le,,.els.
breeding of African buffalo free ofFJ\.fD. theileriosis. bovine tu- \\'bile short· and medium-term disease control and long-
berculosis and brucellosis. for example, is becoming an eco- tem, eradication goals can be, and have been. attained in do-
logical success and a profitable enrerprise in southern mesric livestock using judicious vaccination programmes,
Africa.12 vector conrrol and test-and-slaughter policies. these options
That serious disease events are still occurring is amply and techniques are frequently impractical, difficult to execute,
demonstTated hy die currem pro~lems associated with and mai be culmrally or morally unacceptable in free-ranging
FMD, MC:F and bo,'ine tuberculosis control in sourhern pastoral livestock and indisenous wildlife populations.
Africa.3 28 and the issues of RP (see above). .PPR and ASF In Wirh regard to the multi-species diseases. such as an·
eastern and Central Africa.60 thrax and rabies. many ecologists will argue that these dis·
When dealing ...,1th the threat of certain endemic African eases are also endemic and part of the greater ecosysrem.
diseases. such as F.vl.D ••<\SF and buffalo-derived theileriosis, and because they function as namral population regulators.
the containment option has given best results to date. This is rheyshould not be conrrolled. However. both these diseases
usually effected by mean~ of control r.ones /areas, game- also affect domeslic animals, and have signi.ficam zoonotic
proof fences. physical barriers. cordons and movement potential. There a.re now few if ani ecosystems that are iso-
controls wh ich eftectiveli separate the wildlife from domes- lated from humans and domestic animals, and modern so-
tic lh·estock, thus preventing contact at the interface. and is cieties will no longer accept che impact of disease as namral.
characteristic of counuies with an advanced land-use This renders the laissez-faire argument redundant. For
policy. It is not an option in regions where pastoralism with these reasons. conrrol of anthrax outbreaks in wildlife have
free-ranging livestock and subsistence agrlculmre are the been attempted, using various 1echniq1tes including burn-
nonn. ing/bmying of carcasse5. sua1egic veld burning, waterhole
\\'hen dealing with endemic arthropod -borne infections, disi nfeclion and remote vaccination by means of disposable
such as c:rypanosomosis. Afrkan horse sickness (AHS), RVF dart~ or bio-bullets. These attemprs have mer \\1th \"a~~ng
and bluerongue. containment Is less likely to succeed. and success. On the other hand. mass vaccination of foxes
vaccination and vector control. v,here possible, mai be in- against rabies in Europe, using oral bait techniques, has
cluded to reduce infection andtot tra11smisslon. been highly elfoctiveand successful. 20 Central 10 the reduc-
With regards to alien /exotic diseases that threaten free- tion oflivestocklosses and human exposures during anthrax
ranging wildlife populations. such as RP. canine diMemper or rabie:; outbreaks in wildlife. are large-scale public aware-
and bovine tuberculosis, comainmenr and control can best ness campaigns and mass \'l!Ccination Of livestock.
be effected by addressing the disease in the domestic host by In the case of multi-species producrton syslei.~~. where
mass vaccination and /or test and slaughter respectlvely. In livestock and wildlife are farmed together, certain precau-
addirion, prevemion of contnct becween infected domestic tionary measures may reduce the possibility of ~ignificant
animals and wildlife is desirable, but w1fortunacely not al· disease events occurring: rhese include:
ways feasible in many countries in sub-Saharan Africa. In • avoiding_ inrroduction ofcerrain key wildlife species. which
tl1Cl'e siniations. impro\·fng the delivery of animal health are maintenance hogs or 'carriers of certain indigenous
services (e.g. through vaccination) to domestic livestock diseases. e.g. FMD. theileriosi.s. \!CF. AHS and ASF: and
sharing ranges with wildlife and/or adjacent 10 conservarion • screening all canlc for bo,1 rle tuberculosis and brucello-
areas would probabl}' be most beneficial if a suitable \'l!CCine sis prior 10 introduction onto a ranch.
is available. Rinderpest control has been based on vaccina-
tion and the benefits of eradication are economically signifi- Some positive aspects of such a mixed farming enterprise
cant even wi1hou1 calculating the beneiilS to ,\ildlife should also be hlgWighted; these include:
populations. There is the added benefit that residem wild • the ·vacuum cleaner' effec1 created by the regular dip·
animals may also act as sentinels, providing spatial and tem- ping of the cattle component in order to reduce environ-
poral (based on age srratification) dam to facilitate disease mental tick burdens;
surveillance and identify existing and new fod of infection. • 1he reduction of parasiric o,-d and larvae when ingested
\~bere an alien disease hl!S become t>.stablished in a conser- by non-patent species: and
vation area, the situation Is serious and control options are • an appropriate comblnarlon of cattle and "~Id ungulate.,
limited. and frequently comemious or unpopular. 10 Thebo- for a particular veld-type will ensure that a wider diver·
vine tuberculosis situation in buff-alo in South Africa and sity of namral plant species will be consumed. which

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 l11fec1ious disease, of animals in sub- '-aharnn \frica: 1 he" ildlirtr-Uvesrock ,merrace ~5

will op1imi1.e 1he utiliia1ion or different strata of the The creation oflargc wildlife land-h<>ldings (e.g. transfron-
\'egetation from ground level to 1he canopy. This may ticr parks and conservancies) accompanied by the intro-
effec1ively increase produc1ivi1y for the ranch. duction of species such as African buffalo, "11.d suids and
wildebees1 into cenain area,, may ha,·e a negative impact
on 1he regional lh·es1ock trade as a resuh of disea~e risk fac-
Conclusion
tors. With 1he diversion of finance from namral and animal
The long-running conllic1 b<.'1ween livestock owners and re5ource ,ec1ors ro human \\1elfare, health and education
animal health authorities, on one hand, and wildlire conser- (a tr<md ii, .\frica over recem decades). 1he risk of losing
vationists on the other, Is largely based on differing anitudes control of animal diseases is increasing, and this mar com-
to controlling diseases of livestock associated \1ith \\ildlife. pound the issue. The increa.~ed use of land for mixed \\'lld-
An au empt is made in this chapter 10 highlight the key dis- lifc/!lvcstock systems in areas previously subdivided by
ease issues and to emphasize tha1 these diseru.e problems fences may also increase levels of epidemic disease. The
are frequently bi-dirt'ctional at the interface. A new dimen- question is whether co11n1rie~ can afford co ignore thi~
sion currently faced by veterinary regulatory authorities is problem. however ideal and a11racti\·e 1he theories.
the spcc1re of new syl\'atic foci of diseases. such as bO\ine If pOH?rty alleviation is a goal and a necessary step lO
tuberculosis. bmcellosis and possibly RP. 1hat threaten 10 ensure consl?rvation or resource~. a pragmatic approach to
derail national and incemational eradication schemes lhe economics of land use may in fact be more lmponam
which ha1·e been implememed and executed with signifi- to consen·ation 1han recreating 'Eden', 11 should also be
cant success and at great cos1. kept in mind that until rdatively recently, man} of the de-
Conversely, wildlifo-based ecotourism in Africa has ex- velopment~ in wildlife con~eT\'ation and utilization were
panded rapidly ovenhe past few decades. and Is the source initiated and comrolled by communuies of European on-
of much-needed foreign re\'enue for man> •.\frican coun- gin, and that the!>e may not be su~1ained with the changing
tries. Traditional subsi$lence farming l~ still the largest social and political s1aw s of countries concerned. Equally,
source of dietary suslenance on the continem, and this 10- the!re appears lU be a n increasing realization among world
gerhcr with the growth and htUlger of historically disad- leaders that globali1.a tion b perpe1ua.lng the dominance
vanraged communities for land. fs forcing enterprises and of the developed world and that there ls a genuine desire to
communities with marked!~ different objccrives and land a<ldrc:;s the issues of poveny in Africa. This may lead 10
use practices and priorities to operate ertectivel}' in close change~. which will facilitate trade or livestock and their
proximlty. In sub-Saharan Afri(';t the interface between products.
"ildlife and livestock 1s declining overall as a result of pro- Whatever the auemp1s at global solutions. the changing
gressive decimation of wild animal popu la1ions. :-.:e\·enhe- land-u~e pauem~ in m11ny area,, of Africa will require ongo-
lcss, in cenain areas there is an incensification of rhe ing and focu<ed resear('h 10 further elucidale ihe cpidemio·
imerface. as shared resources become scarce. and rhe de- logical decerminants of many diseases at 1he wildlife/
1-elopment of wildlife and livescock enterprise3 Increase livestock interface. in order 10 develop inno,·ative comrol
1he shared use of land. In southern Africa, over recenc s1ra1egics rhat do 1101 limit land u~e options of comm uni tie!>
years. the trend has been contrary 10 the rest of Africa. \\1th ora countrr's ability to market animals and animal products
an increase in use of land for wildlife or mil1ed enterprises. profitably.

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\n,\.,sS. W,M, A nuL,1F.., u., :mn1 If tnmrnunhy r<m,'er,.,·a1lnn is che ,J.n~\\'rr ; 1u•,,\Jlo, a.1.11. t- 11 "Ml. " "·· 19111. R:ibl"' In ~udu, trrfl//<k1p//111
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236 ,r.cno» ""c Aspens lnnuencini; the o,curren,(' of infectious dlsea£ef

~ ut.,01.i.., H.\o. • Ct1.0Ku:R. a.r..... 2.000. Reseean;h iruo the bre~din~of 3u r>t- \O!'> \,,,. a,,m. ,. 1u•.. s~6. Amhra.~ in the ~ncg<·r Notlonnt P.;uk~
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Pt'Oipcctfw, fir. C;tj(')\\t)HURY.:,.. .{ \CUUlkl.. .\ .A, (l·dsl llt!lmimhs oJ
wildllrc:JU\'e!.tock intrrfacC". fn: u.1,r.t:ri, tu;. (co,ordinau>rt. /nfetr/orrs
l\'i/1//if,•, :-21. Pl)moutl:, Unit,-d Ktn~nm, ScH.>nec l'ubilcutlon<
Dl$1'i/Sl!H)f W//i//lft, lNNaiftlC. f):(l}ll/U;/$ 11111/ ,1tn1111i,wmMr, 0./.E.
lncorporrucd. pp. 7-21.
Scienrt/kmul 11!1'/mic,1/ 11,,-ve. 21, a:1-tiS,
3.J GAI.L\.Glf!:8.. J,. \1.A,(.Al),.\:\1, I . ",.\\'t-'R. J. t..,\\ ffRF..~, LP., 19?2-
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Trop/ca!Anin:t1/ 1/tnltll mu/ J•r"'11wio11. ~. ?IJ.l- 213.
rree,U,1ngMrlean buffillo CS)'""'"'""ffcr-Sparrman> P<'PUlncion ln
tho Krug,•r Xatlo1uJ Palk. a prellml""'1' report. Omwrsupuori Jot/TfW/ oj' 3S """~Ol'SI, <., i.,u,,ri-~~.
,s,,:.. t.0.0.,;., no""""• ""
1993, Robie-s Ln
v,wrlllf/1)' fl?,.,,,n·/1. 63. !~18. Al'rl1::m \\ild dO!;' ,Ly,w,11 ptr111.,1 In 1hdt'«'ngc:I rcgion.J11umal of
1.6 at:Nc:m,. k..o.• 't110~1so:s. c;-..R,.,. a.:ttT. n.r.. 199J, Foot•and·mouth dbcasc 1n : \'il1/llfe Dfwnm, :ill. 3911-402
tmP<1!a ~·1Dpy,wotmrlamp11,1 Pr1J<betlm1,.• aftll" o.u,.
S<itm1fi, .vs l99Ci. Proposed
t.a:1.D"ERSLO\I, G.. \',\S Wl!Wt:£~ l:l.\\ &M.O~~OlJ\\" ~-
Co11f1mmc., tm /tJOt·ntuf.mo:ui: dlJNl.SC, .-\[dean 11ors:t·S:d:1it~Si, llnrl rran<!ronclot C:~n,elv:ltk>n Are,i<; ~l•p< and l'rolimina!')' Dor:t Shoo:,
cnmngiOU$ l>ot111t p!tumpn~wnomn. 2().,,2.J April !99.i. Gab:i.Coml. CSJR. Dki.llon o/War1 1r r:,wt1vmm-1m tmd Fau$/T) Ttl(.·'1,u,lt>,01..
801$\'"'"~ pp. 8-9. ,,tllrnl,o..ch, South Mric.\.
17 ao~S~tA. r.c.. t94-l, I Jtc~itary hean·water res~ia.nt charattN> m ca1tlc 3; c.c.w1u>. '- A, .,wn.. s,,j 1m, Ffc-ld Rcpor1 Ol"I Kudu 1nort3llt}' in M:;~o
Pamu'11g itJ Soud, Afri,;n, 19, t 1-96, ~a:lon:d Park. Sou.th Omo Zon~. l11tt•r1t.nl Publ:'tu1iou. .Nt11(ou11/ ·\nimaJ
,s 1t0!!,1A'S. o.e. & THOM. tu1.. !!l52. D0ghrrgl1-u:rgehoud1:n bf dc..n f/,'U/1/t C,,111w. Sllb<1u1.. Addi< ~/)(UU). Elf:lopla.
appercoopmun )an Arnhonlx ,~n llic'bet:d,. Dttl I Capt rown: A.A 3tJ r.u1LnflA10F. P.u.t.. no11.1S<'>'.\, 1>,H,L ~- ~H:.,,ut·n, 11..r. .. 1963. rubercufos!s
Salken,n. in 1hc fri:c-U,•i11~African tC::ipc · buffolo ·'S)'tt~trus tnfftr-S;,~mn:11il.
t9 ftAC!le>:>:.,MP, u.c.. ,~a~. Htani•t·1:Dtho:or.. lwrdt'rs and fannor<. Th• /01/J'tutl ofCr>mparut/t•r Pmlwlofl:,' and Thcraptmtl<•, 73.33;-3.;8.
origin of Sou1h0:m Afrtan Kok.in communftle:-.~ Ju: <-"'~n,o,. r. '"''\l'tt.,., :\9 IIW<lE~. R.S.. 19~2 Foo, nnd n101111t dlst<><e nnd theAfrle;(n Buffalo
<,a. (cd,). lll11srr111td f//s10,;•o/So111li A{r/l.'ll Jt>hann...burg: Jonncrusn ,S.i•nr,•ru::1.:nf!,,.-r}. /t,11r,u,J ofcompc1mti:"1:. Pathology. 82. 19-213.
8:,JJ Publl\her,.
10 1<£0GtR. n.s... 19;b. Foot And mouth diwa"' m "ildlifewlth parucular
!!O !J!tQCIIIEft.. ft,, AU!lU\l'. ~l.P.A.. P.\$l"OR.1T. p_J',, ~tASS0!\.1!.• 1'CHD:\ J.•
reference to the African buJTrtlo •.s,,·11c1•t1ts tnffi:t). ltt: r Att, A..l., ftd.i,
LO\-rK+\Ri-'I, "'' tt1M 11•ui~. r... l..',oun n.,. ot...~"'n11u v_ 19rf6.. Fl~ltl u~~
1\1/,lllf~ Dm:11sr.s. Xew York. Plenum Publi,;hnti; Corpora<lan.
of,'llC:dnia-rab!e,, re<:oniblnam ,·aec!M for tho oontro! of s,'h'•tle rabie;
In Eu1opc nnd North America. R<•:•11,• sclrmfjlqu,., l<'<'h11iq11, d,I offe"' 11 HWt•E:R, tt.~. 1.9;6. n,u maum:nance of (oat and momh dtse~e. 1n \frlra.
1mvr,uuimwl 1lcf$ l'l'J;(X)r101,. 15, 9~-·970. PbO ch<'>IS l.lmdon Scboolo(Jlyg1c,n~ and Tropical M,•d{etnc.
Univers:<, of!Dndon,
21 c.,n.:tY, A.. ~oo. The use of p.artidpruory apprais.al by \'ett-rinruin.ns in
.\fticn. Olfic<t lt1t'-•11uuto,ull tft!t Epi;oori.:is Srfumifl.< muf T~dmitt:i 12 m x... tsc;. ,,.w.. 19.56.Ammnl DJ.rMs..~ irt .SC,1111, •.\(r,,:a. 3td «In. Pre,octu.:
R,•1•i1.)IV. 19. 702-714. Centre! :\t·\~~ .-\gene)',
~ CAilfY, ,. tit I FYI. ,so,,_ 2001. Communit} pamap;nion and Lh.e deli\et) .a,3 ll1,1L\tL o. • ,micv.HHt~. :-.t•• lS!J19. Communh1<·, wJldtlft= nnd tht: 'the ne\,
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23 OJ!.\\>~\.\ .n, $,, IIC..~ ~. lX>SSD;\, A., t.AVJU;.~:,.OS, t,:•• :\1CC.U.LU;\I, ll~+ 4~ mnn1ro~. o" t1'0:t, Rindnpc'.'t ln South ..\!rica.Ju11rna/ Q/Cornpamtl:1-
RO»urs, >1 .. "\\OOllN<IHI • .. :wo1. Th~ role of pathogen~ In blalo~lcal Pmho/ogy, 15.300,32•1.
con.~t\"adc>n.. /Jt: 1muo;o:-., P.1 .• 1uo.ou. A.. t'.Hrrsn:u. a.,. na:,·nucc.:. 11.
4,; l~.KJLP. R.)t. 1g78. TlurP•.,l/lllttgQj)o111ir"mA/rtt.'tl, htedn. CnJk ro\~n
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24 (:u,1u1i,.;r., 11 11,M." n<>'-·n, s.. 1992- ,\ ft"\1.cw nf anlmitl production In
S0mhomAlrlr11. /DRC 1/epon HMmv. !\'orl(J Wild/If,• Fund .\f11/tl·s;>«fes
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27-29, l>on!clpotol) ~pproathcs to vct~ruw} opldcmio/i,g;- fn p1,1>toral At~•• 1>1
the Hom or o\fnrn. D,;•/,mrl, Pt0pnmnir..
~• Ill! 1·0~. c.. 1993, Pe::socml Communfcarlo11, Prl\'ntc 113gX~oz. ~kukuzn.
13SO South ,\fTicn. -18 r,·nff-...\'JIQ~M. IJ~IU' l t)k(.'.0"·Ut\'All0'.\ t'lr NAnm£M•D ~.\nJR.£
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:.-; ur ,o::-. , .. 1!194, Andu·.t..,. ltl:c..un1:YH. 1..,.w.. 1110M~>,, r..,r.,. 1u::-u,. ,u: ..
ti>T Gnour. !•ell',. Ckmslor,nl Paprr of th, /UC\' Sperle, Stm1ml
ecl.<.). lnfecrlou,J)~ra,s ofLin'Jto<k 1ultl1 S1111c/nl fl<'f"7m<r! to~omlwrn
Commll:$io11 TUC~. Tilc Wodd Con.,;cn·.utlt.m Uo!on
Afrka. Cnpc To"11' OlCford Un:\cr.lty l'rc,9 Southcm .\frtca, pp. 1262-
1289. .;g 11".'1'-', -\.0. 4 t:UN\1NC:.ll,,\\l. ,,..r., l9RI, lia'lt Coa.-., (ever In: ftf'i.T!C. ,,. I,
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ill ,uan L.,,.. >..1.111. o.i , R.\Atu. ,,r. •
or: ,,01,, v.. nr.:scl5, n..c.. ).'JIU:._,, .1>.~.
and Landun: Maninus l\:!Jhnff Puhlt,hcr-.
Kl/017~,n,n,i:zt, 11.•.~···· w01. Th< rpidMniolog)' of 1ub,:~ulo,1sin rrcc-
rn.ngmgAfncan buffalo .SJnr..nucnff•t'J In 1hc KrugcrNa.thmal Pnrk. .50 fA'4-r,·. o.. uos'J>, ,. a-cmw, a.. ;992:. Cattle. WIWllfe:. Both, or ;\clthcr.
SOuth Afrlell. 011<1,mepo,,r1 Joumt1/ o/Vdter/11,1/')· fl<'"'"""·
68, 119-130. R~<ult$ of~ finandal and F.eon~cnlu 5urw)' of Commcn:lal R•ncho< fn
~ufht•m 7jmb.thwc. H1't'f .\llJ(ti~p.'t:!n Proj,YI .\'9. 27. t l.vari•.
19 ot. ~·o:-. v. "t&lt\'OY:'\, 11.e.. 1995. Th~ epid~m,oJom,· oi a major :uuhrnx
i':imbah,\e.
outbreak in 1hc !(roger ~3tionnl l':lrl.. t,\hs11~c1l. lttte•11r,r1t111al
Worl.:sltop or, A111/rro.,. 19-21 Sep1<>nb<:r 1995. IVii:che,m, Unlc..t ;1 ~=· o.r. ,001. rcr-:on:slcommunlrncfon. l'.O. llnx 12, Skuku,a u;o,
!:lngdom. pp. 25-26. ~Uth .\l'rlc;,.

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 lnfecuous diseases of animals in sub-Saharan Africa: The witdlife!livesrock interface 237

so "'"· n.,." sr,.r.1<. n.c;,. 20-01, Unpubllsh,d ob>cm11lon,. P.O. llox ;? 72 ,"Err,-.. w.o.. 1~33. <:urridor ,.h..~a.'-C: ,. (rual form of bovine th1,:.Hen0:).u.
ili><uza, 1350 :;ou,h Afrlc"- cntounc~_n..'<f u~ l.ululand. Journal of th,, Sou:h ,•1/flWfl Vi~rcrrimuy
S3 ~U't. D.r.. ~,uu,. l<.P.1 .. a~~(.I< R.(... C:RnRlfH, "·"' t, ,11011 l,A.20ll0. Tho .\ledlcnl A~mrlan, 26, 7~-e;
rbe ~nd fall or rubcmilo•u In ;i !rce-111n11inc rhacn,a b11boon noop in 73 ~CJStt. ,.11. ""n:R-7.o. R.. 199;. rorcst as human dominnted ecosystems..
ahc "°:ugt.'1 sauonal Park. Omlur~,,JNJ(lrt Joumnl u/1/ettrJnarJ Hh<ur'f'IJ. se,.,~u~.
2:- 512-7.?5.
67. t JS-!:?2. i4 S011VAL R.A.I .. l~g..;. \ L'Ctor..-: Ttck.... ,,,~ COU/.t-lt, l~A.,\\ .. 11il)+\t.(;r)~. (;,K• ..,
S4 ~,.-.. 1'.f., ~"Ill~~. ,.,,•.J .• ••-~Gt$. R.t.. ~ >11om.. u., :!001. Tu!lt!rculo,,1, in n,-n:<, ILL. (~d•,. lnfi•<1/a11J Dl<t'llo~ ofLJ,.,stoek uf1/1 SJh.'Clnl R,f<Ml~
kudu. 1mi;.di11,lmurre1Mfr.ro,, In 1he l(ruger 'l:ntion:il Parl:. 111 $0111ht•m lfrim. Cupe Town: Oxford Unh·crslry Pre» Southorn \frf<'.1,
011tlMt<'fXJ()rt Jo11m11/ of! ,·rerinttt;,· Rt,<t'tUch, 68, 225-230. pp.J.-2,1.
ss s..-n-r~ o.P.. l;RJFK. ~.i•.1 . ., ,ucuu ,. 2000. Tuhctcul~ls ;v,d Its ,s OBtfff,M, P, r.;.. Dt.ZUIDf~HOHl, J.D•• :987 llitd..'"t\\.'3(\?t' hOM~ ('Jthcrchan
!!<Oi:tnphlcoJ di<1ribu1io11 lo fn,e-ra,t~ing lion, In Ult' Kruger 'l:a110nlll t.lomCJ.\lc- rumin:tnl:». 01ul~rstt'p('li)rt Jourm1/ o/V~trrinary R,,runreh. 54.
Pork. Proo,,'1/11g;. oftli, ri,lrd l11tmltlllo111tl C,,,,f,rmt·eo11 271 .. 2;5,
~l)'CObacterium bo,1,. l,l-.16 AUl,'U>t 2000. ('.amhridgc Unf:cd 76 OH-ta rr-. n.R'.\'.:\TI0~.,1 nu t!.PUOOTIE~ to rel. 2000. /m(fmfllfoual A.1rfmal
Kfn_gdom. /-1,;ddt Cntl, 91h ~dn. !000. Pnri-.
,;6 ~l!ET. DrF•• J,.'JUl-.1:. ~.P.)., P}:-0:Rt OI, \M,., )1KHtL A, & HUCUY.t:S\M£'YER. n nvn:4it ,"'t m-vnovMt.,r 1,°""rrnrrF (001). 1~9-9, Can 1.ivcstod,antl \\1!dllfc-
tt.F...,.lt. 1996. ·rubercuJasfs, in butT.a.lnh [Syna•rzl~ c:njfi•r) in the Xru~cr co...~.\i~1? .\n intl"rdi.\-Clplin.at)· :spproach. In: bOIJk"' o. A. uu -.:cu. M
:-.:n,lon:,I l'llrx: sprtad of the dl>C•<f.' ,o other ~peeicS. (),ulcm,poor ·~d,,. s~. i>ublW>ed ~·001, l'onl.u,d House. StagPl3<:.(r. l.ondnn ~\\'It.
Jounwt of\'ctt11lmll)' Flt·senrrlr. 63. 239-2-14. .mP.
s-: ..:ur:-., tt.r... 1993. Humcr gathercf'li arid forrno~ ln Africa. In. DURE..~H.AJ..T. 78 01\"F>. "· • O\\}.'- u. 1980.1111• ft•nc•, ur donch . .>,fr/tan WIid/if,, :J.;,
c;., (ed.•.). Pcopt~ OJ 1Jr1: Stont -~~ lllmt11r(;(JJ/wrrr::. mu/ &,r(v Farmrrs. 25-27
San Fr.u,oisco: Harper. pp. 3s-..;- o.: ~.S.
7~~ J-1·.Nt.\. T- , •• ltOKAX. I,(., ,.. ttl-CH.\\, '( 19{\;. Th!- l."CUlog)· c,f 1hc t\frlcnn
S8 KOCK, :\1,1>.. MUWXS·, C.,Jf, 4, PJ-;"Ri-JX~+ f,S. 2002. Vc.tcrlnntyd:se.ese co.mroi
\'t'l'C:IO:'< of hc:.artwa11.'lf, with p..mJcular re!trt•n\:e 10 •.\mb{t't>1111ur1
in Botswan•: Impacts on \\1ldllfc hc:ihh. etOS)'SlOOI> and niml lmbrtt,.um .lnd --\mhlyomnu, hlrivgmum. ()11dru1apoort Jounml of
lh·etlhoodt.. l,r; /\Cill!HIH!.. ,\.,\.. OHnw. i(,,,, HUU-!,1;, 1,;.,\, , T,\ISOR, G,)I, .,.
h·~11muy Hts(ttlrrh. 5.;, '.\81-395.
1•r.ARi... M,c;.. {eds). Cons,m'l-l_lion ,\/Mlr;me: l£rologtral 11,..-,1111 m ;>rr1~l!C't!.
New Yor'~: O~ford Unl\'er<11y Pre,, In 1>r1:>s. nu VLOW1<JG111." 1967. ~l~Ugnanl cainrrh:tl fewrin l:a;i Afru:a. !II.
'\outr~ll,ing anr!budy fn frco-11,in~ wlldoboo;t. 11,•mtrrT, In l'r1,•,/1w;
.59 a. ltl't'\'Q,,l\, "· .. llll>A " · II., ,vu,. Wlldllfc and
S()Cj; .••, •• Kt~IIJIIA,
'ki<'11u, 8. 1!5-Jlll.
p ..,toral sutlcr;·-shlfdng parndl!llns in di,,.,..., con1mLA1mals of tll~
\',.,.,. York.Am//Nrryo/Scle11c,-;-. In pre». t9ftO. Blot• wild-c~c.•$t .1ntl rhc
81 Jtl,..()\\'lllGHJ. \•, •• f•LAAI~. tLI), ,..,con' ("... ft..
,totfologj<UI 11t1•n1 of bo.in• molltw•nl c:mmnal trvor. Nt1t111V, 18".
60 J;OCli ll.A.. W.U!UU,\, J,:'-1,. M\i..AS.I.H. J. \\A\: \UV1. H .. ',:f,H,I ~G,J, l,);..
1167-1169
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wlld fUlnitwnL> In K<:n>·• 1!19J-l99i. T/r, i ·emi11nry R11t'Qtd. n; 2-::,.. Ull Pl,OWntC.111, \\' .. J',UU:nn, J, _,,. PORCL M~.\ .. 1969, Thi." t!plinodolom· n(
283. Mric:,,n ,wino !'...·or In ,\(rf,;a 77i, I \WrilUll)' R,'(r}f(/. 85, 668.l,"74.
Bl ,.~oc.nt. M., 1g..81. Chnracit"rb,tit,. ot l}'PC~ :md breed~ ofc..1ttlc i.n its PW\\tumn. ·., ., 1•.\ki.:1.11. J,. • 1•1 tw:.'.1, ··'··'-· 196~, ..\frtc"n s.....1m! !~v,~r vhu')
the tro_ples. In; a1,;nc, ~t.,.
,to~f\'tlli. ,.. Cr:ds}, D1s.CVl.W•l a/Cnn!t In in tkki: 10mfthodoro1 mn11l.Jt11a, Murr..a~ coll~~d Cmm nnlmal burro,,·.,
th• Trt>plr, n,e Hague. Ro,ion and London: M3rtinu, :--tJltuB ln T>Cl2ilnla .\nwr~. 221. toil-1073.
Pubtt,hers.
81 Pl/J\\·rucUT, ·,, ru,nr. c:r. Lcm.tc., .\.. 1~-:4. Sn-unJ trammi..;s1:on ol
~ '·"''8'JliE. 11.~,. lr l'UR~DUl..L P~c..o•• 19:~
Ecology and cp1drmloltt\,t)' of \frk:m ~\\1m~ rcvcr \1ru~ 1n the- argasJd tick. Omlll:Mor11$111J:mbara
anthra.< in the Etoshn 'l:,rion•I PorL X3mibia. o,,dl!r,i,11<11m/our1111/ of pt)rl'im,$, \\'utum. n,,sq"rc,1t In V~i.-ri1:t1ry1 Sf"''""·
11, 582-5.ff.:.
Ve1~rint1f')•Rt•.>t'iu-ch. 6t. rt~l.
O:, nowRwm. w .. l't.A.k'i c.1. & P1!.lffC'J.. ,t...\., ~-o. Transo,·ari-al infcc-1ion
&3 ~c,cs. 1<.. 19;0. 1h~ 11re:11i,fn<an c:i.ule plague ep:domk of the lb'9l.ls. wflh :\trlcans\\ine (!;'\'Ct vitu~ln the- ;,r,g.,~id dtl, OmithmftJt'11. 11uml1'l-ltt1
TroJ.iiettl .,11:fmllf Tl('i't/th mul Produalon, 2. i!.10-219. """"'"'· \\'A!lnn. Rrs,arrlt /11 I 'emi111uy 5':!rn<<', 11, ;s:i-sii.i.
o_. c.s.t,, 1995. l'h1.• cff~('t u! 1ron...Jiumnm,:c on th~
:...uorrt-R.'\QZ\ 86 HOD\\'LU. r.c.• -.'-"llYrl i.J. llJJOY<:1 w.,,.
2001 F.\"altt.1t!onofpopuL.ulon
~p1dtmiology or u.nim:d dkease-s. Pur\"'llln'ti \'eu•rlnary .W,trlinn~. ?!». cft'cru ofbO",inc tub<•tculo.sls in {reo,ranjt:ini.Afrlean buffalo {Syr.ct1rus
21'.l-!24 ttljft'l'I. /mmu,I of.\f11m11mlogy, 82.231-m.
~$ ', t..Uh(),, t'., 1999', Chronic Wa>tin_g 0i,t'Ob,c, /t1/ormfl/lOII /kporr bJ' 87 111n:u.'Y.•r'AnXEB. M, "uxso~. 1.. P>.C..u:tt. C ).O(;J.. tt.A.... <'.:.LL\\ li.U,,I}, i.
Ct,Jortula (lud \\~'('1mlug<i,/111;rm1d FisJi Commls.tiOJts. CA.1w1.,nK, ,1 ... 0·,uuF.,.Sc.J.• POS.PISt.Mll. \ ,1!001.\f':~·LF.H'.ltll\N~.1\.. l.UTl".
GS ,L,c<.s. r ., 198(,. The """1· histori• 1)(1he block ll<'Ople In ,outhe:11 Nrfca. It .. ~\\'.\!°•SE...~CiW, Ci..1,, ~H1t\i;;\, M.:\, !,I.\C:ll.\)r.(;\., h,A,. S0\1\11•~:t. it.A,#.
In: CA,rn11<,s, 1 .& "'"'• <.a., ;cd< . /1/11,rr<,te,/ 1/111nryuf.w11tl1 Afr,w. ,\PPrt. ~tJ., 1996. A C':tnlnc tt:Mt!mJwrvinr:-- t:pitlcymft• in Sttrongcti lfr,n ....
lobonne<bwi; and l.(1ndon; Jonathan llllll. (Pdntllt'l'c'l. ltV>), S't1Wtt, Ji9, ,f,1 t -t45

b';' .,1.orri,.:, ft. t.~86. ConununaJ ..t.ruai m.utugffl,~nt plan for tndtgonou.s. 88 ROSS1 nSt l1.8 .• 1983, Rolf c;f wfldcbe~t !ct:d mcmbmnb a.nd Ouid), 1n thr
r,:,,ourcc< !Co.mpfircl. Srartch oiTcm>Winl 1,col%')', Oepanmont n( 1ransmi11-slon ol m:.11l~:tntc:thll'th;1I (1,."\'t'f \inJ~. Tlw Vcttri,u,ry R,"Cerd.
1">1iMol l'~1 lcs ontl Wl1111ll~ ~lam,gcmen,. Hamre, llrnl!ah\\ e. 113. 1;0..1s2.
68 \tU:~:<,"t.YA, T.• t998. T,\~AP,\ Vctt-ri:n.ar,.•O-t•panmcnt Annunl "Repon 88 uos~rrER. P.a. 1994. ihnderpe~1. hJ' C()ITLJ:lt. ,_,,w
1·uomn~. c;.M,.;: n,:,...
l89i--8. Tttn1..tnfu Nah1:mal hrir...'"'i. l•.O. Box 3J3,J. Arusha. Tan,1'.ami+t 1,s. ,u: 1:ds·1. Jnf,.-i.·1!011.< ()f.tm.~ Q/ Lf:'t.s:o\·k with Sp«iat RtNr,11<~ to
Southf!T'tt.-4.Jriro. C.ipt Town: Oxford UntrcnJ..ty rte.s.5SOuth'1m i\tru:a.
69 :,i.rt.t'.\Gt;.YA, T., 2001. T·\Nt\V.n \.cairinary OC'pa.n11wn1 Annual ftt-pon
Pl>. 73~,;;
2000-2001 TQnr.anin X•l:omil l'orks. r.(l. n.,,313~. Anish,
Tan;,..mta. go 't(:<.k>s•..., , .. i99.-. :cdJ llving with Uncatntnry.1.(>ndont lnl.Nmt:diatc
Iecbn<>loi:;.
;-o .MOflJ\J!IOS, \\',I,. MUJtAAY, "·· lt'\ICJ,n1u, \\,,.,, .• 1g$-1. RU\1nc
nyp.£no:,:omi~t-s.i~ /11: ,u~-nc. '-t. 11.,ttJ,"T\'DL t., (edsJ. 1>11eases o/CimJ, in !)t ~:on ,u,..sos RES-ounc;a: co~.>-utr,,!'l."T\, ~aoo. EJ'l\1runm~tttul a.:1,st-,,nwnt
rl,r Tropks. The Hai:w. l!9>1on ru,d l.ondon: ~!minus :>.'i!hon o! \-Ctcrim,ry fc:nr;~ in Ngoinlt.md. Scan \\'U~on Ra<>urr;c ConsultMt,.,
Publisher<. :!3 Che>«r :.ueot cdlnburgh. m13 m ui..
~1 ,w,111, 1:..;,. >o.,'JC>rA1>, 1.. $tJE.Un. o.,t. 1~. l~ndon o.fb<wUl1.• maUg.nunt 9:.t ~o:tftA\\ r-...,aoncuo . .:... GOPno. ,\."' L,Jo1r,,ox. ,:.).., ~,~. Amhr.a.\:
c-w.rrhal (t:\'Ct \liOJ!i (ram C>CUL.ar ~md m~ s11Ct<'tfon~ of,\·Ud~hebt ttutbrcrtl< In \logo '.'.otio11ol Pat~. ,ouU,cm Ethlopio. /lie Wtmnmy
('a)ve,., Rc-s.;itlTtlt in Vttctrlnary3cff,:N,Z£•. 16l-171. R,'COrd. 1~0. 1111-.,zo.

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238 "'mos'"'-' Aspects influencing the occurrence or inrectious diseases

93 ,in:vo.-sm,;•1J.iu111.1'0;\', J.. 195:. \Wldlife i11 S011rh Afri,;o. London: and.\Jat:Qf.i!ll•nr of Frre,rm11(i,ig ,\(r,mm<>I<. Ri,·11r S<:1Mt/fi<!1<f e1
Hamilton & Co. m:l1nfq1ieOfficc lmrmar/ona/ Et>i:oor,,:s. 11, 1097-1107
g.: <UTMQllhR, P., TIIO~!ol<)", r,,n.. IIARr.hl~\\TcS,_,.~ .. fOCGI", C.'1, I, 102 TIIORSUA~. I.A& TIIO~tM. A.D~, lfl-lO, Tuburculo.sb (nth(• C.ipc kudu,
.st>~1t,01<. e.c:.. 2uqo. The foo,·and,mourh di1e"80 rlsl. poied b)' A!rkon /011nl(I/ a;'rfi, S0u11l,Vrlcm• \'e1,rtnn')· .\/edlcr1/As,0<·intlon. 11. 3-l Cl.
bull'nlo wi,bin wllt.llifetonse,vanclc, to 1hc ca1tle lndu,try urlimb,thwc. 103 ntRUPr. L-A.. 19(),9, Critical links: Food 1::C('1,.1.Jit1,·n11d U,1.\ cnvironmi:nr ln
Pn:1rqmf1'f' \lc1eri,uu7 MMidnct, •H, ,1.3-00, the greater hom ofAftic:i. Bl1ckvr11111d pnP"r SJ(IL!/w/tl,r A1uY.J1U o,;d
9~ "'\'A.~ffi'.,OtrJ... n., S9.9-1. Rabif?'.'... /n; C:Oli'Iz.tn, J.A,W, TUC)~tSo:;. r..n. • TIJSTl~. D1i1lagui \'torld R£>.sourtt"S Jn,-tinntand l,U.CS. East Africa Offic-e-.
•.c.. ,eds). lnftttforu f>iseasesof l.f1·<St«k u•tlt Sp«ial Refermu to ·'-airobi. ~ova.
So111her11 Africa, Cape Town• 0.1:ford Unl,ctsit) l're,< Soutllm, ,\frfc;,. 1Q..1 n,x 01;11 MERWt v.1., 19e-1. Ken 01\.6 Perdera,s$C. C~mel own: Human ru1d
pp. 493-552. Rousseau.
96 nt.\~1'81. t.s,. MAISA. o.w. & Sl!SSIS. n.. 2002. 1\nimtll and nnimal produces t()S VfRBfFX \\'.,\ .. l971, Cattlf:. /1': vo1r.1Fn..R..1>.r.. 1,u fltr:...(SLli, ,.r ·" t-tu,,inu.
rradc In Africn. N~w Dt'vclopmtmr Pt1r511«rh1et In I,mm1mio111d Tmtfi.•/01 ..,.,,. (L'<i.s). <1mu/11ttl En9~/Q,x1edia o/5,Jut/imi ,i{rica, South..\lric,,
Nrictt. OAIJ.18/\R, P.O.Box30,S6. ;l:.lltObL ht Press. :-.ntlonal Commeralal Printers.
9; nw.u., G,M,, 19"2. Etlmograplly and Contllr/011 ofSourIi 1\frlt:n JJrfore ,ID tt16 .._,.A\:V-A,ti\. J.tt,, 1997. Conftd1:1ul)· uSL"CI to1hno,•t!1-erina1}' k.rto\\'ltdg~ .uno1:g
/SOS.3rd <·dn. l.ondon: George AIIM and Um,111, 1i.i.•1orulis1> t\f ~rnburu. Ken)'11, Book 1. MJttltDMlogy11111J R,sultl.
g8 nit HERAU>, Harart, Zimbatn, e, 22 .~pril 1§97. /,::,m1«llnrr 1',xflnology. Sn!rob!, ):tnyn.
99 THO,M sos, c;.1~.• t98S. tbc cpidtmiologyofAfrican S\\ine fc\,:r: Tiic role or ur, wooof<)Rn, ,uL, 1.so:1. ·ruh<m:lih,sh in ,,ildlif~ in R1,.1\,'<rm:OC1 ,·et1ono.i
free-thing hosis. in.Africa. Omftt.r$1¥JJJ(JOrt}uunml o{Vt1tt:.rfnury Rtscmdt, Pmk. Ug;mda. CPan l 1. Tro11irol Animal /t,,,1/J/11111d l'!Otiu<11011 14.
sz. 201-20<J. 8Ml8.
100 TI<O»}C>N, "'"" 1999. ,\iU:m•tlve;; fc,r ~n1rolling anini•l di'Ca...U, 108 ,,•oouR1Jtn. ,.u, ..
198:r:. Tube·rc.·1.1lo~i, in wUdJirf In Ruwenzo:i i\·auonal
resulting from tho intorucdon between llve;1ock and w!!dllfo In l'a,k, Uganda. tP•n II I, Trop(MI ,\11/mnl Htt1lrh and Pro<l11c1ion. H.
Sotuhem Afrfc,i. SouthAfrlcnn Jouronl ofxlrnc,:, 95, 71-,o. 1,s-ttx>.
101 ntOM~ON. G.n.. \ 'O.SlO(). \\'.. UTJllllr'OYS:-U:. ,.,. t, eo:c.n~ It(;.. 199::. 109 woonrom:>. 3,1.11., .oo.). 2001~ Qu1tra.11rin tmul Ht1Jlth-S<>i'tw1lngP101oro/$
~1alntemmc·e ofroo, ..And ..mourh dl$C,8j!~ ,'lruses-in buffitlo ,Syn«rrrs ,,rn,r ((/
for !\'/Id/If, TrtlllS/O<'tll/011 ""'' /Ml•(l,q? fnro ''" WIid. P•ris,
ra.fferSparrman. 17791 In SouthemAfrk:t. /11:AIITOI$, M-, Ctd,l. N;a/1/1 fmuce: Office Jnrtrnodon~ dll< cpu:ootJes

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 11
Vaccination: An approach to the
control of infectious diseases

BA ALLSOPP, LA BAB1UK A;'l:D SL B·\BlUK

lntroduction cals into their <urroundings. TI1e same is true for anhropod
vectors or disease. These facts rule ouc any long-tem1 dis-
I! must ha\·e been obsen·ed for cenruries that people ex- ease comainmenc s1rate~· based solely on the use of phar-
posed to a pathogen were o~en resistant 10 reinfection by maceuticals. acarkides or insecticides. however successful
the same or a related pathogen. but the first rncord of an)· these may be In the ~hon term. The develc>pmem of new
one acting on this obser\"ation is that of 1hc English physi- ,•accines is the onl~· long-term strategy that can be foreseen
cian Edward Jenner.52 On 14 May l i96 Jenner inoculated at present for the c;on1rol of dil.cas.t>..s of animals or humans.
James Phipps, a healthy eight-year,old boy who had never Mammalian immunological defence mechanisms against
been exposed co smallpox. \dth cowpox \1rus. and chal- Infectious agents have nlso evolved over millions of years.
lenged him •18 days later 1,1th a \'irulent dose of smallpox and must therefore be high up on any arbi1rary scale of op·
'1rus. Phipps did not react. and ,,1th th!~ momemous med- timal!ty. Finding new ways to specifically s1imula1c the im-
ical discovery was born the era ofimmunologyand vacci110- mune system has to be the most likely way to achic,·e lhe
loro. Since 1hen smallpox has been erudicaied by lasting control of infectious disease.
vaccination. and human and animal losses and deaths due Vaccines lirensed for use in humans ;ind animals at
to other infectious and parasitic diseases have been dra- present are primarily either killed vaccines or auenuaied
malically reduced by the de,·elopment of a range of \1IC· vaccines. produced by one of rwo con\'entional methods. In
cines. F.ven i.o bo1h animals and humans c<lminue to ~\lffer the case or killed ,·acci11es die pathogen is grown i11 large
from man) common diseases. as well as new and emerging quanti1ies and then inactivated by some method that pre-
diseases. Among many reasons for this are: vems its replica1ion but does not significantly alter the anti-
• sub-optimal managemem conditions: genic propenies. Auenuated vaccines employ a live
• expo,ure to pachogens at a young age before vacclnalion mutated agent 1h1.11 has been selected in ,,itro for reduced
i~ recommended: pathogenicity but chat is still able to replicate in the animal.
• unusual or unexpected panerns of the pathogenesi~ of Each of these conventional ,•accine types hru; inherent ad-
certain organisms: vantages and disad,•antages. and there is a need to find ne\,
• the emergence oi new fom1s oi old diseases: ways to make safer, cheaper and more effective vaccine~.
• the emergence of totally new pathogens and disea<es Advances in genomics, biotechnology. immunology, and
(<1.g. ~ipah vims): and w1derstandtng of rhe pathogenesis of diseases have rcsul ted
• the relative ineffectiveness of some exi~ting vaccines. in the development of manr new types of vaccines. such llS
subunit vaccines. live geneticallr modified vaccine(>, and
Despite 1he currem shortcomings of vaccination it seems nucleic acid (or ·genetic") vaccines.
likel~·.tha1 it Is, and wiU continue to be. a more durable and
cost-etlectfre method of combarting bacterial and viral in-
Principles of vaccination
feccions than all 01her 1herapeutic and prophylactic meth-
ods combined. Micro-organisms have survived for millions. The primary mammalian immw1e response to an imading
even billions. of years in the facr of a conri nuously changing agent, invohing the clonal expansion of small populations of
chemical environment. and chey are only going to be brieny ,irgin T and B cells, typically takes two weeks to develop. and
inco1wenienced by the human introduction of nt.' \\' chemi- this is often 100 slow 10 pre.vent serious disease occurring. The
239

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2-H) w.cm" .,.,; -\spens inlhumcing 1hc oct'um:nc-c ol inleclit>u, dl~ca,..-s

secondary responsc 10 an~ subsequem rei111roduc:1lo11 of tht and then ~ele('ti11g variant~ that ,how reduced virulence but
same agent. however. involves the siimulation oi n population which still induce a prowc1ive immune response. Over-auenu·
of specific memory cells thm \\Cre produced during 1he pri· aiion ma1· resuh in suc;h limited replication that the immum• re·
ma~ response. TM, is a larger populn1fon than tha1 of the sponse to a \in1Jc111 challenge i, no1 protectlw. and unc!er-
original virgin cells. so the secondary response is faster thnn :menuation will rcsuJt in the 1accine itself causing clinical
the primary response, 1ypically being less than one \\'cek. 17· ~;. di:,ca~e.. khieving the required fine balance between O\'l.'r· and
37 m The seconda~· response will usuall) dear thf.' pathogen und<.'r-att<:-nualion b, howe\'er. pure!~ empirical and there are
before O\'ert disease occurs. Successful ,:accination depends many practical problems "hich make this conventional ap·
upon exploiting this mechanism by specific stimulation of the proach bo!.11 tediou:: and expensf1·e. Each mutant mu,1 be
primary response with an agent inherently unabJ(.> to caus(.> dh· t~ted i11 1•h'O under a 1~ide range of circumstw,ces 10 ensure
ease. During any subsequent exposure 10 a ,·irulem fom1 of the that it cannot causedisea~e. \\'hile still stimulatingeffecth'C pro·
pathogen the second~· immune n:spon.)e pre\'cm:. the oc· tection against ,~nllent challenge. Special factor., may haw to
currence of disease. This seems 10 be a simple concep1. but be taken into account. for example. some animal~ may reaC't ad-
•
mammalian immune systems im·olve inordinately complex 1·ersely to 1'llccina1ion if th~ are immw1osuppressed due to
cascades of molecularsignals and cellular rcspon,es. For effec· stress. or pregnam animals may be Induced to abon or foetal
tive immunization we need 10 know which antigen-, elicit the defect~ ma~ de\'elop when animals are vaccinat~d panicularly
appropriate specific protective response. and ho" best to de· during the first trimester of pregnancy with. for example. thea1-
li\'er those antigens. for example, administmuon of vaccine~ tcnua1C'd Rift \'alley rcwrand bluetongue V-dcclne.'i tsee Chap-
via the most commonly used routes. either inrramuscular or ter 95: Rifi Valley fever and Chap1er I02: Bluetongue;. 11)(1
subcutaneous. may induce good sy,-1emic immunity but may 1'1ost imponnnt. however, 1,; the tilct thac in the past there has
not induce mucosa! immunity, thus failing to protect at the site b<.>en n<> informntion ;1bo111 which gene~~ have b~-en mutated
of enoy of many patl,ogens. 51 -~. 7J If the ,irulence factor of a and du:rt> has long been a iear that 1.he mutated genes could
bacterium is an ex1raccllular toxin, it will be inappropriate to back ,mmate, or recombine in 1he field wilh virulent alleles, and
use whole cell bacterins 10 protect agains1 tl1is panicular infec- cau,e 1hc pathogen 10 rcwn to ,irulence.3- 7' 1 Recent acl\'llnces
tion. Until recently few oi the infectious ngents encmmwrc.'<1 in in tht, genNic analysis uf auenuated ,-accine agents have shown
veterinary medicine were chatactcrw..cd ar tl1e molecular level, char th<.',;.c 1ears Wc!l'l' well founded.l.iS· -
nor had tl1c host Immune responses to tl1em been well charac· L>isadl'antages of1i1·e vaccines are 1he pos.~ible presence
terized. The molecular genetic advance~ made in the last de· i1f contaminating exm111eou., 1imse~ in vaccines grown in
cade have begun to produce sisnmcam changes to this cell culwre. for exampll• bovine 1irnl diarrhoea (B\ 'OJ inro
situation, and we may expect that the next decade will see \'llCcines grmm in bovine <:clls. 51;· 103· 1 11 that must repltca1e
many imponam advances in vaccinology. the vacdne Cllndidate In onler to induce a pmll!Ctive re-
sponse. ond that cbe presence of passive antibody such as
rhat from colai)~trum. \>\ill limit rhis replication. In many in·
Conventional vaccines
stancl'S the level uf passive amibocly st1fficicn1 to interfere
The majoriry of vaccines used today in veterlna~ medicine are with vaccination is bdo\\ th:n which will prevent infeclion
either live auemmted or killed vaccines, produced according t<> \.\'ith wild types of chc pathogen.;;; In such cases \'accination
principles first developed by Junr.er and Pasteur o,·er 200 and can only be pcrfom1ed after pa.~sl\'e antibody has decreased
I00 y<:!ars ago, respectivcly. 101 Bt)th of these type, of \'nt:ci11cs w a predetermined level. which moy leave voung animals
have been effective in reducing or eliminanng clinical disease vulnerable to disease for long periods It may also lead 10
following exposure 10 virulent field strains of \'ariou~ patho- ma11y animals becoming su,ceptible at once. u1 1he ~aim.>
gens. Equally imponan1 has been their role in curtailing the time thot thlly are expo~ed co multiple diseases simulta·
spread of pathogens to other animals by reducing the quarniry neously. \nimals arc thcr.•fore often var:cina1cd with mul·
of pathogen shed imo the emironmem. These effectii result in tiplc vocC'ine~ at the same time, but chib pro,ide,- the
what is known as 'herd immuni1y· to a pathogen 1• and If the opportuniry for intcrfcrcmce co occur among the r<',ponses
rram,mission is reduced sufficiently the pmhogen i~ r1·enmall>• to different components of the cocl-taiJ.39 •17 92
eliminated from the herd. :?II I.in? , accim,~ also require aµpropriat~ storage l'Ondi·
tions bet\\'een manufacturing and administration. and the
Live attenuated \'accines maintenance or a cold chain is often a critical factor. This is
Llve anenuaied vaccines are bo.,ed on pathoge1,s which haw a serious difficulty in remmc areas, particularly in rural
mutated sufficiently not to L-ause disease, and the two most im· Africa, although it is a problem which also affects inacti·
ponam considerations are whether the le1·el of anenua1io11 i~ vatod vaccines.
sufficient 10 make the 1.1,cine ~afc and whe1hcr tht• potential Despite all these caveats chere ari.' $Orne real advantages
el\ists for reversion to 1irulence. Ancnua1im1 hu,- ('tm\'ention- to attenuatetl \'accint1s as compared to killed \'3Ccines. The
ally been achieved by passaging the agent i11 irirro. either in the live agent replicates in the hose. ~o much less material has to
presence of muiagcns or under ~pcdfic culture cnndhions. he incorporated into lft1Ch dose oi \'acclne than is the case

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 Vaccination: An approa~h to 1he conLrol ofinfectious diseases 241

for inactivated vaccines, making anenumed vaccines less to MHC recep10rs, and the development of algorithms which
expensi\'e ro produce. The immunity engendered by live can predict potentially protective peptide epiropes.102
vaccines is likely to be similar to that indttced by natural
infection. and of a longer duration. and broader, than that
Modern vaccines
induced by a killed vaccine. This is especiaUy true ifthe live
vaccine is delivered via the mucosa! rou te, which is the natu- Fundamental changes in biology are being brought about b)
ral mode of infection for more than 90 per cem of all patho- the era of genomic data, and this is bound 10 impact power-
gens. The resu lting mucosa! immunity pro,·ides bener fully on vaccine development. Currently in rhe public llO·
protection at the likely site of enrry of the pathogen, and main are 860 complele vital genomes, and 65 complete and
should reduce shedding of 1he pathogen into 1he em'iron- 264 ongoing bacrerial genomes. Volumes of genetic daca
menr. This results in better herd immunity than that pro- such as these were unimaginable as recently as when the
vided by killed vaccines, which do 1101 generally induce first edirion of this book was p1.1blished. and we wW give just
mucosa! Immunity. one example of the sort of work which is being facilitated by
these changes. Bacillus Calmette-Guerin {BCO) is a live at•
Inactivated vaccines tenuaced mixmre of Mycobact<'rium /Jouis strain), originally
lnacrivated, or killed, vaccines are produced by inaciivatlng de,•eloped arthe Institut Pasteur in Lille, around 1he IUrn of
the infectious agent. so tha1 it cannot replicate in the host. rhe rwenrieth cenrury, by Alben Calmette and Camille
while still retaining the immunogeniciry of the specific pra- Guerin. After extensive lesting. beginning in Paris in 1919,
tection-inducing proteins. MOS£ inactivating agents do, Caimette began to use BCG as a ,·aceineagainsl lllberculosis
however. have an impact on protein inununogenicity and a in France in 1921 22 and it has been. and still is. used for this
bala1tce must be sought be1ween inaclivarion and reduction purpose around the world. The effecliveness of BCG in any
in immunogenicity.31 • 3-4 Oulbreaks of disease have occurred area is roughly im·ersely proponion·aJ lo the distance of that
as a consequence of incomplere inaccivarion. a problem area from che equator, and this is thought ro be because in
especially with pathogens that aggregate and thus limit the the tropics there are v,idespread infections with other. rela-
penetration of the inactivating agent. 20 Beuer Inactivating tively benign. mycobacteria. it is sunnised that these gener·
agents are being developed. but tesring the level of inactiva- ate enough cross-immuniry in the exposed human
tion during the quality assurance phase is scm vital. Even population that BCG does not develop sufficiently to stimu-
testing, however. would not have prevented the introduc- late protective immunity against tuberculosis. Ln order to
tion of scrapie into sheep following immunization wilh an unravel the moleculargenelic fundamentalsofrhis problem
inactivaled looping ill vaccine.H ObYiously one cannot test a comparati\'e DNA micro-array expression analysis was
for extraneous agents unless one has some informarion performed on 13 strains of BCG, and on l'irulenl strains of
abouc what might be present. M. bo1•is and ,W. mhercu/osis. ll Subsequent genomic se-
A problem with inactivated vaccines, especially those quencing revealed that 91 open reading frames (probably
produced from bacteria, is that some protective antigens are genes} of virulent M. tuberculosis were absent from one or
either nor produced in vitro at all. or require special culrure more virulent strains of M. l1ovis, and an additional 38 open
conditions for their expression. For example, some iron- reacting frames that were present in virulent suains of
regulated outer membrane proteins of Mannheim/a (Pa.f· M. oovis were absent from some or all of the 13 strains of
reurella) haemoly1ica, known tO stim ulate prolcction in RCG. This escabllshes a firm molecular genetic base from
11ilro, are on!~· produced under restricted iron growth condi- which 10 begin to analyse the reasons for the variable effec-
rions.2&, 27 B~cterial preparations may also lack sufficient tiveness of BCG vaccination. and to design be11er vaccine
concemrations of proteins which are secreted into rhe strains. In a way that Calmetteand Guerin would not ha\·e
gro1\lh medium rather than being retained inside the bacte- begun to conceive.9 • to
rial cells.95 An example of11ew ways of overcoming this defi-
ciency\1~1l be dealt with under ·subunit vaccmes' below. Subunit vaccines
11ie main ctisadvantage of inactivated vaccines is that they Subunil vaccines are those which contain one or n;iore pure
are relatively in.effecctve at generating p(o1ecdve immunity, or semi-pure anrigens, and chey can only be developed on
particularly against intracellular bacteria and promzoa This the basis of an understanding of the mechanisms of patho-
deficiency may be mitigated by the use or strong adjuvants6 genesis and a molecular characteriza!ion of the gene prod-
(see further on), but In some cases it is the resull offundamen- ucts which Slimulate protective immunity. Most of 1he gene
.al differences between the immune responses against live and products in any pathogen are irrelevant to the induction of
killed organisms. For example. vaccination with heat-killed protective immunity, and others may be immunosuppres-
listeria monocytogenes is not proteetive because it stimulates sive or even enhance l he disease process, so the critlcal
the production of memory T cells which fail to develop imo ef- eomponents of the pathogen's complete translation reper-
fector COST cells.61 This rype of problem will.be addressed by toire must be identified.
advances in the understanding of how antigenic epitopes bind Subunit vaccines can either be produced by in 1,irro

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 ..
__
') ,•? sr= <N: .-\spects lnOuencing the occurrence or infectious diM!a~e--

expression from DNA recombinant construcis or by purii°)'- concentrntion of a compound to allow it 10 replicare.30 This
ing a specific componem from a conventionally produced compound ca.n either be introduced with the vaccine or fed to
vaccine, and the products may be used alone as vaccines or the vaccinated animals to allow the bacterium to replicate fora
may be used to ·spike' conventional vaccines to increase the shon time. They will thus be able to stimulate immunity but
concentration of a panicular protective componem. For ex- will not be viable when excreted inro the environmenr.
ample. an extracellular leukotoxin of 1\;f, haemolyrica is in- Genetically engineered vaccines can also be ~1sed as vec-
volved in pathogenesis and causes tissue damage in the tors of genes expressing protective antigens from other
lungs of cattle. 95 and immunity co this coxin is critical for in- pathogens. One of the first ,·ec-cored vaccines to be tested
ducing protection against the organism. It is possible to iso- was recombinanr vaccinia virus expres~ing rinderpest
late (tnd purify the leukotoxin from conventional cultures genes118 and funher developments have led to the develop·
for incorporation into \·accines. but it is more economical co mem ofa recombinant vaccinia virus vaccine that expresses
prepare the leukotoxln by recombinant biotechnology:t 7 both the fusion and haemagglutinin genes of rinderpest
Subunit vaccines are safer than whole killed vaccines. and ,irus under the control of strong synthetic vaccinia ,irus
may be more effective. There is redi1ced antigenic competi· promoters. 111 This vaccine proddes cattle with long-term
lion between the constituents. one ma)' target the vaccine to sterilizfag immuniry against rinderpest, it is safe. heat·
the site where immuni-cy ls required. and there ls the poten· stable, inexpensive, and easily administered, and it allows
tial for differemiating vaccinated from infected anlmals (see serological differentiation between vaccinated and naturally
Marker vaccines, below). infected animals. /\nother vaccinia-based vaccine. contain-
A novel application of subunit vacclnes has been their ing the gene which expresses the immunizing glycoprotein
use to reduce disease transmission by iargeting the ,·aC'cine of rabies virus. has been incorporated into baits and disrrib·
against the arthropod vector.79 · 115 When animals are immu- uted In areas of Europe where the fox popularion serYes as a
nized against a specific glycoprotein presem in the midgut reservoir for rabies. 15· 19· ei . 113 The use of this vaccine has re·
of ticks, che antibody interacts wirh the protein in the tick suited in the reduc1ion or elimination or"~ldlife rabies from
midgut when the tick takes a blood meal. This interferes large areas of Europe. ,\ number of recombinant vaccines
with the digesrive process, thereby reducing both reproduc- based on avian pox viruse, are also being tested or marketed
tive capacity and the viability of the ticks and therefore dfa- for use in mammals.91 Since avia n pox ,~ruses cannot repll·
ease rransmission. t.-ate in mammals these vaccines deliver the required anti-
Yeasts. mammalian, plant. and insect cells ha,·e all been gens while undergoing an abortive infection cycle i11 11i110 . i\
used for the production of subunit vaccines 13· 611 ro. to, and comprehensive demonstration oi the immunostimularory
of particular interest currently is the possibility that genetic effectiveness of such \'accines comes from a recent study
modifications can be made to food plant~ that can then be wilh an experimemal HTV vaccine consisting of a canarypox
fed directly to animals for the induction of protective immu- vector co111air1ingE1w, Gag. and Pro genes of HJV. which was
nicy.'1Ah hough theoretically attractive much work ,,ill need used in combination with l>oosterinjectio1,1soirecombinant
to be done before such a procedure can be demonstrated to HIV gpl20 subunit protein resulted in neu1raliting amibod-
be biologically safe and effecrive. ies in 91 per cent of subjects and COS(+) T cell responses in
62 per cent of subjccts. 2
Live vaccines /\nor her live vaccine vector technique which ls being de-
The conventional method of producing live acte-nuated vac- veloped is the use of attenuated suicide strains oflntracellu-
cine mutants had no molecular genetic basis. That situation lar bacteria carryil)g recombinant eukaryotic expression
has now completely changed. and it is now possible to pin- plasmids. or naked DNA vaccine vectors, into host c.ells. The
point which genes are involved in virulence and stimulation of technique has been most highly studjed using li.ceria
protective immunity. The degr<.>e of attenuation can therefore mo11oqtage11es, and the \'ector bacteria have been shown to
be concrolled by deleting or mutating the appropriate genes.33• be lysed after entry Into host macrophages and 10 release the
35. 80•8' The deletion of an entire gene makes it unlikely that the recombinant plasmids.JS, !!ft, 1 tJ This ls an alternative way of
pathogen can revert to \'irulence, altl1ough recombination delivering DNJ\ vaccine construc1s (see Nucleic acid immu·
could theoreticallyoctur wirh a ,,i ntlent wild strain in the field. nizalion, below).
If an essential gene is deleted or inactivated from a virus for ex-
ample, it is necessary to complement the gene function in uirro Nucleic acid immunization
to alkl\v the vaccine to be grown to high titres. After introduc- ~ucleic acid immu ni'l,ation is 1he title recommended by the
tion of the vaccine into an animal an infection is initiated and World Health Organization for a procedure which has also
protection is stimulated as the immune system sees almost the been called third generation , •acdnation, genetic irnmuni·
full compleme.nt of viral proteins. The lack of a replication zacion. naked DNA immunization. and pol~'llucleotlde im-
cycle, however, means char the virus is not shed into the en~i- munization. The basis of this approach is to vaccinate with
ronment.24· 33• 35 · 81 Asimilar effect can be achieved for bacte· recombinant expreS!;iOn plasmids. comaining prorecrion
rial ,accines by modifying the organism to require n specific stimulating genes. directly into host cells.29 • so. u - Various

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 V3ccination: .\n approach to the control oi infccuous diseases 2-13

delivery mechods have been used to introduce c.hese plas· the ndoption of this technology.9 ~ There is concern over the
mids into c-ells, including injection. panicle bombardment use of ant!biotic-resiscam markers in plasmid selection. but
('gene gun') and direct application 10 mucosa! surfaces. all since other selection maJkers could be used this problem
of which result in nansfec1ion of cells and expression oithe should be easr to o\'ercome. Of greater concern is the possibil-
foreign protein. which then Induces an immune response. ity ofimegration of plasmids into the cell nucleus. which could
There are many inherent advantages to nucleicacid,•ac- lead to activation of endogenous re1ro\;ruses or 10 deactiva-
cines. As a result of the endogenous production of the pro- tion of tumour-suppressor gene~. 76 To date, however. the Fre-
tein the imnrnne response re.wmbles that observed c1ucncy of int~-gration 1s con~idered 10 be less [han lh-e
foliowlng a natural \1ral infcc1io11. Typical responses have plasmids per 150 000 ceUs?1• 76 which is cwo orders of magni-
been shown to include both cellular and humoral immunity, tude lower than nonnul mutntlons. t\n.other concern b the
producrion of memory cells, and prote<:tive immunity of production of anli ·DNA antibodies follo\~ing hnmunization
long duration.A,. ~ 1The D:S:A plasmids namraliyconrain spe- with 0;'\A. although experiments suggest that this possibility is
cific CpG se.quences and 1his helps to crea!e a cytokine micro- ahnoM non•existent. The induction of anti-DNA amibodi~
environment which enhances the immune response against against naked DNA is known to be very difficul1~:z. 43 and i[ ha;.
the no,·el protein.z.s, ""· ' 9 • 60. 99 By delivering the plasmids to been obscn·cd th:u milligrams 01 D:S:A injected into cattle on
mucosa! surfaces. it is possible 10 induce mucosa! immune numerous occasions over a period of more than a year failed 10
responses and immunological memory.:;. J6, 58· &i Since a produce any de1ec1able ami-DXA ancibodies. 108 Only when
number of plasmids can be introduced simultaneously it DNA is delivered in Frew1d's complcceadjuvanr is il possible to
should be possible to immunize animals against a \'llriety of induce anti-O;-.:A antibodies, so regulatory agencies are begin-
diseases at once wiLhout the concern of interference t.hai is ning 10 accept that this should not be a concern. especially in
often observed with conventional vaccines. 18· 4 - Also of the case of li\-.::s1ock. where their lifespan is relmi\'cly shon.
importance is the fact that anr es-semial post-translarional There 1s still a concern that humans consuming animal tissues
modification of the immunizing protein which may be derived from lives1ock inimunized with nucleic acid vaccine,;
required ro i11ducc specific immunity will occur namrally•. may suffer some untold consequence, and ftuure tescing will
DNA vaccines can induce immunity in neonatal have to be performed before using such vaccinated animals for
animals. even if they have maternal antibodies to the immu- food.
nizing agent, 16 and it is even possible to immtu)ize foenises.·• 0·
·11 tr animals were varcinated at binh thei•would first be pro- Marker vaccines
tected b)' passive antibody acquired from the mother. and as The ability to di!Tcremiate animals tl1at have been vacci-
this protection wanes active immunity would develop as a re- nated from those that arc potential carriers of an infection is
sult ofvaccination. TI1is will ensure that there is no ·,,~ndo\,'Of an impor1am requirement, and marker 101\'A - Differenti-
opponuni1y· for the pathogen w establish itselt:33• 10• ate Infected from Vaccinated AnimalsJ "accines have been
Even with these many advantages no nucleic acid vac- developed for this purpose. 109· 110 Efforts are underwny to
cines are licensed for any animal species and the main rea- cradica1e Aujcs;d..,'s dlscase from the '.;etherlands by using
son for this is 1he inability 10 deliver the ,-accines effecrl\'ely. a 01\'A vaccination s1ra1egy.'i2 and similar ,•acC'ines are pro·
DN:\ vaccination is very inefficiem at present. ,\;th as much posed for 1he eradication or bovine herpesvlnJS· l. The con-
as 95 per cent of the plasmid being degraded before it enters cept of DIVA vaccination can be applied ro any disea&e
the ceils. and there is unlikely rn be any major commercial situation and ~uch vaccines arc certain co become impor-
uptake unless better delivery systems are developed to tar- tant In 1he fu1ure. 1 2 • 1·" 5'· 116 Marker vaccines can also be•
get the DNt\ into the nucleus of antigen-presenting cells. used to complement quarantine and slaughter-om policiel>
One improved delivery system that is being investigated in countries where an exotiC' disea~e b introduced. ll is pos-
involves an RNA replicon vector derived from an artenuated sible to vaccinate animals at the periphery of an outbreak
strain of Venezuelan equine encephalicis (VEE) virus and area nnd then 10 differentiate the vaccinated animals from
packaged into VEE-like parncles. 'fhe genes of interest were those iliac ha,·e been exposed to the 'wild' infection. This is a
expressed from recombinant replicon R>IAS that also en- far more economic and humane method or emdicaring a
coded lhe replicase function and were capable of efficiem disease than the conventional slaughter-out policy.8 · 48 · 11 ~
imrncclluJar self-amplification.-1 6 . •o. 85-117 Recem develop- The basis for DIVA \1ICCfne:. is thedevelopmem ofa recom-
ments in replicon technology. based on the fact that the binant \'3ccine from which a gene or gene., have been deleted.
genes can be delivered in an alphtl\1rus coat, ensures that and then 10 develop a parallel diagnoscic kit which detects and
,,·ell over 90 per cem of the plasmids emer cells. and the compare~ the amibodic, developed as a result ofinfection or of
gene is not onlr protected b}' an artificial viral coat /enve- immunization. For e.-<ample. animals infeeted naturally \\ith
lope. but the envelope can be engineered in such a wa} that ,\ujesi~1 s disease or bovim, herpes\in1s-l develop a.ntil:'odies
it targets the antigen-presenting (dcndriticl cells. to all of the proteins of the viru,. However. if the animal i, var·
If the problems of delivery of nucleic acid vaccines are over- cinated with a ~ingle glycoprorcin rgD' or with a gE gene-
come there will still be regulatory issues as potential barriers co deleted live vnccine. they• wilt n()t de\1?lop antibodie!' to iE. An

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2-!4 "mos tlsr: A$pects influencing the occurrence of infectious diseases

animal with antibodies lo gE would ther.efore be considered ro containing CpG sequences.25 Adju~-arus hav.e to act as delivery
be a latent carrier of the virus.s5 Curremly this approach is used vehicles for tJ,e antigen, and the \'Chicles 1101 only affect the ad-
lo dlfferemiate between animals thar have recovered from juvandng propenies but may also have adverse effecis ar injec-
fool-and-mouth disease rFMD) and may be carriers and those tion sites. It has been shown that injection site reactions in
lhat have merely been vaccinated. Modem FMD vaccines are cattle resulting in tissue damage cause significant economic
inactivated and do not contain significant quantities of non- losses. 106 As a result effons are being made 10 find alternative
structural proteins ofF,\111) virus, so vaccinated animals do not melhods of immunization or to find adjuvants 1.hat do nor
have antibodies 10 the non-sm1ctural proteins ofF:VIO ,irus, in cause adverse tissue reactions.
contl'a5t to infected animals.0; , ti8 Readers are directed lO specific reviews for a more in-
depth treatment on adjuvams. 53 711• 93
Adjuvants
Passive immunization
Bolh subunit and inactlva1ed vacciqes generally require for·
mutation widl an adjuvant to enhance immune responses in The new-born of livestock acquire passive antibody from
order to be effective. Puri.fled antigens, however, are generally the colostrum of their dams but this immunity wanes rela·
inherently weaker immunogens 1hru1 whole killed bacteria, lively rapidly, leaving the neonate fully susceptible 10 infec-
since the latter contain components such as iipopolysa- tion by a variety of infectious agents within a few weeks or
charides and CpG sequences which act as adjuvants. An ef- monlhs of birth. Neonatal immunity can be ex'tended by hy-
fective adjuvant is therefore particularly imponanc for per-immunization of the mothers, so that higher levels of
successful subunit vaccination. few e.xperimemal adjuvants passive antibody are transferied. a technique which has
have been shown 10 be effective and mosr oflhem are e."pen - been used In cattle and pigs to increase tl1c immunit) of
sive and often lead to adverse side reactions and tissue dam- neonaces to diarrhoea caused by Escherichia coli, rotavirus.
age. For example, mineral oil-based adjuvams are often or coronavirus!· 75 The druns are immunized priono partu-
effective. but they are not metabolized and remain at I.he in - rition and the elevaced levels of antibodies in the colostrum
jection sire, and for lhese reasons a number of countries in and milk bathe lhe lumen of the neonatal gut, thereby pre-
Europe are threatening lO ban them. Similarly, aluminum hy- venting altachment of E. coll. rotavirus, or corona,~nis to
droxide adjuvants are associated "~th fibrosarcomas in the intestinal villi. It is also possible lo feed animals specific
1
cats.2 • ll"l Finally, even if an adjuvanr induces good humoral antibody for a short period of time when they are mos1 sus-
immunity there is frequently very limited. if any, cell-medi- ceptible 10 disease, an example be:ng the use of monoclonal
ated or mucosa! immunity. Since these latcer types of im- or polyclonal antibodies to the K99 antigen of E. coli co pro-
mune responses are critical for protection againsc many rec1 calves from E.coli induced diarrhoea. 94 Since the infec-
infections this is a serious concern. tion only occurs in the first week or two of life such an
Although the molecular mechanisms by which adjuvants approach is economically feasible.
e.xen their effec:1 are not fully known. ii is believed chat they Passive amibocly ro a particular infectious agent tends
help create a micro-environment where the cells of the im- 10 prevent effective immunization against that agem. espe-
mw1e sysrem can interact and create an appropriace cytok- cial!} when using vaccines in\'oiving live organisms. Repli-
ine caseade for auracling those cells of the immune system cation of lhe vaccine agem is inhibited, and a protective
111hich tead to Immunity. Since most adjuvants help retain immunity fails to develop. In moslcases rbe le,•el of passive
antigen at the injection site, some believe that maimaiL1inga antibody which will interfere with vaccination is higher
depot of antigen ls critical for the establishment of the re- than 1hat which \,111 prevent infection with field s1rains of
quired microemrjronment for cell and antigen interaction. the pathogen. We have noted above the closure of this
Many natural and S)Tithetic immune modulators are used 'window of susceptibility' by using nucleic acid vaccines.
or have been tested as potential adjuvams a:nd it is convenient Since amibod}' in animals with passive immunity is nOl
to classify tbem into mineral oils, plam, bacterial, or chemical/ present at mucosaJ surfaces it is also possible 10 immunize
synlhetic adjuvants, although many adjuvams comain a com- them in the presence of colostral amibody by ,·accine
binalion of these components.JOO For example. Freund's delivery to mucosa! surfaces and 1hereby induce mucosa!
complele adjuvant is a mi.\,u.re of a mineral oil anc,! bacterial immuni[)•. 119 Finally. in ,.aero vaccination wilh nucleic
components, and a newadjuvant is being tested which com- acid vaccines has been mentioned. and in ovo vaccination
bines aluminum hydroxide "~th a synthetic oligonucleoiide is gaining more popularity.37· 88, 10 1

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 \'acclnation, An oppro.1ch to tlw ei>1mol 01 inictliorn, disease;, 2-1:i

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:s• ouQtrt o. ~,.,,.1lH\U... (t.1-, lfiiMl!l.. tt..,\. 1. u rc.m,·uRnL c;,J., 19a9, F.Cfccu;. Q(
fnnnclin ,nncn,,mon Qn bO\'ilte h~tpos,·,rus,J glycQpro1c111s ~nd
12 nROMA'X~. f,Jf.~ nrMf.UQ, I'--\., ~UURT. t Uta:. r .. I, (".O~b. ... 1~9~ i.\J\llbotl)' r~~pun,e thciced h~ rom1J.lin·Jnuc11\.1({-d vn.c-cuK-s in r,lhbit:,..
O!ogno;l~of pc,~b1tn1 aphto,·lru, infettion, nnd Ir, dill'<..,ntla1l~n from \ttl'(itl(•, i. ~13,-.S:?O,
\~ttln:uiou re.,µ(>n$t>- in ("\,'lttJe-l)y u~ ur el\Z)'nlt"•hn~cd tn1muno·
32, tJ..Of"AS, .\,k.., (l'RA,\.\l'Hi:,utl', ·1,. PU,.'(fl\S. l-J., H>CT>, R,, OUl~HOf.. t.,
cloctrotrun,kr blm un•l}-.l.< "1th blocnginccn:d nan,,1ructur.tl ,n•l
uu~~1.,.,1.\s. w \." ,.,,unr,1A~. '""" 2noo, ;\L,1jt.'i1J;\··s dt.',-ea.,c \'lrus
antigens. Nnorlcnn Jo11nt«I of\lctrrinary R,•,wrr/,, ,~. 82...aJI
oradrc,ufon r-~mpafgn <ucct~<sfully hendln~ "" la,1 s1oi:e rn l"hc
u atA:.tA~. r.w.. r.ru:conv. 't. ~\\1n1,...i,. D." u,...:v. 1... 1~8 Pm11J"Cr!cm o! Nt1in·:hmd~- \ i,u1rim1ry• Qt1a11.:rl}~. 22. 103'- tor.
,ir:11 til}tOJ>ro:efn;. in ge11ctl<allr enll'""""'d nU1mrn:1Unn cdl line, for
~ f',\PI\HL )U ,, \.l•~U-\..~, (; ~.. UMU.{' C CT~AllUOU, 5-. IX'qll~1 ~ ill
C\e u., \":tC'C'tnl~ aguh'IM fmmunt- detirft'nC)· rulf'0Viru~ :tpplf1'tl \rlr,>101»·
R~('1T(h, l, 1;...24
Ml~"(')'i .-,.,.•• 19-9,1. Vracc:inc 1mtc11tlftl of 3 hcrp-e,. ,lmpltx Vlru-S type I
mutam \'..ith un csst'mi11J gl}c·upruunn deleu.'<1 Joi,rmll qf\lirqf~·. ~ .
W amr.11-~v.,u:m?\, I,. ltO~,-A,. ,\.. l'IC~. J•• MUMFOHJl, •• II Of~'"· M.. 1997- S:?':'-!l3;?.
Expn,s,.ion of 1hc non•'1nic1u:al pro1<1n ,s1 of cquin~ inOu~n,.1 A
,1rus: Ort<'Clion or and- ,SI antibod, in posi ,inf,'<llon ,'<lulni, ,~r., J·l run:uso,. ,, • woon. n,f,-&1 ,u.~mt. r,n .• 19~J. Antfg,mk ~~ruc1urc ol
JoumaJ of\:irv!ogy M.,1hr11J>, ,;s. ?:,:;...:ml. pohonru~ ,n in4Ctl\1ltcd \'ncc1nc;.., Jm,rnal q/ G11mtrnl Wrolog;·, 7 l
ti85-600
lj !U,.\SOO, t., L~1.t«h. ,\f., 0,\111\AT J, .. CUQtlC I. t-.. JJkOtUIU:.1 8., flt ,oy,s.. C.,
J!i EOkMl,Ml-1¼. \,. 1' ,tuu,u... H .. "'11.1,asso~ G.. f,,;,Ayt, , .. 1),\\TIS.•PO'('Vfl'R. '\ , &
K.P.AH't", s .. U,\lU.'r'. , •• ,10,,LR. '4 .• PA~ranrr P-.?. 4.\UDFJtr. ,u., JOOl.
Klne-t~o(humoral immune ,~pon4.c.' nm:r r,1b1t< \~..c ond ~11:,,.~-.; . T t9,2:. Con~tru(:mm and propt•mC°'.'of a mut,0,mt ofht'rp~
,11ecin•t1on of rnpth't' In, cub, I\ ·u /rm i•u/11,..\ \\1th or \\ithou, ,implex ,1ru, :ypt, i wltll glycopmlell1 II (odl"&••<tt•t'nc~, dcl~1cd.
m•1emall} dcn,·cil anul>odie,~g~lnst 1h~vai:ci1w l'11<Tl11,. 19. ~805- Jmirnal of V/rolog>. i;t.,. 3-1 t-.148.
..3!;. Jfi ..UUJ'M. fl Jr.. k'i.f~t,;U,Hft, r . .\., \\'U.:WS , L.A •• niu.:HI-L \. ,,.. l·ULU.R, f.T ..
:.tllPH\, t .. \\II,,••, WU:'<,~ .• H1,u.oo.c .A., tO.\''.\b, J.H. fill
16 80T, .,. ~ 90'."o:\. eu :zooi.. Cenetk imm.unludon of nt.'On.~ll... •'illc:robc.•s
ln[<•·t 4. 511-320. ,1u1wun··r.ort1l. M. :roo:t rnduC1ion of \lui:w~J ?rorccc;nn i1pin:t.t
Prlm1r, . Het<rologou,Slmfon lmmunodeli<:1ency\ irus hy n ll:-A
s; BJllV>tr::T, :".~.o.. aon, '1,"' "'''-O'.'\, ~.t .. rm, T ct.tll mcnt.ory: Xew \°amne. Jo,rr.al o}I im/Q~·. ·o. 3.109-33!;.
perspectives. lnm1w10To,rJ· Todt,;•. ! ,; 197-J 99.
17 GM;1c ~t.. ,-r 1uu.. u.,. ,u'\RMA, , . \1 ,. 19~ Tn ,,;,u \·,1«im,t1nn (Jf !o.pt."\'ril~
lff .liRAU-S. Jt. B.\8JUs,.'.. I .\. '1 \· '\ "• ORU-~P.: UTIUA' >\:\' Dl ~ UIJfO, , ~. 1998.
p;a:1hugt'fl• fnoe dttci(tn\ \\ilh \·,u:-cult" .. <ont4inlng multiple ng<-'m,. ,\r!,m
C4mpatlbllity of plasm,d,.-.,pr,.,_,,n~ difrc«•m ••1111,ci» m 11 ,ml(lr 0~
!ormut.thoo. Jm,rnol r:f<A-ntral \ 1re,,lf>K.\•. 79. l!l65-2970.
v,,.,..,,,,,, 43. 2,1-301
38 ct:r,;t'<' llt\·, •·~ mn,ur·u, , ... ,p,u,,t, ... .i.111.c .~\1, .... -.!"Ml'flf..).JJR. J.,
i9 1aocm0:. fl_ & P,\..;;JORF.T. f'.·P•. ;g~ Rahat,\.etudlcadon In Selltl,um by (o.,
Jr.:oua-,1.,untA, A. " t.Ot:llu w ZOO:?. O!!la\'('r') c,f prou:ln .inligcn., i\nd
\.lltcina-tian ui.ing v:n:cinia•mbte~ ttct1mbln.in1 \iru11.. Onll&'t.f(1,•poo11
0:\A h)· .a.ucnu,nt.-d in1m,t:llulur b-.1c1i.!n11. lnt,~r,u,1it.Jnn//Qurnt1/ OJ
Jnunml ofl'tLl>f'i,uuy Rcs(mch. 601 • .;&~i:5..
M,..t/<11/ .11,a~/1/n/lJ/{)·, :?lit.;;; ·iitt!
!O BRO\'"'··•., 19"--Jl, fte'\h.-t,· of acddcn,... c~u~ by fn1.-omple~e- in,1cti\·ation
l9 GtOkGt . L\\ • ,\hf),\, .. ,-\., \.Ull.\1ll, 1.• C,UUCM ,u1..198fJ. Enh.1.nc,('fflCnl
of mu=. ~..,/01,mmu /11 /Jfo/1>g(ut1 ,111111/m//1:,111011, ft! 103-10,
of 1n1ec1lou, llo.-lnt ...,..,.,o<onjun,mitl, by n,ndfficd-11,·c in(«:114ui.
21 bH~TO~. C. '- MA'IOS. Ir...\',, 1997. 0o po~nJ.C.::in.nl ~tC'cmHt., tl('C:u(itt br,tine,rhlnotrarh~ns \'lru1i1 vacC'lnc·,Am,•ri,mr /uunutl OJ\ ·?:ffim1ry·
,\.U~trn.11:m t:.att'? A11..$trt1liun t'eti..,rirutry•Journnl. 75. 100-10&. Rw,·ar<h. 49. lft00..le06
22. C.\L\ft.Tft, .,•• 192; La \':,.cc1ni.1uon ?rc\'cnu,·r romre lu lubcrculO)~. ,IU r..UU>T'\, \' "4 1'J\81Ut:. I \.. \:A\ OUU'\:I, I lTffl.-\'AS t,i-,, HUAJ.... ~ lo (,FUUU:.l,
:\la..•s•m 01 Cle. Pnns, P,, 2QO(}. I ei~l lmmunil"tinn by• ll!\A vaccine d~li1 ,ied or•ll)· Into the
:f3 CL\t\»f.1',J:., bl f (!.UJ\\, \\ .• pt! c;,:u,·1· 'P,, U.t~01Uli5f~'\, C.' bOtA\.\.li\, \\ ,. :unnJocit rluld. Sntur•· .\kdin11r', &. 9?9-1•:t!
1S9>. Freund\ complete ndfu\'nm: An e/Tet1h-e but d15ngroenbl• .fl (.I RUT,,\·,. ,,1rne1, ~,. llkOW'\.'Ulc. JI fl,\!tlU);, l,.\. I. <.iA.tlJJCL. P.f~ 2002.
formula. f,r:s,-arch lrmmmo/og; 143, HS--183. 01.,l OX,\ ,·act,lnotlon In ulcro lmluct< mu,o,"1 lmmunliy ,nd Immune
2..-1 D.\ C~IA. X•• ~,U,'A, M.1',, ~Hl.l J. ttk,OC, ..,ti-\S, \t.,\, t, t.:.,\1PI b,'.\S., 2000. mmnol')· in tlwnl·on;ice, Jcmrnnlo/ Jmm11111>/oro,, IAA. tft?i-lAA5
Carhtructlon, phenorn.>ic .tnal)c,ls,•.n d fmmunogonici~ of a 111.5 Ul.:!9 St: t,1tXE.~a-,. G.~.. f\Ull, )I .. tli)\\'I U.. ll,, LI J,1;:()\\'nu. 1.1t. t. l•hl\~t,;\·, I).~ •• l!f~3
double dt!!t>lion mutant oflw~ simplt:'< ,1ru.... 2. Jounud u/t irut,~. lndurUon ofimmunt·medlotl'd glomtruloncphmi< In nonnal mire
14. i9G3-7971. ftnmun~h-d ,,·ith b:u:u.•rlod D~,\. C..'ll11ttttl lm1m,1mlo,r.•1m1I
2-5 IM\1~. 1-f,t... \\1:Ck,\TS,\.i 11,. WAt.n~cu,111)1 . , ., •• ~Clttlktt, J. t, ~,m1.. A,'\I., /11111111110Pf1tl:0Tog)· 1>8. 283-292.

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246 '1t:tlC" <>M~ ,\~pects inlluendng the occurrence ofinfe,1ious di~eases

i3 GIU.'L.;()~. t.~,, kUM. P., 1'1Pf't.,. )t•.'f,. \U,.Xl\~OJ!k, A.I.., u.FJ:0\\1Tlf1 J.~. .&. pa.~l\'cly lmmm1c mkc a.ftel' lmmuniz.lildon \\'Ith n f ) ~..ba.-;ed \'ctcclne.
rJSrrs,:,·, o,>,, 1996. )..lodul11tion <•f renal dls~asc In au,olmmuno /oumal nfG,•11,r,1/ Vlmlt?g)·, 80. 282~2B37,
:'>Z81~/.\\' mlct b) lmmunl1.1nion \\i1h b.1e1crinl I);<;,\, Joumn/ of S... u,·1:-..c;~i·o:,., ,.a., u,,s.• RotUS"SOS.11.4,\...'-:UJ'iJlSO?>,, O.J .. 1998.
r.:.,CJNrimmtal Medl<ine. 163. 13R')..1 ~97. lmmun17.;,Llon ofthe fom»le gen11.i tmct \\ith a 01',\·bns,:d vatc,nc.
.;..; ~tcoos. \\'.S.. 19..;6, ,\d\-anc.c:i. in V~t~rinary RCSt?arth. '171t• Vtt¥d>till)' Jn/«11011 mtd /11wumir.v, tl!l. 322-3:!9•
R,·,ar,/, S8. 516-"25. 0$ ,ur.m."'· ,.,.. HUCIISOX, G .. S-T<>S!'.. D.>!.. "El\E!>JTH, , •• ,\L\IOSD. ,.w.,.
.;; GR."'· o., 1~~- lmmu,,olol,!lcai memo~·.Jlnnuo/ Re,,rtwoft,mmmolo1,>'. ,n~oll, P.O •• 2001. U\'C•31ttl1u;ued 51-r.,;ln;, nf improve.'(! gencr:c :\U>:hillr~·-
11 ·IS-77, ~, ..!opm,•11111/ Biology l/JIJ$,IJ. 105. 17S-J 8;
.;& t.lOflUN, f.H., UA\'1$, ~-l., \JIO:"-SO~, J,f,'., UIAHl.tS, 1•.t;., SL'.UO:\, (J,C,, 66 MA.<:lv\Y, C.R.. 1993, Jmmunologlc;il memory. A<l1'<!t1<1'S lt1 h>m111110/og;,
:m7.U>;J, K. a, 1011:0.'STO.~. nx.. 1..99.;,. Specificrestri,c.uons m the-progression 53, 217-265.
or \'enewelt111 equlni· enceph,tliti> ,irus-,nduccd dbeasr resulung G; )lAt".MY, I>.$.,. FORSYTII. )I,.\., D,M1t.s. P.R.. BtnU~1AN1, A., ff.l:LS-HA\f, <,.J
uon, .sintt,Jc an1i1)() acid change~ in lhe gtycoproteins. \'lrOloS)', 206,
ru~n. ~.«RY""'· M.r,.. 1998.\. 01tTrrcndatlnginfcctlon rrom \-acdnaH<.1n
99.--1006.
m foot•und-mouth distasc \1)1ng u panel of recomblnam. oon•nroc-mtal
l"i K.\'RLA('.IJ, ... , •• N)JTfR. ,. \. \',\l\ DRU:-.t.',. I ITTl.P ..\'4:\ ur, UCJRJ.'", 5 •• VA"!\ prou~ms u1 EUSA. Va,,,·mc, 16. .;.i&-··t59.
l;lQ,:,O)i.l._R:,<;01.u, J., l'ARKlill. M.P,, '1~'6. "f,J. (I( rh~ effcc,
IASr.t.~. e..o•• l992
U8 :\l,\CJ,...\\', o.~.. fOR~,,11, M,1',. t)\\U;b, P.R. ftMtT. J,1' .. l9'9Sff.Antfbodyto
D1 subw1i1 or modlfted live bo\ine herpcsYirus•l \";JCCin~ on lhe cffic::acy ;he nonstructurnl prorcln,.<J( foot-en,t.,1nuuth dl'\C!Wu.: \·[r~in
o!-o n..'Comblnant P<1s1cu~ll.a hr.em(J/J'llt;,(l vncdnc for 1hc prt"\-enuon of vacc!n:itt.'"d antmiil"> c>.po.scd to inftc:t!on. \ '61..•rinmy Qunrrcrly. :?O Suppl
respirwcu)' dt.se~~e in ftt<..'<fiot cah.'i"s. 011mdia1, v~:,ritr.tlrj· Jnumnl. 33. 2,S!>-H.
73, -i~ l.
0g i\1.\IP\, ~ 1~~. f..."<p~~lon of (OrC'18rl gon4!3 in lnS4.'Ch U-"ing.b.-u::utovirus
;8 llM'\\'OOJl, ~., IUN~'$. S..1 CIU'!iP1',, !-,, fCOGf-..A, P, ": WHEl!l..()('.1',, \·,, 2'0Ql, FMtl ,~no,,. 1~89.Antwal 11.itil!"'"fEmomalog:,•. 3 i, 351-372
Cf.lnUOl !>U11l~~. 'I'll•• \-'rlt(UU11J' Rt.YOTd, 1 lS, 530-S.11.
4.9 Uf{\'llY.. u .• Nu,u,. u,. PU!ilCl-:0, •••• ~,,m,. j, 1/~H\'I.AI 14.'.»H:\, ., •• 1998.
10 '-1.-\RC!l,\Nl. f).J.• tt•~M4 LR~, ff.HT.I". G.\ •• MU~c;. C.H., CJIONltB.
,., 1991. Gt:ntttlcitll)' cn~iltL\d'ed ~ubunh vn.cdnc- against fttlJnl~ ll'ukemla
J, MAU..li.t..Rt,

Mnrburg ,iru, ,••c:cln"' ba5':d upvn alpha,1rus repllcons proiet:t g)>lnea \'lrLu: Prou.'Ctl\'c immune re,~pollii<: in cill>- Var:cilw, 9, .89~a.
p£s- ond no11luun.w primates l?rolO&)', 251 1 28-37.
'Ti ~Mk11.S, r .. )l,\ft~FA. ~.f'.. Jl[0~11U)M, a. UL. T'•• UOFFMi\N, S..L. ~o.a.,:AX', J••
,50 HUfif.MA~ ........ "\UlF.C,\H, :,.,, 1. lfl!.f.1.\"l'ltO~J. M.C:.. 199.;, PrCl.h-'1."tfon agalnu
H08AR1 P.<6" Ltw. o .• 1999. Pia.-smid !>~A mntruia \'.lrctne: The potentl.\l
malaria byimmunSu1inn '"ith a pl~rnodl..tun ~10elttt drt'ums para2oitc for i:t,ertomlc in1eg~tion after int~mu.sculrtr injection. Jlunum ~ne
:uowfn nur.:l(.>icacld vaccine. \,flccme. J2. 1529-IS33.
T/t,rctJ))C 10. 75~768.
;1 ~ r t . ttA. 1-tll"JtRPM.- R•• <...1,,0, V. ~ I.FTttmOltTlt. (;.f.. 1~2- lndu<:tlon nf ~
72 ,u;cu... J.K., \IUTCC:.J,.\', J,, O(R11.U.\UCII, M. r .. f~l.)fUOCE". 1,ft., lllk,\S.J,.\\,\. ,:
mueoW barrier to bo\intr hc.tptrS\-iru~· l rt.'J)Ucatlo.n In ('3.ttld \ 'imto,ro•.
,. ~\'O~O. H.. 199:,t. Th<' mucosal tinmun<t. ~)")ftm: fl'(.)m fundnm~ntal
188. 256-26:l.
concepr.s to ,·.1retnc dc,·dopment. \1at:cmct. I0. ?5-88.
6-~ 11:>.:\tff, t., l79B~An inquiry ir:ro 1h,· r.nrt..tes tuul-ejflxis tJ/the ,,anoltt"i!
73 )lESllCA"'L f.,.\DRA.lt.\M, IL &WM. P,l- 199.&,. Common l'TIUC(>.'kl.l immu1w
r,•u,·c11u,t•, a 1li.'<,'U&1 tl(\l,!.01,'t'r.:,! in WmtO[llJJ: U.-'t!Slttm Ct)lmlitsofE11stond,
sv,;1om nnd .stralcg'f"5 for 1hc dcvc!qpmem q( vocciu"'I ai the m~COS1U
1u1rtieutarl)' GIDUW!.lfflhtU. tuut Jmow11 hy 1/ur ii.am~ oftltt row pox.
.surface~ b,:oGrt..\1 r.1.•• l.,\.'>IM, M.£.. ).tGGtt. J.Jt.. MA-if.Ct::\', 1.. ~'Til08lm. w.~
Puhlhh~d by 1h,, amhar. ptinti:-d h~1 S:,:mp~c,n LCJw~ l.onclrm. I ;99_
nu:x•sstOO:, J•• (c..-ds) .. ffmuJt)()()ktJ/.\1ucosal lmmuuologr. San Die.go
;)l ll~M~c;s, n .. SIM~S!lo. J.R.. '- 1a.,\Tll. A,\\'., 1.99,, Ad1urnncs Jnd r;ll'h\"i.ll'Y CA Ac:adcm.rt Pr'l"~S. lnl:', l)J),3Si'~372.
'\y~tcm~ for \·ital\ 11ccincS-m-c(hamsms and pou:ntia1. Ott, tlopmcms fn , .. )lL"iOM, ••.IJ,.JOH.\", ., •• nJt<lUSQ:\, "# * {Cl!:\('ICU.. l~P,, 1966.1\n11genic and
B1olCJgftuf Slom(urdiuuron, 92. 19-2S..
nmleculD.r evoluuan of 1he v:iccme st.man ot ()-pt1 3 polh>\1fU~ dunng du.•
~ J(.):\t;:,,, T.n•• UUAWV.. X (,M'>1l.iX~I. Ft.ft., c.:tlAkOl:'."\'ft. r .. ),,-Ot..OfJN\.. .... .. period oi t'~crcllon of• prfm•ry snc,:lnr. Jo11r,,11/ ofGmwml Viroloio·, 67,
n"n" u.1 , t:mY.<'",, ,- "·'" nou·,1Ax. ~.1 .. , 1999. S)Tlthetlc oUgonudeptJdc-s 69}-;()6.
c:onld1n111y, C:pG moufs cmhunce unrnunoitohicy ofa pepudt m;lhuia
\';&ecine in Ap111s ruonke.n. •!tzccim:. l 7. 306S-307 l,
75 ,,oox, n.,,." BO'S:"(, r.o., 1993, \~ac-cin~ for prt,·eming 1.-nte.ro,oxig-=n1c
Es,·herichiacolf inft..ction in form nnunab. Vacriuv. ! 1. 213-!?0,
5l lr,J..\SHUtll. \l,J. Ii\ \N OIK~CHOt. J, T , 11:)96. Eurly tmrnun{~ b)' B l!t.·e
76 StCHOL<. \\',\\'., UitJWtTH, 8,f .. M.\~.\M, s.v. ~Tl\011.0. 11.J .. lggs. Pou•mlW
gE•nt-g.uh·t! bc,wioe heq>c-,Jw.. , 1 mn.rkerv-.i<..•c:h,e.. Vctr,rinar,·
0)1.,\ \'Occlnr lmcgradon into h,m c~ll iennm•. Ammls of r/!e Stw York
,\lf<'l"o/JiO/(>gJ', 53, 1'1-19b.
Ar,t1tlw1; ofSden""', n2, 30-311.
51i ~st. ,.A." HAHJ.:.~fS..'\, ,.,\,, 19,;-.;. \1rillcontumh1a1jon oHt!Utl bo,inl!
77 !1.lf.J~-..fi:.., U.~ .. Ol l -.:~UW.'l(:i'.. '.\J.B., FOR<1\fUtr., Jt.. "T'ADUJE'):, T., BOT'.'1:1!1\,).."
"t.•rum. n,e Vett!rillur,· Hrcord. 97 16.
qon(".,uJW, l .. 2001. Revcnlon of a Uv~ porcine- rc-produtd\'c :i:id
:,, IJ:\'\tAN, T.<;.... ,·.·~, i:- ,auu ....~ J. ~ :;,-""vat. $t.J,. 1989. Priming f0-r !oral nnc.l re.1plratory ,yndrumc ,iru• ,'llctlno lm·~ugated by r•n•llcl rnu1"1.!on.,.
,~'>tcniicamlbody nwmol)· respou,.s 10 bovine respirutO:)' <)'llC}'tW /01m11,I of<;,11,ru/ Virolot;>·, S2, 1263-1272.
\i~ i.•!l'ect or-amount or \1rus. vinli replicntion. route oi Qdm(n1s.mnion
78 o'H.J<ii'", n.'I. 1998, Rtoeentd.dn1nC'h in ,·acdne ;,Hlju,r.,n1.sfnr S)'NCmtc
:uui mau:nhtl .miibodies. t 'tn•,-f,uuy lmmu11vlugymul
,111d muco.al adn11n1smu1on. Jormlhl ofPlwrmat.)' 1111d Pl1rtmr11.:olog;·,
Jrr,munopa11toltJgy. 2:l. u;.-160.
;(). 1-10.
58 tR,\L11r......:out1L, J.P.. 2002. ~lucosa:~targeted UNA \':l(cinarJon. TNtttls 111
79 OVDR!lf..£0., J,I'., WOS(:,J.Y ,).S,, JACJ..$0.X. LA.. & OOllSO~,·. c., 1988. Htucford
[m1111moturo·. 22. 646-648.
rwtw im:uunlt•d nnd pro1<!C1ed apl11<1 Bo<Jµh/h,rm/ero1//l"'''·;,h
59 );.1t1n;, ,\,:\1 .• 200~ <~p(J motJf, m bme-tla.1 D~;\ ;ind th1nr 1mn1u.ne effec,:1.._ $Olubl~ and metnbmne a»ocla1ed nntigen, rnrm 1hc mldgi,1 .,, tick.,.
nJ11w,ll lk1·:11u,oflmmuno!ogy. 20. 7o<J.-7f,O, Parw//o1 /mm1111ofogy. 10, 40~; 10,
uo KtUtti, .\,:\f.,.w·.t;;YUSG, Y.• ~C".HOHH. f, ~ U;'\V1S, lf,l-. , ~ . 7'h(' rolt- orcpc; S(i O'S'tl:JUU.1!0611. ~ •• Sl'VJ'OI.D'l. c. 6 liL1U RS ~"·· 2.()01. ()cl(!tion of g<•ne 5:?
dinuclcotidt!..~ ln DN\ V.:l:ccine,,. Tfl•mts iu Mi<robu1/ogy. 6. ?3-26. 1:ncodlng gl)"tX>prutelt1 M ~i equlncherptsvlrus 1)-1'• I strnln RacH
~J l..\\J\',\U, <,,, \1JU. (,,. KOS'G. r .. HORSC. T .. ~llK~if..t... );.. :.eruUNi\, ' ·· TU~,t~ resul,s in lncr,ascd immunogmitl~ 1·,tu,'illaryMJcroblolog;· 81,
IILA. t.. PAM LB, ,u,..
:tOOl, Ptitnlng of miemory but not eftte'tor COS 1' cellc. 219-226.
b~· al..iUL'<I bac1erlal\'2cclr.e.Sc/e11c,. 294. 17$-17';19. 81 OUAUK£.,a1, \\', , I.A\tOUllEUX. L, Wl!8at, J,M, & MA'SSUl, U.. 2000.
6;! Lhnfl, ):., CU.Mt rr. T. .t. .RURt. H., 1996. Vaccine sltc·~~ated sarcomas ProlC1t:i,i!•rlcle1(!'(j adcnoviru~ ,·t:ctor;. and comple1~1enHng ceU Jin~
,n cow Clinical e:q1<,rkn"' and aluboramt)' rc,'iew. (1.982-1993), po:.nllal appU~mlon.< of "1ngl.--rowtd replicotloo mu tu.ms for
/01mw! of1Jou,\muko11 A,1fmn/ Ho,p/tal/\:lsocla:/011.32. 91-93, \'ticcin:uion 3nd gene thetap)'. Hwmm <AnL- 71,~rtJpt, I l, l.3~J-l3S3.
6) L.£",\'1-S, P.J .• V.\N DRUXES UTnL~\"AN' O.l!X uui:o.::. s. t: 6..\.ftwg L.\.. ;999. 82 PA~TOAL'I ,.,,, *<8"0<'~111,n, &-.1998. Ep!demiologyand elimina,ionof
Induction ofimmun<.• r....-spon'Sc.-s to bovin~ h(.'rpt'w;ru.,.. rypc 1 go in rnbih in \\1.'Sl~m Europ,. VertJrinaryJo"r,u,,/, 156. 83-90.

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 Vaccination: An approach 101hc control of mfocrious disease~ 24,

83 r.,\10,u,r. r,,. "•=•m•. a. 193~ Epidcm,ol~gi· 4nd oonuo1 of t1>X 101 st'ONtr~ H., ..,11tc1-1ru.. o . ..- ttKucn, ~: 19g;. In t1i<a\'accln:1tton oi chicken
rabies m !:.urope. Va«ine, Ii l";'.38-J75-4. embryos \,ith c-xpcrlmcn1W Newcastle dL~ca54.? and A\1an oll•cmu4fon
8.i JtRffJU;., c ...... 81:UNIG. Mi:., .• H.\T.\.,o,,:. GNOUl. \t .. C()U,'-IA.lrri:, ht PlfR, G.a
vncc!nes.A1·,a11 l>tsea.Jes, 11. 856-863
"GOLDDEllC:. i.o•• aoo:i. (",01u1ruclio11 and chanic,cru..11ion nr o U,e. 102 SfURSIOLO, T., no~n. 1-.. UIN(,, 1.. Mf>OAJ7.Zi\M, 1... nu Jt.b'.CI, (>., ~.\111:-i;, U,,
aucnua1t!d nroA tldction m1,uum c,f P~:ulo.mo11111 nt1n1ginmo ~ .1 IUtAXl!..'71(.M.lR, ,1,, C:AU..-\7.l.J, f'."~, PNOTrt. M,P., ~iNIV,\UUA, I', Ft/ 11,\...tMl!.R, 1..
c11ndida1e Intranasal ,•accln~ ltl/«1ro11 n11fl /mmu11fry. iO. 1507-l~li :999. Goncra<lon o( tis,ue-spodficand proml,cuou, HI..-\ llgond
85 l•USUKO, P,. HRAY, ,\1,. LUO\\'IG.G.\', P.\Rlr.~ '-t , <(;.U.>WJOHS',A., S..\.'\'.Clll:t.. datnb.s•• u.,lng Dt\A mkr~rrny, and ,imml ML\ c1n.,. II matrkc,.
,\., f.\UWX<i , P.ll- it ot..Mrr11, J,1 .. 2ono, R~combinlmt R~A rcpliconi Nan,,, Btt>tt.:bof<>gy. 17. 55S-S6l
dorh-.d frnm atte.nuoiNI \"ennucl:in "'!Ulne cm:cphallti< ,1ru, pro10<1 103 nmR:'1,'TOS, o.lf _19&6. A ..ul"\'tf of m)'tOpJU~m-a d~(t.'Ct10n In \~terinil.t)'
guinea pig. and mlcdrom Ebola hrmorrhagic fovcr viru ... ~ncnn,. l9. \"i.lcdnui. \·f l,am•. 4, 237-240.
142-IS3.
10• TIZAllD. I •• rm. Gre,,c. on1hrrucgaie, and loonnoi cough· A rcvl<ionl<I
86 PIJ\IIK<l. P •• UfitSB61CT. J .. PA'IU\ER. ~S,, ,.,unuso. J•. &,-$Mil It.,.. :WOL hf.MO')' of earl~ \'tzrnnnry vac:dne>. Adt'4UJCtS (11 Ve:trlnnr,· ,\!f'dlrinf', 41.
lnd..i\'idul'l.! nnd hi-vn1em ,-ncc:in1.,~ b.tscd on 11.lphavirus repHeon~ protect 1-24
i;ui1w._ pl~ 3gru11>1 ilifoction "1th l~<u o.nd f.bola ,•Jru>l'.; /our1111/ uf
105 VAt.l!\.7,UHA. P., \1ElJl'lv\, /ti.. RlnTl:lt, \\1,J_ .\.\.t\U·.Rllt. c;, 4' RU1Tfil~ 8,,0,,
brul<>gl•, i5, 11677-l 1685.
1982. S)1l<ht>i< Jnd A>S•mbl)' of hcpatllb B ,urfoce an1igon part1cle, In
87 JllJS.t«){O~ 11. , 9".\1U{fA, M.. LUP\\1G, C .. l),\\'IS, !\.,I,•• JOUSSiON, A.~ <ii: £\1mt.
)'Cast • .\"atut('. 29S. 3-i-:-35().
J..F... 199;. RcpHcon,holpcr~y.s1om...1, rrom autnu::-ncd Venc7.ueJon i?quino
onccpbalitl> ,1rus: Expr=lon 01 huierologous l(OllOS in ,·ltro :ind ,ot. VA:,. oo:..all>GOU>. ,•• OUDHl{1, P,t.., A.\~U~. J.f V.\:-.nL'"'OP. M- DIXON", 1'a,
lmmunwuron "!lnll'!S! heierOlbgou; pat<hogcns /11 m.o. I ·1,0/og:,. 2.39. ~ <rr••"- "·'·· 1g\j0. Tho effect ofvaccine. Jnd antlmlc:toblal; on <he
3119-101. rormudQn ofin)l'C"llon ,irc 1.-lon, In rubprlm•l> or c,p~rimentally
injected beef C11J,·,,,. Cnnndfn11 V.-1eri11a,yJoumn(. 40, i>1S.-2Sl.
88 RAUTO.'SCUU!J..-.;, S., SllARMA, ,,,, .. \\1:.StO\.... 8.1 .. "lC.:MIU.L.:\:, J,, ,u:-.>.ut. n• •
C(K:1-tAAN, ,1., 1.999. Embryo \-accinaticm ohud.~ against ~ewQUIC! 10'; V,\,.°' l>ll\>:-.t-, UrtH-\ ,\"/ nf-N HU~. S., BR.II.US, R,Jt., U,\\•lS, 11,J.. K.<\,H-\0:,.lES,
d!;i,o.re lnrcctlon wuh recomblnan1 fo\\lpox ,·1=
cons1rutis coma1n!ng a,c•• BAn1u>.. L'- « f.l{ttnl L. 1•.1 •• 1~99. lmmunll:uion c>r ncun.11ti. with
inurr(cron$ as,adjt:vanl">. Vaci•:n,•, 18. 42~33, PNA cncodinR a bo\'ln• h,.,,..Mrus gl)topro1c,n b ull'cc1l\'t In ihe
prestn<e ur mutcn\:tl omlhcxm~s. V,ral lmmwrolog.•, 12. 67-77
89 fltlMA:\~. J k~lllR..\111;a. a., ~'(>OU. Modula1Ing spt.-cific primh,g of
Immune ell'totor funttions by 0:SA·b•><d ,-.cdo.1d1>n ,a..,,cglo>. 108 VA.., (>RUS[S Urrtt.•\'A.\' nF'.\' UUfl)r.;. , .. c:;umn. \', wnm, t,.,, PONT-Af\f)Ll.O.
p,..,,./opmenu,f Hiol<>gy f/Jas,f/. 104. 15-"!4 It., AAS...-lS, R., \J\\lUt.h. A,,I. IL\tlUI.. LA. :woo. RC'<'tnt ;c;l\·,,ntt-s. ln Ult: use-
90 RODRt(,UE$, )L\f., Rltlt:IHAO, M,. i-E.ttElR,\•CHlOCtOL-\, \'., IU';.~IA.. L ~ (.():)."?,\ ,
or ONA ,·uc,:lnl'> for the trea1men1 of dl<ta,.,s 01 lonn<:d •n,mbl>.
, .. 1~(1.. Pn:domlnan<t'orC04 Thi
.,na C!)8Tcl ceUs mealed b) -~di,mw m Dr11,; O.-ll1Wy. /lc1,/l'IJ• 43. 13-28
chancwri,.ahon ofihe c,•llularimmunt n,,.pon,c l!\'Mtaocd by 109 ,.A., O)Jt)QC_(J"T. 1 t , ,m. l)l\'a \'ttt.-clnC41 1h..1t rrduct \iRJ.\ 1r.111,ml\"l0n.
immunJ;c.amm \\ h?1 a O~t\ ,·acrlne coma1mng. a rryp~mfh,>ma cnui /oumnl ofBiot<'Clm~f<>gy• .J, 19S-20&.
gc11e. t11J,-c1um and Jmmum't)·, 6':" 3855-3863- 1H.1 v,,N otuc:.Hor. s..t .. ~,. Pn•.~cmr and fuiurt ohewrinut) ,.-u•:d
91 ftOl.jUU), C.11,. AARJU!TT. T., E\'A..~~ ·s.-"., ~tf(.IUN(j,, rt;S•., ct!!tS1U)S. P.O., votclnol11gy:,i miew \lcwri1:t1r)'Q1111r1rrly, t3. 100-108.
BOST'QCl,., r.. ,,,.tu.,,ci,;, o.~ .. t99-.i . A~U)glc tapripo>- recombin:mt \'Occint
UI \"lft..\kl)I, 1°.11., .\I.U:, hll,, AHMAD, 'i:,. JO:'o"B, LA., ttE.'1-i\l. n,, N(i.()llUJ, ft..~ ..
for the pro1e<:1Ion of C'1t1le ngalnM nndflJ""t nnd lwnp)' <kin dl.e.,...
\\A)(\\h\1, ll.M ,u.. I .CZ&A11HF.R. 8.C, . .,_ VlL''"' t ,D .. :i:no2
V'l;$U<..
\acctn.-. 11. 737-742.
l.ong·l~rm ,1vnllzing lmmunfl) 10 r!nd•rp,.,s: in canto "~ctina1cd with a
92 Mlll. l -'-;, ><,\fllllltl.8• .,,,_ 1983. Suppre,,ie,n of n~urrophtl ond momblnant ,.,cdnl• v,ru, c,cprc>sing hignle..-<l,a! 1hr ru,lon ond
l)mphOC"}'tC Cunc"lfon induced b~· n \'llc:t'inalstrain ofbo\'ittl· \'frnl hcmawuumn gli·copru\eln, /mm11,( o/llro/'1&"! i6. -HH-191
dillrthta ,1,,.,, "11b~d "ithout <hi.' concurront adrninistnttion ofAC!lt.
Amu,i,nn ltmflwf of\',•hrit:al)· Rt•SMrdL 4-1. 2366-2.372.
n~ \',',\:\(;, C.,·.. Cl-tASG, T.\',. \ ..',\1,f lll.D, ,\,.,\.,, e.... ~Ht--,.. ,,•• 1H/tSt.. M.l-.
LV.U.ROTJI, J.. f.Jl1.~. ~,p., 11. )1.C.. ux. 'W'.L.. jOSf,, \1.JI .. Y.\:\C:, ,,.c;., on11, ~ ..
9.."l "'iH.OX.11, S~\., ,\1,•S11,;\MISJ1 M, & M()RMO\\', w.,..
:U,()O. Oclfvtn:ysys1ems lOr
•li.\MER. 11. • ,now"· , .• :1101. Synohelic Ptpude-b.1$td Vaccine nnd
malccu13r ,.iaccimrnon. C«rrrnt Opmi(ms fn Mol«ul((f T1:erapy. 2~ l)lajjnO>tk Sysi•m rortJrocdwCnnunl of FMO. 8/Q(oglmls. 29, 221-228.
37-S.I.
n3 , ..•tiss, i;,
.. c;1t~~.\HOITTY, r .. ~001., transfer of cuka~otlc cxpre\$fon
,., .;i,Hf..R.MA..'· D '.\t,. .\CR~ s.o,~S..\DO'f.St,;1 JI',! ,, srrus<;;n1. Jo<\., BRA\.. B.,
plasmid, 10 mamm•li~n ho11 cell, by bacterial ca mer,;. C11,.,,•111 Opfnto,:
M\'SOVLD. t .J.c;::,,. MU~Pt.At, c.c 198!', Prorecnon or t'o.1.h'~ ~s-uin5t
in Bwrttlmofog;•, 12. 4&7-IT.!.
fe:al colib.lcdle>s,s by o,aily ~dmim,1t:cd F.. ,oH 1:99 ,pe<i1ic mono<:lonol
Jnti:bodr /11/«rion and lmm11mry. .f-2, 6,..iJ-65'8. 114 \\'E,:>,,S\'OORT. c# .• TUt,....rtt.A, r... 1988. 80,-tne ,1r-aJ dianh~a viru, in(ecuon
In ptg.l~t) born ta ~w, ,11ecin,.lu:d Jg4ln!rt S'+..1nt !c,-,,r,\ilh
95 ,11c'IAS, P.t ~ \\11.Xlb ti.'.\., 1982.. C)'toro>:ln or Pruh!umlla hfb!nu,J,ittrn
conum11u11ing vim>. R,'1't<11rl< \'et.mna,:.· ~.,,~. ~;. 1~~148
ac1mg on bovine leukocyte,. fnf,rnon and (mmunlty, 3S. 91-9·1
115 \ ..11..L\Ott:-., P• ., .a-.:-.u\ t>.H .• 11)88. Vaccination \\1th ·co:,cc.11cd' antig~ns
96 ~MJ;J)t,.G..\ARO \t.A.0.b'Ell:, &, MA{)SL'l:, 11:..C.,. SJEU""EX, I., HOill JhSS!l."1:• .M., U.1,
for lie~ conirol. Pnms,10/tl/(y 1'11dn)•, ~. 1%-ISB.
1.r.... "''"'· v .• 1997. rl0<e<tlon of :smlbodle< "gnlrm p<>rctne pnrwvi,·u~
non-wuciuml prot<'1n ~Sl m•r dl.s!ngu~h ~e,wecn vacclnnll'Q and tlli \\ln,:,,..:-:x. G.., OHUX(.11.H \'. k Ht'HL\., "" 198-2. \1ultiplrcath;n o( Auj~k)'.>
infeetcd pigs. \,~ i,ritta()•M/a'oiJiofogy. 5-1. I-lb. dlseMe ,1ru, afuir cxpcrlmeno31 lnfoctlon \\i<h high 3ml low <l0<es or
9; s-rrn,. 11.,.• ., ~u"""-~· D.M.. 2001. The rcgulouonofDNA ,11ccin<> ,,ru..~ LtmtralbftJII fm \'t'tllrinnnm:.:JUln. 29, SGl-675.
Curum Opin/or, In Blot.-cluwl<>K)'. 12. 299-303. II';' \\()l..ff-. J..A,, \L\LD~t. )l \\',, WU.UAM-11-. P,, A:,(;Anr. (i.., IA,-;J, , ..... C\C.'\:rN. P..L.
98 "$1'R.L-...f,. ~. Dlf."1ft1CU, G.. ~lt.'\\'l~, !>,. U.Jl \U:Ul.t...''. ..... f.tt...,t SCJ:ll.\', I,., 1990, Din.~ atene tr.in-srtt Into mo1tse mwcJc in ,1,0. :kitwct. 2.;-
GOUlL ._....• :woo~ Novel bnr;umal )Js1cr1l$ for the. dch\'ery o( MGS-14611.
rt'COmbinan1 prolcln or DX,\. nMs lmmmrologymrll Medfc,,f 118 YIL\t\, t .. 'f)ll. D., fON-£$. L, ()\\1:S~, s.• own~JA'", M,, >?[n.U~. G.,
.\olitr1>b1olog,-. 21, 299-JO,. \'A.\1..\K\~,....,. ;. IMLt. u... 1988, Vrorttdon of cattle ug1Jn-(_;1 Rindct"p61
99 STA.", \.(;., CAS,\RM.-S .. HMUMCA.,'U, T .o.. XUNMA~. u.~,. (,I fJO;\t\, C:..A., aoo:. \\ilh·,,accinia ,iru" recombinant~ cq>rt<ssln~ the I IA or f g&trh:. Scitm:e,
C1>G motifs o( DNA vaccln<> induc,· the cxpre,,slon of chemoldn~s and 242. 1058-1061
~1HC <lass II molecule, on m)'OC)Se,.. F.tlropean Journal of/mm1111olo[0·, H9 Z.>\.KHl\tltOtOUt,,, ,\,-.;., r, \lk, C., ;\IIJTWIIU, C., PAS-P, ~ lt.Af,.A•IJ..,\'fKAl>.A, M .,
31301-310. r,RIMLL, e,e,u~. 1...,. i, TI~OO. , ,x., 1999. ~luco,al immunl~uuon o(
100 \TRAUb. o.c.• 19<90. lnrcctlow. bo\ir'lci :hlnotr.u._ilc-lti'(. t,1: 1>Js-n1.1t "·· \~UR· cah-e<. \\i<h n,c,omb,n4nl bovlrw odceovina 3: lnductlon or protecthe
us. 8., t~,.
l"iru., f11f,xrio11>ofllum/11ams. Arns1crd:ini, EJsc,~r. pp. lmmunlt~· 1nbo\1n~ h~rpe,-1ru1-l. /011mnfc/C•11rml Virq/ogy, 80.
71-108. 1263-1269.

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SECTION TWO
Protozoa[ Diseases

• .... •.. .. ..•

.•••..••
•

... .
•• •
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•
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 12
African animal trypanosomoses
Synonyms: :-agana, 1serse ny disease

R J CONNOR A)<D P VAN DEN BOSSCHE

Introduction by blood-sucking reduviid bugs. certain small wild animals

and dogs harbouring the infection. The second form is
The uypanosomoses are disease, of humans and domestic ani- human sleeping sickness. This occur" in :\frica and is trans-
mnls chat result from infection \,ith parasitic protozoa or the mitted by blood-sucking !lies of the genus Glossina. com-
genus Trypanosomt,. Trypanosomcs parasitize nil classes of monly kno1,·n as ·tse1se flies' or simpl) as ·tsetse·. The
vertebral~ fish, amphibians, reptiles, birds and mammals. majority of animal diseases caused by trypanosomes occur
The parasites, with the exception or Tryptmosoma equiperdum, in 1he tropics. In Arrica, several species ortse1se-1ransmittcd
the cause of dourine. are transmitted from host to host by hae- rrypanosomes cause African trypanosomoses in domestic
matophagous vectors, and usually cause little appreciable animals. which in southern Africa arc collectively known as
harm LO either the vector or the vertebrate host. However, sev- 'nagana'. a \vord derived from 1he Zulu ·nakane' meaning
eral species of trypanosome, which parasitiZe mammals are tsetse lly disuase. An Important form of Lcypanosomosis
less well adapted and commonly cause disease. kl1own as do urine, caused by T. equlperdum, also occurs in
The trypanosomoses form a group of diseases. each spe- southern Africa, hu1 has no arthropod vector (see Chapter
cies of pathogenic trypanosomc causing the disease trypa- 13: Oourine). 'Surra· is transmitted by biting flies other than
nosomosis. The course of a uypanosomal infection varies tsetse flies and. although it occurs in many parts of the 1.rop-
considerably and depends upon both 1he species of trypa- itli, including nonhem Africa. it is nm present in southern
nosome and the host Involved. rrypanosomosis is generally Africa.
characterized b>• the imermim•nt presence of parasites in Tsetse flies have recently been disco\'ered in Saudi Ara-
1he blood and imermittem fever. Anaemia usually develops bia.·· but they are only known 10 beof imporrance in Africa.
in affected animals. and this is followed by loss of body con- south of the Sahara. where dw diseases they transmit are re-
dition, reduced productivity and, often, high monality. sponsible for great economic loss (see Chapter 2: Vectors:
The firs1 report 1hat associated trypanosomes with dis- Tsetse flies).
ease was made from India by Evans61 in 1880. He found rry- The large populatiorn, of wild animals, which have
panosomes in the blood of camel~ and horses which were 1hrived for millennia in 1he 1seMe·infested tracts of Africa.
affected by a disease known locally as 'surra·, and which is have evolved \\1th the-~e flies and the rrypauosomes they
now known to be caused b)' Tryp<111osom« ei:ansi. Subse- transmi1. Hosts and parasite~ have become mutually
quently. Bruce35 made [he major discovery in Zululand, adapted and coc..,ist in a balanced rc,lationship. Humans
South Africa. 1ha1 trypanosomes were the causal organisms first brought domestic animals into the tsetse fly belts of Af-
of'nagana ·• ortsetse ny disease. The early review of 1rypano- rica relatively recently. and although humpless cattle or the
somes and trypanosomoses by Laveran atid :-.1esnn, 155 Bos ra11r11s t}'PC, from which the present-day taurine breeds
translated from the French by Nabarro in 1907, was followed such as the West African Shorthorn. x·oama, Muturu and
by Wenyon 's3 18 contribution in 1926. Subsequent major re- Baoule are believed 10 have descended, were introduced
viewi of uypanosomes and African trypanosomoses were into northern and \\ c.-stern Africa from 4 500 13C onwards,
made by .Mulligan and Po11s. 204 Ford.as Hoare, 111 Jordan, 136 the humped Zebu. Bos i11dic11s type, arrived some 3 000 years
and Stepben.2H The considerable interest in ll)'Panoso- later, and did not reach central and southern Africa until
moses arises from their imponance and 1he intriguing biol- around AD i00.80 Goats and sheep were introduced at aboUt
Og) of lhe causal organisms. the same 1imc. Bee-0use of this relative!)' recent imroduc-
Two forms of hum,m trypanosomosis exist: Chagas' dis- tion, the relationship between tsetse-transmitted trypano-
ease occur~ in Cem ral and South America and is transmitted somes and domestic animals has not fully evolved and
251

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252 w..,tos rno: Prornzoal dl~e«s~fi

infection with these parasites frequently produces disease. mosis in southern Africa is largel} determined by the dist:rl-
However. the West African humpless cattle have had longer bmion of tsetse in three major fly belts. The first Oy belt. with
10 adapt 1han Zebu cattle and this may explain why the~ Glos.•i11a mor$itans 111orsi11111$ and Glosiinn pallidipu as the
possess the trait of trypano1olerance. They are able 10 live main tsetse species. links tsetse foci in .Malawi with the large
,,ithou1 drug treatmem in tsetse-infested areas where other Oy belt common to eastern Zambia. Zimbabwe and Mor.am-
cattle die. Furthermore. some indigenou~ breeds of goats bique. The G/ossilu1 mor$ita11s c,•111ra/f$ fly bell CO\Crs west-
and sheep, such as the Dwarf goats and Djallonke sheep of ern Zambia, pares of western •\ngola, the Kwando River
West Africa a nd the small East African breeds. arc m()re try· drainage in Xamibia's Eastern Capri,~ district and the Oka·
panotolerant than are exotic breed,. 211 \'ango Delta in Botswana. finally. a third Oy belt with the
The ravages of nagana have long been recognized by the species C/ossina austeni and Glossina brerlif)fllpis coven;
inhabitants of southern Africa, and early attempts 10 intro· pare; of :-rozambique·s southernmost Matumine district
duce livesmck into tsetse-infested areas were unsuccessful and norch-eastcrn KwaZulu-Xatal in South Africa.
as draught animals and other stock succumbed to the dis- Concerted efforts to control tsetse over the pa<t SO years
ease. The devastaclon whkh resulted from the rinderpest have resulted in significant changes in the distribution of
pandemic of the 1890s (see Chapter 49: Rlnderpest) de- tsetse and tserse-transmiued uypanosomosis. Unfortu·
stro~•ed almost entire populations or wild animals and mil· narel}. few of these achievcmcms have been sustained. In
lions of canle. Without hosu; on whid1 lo feed. tsetse many countries of southern Africa, the current distribution
disappeared from large areas. \Yid1 the constraint of tsetse- of 1,e1se and. hence. tSelse-transmim:d trypanosomosis is
transmitted trypanosomosis removed, settlement in not muth differem from the ecological limits of the fly disrri-
Zimbabwe was rapid. Some 25 000 cnnle had sunived the bucion.
rinderpest cac'8strophy, and af.cr restocking there were over The epidemiology of bO\'ine trypanosomosis differs
2,3 million carrle in thccoumryby 1929. 11 II01\'ever. at about significantly between counrries_ Generally .speaking. three
dlis time tsetse were dispersing from residual pockets. and different types of epidemiological simations can be distin -
trypanosomosis again became a problem for live,,tock 0\\1l· guished. Fir,1. in Namibia and Zimbabwe. progres,ive
ers. By 1931, tsetse were spreading at a rate of2 500 km2 an- tsetse control operations have gradually pushed the tset-:e
nually.4? and game elimination co control tsecse began in Crom 10 its currem position. The interface between tsetse
1932.43 Since then. strenuous efforts have been made 10 and canlc occurs at the edge of the artificial barrier.
contain the tsetse fly (see Chapter 2: Vectors: Tsetse flies). pre\'enting tsetse from rein\·ading cleared areas, with the
In many other pans of southern Africa. livestock 011~1ers pos~ibillcy of cattle entering tsetse-infested country.
have also had to live with the tSecse fly and its consequences. Secood. in Botswana, ~'1ala1,i ond South Africa, imeraction
Tsetse infest 10 million square kilometres and affect 37 between tsetse and cattle occurs at the edge of the ecologi·
countries. which makes African animal uypanosomosis a cal llmlts of tsetse clistribucion. In principle. ranlc are not
problem of rn1ly continemal magnitude {Figure 12.l ). They able to C'ntC'r 1setse-infe,1ed land. Finally, in -\ngola.
live in frost-free areas that have an annual rainfall of650 mm Mozambique and /.ambia. ca1tle ll\le in c.serse-infesced
or more. In arid. marginal habi1a1s. tsetse only e:..;s1 in the areas. They arc ,urrou.ndcd bi ,~etse flies and are subjected
better wooded and better watered s trips where the ho5t spe- 10 co111inuous diailenge.
cies concentrate during critical times. such as in the late, ln :'\amibia, bovine trypanosomosis is most pre,•alenc
hot. dry season. In areas of mixed agriculture. canlc are val- along the Kwando River on the western boundary of the
ued for the milk. draught power and the manure which they Eastern Capri\'l.29; The deploymem of odour-baited. insec-
provide. They also represent a means of hwestment. In most ticide•treaied targets along tl1e northern part of the Kwando
m.ral areas people rely on poultry and small ruminams for River ha, resulted in a slgnificam decline in the incidence of
meat. caule only being slaughtered for Important soclal cer- trypano~omosi,. Occ.isionally. cru,es of nagana are detected
emonies or to realize large sums of money. )lost of the in canle grazing in the vicinity C>f Katima l\.lulilo. situated
seuled areas of the tsetse fly belts of southern Africa are used near the border with Zambia. These latter Infections are as-
for traditional mixed farming. but the presence of tsetse se- sumed 10 originate rrom the nC'ighbourlng Sesheke area oi
riously handicaps development.'·- The general Impro~·e- 7.,ambia: cattle that graze along the Zambezi Rh er are
mem of agricultural production depends on the ~reatN use probably challenged br t,etsc that cross the rlvcr.
ofanimal traction, in place or manual tillage, enabling larger In 7.imbabwt. tsetse has been controlled successfully
areas to be more clficientl~· cultivated. I Iowev,•r nnimal~ and control effort. ha\'e bc:,'n sustained. This is partly due to
alone cannot ensure this developmem. The availability of the country', fa\'ourable position at the edge of the natural
credit schemes and oupplies of seed. fertilizer and imple- limits of th,• tsetse distribution. Since 1981. approximate!}
mems are also needed before the potential can be realized . 55 000 km' of tsetse-Infested land ha< been cleared of t~etse
The presence of tsetse is. nevertheless. the largest single ob· using a \'arief} of methods.268 ,\ combination of odour-
Stacie to progress in affected area~. baited target~ a11d insecticide-treated cattle consti1u1e a
The distribution oftsetse-cransmitted bo\ine 1rypano•o· 'barrier' to reinvasion. confining tsetse m areas less ~uilahle

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 African animal tryp3nosomt>$t/S 2~3

0 Tse1se lly

(I Cante
al Trypanotoleiant cante
Figure 12.1 General d1strib1.o;,on
oi tserse t,,es a:id cattle mA/r,ca

for agriculture or along internarional borders, and pre\"em- free of cattle. Because canle-flr contact is low, bo,ine rrypa-
ing them from invading pre,1ously cleared areas. By the late nosomosis is 1101 a maJordisease in Botswana. The more im-
1990s. bo,ine trypanosomosls was prevalent only in the mediate risk is to the large number of murists who ,1s!t the
northern and easrern pans of the coumry. In these areas, wildlife areas, as the) face the risk of conuacting sleeping
canle comracr llle disease as they graze within or along the sickness.
edge of the barrier29 ·1 where they are e.~osed to rsetse Oy. ln Malawi, where the human population density is hlgh.
Challenge is generally low and bovine trypanosomosis does the pressure on land and the concomitant clearing of \'Cg·
not constitute a major constraim 10 animal health and pro- etation for cultivation. confines tsetse to game reserves,
ducrion. :S:otwithstanding the favourable dlsease situation. national parks or forest reserves. Consequently. cases of
the trypanosomosis threat Is real. A breakdown of the bar- nagana are restricted to areas surrounding the rse1se-
rier to reinvasion could e.xpose o,-cr one million highly sus- infested zones and areas adjacem 10 neighbouring Zambia
ceptible cattle in the commercial and communal farming or Mozambique where tsetse is prcsem.m The pre,·alence
sectors 10 the risk of rrypanosomosis. Cominuous disease of infection depends upon the cattle-ny contact and varie,,
surveillance is essentl;il. The speed 1,ith which tsel.Se 0} are between areas and sea~ons. ll decreases with increasing
capable of reinvading cleared areas was demonstrated on an distance from the tsetse infestation. Although bovine crypa-
expetimental basis when a portion of I he target barrier was nosomosi~ can assume epidemlc proportions. it is localized
removed in the northeast of the country.= The Impact of and doe:. not con~titutc a major threat to livestock produc-
the breakdown of comrol on animal health was also clearly tion in 1'1ala\\1. The ever-increasing demand for land \\ill
shOl\11 probably further reduce suitable 1se1se habira1. However.
Tsetse flies Infest two areas in B01sw1ma. the Kwando- the expansion of the human population will compel people
Llnyami-Chobe fly belt and the Okavango fly belt. Most of to live at the edge of tsetse infesta[ions. which is likely 10
the tsetse-infested area comprises wildlife zones that are result in an increased incidence of nagana.

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254 s,n10, n,'O: Protowal diseltses

In the lace 1850s. 1setse flies infested the Limpopo, laboratory experiments, it is often difficult co relate the re-
Mpumalanga and KwaZulu-Natal provinces of South Af- SL1hs 10 1he field situation.
rica, t10 but the rinderpest pandemic Jed to the complete
disappearance of tsetse from these areas. However, in
Aetiology and life cycle
parts of Zululand. including the present-day game reserves
of north-eastern KwaZulu-Nara I, pockets of wild animals Trypanosomes are protozoru, parasites ot' the genus Try-
and tsetse SUTVived. and by 1905 nagana was again threat- P((nosoma. order Kinetoplasdda, and have, as characteristic
ening livestock in the region. Clossina pallidipes dispersed organelles, a kinetoplast and a flagellum. Typically. trypano-
i1110 farming areas causing serious epidemics. The advent somes are dJgenetic parasites and thus require t\\'O hostS 10
of organochlorine insecticides provided a suitable means complete their llfe cycle: they multiply in the blood. 1issues
to remove tsetse, and between 1945 and 1952 C. pallidipes or body fluids of a vertebrate host and, wl1h the exception of
was eradicated from Zululand by large-scale aerial spra)'· T equiperdwn which is venereally transmitted. are ingested
ing ope.rations.65 Thereafter. the two species found in Zu· by a haematophagous iTwertebrate vector. With a few nqt-
luland, C. austeni and G. brevipalpis, were responsible for able exceptions. a cycle oi development a11d mamration oc-
sporadic cases of bovine 1rypanosomosis. t43 Despite the curs in the \'ector, after which the parasites are uru1smined
restricted distribution of tse1se, a widespread outbreak of to another vertebrate hos1 as the veclOr feeds. Transmission
bovine trypanosomosis occurred in J 990.13 is ei1her by inoculation of trypanosomes "~ill saliva or by
In Angola, nagana is a serious constraint. Over a quarter contamination of mucosa or broken skin with trypano·
of the counuy is tsetse-infested95 and 1he 3,5 million cattle somes in the vector's faecal ma1erial. voided during the
are kept mainJr in the south-western, tsetse-free areas, al- blood meal. ·n,e type or developmem cycle wilhin the vector
though there is a small population of t.rypanotolerant cattle determines whether or not infective, metacyclic parasites
in Cabinda Province. Control measures are currently limited are present in saliva or faeces. On this basis manunalian try-
to the use of 1rypanocidal drugs. panosomes are classified Into the two broad sections or·sali-
Tsetse fly arl!: a serious threat 10 livestock developmem in varia· and ·s1ercoraria·. 11 t
about two-thirds of Mozambique. More than two decades of In Africa, the pathogenic trypanosomes that cause
civil war reduced the cattle population by more than 80 per sleeping sickness in humans and nagana ln domestic ani-
cent and made bovine rrypanosomosis less prevalent than mals are saJivarian, and cyclical developmen1 occurs in
before. Currently, most of Mozambique's livestock is dis- tsetse flies (Figure 12.2). Transmission of a ny trypanosome
tribu1ed in areas where tsetse is either absent or present at species can take place mechanical!)• without cyclical
low levels. Nevertheless. bovine trypanosomosis constimtes changes occurring in the vector. ln nature, this ls effected
a threat 10 the development of rural areas and could be a by biting flies, such as Tabanus, Sromoxys and Lyperosia
serious constraint to restocking programmes. spp.. which feed on more than one animal before reple1ion.
In Zambia, on the ocher hand, bovine trypanosomosis is The fly remains infective for only a short time, although In
a disease of national importance. Three-fifths of the coun- So.uth America and .Mauritius (see below) mechankal
1ry pro\ide suitable tsetse habitat. Despite i1s long history transmission of Trypa11osoma 1•i11ax by haematophagous
oft secse control. large-scale operations have met with lim- flies 01her than tsetse has enabled the parasite 10 become
ited long-term success. This is largely a1rribmable co the established. E.xperimen1ally, tr)rpanosomes may be trans-
absence of natural barriers, which results in a continuous rniited by·syringe passage· of infective blood. 270 'Surra' is a
threat of reinvasion oi cleared areas by 1serse from adja- disease that affoc1s a wide range of host rulimals. and it oc-
cent infesced zones. ln contrast to most other countries in curs in 11:orth Africa. the Near and far East, Central and
southern Africa, but in common with countries in eastern South America. the Philippines and Mauritius. It Is caused
Africa, cactle are kept in tse1se-infested areas. Under b)• T. evansi. a dyskinetoplasdc form of which- known as
these conditions, disease challenge and prevalence of Trypanosoma equinum - also causes disease in equids in
infection are generally high. Consequemly. bo\ine try- Central and South America, where it is known as 'mal de
panosomosis constitutes a major constraint to rural de,,el- Caderas' or 'Murrina'. These parasites bave adapted to an
opment in large areas of western, southern and eastern cmirely mechanical, non-cyclical mode of transmission by
Zambia.4 '1 blood-sucking flies ocher than tsetSe. TryptmoMJma rheileri
Early work on ttypanosomosis. much of it conducted in Is a stercorariru1 parasite of cattle which deserves greater
southern Africa, concentrated on describing the nypano- mention. lt was first reponed by Theiler281 in the then
somes and studying the nawral history of the parasites, their Transvaal in 1903, and has since been found 10 occur in
vectors and their hosts. The greatest advances in our know- cattle throughout the world. It is ltansmiteed by rabanid
ledge of trypanosomosis over the past two decades have flies and is widely regarded as being non-pathogenic,
been made in the areas of molecu lar biology and immunol- bu1 in certain circumstances it has been associated with
ogy. rather than at the field level. Although ex1remel)• valu- disease. II 1.z80,J1s
able research has been conducted in carefully controlled The following accoum of pathogenic rrypanosomes

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 Mricnn animal 1rypanosornoses 255

deals only\\ith tsetse-transmjned salivarian parasites. unless which e.~tends anteriorly. and which i~ connected to the
otherwise indicated. They are important as they cause severe body by an undulating membrane. Beyond the anterior ex-
disease. Human sleeping sickness is caused by Trypa11osoma uemicy of some species, the flagellum may extend free of a1-
/Jrucei gambiense and 7: b. rhodesiense. Wh ilst these two sub- tachmcm to the undulating membrane. The beating of the
specfes do infec1 some domestic and \\~Id animals. there are flagellum pulls the 1rypanosome forwards. impaning char-
other, more significant pathogens of livestock which are acteristic motility. Wilhin the cell. in a posterior position
found in threesubgenera:Duuonel/a, Na111zomo11asand Try- and at the base of the flagellum. a kinetoplas1 is found, and a
panowon (Table 12. 1). The founh salivarian subgenus. Pyc- single nucleus is located almost halfway along the bod~·. In
110111011as. is represemed by only one species, T,ypa11osoma the 1setse fly. trypomasligotes transform to epimastigotes
suis, which is of little economic importance. (synonymous with 1he crithidial stage in the old terminol-
t\s a consequence of their pathogenicity and their com- ogy) in whicl1 the kine1oplas1 has migrated anteriorly, to a
plex and fascinating biology, the sali,•arian rrypanosomes posirion adjacent to the nucleus (Figure 12.2). Differences
have been, and will continue to be, the s~1bjec1 of intense re· in the morphology of the trypomastigote stages of the vari-
search.111e remarkable alternate adaptations of these extra- ous species form the basis for differential diagnosis. The
cellular parasites to mammalian and insect hosrs are major characteristics are clearly seen in thin blood smears.
reflected in morphological changes 1hat are readily detect· stained 1,~th Giemsa's. Leishman·s or other Romanovsky
able by light microscopy. Bloodstream forms are trypomas- stains. Trypa11osoma co11golense (Figure I 2.3a) and T. 11ivax
tigotes (figure 12.2); from the posterior portion of an (Figure 12.3b) are monomorphic parasites. whereas T. bru-
elongated body, some 8 10 35 µm long. arises a flagellum cei is polymorphic [Figure 12.3c). T1ypa11osoma congo/ense

~
• hx
~ ·
le
>
-% ; ;.]!-::::,
~
T vivax

ph
s
I
T. congo/ense oe Figure 12.2 Life c-ytles of
Afrttao pathogenic lr)'panosomes
ca = card1a f= orovenmculus}

er = cmodutt
r brocei
hg = h1nogu;

hx = hypoi)haryn.1.
= labium
.. -. - •• t -- . - ,. -
mg le = Jabrom-ep,pharvnx

II):., .. ,e =proooscis
ma = Matp1gh•an rub.s
mg m1d9ut
"
c)e = oesoDhagus

Ph = P,'Jar\>TIX
~ hg pm perilroph,c membrane
=
-~
"C

= rectum
s = salivary glands
= ir.:~cil~ metatrypanosomes

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256 = x rwo: Promzoal diseases

is lhe smallest ~pecies and b bet\\'een 8 and 20 µmin leng1h. Tl'}'Panosomes are therefore robust organisms. TI1e cy-
It has no ftee flagellum. and the kinetoplast is usually sub· mskcleron of closely packed, parallel micro tubules. strongly
terminal and marginal.2 ~0 T1yp<mosoma 11i11a.t is between 20 interconnected by associated pwteins. is ve,y resilient and
and 26 µm long. and ha~ a long free flagel lum and a large, helps 10 retain the body's ~hape. The cytoskeleton suppons
often terminal kinetoplast. The long slender form of T. bm- the surface membrane (Figures 12.4 and 12.5), which. ln
cei is characterized by a long free nagellum. the presence or a mammal-infective stages. is coated with a surface glycopro.
conspicuous undulating membrane and a sub-terminal teinsome 12 to 15 nm thick. Within the cytoplasm ofthecell
klnetoplast. The posterior end is narrow and or1en pointed, arc organelles, which arc supported by Intermediate fila·
and this form is ben,·een 23 and 30 µm long in conuasuo the mcncs. 126 A nucleus, Golgi apparanis. rough and smooth en-
shon stumpy forms \\'hich range on average benveen 17 and doplasmic reticulum. glycosomes, lyso~omes, vacuoles and
22 µm. The short stumpy form has a well-de1·eloped undu- single mitochondrion are all presem. :,0.'1 icron1 bules also de·
lating membrane and usually lms no free flagellum. The limit the entral\ce to the nagellar pocket, which is an invagi·
lengths or the intermediate fom1s range from 20 to 25 µm. narion of the cell membrane from which the flagellum
ll1e kinetoplast or T. brucei is smaller than that of either '/: emerges (Figure 12.5). This pocket is a highly specialized
11irtax or T. co11gole11se. Tryptmosoma sim iae is between J2 area or the cell membrane at which endocytosis occurs. in a
and 24 µm long and displays some polymorphism: most manner similar to that found in mammalian celb. 305 Serum
parasites are long stout forms. sometimes with a short free albumin. lipoproteins. iron-bound uansferrin and other nu-
flagellum. Short forms ofT. simiae resembling T. ccmgolense trients are endocytowd from the flagellar pocket br blood-
and long slender forms also occur. 111 stream U')'Panosornes. The exchange of the contents or the
The reasons for the cyclical changes from trypomastigo1c pocket is assisted by the consram. pump-like beadng of the
co epimastigotc and back 10 1rypomastig0te, in lhe course of flagellum. which is made up of an axoncme and paraflagel-
the trypanosome's life cycle. have become dearer as a result lar rod. and which also provides tl1e means of propulsion. It
ofultrastmc!'ural and biochemical studies. The morphologi- is ensheathed in surface membrane and is attached to the
cal changes which are readily seen reflect the tr~'Panosome's body by a series of junctions and numerous filamems. 262
adaptation 10 the different physiological environments en- The beating of the flagellum draws the cytoplasm of the cell
countered in mammalian and insect hosts. Ultrastructural, up into a series of transient crests which form the undulat·
biochemical and immunological smdies 196• 299• 300 have nm ing membrane.:i-00 A1 the base of the nagellum is the basal
only revealed the reasons behind the cyclical changes in body which is associated \,ith the kine1oplast. This laner
morphology, but have also elucidated the parasite·, survival structure consists of bundles of o:-:A fibrils locarnd in a dila-
tactics. Tl'}'Panosomes show remarkable adaptation. They tation of the cell's single mitochondrion. 111• 300
survive not only in the turbulent bloodstream, where they In the mammalian host. the abundance of glucose as an
face vigorous immunological assault, but they also with· energy source is associated with repres~ion of the mitO·
stand rhe digestive en.z~mes of rhe tsetse fl)"s alimentary chondrion which lacks cristae and is tubular. Energy is de·
tract. ri\'ed from the metabolism of glucose 10 p}rruvate. 299

Table 12.1 Pathogen1c1l'i TRYPANOSOME TR\'PANOSOME !CATTLE GO.!\TS SHEEP PIGS HORSES DONKEYS
of sahvarian ll'IJ)anosomes ,o SUBGENUS SPECIES
lrvestock
Trypano:;(1()(1 r t11w:e' + ho;
, e~-ansi1 +, I:+ .++
T et1U1f)erd:sm~ ....
Nsnnomonas T rongolense +4- ~ ~

T. s,m,ae T -·-
Durronells f. Vl°l'IA ~ .... •
~otnon<1s T su,s' -
N:;tes
1 Under usual field cc,ndi~ons. btit which is mc-:!;fied by manv fac101s
2 T bruce, gamb,ense and i btucei thodes,ense cause human sleeping s,cmess in West and East Afnca
respecuvely. and ha\-e animal reservoirs 1n which oa!llogen1clty 1s low Human Sieeo1ng sidness in SO\ltnem
Africa - in the Okavango Swamps. Luangwa ana Zambe11 valleys - is caused by T brocei rhodesiensg
Trypanosoma bruce, brucei'.s not infective to humans
3 Mechanical transm1ss,on nv biimg !lies other 1nan LSelse
4 Transmission •s vene 1eal
5 Rarely entoumeieo
- = not infectrve/l)athogen,t - ~ mildly ;:aliloge~ie -= moderately catnogenic .-... = severely llal/llll!enic

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 a C

f
b d

Figure 12.3 Trypanosomes 1n min blood smears, xi 000 stained w,th Diff-Ou:c~ a = i.",08/'<0S()ma r.ongoiense note aosence oi free ilagellum;
b - T,ypanosama vivex note long free flagellum and large k,netopiast., = Ttypanoso:na b1ucei note polvmorphism. piominam undulaung membrane and
lree flagellum. d ; Tl')-Mnosama /;tr.Jee, d•vid1ng by long11ud1nal buial)" lisSJon (UnpubFsned pfotomiCiogreptis by covte$y of Dr L. Logan-Henfret.
l~iemat,onal laooralol'/ for Re=rch on An,mal Diseases. PO Sox 30iDS. Nairobi. l<~nyaJ

Howernr. 1he miwchondrion transfom,s in the tsetse fly. labrnm wall by means or 1hc tlage!lum.88 Parasite$ tha1 are
swelling and acquiririg tubular cris1ae. F.nzymes appear swept to th!.' midgm do not survive. In ll1e probo$ds 1he at·
which oxidlr.c protinc. the main source ()f enurg} ior both fly tached parasites lose their ~urfacc coats. ttansfom1 10 epl·
and rrypanosome.:i!l~ Thls transformation is evident as the mastigotes and muhiply in the labrum. They then de1ach
kinetoplnst rnigrnie:. and the lrypanosome~ assume the epi- and move to the hypopharym: where. after re-auachment.
mas1igote form In !he fly. l11ey transform 10 mcuicyclics. The complete cycle takes 5 to
Trypanosomcs repwducc by longitudinal binary lis$iOn. 13 days. before glvcoprotein-coated metatrypanosomes can
both in the blood$tream (Figure J 2.3d1 and in the fl). al- be inoculated a, the fly feeds.88 Thi! developmem or mem-
though a sexual process can apparently occur in 1he rsctse bers of the subgenus Vn11nomonlls and Trypnno::0011 is
fly. 131 J"S ;>.tultiplicarion in each host culmrnares in the pres- more complex and goes through an immature or midgut
en~ of mature trypano:.omei,, \\'hich ~lop dhiding and a.re su1ge. In the midgut of 1he fly, pre-adapted trypano~ome-.
pre-adapted 10 the conditions that they will cncoumer in the or shon stt1mpy bloodstream forms In the case of T. E,rucei.
next cyclic,LI host. As II tsetse ny takes its blood mea! from an tram;Jorm to procychcs, elong1ue and shed the glycop rocem
infected ho~t i1 ingestS rrypanosomes. Pre-adapted para- coat. The coat h, progressively replaced in a short time by a
sitei:. ~urvive in the fly, but trypanosomcs that an: not meta· coat of procydin, which probably pro\ides protection
bolically adapted to the new physiological conditions die. against proreoly1ic enxymes of rhe midgm.~~ The prncrclic
The cycle of clevelopmen1 of different ~pecics of salivarian trypanosomes begin co metaboli7.e proline as a $ource c,r en-
trypaposomes is very similar, but there arc important differ- ergy for which they compete \,irh their host. the tsetse. The
enc:es in their localization in the tserse fly during their devel- transformation of bloodstream trypanosomes into procyclic
opment (Figure 12.2). Trypanosoma rilinx has the s!mplei,t or rnidgut fom1s \,ith in the llr's midgut is a crucial tirs1 s1ep
migratory pauern; development occurs onl} in the probo~- In the establlshment of a u·ypanosomal infection. Tronsfor-
cisand the pharynx (cibarium). •11 30 Bloodstream trypano- marion proceeds rapid I>· in !he posterior part of the nndgut.
somes are taken up through the food canal of the proboscis. the first procyclic forn1:. appearing 11 hour, after inge,.
and the pre-adapted. late bloodsrream forms auach to the tion.288 Factors known 10 influence this process include

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258 srcno."<1wo: Prolozoal diseases

Axoneme

F
Paraflagellai
rod VSG

Pelhcular
m1crotubules

Figure 12.4 Elecuon miCiophotograph

of Trypenosoma congolens!1. tross-seciion
$hewing ilagellum (Fl. nucleus {Nl. m110-
chondrion {M} and variable surface glyco-
0,2µm
prote1n coat (VSG}. x86 000. Bai ;epresents
0,2 µm (Unpublished electron micrograph by
counesy of Or P. Webster. Yale University
School or Medicine, Department of Cell
B•o!ogy, New Ha\'en. CT}

trypanolys.ins and trypsin or rrypsin-Jike molecules in the to maturation. Before the infection is marure, procyclic
t1y·s midgur. 126• 127 the t}rp·e of host blood at the 1ime of the forms transform into epimastigote and lhen to mctacyclic
infective feed . 190• 194• 203• z-6 and blood composition.93• 1115• forms. From the midgut, trypanosomes migrate to the
230 The tsetse fiy's immune system also plays an important mouthpans or salivary glands. The midgut procyclics are
role. A Immoral mechanism involving lecrins is implicated free S\v;mming; they move 10 the ectoperltrophic space to
In 1.he establishment of U')'J)aJJosome infectjons in Glos- form an actively dividing population. They lose their glyco -
187
sma.
. !86. Jn mverte
· brates. Iecuns
· b'ind to spet1'fi ccarbo hy- p,ro1ein coat and move forward to the proventriculus where
d.rate groups on cell surfaces and may agglutinate certain they stop dividing. The proventricular ·mesocyclic' trypano·
£ells. This may result in l}'Sis and death of procyclic trypano- sames are longer than their procyclic precursors; they rein·
somes.49• 119· 128• 129- 220 In the midgut, rhe ritreoflec1ins and vade the endoperi1rophitspace and, in the case of members
the degree of inhibition by lecti ns largely detenn ine the ease of the subgenus Na11 nomo11as. move via the oesophagus to
\\tjth which trypanosomes in an infected blood meal estab- the hypopharr--nx where they anach and complete their de-
lish a midgu1 infection. velopment to become coared meracyclics after 7 ro 40 days.
The mechanism of maturation of a midgut infection_ is :Vlembers of the subgenus Tryrpa11owo11 migrate via the oe-
complex and, once established, it does 1101 always progress sophagus, mouthpans and salivary ducts to tlle salivary

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 •\friran nn!mnl trypnnos<>mo,;e.; 259

Mic:oti.;!i'J!es of flagellum

Endoevt1C vesicle
Endoloolasm1c
reticulum
Mi10chondrion

0.4µm
Figure 12.5 Electron micrograph of Trypenosoma t,,ucei section th1augh the flagellar poc',et (FPI reg,01 of the cell M,c•ciub~fes are lo1191tud1nallv
:rectionad. x44 000. Sar represents 0,4 um (Unpublished electron m:crograph bvcour1es•1 af Dr P Webster. Yale Unovers,r, School of Medicine,
De,rartmeN of Cell Biology, New liaven. CTI

gland~ where the parasites form Oagellar attachments to the mour or the eye. myocardium and in other 1issues. 1u:s, 318
epithelial cells. In the salivary glands the trypanosomes go In contrast to 1'. congo/ense and ·1: 11fl'(1x, infection~ "1th
rhrough Com stages of development. The mature. coated Trypa11azoo11 species arc characterized h)' generally low
metacyclic trypanosomes undergo morphological and parasltaemias and a marked invasion of dssues.'>r, 11 '
metabolic changes that pre-adapt lhe parasites for their life Because of the greater accessibility of bloodstream
in a mammalian host. The developmeni cycle of T. bmCPI parasites to investigators. this stage of the lifo cycle has
takes from 17 to 45 days. i-s been intensi\'ely studied. The ability oi trypanosomes to
As the infective tsetse tly reeds. metaq•clic uypano· establish prolonged infect ions Is aurihutable to the phe·
somes and saliva pass l'hrough the hypopharynx and are nomenon of antigenic variation. 22 • G-i. ~i Each bloodstream
inoculated imradermall}~ it is here that infection is estab- trypanosome is completely clad in a dense surface glyco-
llshed.3· i;a, ; , Trypanosomcs multiply in the skin. and may protein coar298 (Figures 12.4 and 12.SJ. which consists of
produce a chancre.248 which is a local s kin reaction that Se\'eraJ million tightly packed molecules. each of which is
deYelops into a raised. indurated, hot, painful swelling. The anchored in the parasite's surface membrane.250 \Vilhin a
chancre may attain a diameter of 100 mm In 10 to 12 days, population of trypanosomes originating from a single in·
but regresses 10 to 15 days larer.3 From the skin, the try- fecrion. almost all hear the same glycoprorein coat and are
panosomes reach the blood via the draining lymphaci~·s thus of the same amigen t~'Pe. :\s parasltaemia rises. a S\\ift
within a few days.3 · l6.. ;1. ;a, 16"· l i 4 Trypanosomes multiply antibody response is elicited against tht antigen type Cl)(·
in the bloodstream. and although initially their lo" num- posed on the surface of the bloods1ream trypanosomes.
bers make detection difficult. the generation time of only a These specific antibodies anach ro the ~urface glycopro·
few hours soon leads to high levels of parasitaemla. Try· tein and produce complemenr-mediated ly~i~ of all trypa-
panosomes may leave the bloodstream to reach various nosomes of that antigen type. However. before antibodies
extrava,cular sites. reach uypanolytic levels. some rrypanosom,,~ - as fow as
Trypa11osomn congolense, for long regarded as a striclly one in 100 000-5,,·Jtch orr the gene that controls the pro·
incravascular parasite, 116· 205 recirculates in lymph4 • 71 and duction of the initial surface glycoprotein and activate a
has been recovered from the central nervous system. 184 gene that code~ for 11 different protein.222 Trypanosome~
However. this species fs mart:! commonly located in capillary which bear the new surface glycoprotein are of,1 different
beds where parasites attach ro the endotheUum.31' · l6b Try- antigen type and are not destroyed by antibodr against the
panosoma 11i11ax showi. a greater tendency to invade tissue first antigen type: they survive to produce another para·
and has been demonstrated in cerebrospinal Ouid. perivas- Sitaemic wave. which in turn is remo,·ed by amibody
cular spaces of the central nen•ous system. the aqueous hu- specific for that amigen type.

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260 ,i.<. rmx nH,: rrotozoat disca~cs.

Figure 12.6 ScaMmg

electron m,crograpn of an
intermediate (bloodst,eaml
f01m cf Trypanosomil b,·ucei
from th8 blood of a mous~
No1e tile prominent undu1at 119
men-brane, pointed l)()Ster•or
e.od a'ld long. free 'lag:eHum.
A ·streamer or f1lopoa1u:n can
a:so be seen. (By courtes·.- cl
Or P Gardiner and reprinied
by k,oo permission oi Vinan;
Nar.:ul,-a and Parasitolog~
Toaay1

By lhis time a third varian1 ha~ arisen. and. escaping the lion is no1 kno\,TI, but h is thought 10 be related to the hose·~
cffec1 of host amibody. it ~u,..,ives to produce Lhc next f)ara· su~ceptibilit~ w infection and disease. 21 · l'M!. • 63 Similar
sitaemic peak. This antigenic variatlon i~ the re,ult or se- prt!-adapti\e changes ha,•e been found in r. 11iPtL\.1111 and arc
quential expression of q1riabte suriace glycoprotei1l< (VSGs) believed to be ncccs~ar. for 1hc cyclical development of
which constitute a repertoire of variable amig!m types T. ro11gulen,11.
l\".-\Ts) lnJecrion< arising from a single trypanosome m.1y fhe ~un<i111l of the non-dividing. prc-a<lapted uypano·
have a repertoire oi more than 100 VATs. 2u Thus shielded ~omes in (he bkmdstrcam ma\' he a~sociatcd "Ith the abilh:y
from total destruction. 1rypano~o111e infection~ usually nm to shed the VS<, ('Oat. ,\lso. rhe 'flow' ofVSG molecules over
prolonged courses, si11ce 1:ach \'Al is present for ~cveral the •urfac11 mcmhraue enable<. VSG-bound antibody to be
days before being remol"ed. Although there is a mt'(ture o( a endocrto,ed in the flagellar porke1. ""1· \Ol• 1n addition 10 rce-
small number of VATs within a parasitaemlc peak, the ~e- mo1ing potcn1ially harmful amibodv and dela}~ng complc·
quence of e,'(pression c>f \' \Ts tends to be quite swblt> in ment-mo.'diated lysb of the trypanosom11, this proceS> may
donall~• derived trypanosomes.io.. This imparcs. lnm1w10- also enable \'<;G to be recycled. Thl' shedding of \'SG by
logicallr distinct characteristics to :i strain of uypanosomes. smmpr form~ of L bruc:ei1" 8 and the format ton of 'stream·
1he dis1inc1 strain being called a ·serodeme . In 1..he course or ers· or filopodia (Figure 12.6). t'omisting of coated mcm-
~ uccessive parasitaem ic waves. some trvpanosomes swp di· brane:100 m1ty represent a protecti\(! mechanism.· <-0 The
•viding and transform to the pre-adapted fonn able 10 sur- s1ifl"cr cell membrane~ of ~tumpy 1'. lmm:i p.irasites may
vive in the tsetse. This is most readJly ~cen in T. lm,cei also pro,ide some protection against antibody-mediated
infl'ctions. The long. slender trypanosomes I apidly divide 10 h·sis
, .2-"J
produce parasitaemic waves. bm some differentiate After ingesuon b)' the tsetse, pre·adav1cd trypanosomi,,~
thmugh imem,ediatc fnnns 10 become non-dividing. short shed tht! gtycopm1ein coat. u-an~iorm, multiply and final!)
stumpy parasites. The>sc pre-adapted parasites do not shed mamre lnfocti\'C tsetse !hen transmit metacyclic try·
thclr surface coa1>.2~i Precise!} what induce~ transforma- pano~on,e, to another host. lrrc,pective of the \',.\T ol the

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 African animal uypanosomoses 261

bloodsLream trypanllsomes ingested by a fly. the metac:yclic nary fissiQn in the tissues, main!)' in the epimastigore form.
VATs of a serodeme are relati\'ely constant. After repeated When multipl}ing parasites are fow1d in the bloodstream.
tsetse-transmission of a single serodeme, the composirion polymorphic 'immature• fonns are present.
of meracyclic VATs is usually very similar. Although the When r. rheileri is Ingested br mbanid £lies. a cycle of
meracyclic VATs of a serodeme are always similar in contrast developmenr occurs in the hindgut and infectil'e meta-
to the large VAT repenoire of bloodstream trypanosomes. trypanosomcs arc voided with faecaJ material. Seven!I
there is nevenhelessantigen heterogeneity, even among the species of horseflies have been incriminated in the trans-
metacyclics \~1thi1\ a single tsetse fly.~01,. lli '.\!any sero- mission of T. 1heileri (see Chapter 4: Vectors: l'a banidae).
demes occur within a single species, each serodeme ha\·lng which is mainly accompJished as metatrypanosomcs pt•n·
its own \ 'AT repertoire. The antigenic diversity \\1thin a spe- etrate intact oral mucosa.i, This occurs as cattle use their
cies leads to the possibility of animals in a tsetse-infested tongues to fend off biting tabanids and simultaneously in-
area being exposed to a large number of antigenicall) dis- gest their faecal materlal. TI1e contamination of broken skin
tinC't trypanosomes, but although tryp~nosomes within a with metatrypanosomes of r. theileri is a less important
species may be amigenicaily dissimilar, the)' are morpho- route of entry. The prepatem period ranges from four co sL,
logically indistinguishable. days.2~ but detectable parasitaemia is short-l ived and is not
The chatac1eriza1 ion of trypanosomes for a long time re- accompanied by clinical signs although infrequent reports
lied on comparisons of their morphology. motility, host lncrimln:ning T. Iheilerl as a pathogen have be'en made. 200
specificity, lSetse tnmsmissibillty and their de\'elopmem In the lauer, concomitam diseases usuall} cXisted and
within the fir, but morerecem charaC'rerlzacion methods in- probably accounted for clinical signs, reduced host
clude isoenzyme typing, analysis of k.inetoplast D~.\ h) resistance and a consequent rise in T. theifgri parasitaemia.
polyacrilamide gel electrophoresis. pulsed field gradiem Similar evems occur in many wild animals whkh har-
elecrrophoresis of chromosomal digescs and D:.S: \ hybrid- bour tsetse-transmitted tl)'J)anosomes. Infected animals
i;(lltlon.88· 91 ln view of the great diversity which e:..isrs within show no clinical signs,w~ btH when they are subjected to the
a species. anempts have been made to standardize me no- stress or capture. for example. their immunity is reduced,
menclature of dis1[oc1 populations of ll')'l)anosomes.323 It parasilaemia flare~ up and clinical disease may be precipi-
has been recommended rhat 11ie use of Lhe term ·strain' tated. The equilihrium can be restored by reducing the
should be avoided. but il is !>till,, idely ust'd. ln rhecomext of stress factors and by providing trypanocidoJ 1rea1mem.
\'llterina11 medicine, certain strains of trypanosomes are Losses in captured wild animals due lO tl'},panosoinosis ma>
recognized for their lo" or high virulence, the ea:.e with be reduced by prO\iding trypanocidal treatment at Lhe time
which they are transmitted by tsetse, or for other broad of capture.
charac1erlstlcswhich may not be entirely srable.
The sequel to Infection with sal!varian trypanosomes is
Epidemiology
not always disease. The ou1come is determined by many
factors. frequently related 10 the suscepUbilil) or the ho;t The epidemiology of African uypanosomosis is almost en-
and the pathogenicll)' of the 1rypanosome. In the case of tirely dependent on tserse flies. 1311 African trypanosomes are
wild animals. a natural cycle of tT)'t>anosome transmission well-adapted parasites of many species of\\~ld animals. and
occur; which b 1101 ~oclated with dist'ase. Similar!~, in sylvntic cycle> oftrypanosome transmission occur through-
some breeds of domestic animals. infection with sali\'ll!'ian out the l Omillion square kilometres infested by this unique
trypanosomes is tolerated. and host and parasite reach an vector. The disrributiou of rsetse in southern Africa is de-
equilibrium. Oisturbancc of the equilibrium may preclpi· scribed in Chapte r 2: Vectors: Tsetse Illes, but wllhin the
late disease a long time after establi,hmcnt oflhc infection. general ecological limits of distribution the problem of try·
Thus. although the tsetse-transmitted trypanosomes are panosomosis is not staric. The 1990 outbreak of bovine
aeriological agents or African trypanosomosis. infection is trypanosomosis in the KwaZulu-:-.:acal Province of South
not alwa~·s synonymous l\ith disease. Africa28 testifies lO the dynamic nature of the problem.
The occurrence orr. theileri io healthy t-attle throughout The natural hosts or salivarian trypanosomes usually
the world II exemplifies a well-de, llloped host-parasite re- show no clinical signs orlnfection, host and parasites being
lationship. This sterc:orarian species is the iargest trypano• ln equilibri_um. 2uz The large numbers of naturally infected
some of cattle. and bloodsueam forms reach a length of wild animal hosts constitute a huge reservoir of trypano-
100 µm. fhis stage is cha.racterilied by a long. thin, pointed some~. Once infected. tsetse remain so for life and thus
posterior end, a long. free nagellum and a prominem undu- they 100 fom1 a reservoir o( infection. Conscquemly, when
lating membrane. The large kinetoplast lies far from che pos- t!omes1ic animals are introduced into areas in which syl·
terior e:..1remity, often in a marginal position. The nucleus is vatlc cycles of trypanosome transmission occur, trypano·
found midway along Lhe body. Ho\\'ever. these parasites are somosis always emerges as a serious disease. 175 Wild
infrequentlr seen since infections are characterized by low animals are the natural hoses of 7'. brucei rhodesiense. the
parasitaemia. Within the bovine. T. tileileri multiplies b) bi· aetiological agent of human sleeping sickness in C'entral.

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262 "r.itt>., 1wu: Pro1ozoal dlsea~~

eastern and southern Africa. Thus, people Living and work- western Zambia'18 and in the Lianshulu/ Mamilll area of
ing in tsetse areas are at risk of concractingthe disease. but Namibia's eastern Caprivi. 295
for animal trypanosomosis 10 occur it is not al\\'ays neces- The pre,·alence of metacyciic itlfec,ions in field-caught
sary for livestock 10 enrer tsetse,iilfe.~ted areas: tsetse a lso tsetse increases with the age of mes. ur.. 158, a;;, 2r;o, llt. :13:i Jn
move. The seasonal dispersal of fly, during the single rainy the case ofT. uivax, which does not pass Lhrough the tsetse's
season in southern Africa, frequently res.u lts in an in- midgut, 1he rate of acquisition of new infections with in·
creased seasonal disease risk to livestock kept some dis· creasing age is often constant. 332 Up 10 100 per cent In fee·
tance from the primary tsetse habiiat.2$ 2 tion rmes with 7'. uiutLr can occur in tsetse that repeatedly
Changes in land use may also alter the e.,tent of tsetse in- feed on T. 11ivax-infec1ed hos1s. 39 For those 1rypanosome
(estalion (see Chapter 2: Vectors: Tsets e flies). The aban- species that go through a procyclic stage in the midgut of
donment of culcivation. for various reasons, permits the tseLSe {T. co11gole11se and T. brucei), the relationship be-
regrowth of vegetation. which may then pro~ide suitable tween age and infec:tlon is more complex and is affected b\
tsetse habitats. Co1wersely, the intensive settlement and the reduced susceptibility of tsetse to infection as they age.
cultivation seen in some areas (throt;ghouc Malawil5• 292 for The prevalence of trypanosomal infections in tsetse is
exampJe) des1ro}'tse1se habitats.Although removal of tsetse also affected by host preference. Two aspects are important
should also remove the problem of trypanosomosis, live· in chis context: firstly, there is a diversity of hoSt preference
stock may have to graze somedistance away from cultivated among G/ossina and. secondly. there is a v-.i.riation among
areas, along the edges of forest reserves, and may therefore different species of hosts in their ~usceptibiliry 10 infection
scill be exposed to tsetse fly challenge. with the different species of trypanosomes. 111 For example,
The epidemiology of tsetse-transmitted uypanosomo- G. tmsreni in Uganda, which has a preference for feeding on
sis is complicated and ts well reviewed. 1,q, i-s. 2 >1, 25s. 324 suids. has a low •r. 1iivax infection rate. since pigs are refrac-
Whiteside321 identified at least 18 major variables in the tory 10 infection with this species of trypanosome. In con-
epidemiology of African animal trypanosomosis. which re- trast. G. pa/pa/is. which shows a preference for bovid blood
late to the interactions of tsetse, wild hosrs. livestock and meals. has been found to ha,•e considerably higher T. vivax
their management, the trypanosomes and climatic condl· infection ratcs. 195 Tsetse species are often grouped accord-
cions. The trypano:somal infection rate in lllerse is of prime ing 10 those that feed mainly on sulds. bovids, suidsand bo-
importance. The ease with which infections de,·elop in ,~ds, and las LI y, those that feed on most available hosrs.
tsetse depends upon the fly's vel:torial capacity and including humans. 307 However, host preference is not a
specific factors relared 10 the blood of host animals. How- rigid beha\~oura l characteristic of a particular tsetse spe-
ever, within a particular tsetse population, the prevalence cies.29· io. i 3 ,. i 93.z~9 The relativeabundanceofa hostspecies
of metac>•clic infection$ can vary significant!)'. For ex- may comrlbu re to observed host preference. For example,
ample. a recent srudr conducted in Zambia showed that within a country a tsetse species may feed main I>• on su.iclsin
the monthly roraJ prevalence of mature rrypanosomal in- one locality and on l.lovids in another. This is the case in the
fec1ions in G. pallidipe.~ could vary from 3.2 10 16 per Eastern Pro,ince of Zambia: on the plateau G. m. morsiums
cenc330 and is determined by factors such as the age of rhe cakes 75 per cem of its feeds on cattle.296 in the adjacent
tsetse population and 1he prevalence of 1rypanosomal Luangwa Valley the same species rakes most of its feeds
infections in the host. from suids. 21>1
Tsetse can only transmit memcyclic u-ypanosomes if the An animal entering a tsetse-infested area risks becomhig
flies live longenha11 the duration of the developmemal cycle infected "i1h potentially pathogenic trypanosomes that may
of a panicular trypanosome species. The developmemal pe- produce disease. The degree of risk depends largely on the
riod of uypanosomes varfos and is parasite species-speci6c 'challenge'. Ahhoubtb Lhere is no adequare definition of'chal·
alth()ugh a high degree of variability within a species has lenge'.255 it Is generally regarded as reflecting the number of
been observed.53 Generally speaking. the du.ration of the de- infec,ive bites that an animal recei\'eS in a given time. How-
velopment of trypanosomes in tsetse increases with increas- ever, true challenge. or risk. Is detem,ined by the interaction
ing complexity of the developmental cycle. The relatively of the number of infective tsetse bites, host preference, host
simple cycle that T. 11i1,,ix undergoes may last only for five susceptibility and the virulence of the parasite. Of parriculnr
days. compared with the 17 10 45 days required for the importance is the relationship between absolute tseise den-
completion ofT. brurei'scomplicatedcycle. Hence, the pro- sity and biting rate. Agood undel'$tandi 11g of this relationship
portion of T. 11i11ax infections in 1se1se increases under is essential when predicting the impact oftsecse control imer-
ad\'erse ecological conditions. when the Oy's shonened vemions. Usually tl1e challenge incre-ases with the tsetse
lifespan prevents the completion of longer cycles. Such population density but 11ven at low densities tsetse can still
ad,-erse conditions often occur al the edge of fly-belts cause a substantial disease problem. This is partly anributed
and result in the predominance of T. viuax infections in to the often observed, increased freque11cy "ith which
caule.~"· 159 A high proportion of T. 11i11ax infections in caule flies that have metacyclic infections in their momhparts
133. l!r.. 247
is. for e."<ample, observed at the edge of the fly-belts in prob e.

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 African nnlmal uypanosomosl:$ 263

There are various reasons wh)' a particular animal inOuence on the epidemiology of tl)ipanosomosis is the use
species may be subjected to greater challenge lhan ano1her. of uypanocidal dnigs. Whilst permining the use of isetse-
11)e beha\iour of a potential hosl can influence the ease infested land. chemotherapr may alter the prevalence of
with which a rsecse can engorge and t.his ma~· contribute 10 1rypanosome species in the area. In Zimbabwe, il was
observed preference. In Zimbabwe. the results of one s1ud>• reported that the use or diminazene increased the
showed that 1he frequency with which blood meals were r.
pre,·alence of uil'fl:r in cattle.n Furthermore. the repeated
1aken from calLle, sheep and goars by G. morsita11.s and G. use of 1rypanocides may result in the emergence of drug-
pallidipes was 500:5: I respecrively. Observa1ions on the resistant strains of ll)'panosomes. and the eptdemiological
relative anrac,iveness of these host species showed that the picture then changes. 2,9
ratio of 'probes:successful feeds' was 3,8:1 for caule, 28,0:l Wild animal hosts of tsetse. such as kudu (Tragelap/ws
for sheep. and 63.2: 1 for goa1s respectively.24~Thus goats are screpsiceros), warthog (Phacochoerus ae1hiopic11s), bush-
atuactive to tsetse but in response to the flies' attempts 10 buck (Tragelapilus scripr11s). bushpig (Potamochoerus por-
feed, they are vigorously defensive; they shake their heads. cus). African buifalo (Sy11cerus ca/fer), African elephant
wrinkle their skin, stamp their feet and even arrempt 10 bite (loxodo11ra africnna) and square-lipped and hooked-lipped
the flies! Nevertheless, goats and 5heep do acquire rrypano- rhinoceroses (Ceraro1heri11m sim111n and Diceros bicomis
somal infections under natural condicions.2 1. ioz. 271 Impala rcspcctlvclr). are \ita.l 10 s~·lvatlc c>•cles of rn,'Panosorne
(Aepyr:eros melnmp11s) behave similarly when tsetse attempt transmission and they form a major reservoir of infection.
to reed. It can be surmised that sick animals. which are less The increasing populari1y of game ranching or farming fre-
vigorous. are a, greater risk ofbelng fed upon by tse1se. This quently emails the translocation of these animals from
may- ex-plaln che increased feeding success of G. pallidipes ise1se habitats to tsetse-free areas. t\s a result, clinical dis-
on T. congo/e11se-infected oxen. 18· 19 ease may occur in animals normally regarded as 'immune'.
Management practices may also alter the challenge to in the absence of the 1sc1sc vec1or. There is also the possibil·
which livestock are subjected. and in this sense manage- ily that 1rypanosomes could be mechanically transmitted
ment is central to the epidemiolo1,•y of uypanosomosis. The from these species to lives,ock.
different managemem of calves and adult caule can signifi- \,\'hllsc 1he bite of an infective tserse clearly represents a
cantly reduce 1he level of challenge 10 which young animals uypanosomosis risk, infection does not always become es-
are subjected. 52 Observations on the grazing ranges of live- tablished. Furthermore. even when an infecLion is estab-
stock in We&t Africa showed that while cattle foraged widely lished. disease does no1 always ensue. A variety of host-
lo 1serse-infesced habitats, sheep. goars and donkeys re- associated propenies modifies the outcome and funher
mained closer to the villages. ,\s a result. small ruminant,. complica1es the epidemiology of uypanosomosis. Estimates
and equids were less exposed to attack by tsetse than of the efficiency of cransmission 10 susceptible cattle \'llry
canle.2li" In somhem Africa. the grazing patterns of commu · "~dely but field investigations suggest that it is very low (2,5
nal cattle vary from season 10 season.:?61 In the cropping sea- percemJ.76. 19t. .!57
son (;"l,ovember 10 March) cattle are penned and herded Wild animal hosts of 1se1se and ceriain 'Nest African
away from the crops. In the early dry season (;\pril to July). taurine cattle are toleranl oi tsetse-1ransmitted uypano-
after the crops have been harvesn:d, cattle are aliowed 10 some.,;. Carefully contrulled experiments have shown 1ha1
roam freely: they feed unanended mainly on crop residues. the uypanotolerance of N'Dama cattle and African buffalo
In the late dry season (August ro October), the cattle have to is an innate characleristic.2 · 1°&. l~li. 208 Th is trait also occurs
move further afield to find grazing and browse, often enter- in i<ome breeds of sht:ep and goats. 102· 123· 211 T1ypanotol·
ing 1sctse-infes1ed areas. Furthermore, the mo\·emem of erance is a much studied phenomc.non. 124 • t9~. 2<17, uo. 211
animals from tserse-free pa~tures 10 r$etse-infes1ed water- 252 Nthough the underlying mechanisms are incompletely

ing poi.ms can create temporary challenge. $edema!)' ani- understQOd, it is generally accepled thal innate and ac-
mals may face a seasonal increase in tsersc challenge as mes quired reslslance. as well as environmental and manage-
cilsperse du.ring the rainy season. Even within the vicinity of ment factors, affect ll')'Pano1olerance. Lower uypanosomal
a village, the actual challenge can be quite variable and can parasitaemias and less severe anaemia occur in trypano-
lead to differences in herd infection rates within a small tolerant livestock 1han in uypanosusceptible animals.I>&
area.302 Consequently, a thorough knowledge of local man- The epidemiological significance of this trait lies in che
agement practices is imponant to understand the epidemi- lower morbidity and mortality due to trypanosomosis in
olom; of nagana in a particular area. tolerant breeds • .:-;evertheless. e\·en \\~thin a generally tol-
Management also determines the general well-being of erant breed some individuals are less tolerant than oth-
liv~tock. If, for example. animals are heat-stressed and ers253 and tolerance may be affected by the degree of
their feed intake is therefore reduced. their resistance 10 try· challenge. 67 Similarly. within susceptible breeds indhidu-
panosomosis is al.so reduced. increasing the incidence of the als can be found rhat are apparent!)' less susceptible to the
disease,.,,., A careful distinction has to be made between an 1.>:ffects of trypanosomal infection than most other ani-
occurrence of this kind and increased challenge. A major mals.~0, 217• 231 Although lrypanosome•infected animals

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264 srrno, 1wo: Protozoa! diseases

may effect self-cure. the usual sequel to infection in toler- mals from succumbing 10 nagana. The introduction of tsetse
ant animals ls the establishment of a balance between host control measures in such areas would result ln a loss of im-
and parasite. If the host is stressed. the equilibrium is dis- munit)· rhat would render che population highly susceptible
turbed and a clinical episode of variable severity is precipi- to infeccion and disease. If tsetse control measure~ break
tated. Stress takes many forms. Animals in late pregnm,cy down. se\'ere outbreaks of rrypanosomosis would ensue,
or chat are lactating are more susceptible to trypanosomo- probably \\ith high monaliry. as happened following the
sis. •-;. :?03. 233 Oven,•ork abo constimtes a stress, which, in disruption of 1;,e1se control during Zimbabwe's llberatlon
infected trypanotoleram stock. may precipilatc diseasc. 175 war.lSl,
This has serious consequences for animal trac1lon and for Ofgreat sign iticance 10 the epidemiology of the disease is
hard-working bulls used in restricted breeding seasons in the nature of the parasite. Xot all host species ate! equally
u;ecse-infested areas. In a similar manner. lnretcurrem dis- susceptible to infection \\ith each species of rrypanosome
ease is stressful; try panosome-infected animals with helm- (Table 12.1 ). Whereas T. 11ivax is a serious pachogen of
imhosb or other diseases are more se,·erelv affected than cattle. pigs are highly resistant and. consequently, T. t·h·nx
rho$e with either disease alone. 104 • 1' 18• 1711• 27 1 lnfect!ons are 1101 seen In pigs. Con\'(irsely, T. simine causes
AdequaLe nutrition enhances the abilit} of infected ani- peracuce and rapidly fatal disease in pigs. whereas ca1tle are
mals to withstand the ndverse effects of a try·panoso.mal in- apparently reiractory 10 inrccdon "~th this parasite. 111
fection.1·15 Hence, rrypanotolerance is grea1I~ reduced by a Within a 1rypano$ome species are strains of differing ,iru-
low plane of nutrition. such as that available during the dry' lence. Usually. T. 11iuax infections in domestic mminanL~ are
season. 17r, Inadequate nutrition results in generally lower prolonged ,,irh long aparasitaemic intervals. Occasionally,
average herd baematocrits. at all levels of disea.~e preva- howe,·er. an acme haemorrhagic syndrome is seen in caule
lence. In the dry• season. animals may also have to trek long infected ,,~th some srrains of T. uit·ax.33. i;,,, :uo Infection
distances to watering poims. The combined effect of poor \\~th a single species of ll)'Pa.nosome can, therefore. pro-
nutrition and increased exercise Is often associated with an duce quite different signs in the same host species.
increased incidence of rrypanosomosis in the dry season. An interference phenomenon also occurs. which either
This is not necessarily due to an increase in trypanosome delays or pre,·ents the establ!£hmem of an anLlgenicallr dis-
transmission at 1.his rime oi I.he year but could be explained tinct uypanosomc in animals with an existing trypano~omal
by higher parasitaemias in animals with reduced tolerance. infection.''"· 70• 200 The mechanism of this interference re-
However, dry season grazing areas or watering points are mai11s unknown. bu1 if the phenomenon occurs under nam-
often locatod in tsetse-infested areas. in which case the ac· ral conditions ii would be epidemiologically Important.
wal incidence can rise. Similarly. such a rise of incidence since ii could limit che numberofinfections 10 which an ani-
has been observed when cattle graze beyond the boundaries mal is subjected. However, by testing bovine sera with the
of a tsetse control area during the dry season.~kJJ highly sen,irlve polymerase chain reaction (PCR) rechnique
Age also has a significant effect ott resisLance to trypa- it has been found that mixed infections arc far more pre\'a·
nosomosis.211 r1 is widely recognized that canle born in an lent thnn had previously been appreciated?15
infested area do not immediately succu 111 b 10 disease. even The occurrence of trypanosomal infections in areas ap·
though they acquire 1rypanosomnl infections \\'hen young, paren1ly free of tsetse promoted the theory tha1 infectioru;
whereas caule brought into the area readily succumb. can be mai111ained fn namre by the mechankal transmis-
ThTee issues arise from this observation: firstly. young sion of try-panosomes by other haematophagous mes. In
animals may be protected by maternally derived anti- Malo.wi, a firm opinjon developed that tserse-transmltred
bodics.:170· 21 1. 312 ~ccondly. tht> ability of animals to acquire tl)'J)anosomosis only oc1turred in close proximity to ,,ildlife
protective immunity is implied; anti. laslly, young animals sanctuaries. \\'here the flies were readily caught. Elsewhere,
may be less attractive to tsetse and, thus, face a reduced mechanical transm ission alone was believed to occur.6 It is
challenge. only relath·etr recently, witl1 impro,•ed trapping methods
Despite the amigenic complexity of trypanosomes, in- that use attractant odours. that it has been possible 10 reveal
fected animals do mount an immune response which. espe· rhe presence of Glossinn at low population densities. which
cially when supported by chemotherapy, can co1uer $pecific were hithrrto undetectable. The recent discovery of G. µ11/-
protection against homologous serodemes.$; w:- 3.."9 Thus. /idipes by these methods in the areas of :Vlalawi thought to
\\ithin a defined area, animals may acquire protective im- be free of tsetse~5 may ex-plain that part of the epidemiology
munity against locally prevalent serodemes. However, the of the disease in which mechauic.u transmission was
movement or these animals to another area may expose thought to play a role.
them 10 different strains, or serodemes, to which the)• may Try,panosonui oivax has become established outside Af-
succumb. 175• l 11 This type of tolerance requires continuous rica, and especially in Somh America and Mauritius. In
challenge and explains the low mortality of caule in high South America it !s believed to be mechanically 1ransmi1-
challenge. endemic areas (such as the Eastern Province or red.31s The cyclical development of T. viunx is confined to
Zambia) where trypanocidal drugs are used to prevent ani- the mouthparts of tsetse flies and there is limited migration.

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 Al ricau ,uurruil ttypano~omose, 265

The clo&e similarity of this mode of transmission 10 non-cy- accompanied by a febrile re,ponse. which I~ 1hen follow~'<!
clical. mechanical transmls~ion of T. 11/r.'(I.\ on the comami- by an aparasitaemic. afebrile period. l11e 1ime between
nated mourhpans of other haematophagous files is widely parasiwemic peaks ,:arie;; from four to seven days in goats
as~u111ed 10 have enabled this specie:. of trypano,ome to with r. 11il·12.,· mfcction~.11u 10 12 days in <"aHle infecwd with
adapt readil)' to acyclital transmission. I iowe\·er, the possi- T. co11gol1msP.t1:1. In long-st.tnding infec1ion~ parru;ires may
bility that cyclical development might occur in a vecior. or not be de1encd for months, and when parasitaernia doe?~
,·eccor~. other than G/ossim1 spp .. has not been completely occur. it i~ of1en low grade and the animal i~ afebrile.
eliminated. :"severtheless. there is some evidence that me- After an infection has become cscabiished. a protr.ic1ed
chanical transmission of T. i,/i<t1x can occurin areas adjacent battle ensues as the parasite pro,·okes an immune response,
t0 tsetse-infested areas. Reports of fulminating r.1•fra.1: in- only to evade iCl- full effect. 'rightly clad in the thick, dispo~-
fections which spread rapidly in the apparent abse1,ce of able protecrive glycoprotein coat. the uypanosome popula-
tsetse. leave linle doubt that transmission can be mechan- tion is ass.wed of conunued suni~al in the face of the ho~t·s
ical:'6· z;, and the rapid spread of T. simial.' \\~thin a pigger) vigorou~ defence. :\nti-VSC. antibodies destroy large num-
i~ believed to be due tu mechanical tran~mission. bers of t1y1)anosomcs, and dead a11d dying parasite, and
A wide variery of factors are involved in 1he epidemiology host cells are found extracellularly. These are removed b~
of u,'Panosomosi~. l.r I~ their intemction which detennine:. phagocyte~ and presen1ed a:, antigenic evidence 10 c-clls of
the clinical piclUre in an area. and, even 1hough mathema!l- the l)inphoid series. In the course of the comes,. many
cal modeb have been developed to quantify the relati\'e im- harmful substances arc released from boch intact and dam-
portance of these variables,256 much remains 10 be clarified aged parasite, and host cells. The destruc1ion of large num-
and the models have yet 10 be applied to tlw field. bers of parasi ws releases lysosomal and other cnz)'lllC.\ as
well as ~1ruclllral proteins. Some of these enzymes huve
been identified and are thought to be directly harmful 10 the
Pathogenesis
hoSI. ~lany biological mediators, such as the vasoacr.i\'e
The precise pa1hogenesis of the trypanosomoses remains amines. are released from ac1ivated or damaged host cells.
far from clear. Review:. of the subject t&, lw. r-, highlight the Activated manophages release powerful cy1okine1>.i:1 in-
cumples.ity of 1his group of diseases: the vorious species Clf cluding lnterleukln-1 and cachectin (tumour necrosis f:lc-
hosts Involved differ in their $USceptibil11y: there is great di- ror). but their roles in Lhe pathogenesis of' 11)1>Unosomosis
,·ersi1y among parasites within a species: and it b dffficul1 10 have ye1 to be invest igated fully.
e.-.:m1polate the rcsulcs obtained from cxpertmemal infec- Much work has concentrated on comparisons or the re-
tions in rodents, ruminams and other animals. w explain sponses of tl")'])at101oleram and u:ypanosusceplible animals
the pathogenesis of natural infeclion.2;., Three fea1ures - 10 infection. There is clear e\'idence that tolerant animals
anaeml.i, tis~ue damage, and the suppre~on of immune achieve better control of parasitaemia tha11 do su~ceptible
rcsponses20"· 189 - dominate che pathology of uypanoso- animals. and this ability is correlated with less severe
mo~is, and re~earch aimed at elucidating the pathogenesis anaemia and le~ severe disea~.:-. 66 Anaemia is a cardinal
of lhe disease h,is largely addressed these a~eas sign of 1rypanosomosis in man) domestic animals, .ind the
Infection becomes established at the site of inoculation aetiology is probably similar in all species. There is nosingie
of metacyclic ll'ypanosomes in the skin. where a chancre cause of the anaemia in trypanosomosis: che pathogene~is is
may form. Multiplication of tht? parasites Induces an innam- complex and involves 3 variety oi mechanisms. some of
matory response. the severity of which depends upon the \\11ich are better unders1ood than ochers.
size of the inoculum. the species of parasite and 1he breed Comprehensive rC\~ews of anaemia in trypano~omo-
and species of the host. In cattle. T. congolensu ls located in si.~09· 2 : 6 form the basi1> oi the following account.
dilated lymphatics of the papillary dermis and hypodem1is. Tl)'Panosomes release haemolysins and enzymes (pro-
in which the parasites form nagellar anachrnems to the en- teases. phospholipasei. and neuraminidases) which directly
dothelial ceUs.3 Dt:ad and d~1ng parasites are also present in damage red blood cell membranes. The fragllit) of erythro-
the lesion. mid various proteins and peptides have been iso- cytes is also increased by fe,·er. and afiec1ed cells have a
lated from 1he chancre. These may originate from ei1her 1hc shonened lifespan. lil")'lhroq·t~ are also damaged by 1rrp-
host or 1he pnrasites. 12·1 The chancre reaches a maximum anosomal antigens, which adhere 10 the red blood cell
diameter of some JOO mm. 10 to I l days afrnr an infective surface labelling it for removal. The a1tachmem of amigcn-
1set~e fly has fed. ils development preceding im•asion or the antibody complexe~ to red blood cell membran~ also re-
bloodstream by trypanosomes. and is ac<:ompanied by en- sults in damnge. Complemcmt nm be direc1lr ac1ivawd by
largemem oithe draining lymph nodes.a. 72• 7" uypanosomal \'SC. as well as being act!Yared by antigen-
Tr)'Panosornes are detectable in the blood 13 to I 6 days antibody complexe~. The aHachmem of cornponelll!. ol
after an infective tsetse fly has fed. 170 .\t thh time the chan- complement or complexed complement 10 erythrocytc1,
cre begins 10 regress. and the charact-eris1ic series of mter- produces damage and promores erythrophagocytosis.
minent para~irncmias begins. Parasitaemia rise, and is Anaemia occur~ largely because damaged erythrocytes ar.-

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266 st<:TIO>< ""': Pro1ozoal diseases

sltaemias.J\nimals lose weighr and condition and. as a reitull

of dyshaemopoiesis. remain anaemic. fa.'tensive baemosid-
erosis occur, as a result of erythrophagocytosis, and the
trapping of Iron in phagocytes is believed to contribute 10
the failure of erythropoiesis. Despite the apparent absence
or parasites in the circularion, red blood cell des1ruc1ion
continues. and insufficienr eryrhropoietic compensation
results in persistenr anaemia.
Trypano~omal parasitaemia con-elates with comple-
ment activity in caLtle. In the sera of N'Dama (trypanocoler-

•
am) cattl(> which develop lower paras!taemias than Boran
cattle. complemem activity i~ higher than in the sera of Bo-
rans infected with the same serodeme. 125 Hypocomple-
111e111aemia is a frcquem feature of the disease. \\/4th levels of
component C3 being marked!) reduced: this may be asso-
Figure 12.7 Bone marrow mpress10n srnea; from a ca,f w,th an acute ciated with reduced resistance to 01her infections.2;-; Vari-
Trypi!nosDma vivax infectton. sllo\111rig .; macrophagewh1ch ttas ous changes have been noted in the composition of
phsgocytosed several ervtluocy1es ~ t 200. Stained w,th leukocyte populations during the course of infection, but
Wrights·leishman (Unpubl shed ohotomicrograph bv coui l!sy o" beyond the usual occurrence of leu.kopenia, other reponed
Dr L logan,Henirey. ln1e111at,ona l aooratory for Researtll on A.nimal
changes are equivocal.
mseases, PO Box 30709. Na,rob1 Kenyal
The suppressive effect of 1rypanosomal infections on the
immune responses of laboratory animals has often been
removed from the circulation by cells of the mononuclear demonstrated experimentally. but its significance in live-
phagocyric system (MPS) in the spleen. bone marrow stock has 1101 been determined. Trypanosome-infected ani-
(Figure 1;11, lungs and haemal lymph nodes. To deal with mals do moum protective. amibody-mediated immune
the abundance of cellular debris and amigen -amibody com- responses agains1 the parasites. 198· :l!Hl and both lgG and lg'.\.!
plexes 1he :Vt PS becomes greatly hyperplastic, which funher are produced. 111e lgM levels are consis1en1ly elevated dur-
enhances erylhrophagocytosis. lmravascular haemolysis is ing inrec1ion and appear co be directed mainly against \'SG
thus not a prominent reature of lrypanosomosi.~. A hae- anrigens. Whilst some JgGs are directed against the com-
morrhagic syndrome is :.ometimes seen in acute T. 11/va.t mon. somatic trypanosomal antigen$, others appear to be
infections of caule. in which sudden and severe lhrombocy- directed against some of the host·~ own cells. An1i-ery1hro·
toperua occurs. This is associated with fulminating para- C}'te amibodies contribute 10 the anaemia of uypanosomo-
silaemia and exten~i,-e petechial and ecchymotic sis and il b possible that a similar mechanism operates
haemorrhages. The thrombocytopenia and resultant clot· against leukocytes, contributing to teukopenia.209
ting defects, as \\'ell as the presence of circulating immune Some animals arc able to control parasitaemia quite ef.
complexes, damage the capillary endothelium. leading 10 fecth·ely. and this appears to be antibody mediated. at least
disseminated inrravascular coagulation. In pan. Trypanotolerant Baoule caule mount earlier and
The removal of large numbers of red blood cells from the greater amibody responses to the first parasitaemic peak of
circulation occurs soon after the onset of parasi1aemin. and T. co11go/1111se, compared with similarly infocced Zebu
produces a fall in the packed cell ,·olume (PCV) -haema1- caule.2 • ~.,~ This ability to control the firsr parasitaemic peak
ocrit. Packed cell volume is a reliable indicator of anaemia is associated with the less severe anaemia occurring in try·
and is correlated with parasitaemia-the higher 1he parasi- panotolerant ani mals.
taemias. the lower the PCVs. The anribody responses of trypanosome-infecred cattle.
Accompanying anaemia is leukopcmia. which possibly sheep and goats co non-uypanosomal antigens are. never-
arise~ from direct inhibition of stem cell differentiation. Al- theless. depressed. 173 The results of one investigation re-
though I.here is an erythropoietic response 10 anaemia early vealed tha1 1·. co11gole11se-infec1ed caule produced poorer
in the course of infection. rhis seems 10 be impaired. anamnes1ic responses 10 secondary vaccination with a poh•-
Whereas anaemia is largely attributable to an increased rate valent cl0S1ridial vaccine than did uninfected control
qf erythrophngocy1osis early in infection. the anaemia of the cattle. 112 On the other hand. cattle experimentallr iniec1ed
late phase of infection has a different pathogenesis. with r. congolense and vaccinared against foot-and-mouch
There are se,·eral possible sequelae to the early phase of disease \\'ere considered to have developed pro1ec1ive anti-
infection, which depend largely on the nature orme parasite bod~· tirres.~69 However. the effect of nypanosome-induced
and the susceptibility of the animal. There ma}' be sponta- lrnmunosuppression superimposed on the stresses of mal-
neous recovery or death , but very often there is a chronic nutrition, uekking, prcgnancyorlactatlon may explain \\'hy
phase which is characterized by infrequent. low-grade para- affected animals frequemly succm11b to o ther disorders. The

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 African animal 1rypanosomoses 267

precise acriology ofimmunosuppression in trypanosomosis cular permeability. In T. congoleme infections a generalized

is obscure. Ir may involve B·cell mliogens. facmrs which dilatation of capillary beds. which alters the haemodynam-
block the release of amibody from plasma cells, reduced ics, is obsen•ed.1&-l The concomitant anaemia and more
T-helper cell funclion, and depletion of lymphoid elements sluggish tissue perfusion affect the exchange of metabolites
in che spleen and lymph nodes. 290 Pan-ides from dead and and are associated with intracellular oedema of cap.illary en-
dying trypanosomes stimulate the :"l·fPS. and the cytokines dothelial cells.221 Fibrino~1s micro1brombi form in response
released by cells of the MPS may conrribu1e to the lmmuno- to endothelial damage. These changes can be prominent In
suppression associated with the disease, as may products r. uivax infecuons. with which disseminated intravascular
released by disintegrating uypanosomes.12· = coagulation is more commonly associated.
There is a growing appreciarlon of the close functional re- Alterations 10 lhe microc1rculation produce secondary
lationship between the imnnmologi.cal apparatus and the en- degenerative changes in tissues. As capillary permeability
docrine system of the hypothalamo-pituitary-adrenal axis. 1i increases. phagoq1es and products of the parasite extrava-
Both systems are characterized by delicately balanced regula- sate more readily and are responsible, in part, for some of
to1y mechanisms. mediated by potent cytokines which have 1.he tissue lesions. Cytokines and parasite-derived sub-
short half-lives. Interleukin· J Is a polypeptide released by stances, such as proteases. may directly injure host cells. an
macrophages and has a wide range of effects on man)' tissues. effect that is exacerbated by lhe formation ofimmnne com-
As well as stimulating T cells 10 produce interleukin-2. iris a plexes in tissues. In the case or T. bmcei. parasites localized
potent pyrogen, stimulating the hypothalamus. the piruhary in tissues cause mechanical disruption of host cells and
and the adrenal glands. Locally. interleukin- I increases vas- probably also ha,·e some direct toxic effect on host cells.
cular permeability and is involved in the acute phase changes possibly mediated by their enz)•mes. The cellular response
of !he inflammatory process. The effects of trypanosomosis provoked by parasites within the tissues causes ft1rther
on the endocrine system have recently been demonstrated in damage, and au10-ancibodies are also thought to play a role
canle and goats.94 Trypanosome-infecced animals show in inducing lesions.
abnormalities of the thyroid gland. ovaries, testes. adrenal The influence of all of lhese changes on the course of the
glands and pituitary. To what extent the changes are medi- disease depends upon their severity and upon the degree of
ated by the direcc action of the parasite, the hose's immune impairment of the affected organs. An importam feature of
response or by imbalances in the endocrine system. remains the pathogenesis of rrypanosomosis is the effect on lym-
to be determined precisely. phoid tissue. As the disease progresses, tJ1e hyperplastic
A parasite peptidase has been de.monsmned in the MPS grossly increases che volume or tissue in the spleen.
plasma ofT. congolense-infected helfers. 151 The possibility lymph nodes and bone marrow. This hyperplasia of reticu-
e:>.ists that this enzyme might inactivate certain hormones loendothe!ial cells reduces l)'Jnphold cell density, and even-
or imerfere with the host's enzymes that are responsible mat lymphoid depletion can occur.297
for endocrine regulation. Such disturbance of the homeo- Chronic disease is associated l\ith progressive emacia-
static mechanism would have serious consequences. In in- tion and eventually cachexia. This is usually accompanied
fected canle, a lthough levels of adreno-conico1rophic by low levels of parasitaemia, but the pathogenesis is poorly
hormone are similar to those in uninfected controls. corti· understood. The reduced lifespan of red blood cells, the in-
sol levels are lower. and this might indicate adrenal creased catabolism of many proteins?·; and the associaced
dysfunction or n decreased half-life of cortisot. 125 The in- negative nitrogen balance of clinically affected animals
rerprecalion of research findings from laboratory-housed would appear to be largely responsible for the piti-
animals should always be made bearing in mind thac fully thin condlrion of animals suffering from chronic
chronic infections in stressed animals are commonly trypanosomosis.
encountered in the field.
The pathogenesis of !'issue lesions varies with the species
Clinical signs, pathology and diagnosis in cattle
of trypanosome. Trypanosoma congolense and T. ufoax are
main!} inrravascular parasites: they induce changes in the African c:rypanosomosis is. Lransmined by tSe1se nies and is
endothelium of capillaries. and so indirectly cause damage primarlly caused in cattle by T. congolense and T. 11i11a.i:. ln
to adjacent tissues. Trypanoromn brucei, on the other hand. endemic areas. the disease causes economic losses, but it can
has an affinit)' for tissues. Its presence in the e,mavascular also occur in areas remote from tsetse infestations, as a rei;ult
compamnent is associared with marked lesions in parasit· of1he movement ofpre\~Ously infected animals which, \\1hen
!zed rlssues. suessed, show clinical signs oftrypanosomosis. Additionally.
The severity ofendothelial injury also depends on the in- small numbers of infected tsetse flies. carried long distances
teraction of host and parasite. Tryptmosoma congolen.•e by vehicles, can transmit uypanosomes in areas othem~se
often attaches to erythrocytes and capillary endothelium. free of crypanosomosis. Becognirion of the wide range of
Damage 10 endothelial cells by parasice productS. lmmune clinical signs associated with the disease is thus important,
complexes, vasoactive amines and cytokines increases vas- both within and outside endemic areas.

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268 ,,,r:u<>s """ Proto;toal dbeases

...
i_ .,.
•
,.
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•
• /.lit> •
.. • •
~

Figure 12.8 Subacute tryµa!Y.lsomos1s rr ao ox: the an1r:>.al tias iost .~e,ght and condi.ior. aro It 1s deiected, wH.n oroop,ng ears a~d flacc,d .ail
Repnmeo di kiod perm,ss·on o! FGU Consulting ano Enginaerin9 GmbH. KOnigsrn,n. Ge,nanv. and tne Regional Coord:nator. AnCP Harare. Z•mllatr,vel

Cattle living in isetse-infesred areas frequemly have AcUte 1rypanosomosis causes sudden reduction of milk
mixed uypanosomal infections whid1 cause more severe yields and also abortion.116 Oxen, bulls and young stock
disease than infections with a single species or trypano- with ncute infections are reported 10 be suddenly 'off-
some. Generally. in eastern and southern Africa. r. co11go- colour'. The)' are generally in good bodily condi1ion but are
lense is more pathogenic to cattle 1han T. ,,;,,ax. and dejected. their ears droop and chey may walk stiffly. If ani-
produces serious disease. where.as T. /irucel is ,ddel\ re- mals are febrile at the time of examination, rectal 1empera-
garded as being of minor clinical signifita11ce. 1ures may reach 39 •c in T. congole11$e Infections. whilst in
acute T. vii,ax infections the)' may be as high as 41 -C. Pulse
Clinical signs and respira1ory rares are raised in febrile animal~ and there
The course of disease due to infection with salivarian uypa- may be piloerection. Conjuneth'al mucosae mar be con-
nosomes is variable and 1here are no clinical signs specific to gested early in the acme phase. bu1 later, as anaemia
bo,~ne uypanosomosis. The manifestation~ of disease de- develops. pallor of the mucous membranes is evident.
pend upon the degree of ctan1age to specific organs and AcUtely affected animals quickly lose weight and body
upon the degree of anaemia. Trypanosomosis in caul.e may condition. although they continue to eat. They become weak,
be acu1e. subacure or chronic; acute disease may be fa1al dejected and lethargic. often standing alone, away from the
after brief Illness lasring two to sL-: weeks, but chronic (!is· rest of lhe herd. not seeking shade. E.xcessive lachrymation is
ease lasting many months or even years is more common. sometimes seen at 1his s1aie. After a short illness some
, Chancres are rnrely seen in canle with naturally acquired acute!~ affected canle become recumbem for a fc\\ (lays be·
infections. The first signs of c;llsease are due to the fever which fore death occurs. Corneal opacity of vari~ng se,·eri(}· has
accompanies the onset ofparasimemia. As parasitaemia falls. been reported in some T. brucei and T. 11it1a.x infections.2 ~4 A
so does fever: rhe course of infection is therefore character- proportion of acurely affected tanle gradually impro,·e to
ized by Ouctuaring parasitaemin and parallel fluctuations in enter 1he chronic phase of the disea.~e. Occasionally, acute
body remperamre. lmem1irtenr parasitaemia and malaise are 7'. 11i11ax infections induce a haemorrhagic s~'lldrome.aa when
followed by increasingly severe clinical signs. animals bleed spontaneously from the ears, nose, mou1h,

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 AfricM animal nypano,omQse; 269

anu,. and ~kin. Pcteci,iae and ecchymose.~ may also be evi- stray from the hNd and became lost. This weak, emaciated
dent in the conjunc1ival mucosa:n. 179· 1 1Q animnl displared periodic generalized tremor. whereas an·
In contra,t with the acu1e drsea5e, subacute trypano$O· other similar!} affocwd animal in good condition showed
mo~!, Is more common and rum, a more prolonged course, hyperaesthe~ia and agitation.
wilh many :inin.1als apparemly making a spomaneous. re- Whilst ior descriptive purpose.sit is convenient 10 clas-
cove~· However, they remain infected and frequently ,uf. sif)• trypnnosornosii. as being acute. ,ubacute or chronic
fer rrom chronic trypanosomo,is. rn ~ubacute cases the course that an infrction actually follows is modified by
(Figure 12.8), animals are intermiHently 'off-colour". be· many factors. ,\ subclinlcally affected animal may. when
coming weak and dejected, \\ith drooping ears and flarcid stres,ed. suffer an acute clinic.ii episode. while acutely
tall. \\"!thin a few wcrks they lose weigh1 and condition. the affected animal5 may mal..e a gradual recovery onl} tO
coa1 becomes dull and a marked jugular pulse de\·efops. remain chronically affected. Ym orher animals may effeet
On closer examination. although intermittent fever occurs. ,111 apparent self-cure. \\ hilst continuing to harbour
it is rarely marked; mucou.~ membranes arr pale, and pulse 1r~rpanosomes. The orcurrence of concomitant disease fre·
and respiratory rates are slightly raised'. Superficial lymph quently masks the underlying crvpanosomal infection. but
nodes and haem.ii lymph node$ are frequent!) enlarged on occasions may exacerbate the clinical signs of t:rypano-
and are readHy visible. Gradually. thll frequency and inten· somusis. Tlw imrnunosupprcssion induced by trypanow-
-:ity of parasitaemia decrease and fe\'er subsides. Depen- mosis renders infecced stock more su~ceptible 10 ocher
dent oedema. pnr1icularly in the subrnandibular region. cli~ca,es. In many cases. the underlying cause of disease or
may develop. Thin. rough-coated. anaemic:-. lethargic cattle death may be conc,•aled by the presence of 1hc more
\,11h generalized lymph node enlargement are said to ha\'e reaclilv dC'lectable, secondary Infection. such as anaplas-
a 'fly·-~truck' appearance. Death may occur between four mosis, helmintl10,is or lumpy skin di~ea,e. The conse-
and sb: months after the onset of disease. but many ani- qm.•ncl' or thi, is that the true extent of t~'Panosomosis b
mals make a grndunl recovery. \\'hich is assisted by a good often underestimated.
plane of nutrition.8~ In endemic area$, t1ypanosomo~Is m canle is a herd prob-
Chronic bovine t~l)anosomosb is by far the most com- lem. 111~ general condition of the herd is poor. although some
mon form ol the disease in endemic areas. It may arise.either indi\idual animals are in surprisingly good condition. Trypa-
asa result ofpariial recowryfrom acuteorsubacute disease, nosomosis causes !Serious reduction in producthity, and care-
or from an Jnitial suhdlnical or mild infection. Severely af- ful annlyi.is or records reveals its impact on fertility, which is
fected cattle may be extremely emaciated, having lost a large reduced in both males and females. 121 · 13 1. m In addition to
proportion of their musde muss: they are often just 'skin their being generally weak. the semen quality of affected bulls
and bone· Their gcncr.u condition i, very poor- the hair is ib reduced.~"> Infected cows lose weight and their orstrus
sparse and the coat is rough. dull and staring, The skin is dry cycles become irregular and 111aycease altogelhcr. 16:! This may
and scaly, and in long-standing infections Lhe hair of the tail result in a significam reduc;tion in cah~ng rates. In the Eastern
S\,itch may be completely absent. ll1e precrural lrmph Province of Zambia. for example, the calving rate in a c.rypano-
nodes and haemal lymph nodes arc usually visible from a somosis endemic area is 44.1 per cent compared ro 60,4 per
distance, and there is a marked jugular pulse. Although cent in an adjacent t~etse-[ree area.61 Trypanosornal infection
chronicallr affected animal& continue to eat. they are weak during pregnancy· ma}' induce abortion. 16· 131 · Zl6 but when
and lethargic and arc unable 10 keep up with the rest of the pregnancy is carried 10 tern,. the binh weights of cnlves are
herd. Geophagia has been associated \,ith bovine uypano· low.b.l 2.~, l~'Panosome-infectcd young stock grow poorly.
somosis. maini)' in West Africa.' 08 but careful anamnesis they show signs of ill-thrift and remain stunted. The com·
oilcn revcllls L11a1 cattle with chronic 1rypo.nosomosis in powtded losses from tower fenilit} and milk yields. increased
other pans or Africa also ha\·e a tendency to eat sand or lick mortalit)· and reduced growth rates are reflected in low herd
anthills. !1,e chrnnic ~,11.drome is characterized by infre- perfonnance and often in drcroa,;ing herd size. A composite
quent. scanty parnsiiaemias, which arc not accompanied b~· productivity index incorporating these parameters has been
fever. ,\naemia persists and is usually severe. Mucous mem- found uscfui when a~essing the impact of bovine uypanoso·
branes are ve~· pale. the pulse is rapid, and breathing is la- mQsis and ilS comrol.1~a
boured: in resisting restraint, 1he animal may collapse in The clinical signs of trypanosomosls 1cnd 10 be more se-
respirato ~· dis! ress after brief ext'nion. The combined effect vere in exotic breeds of cattle than in the local brec<ls which.
of aJ1acmia. circulato~· disturbance and myoe-ard1al dam- in pan. reflects the adap1arlon of local breeds to their emi·
age frequent!>· produc<lS acute cardiac decompensarion ronment. :\d<litionally, exotic breeds are often under greater
which leads to sudden death from congestive heart failure. production Mn>s~. which further exacerbates the dis.ease.
-\!though r. l,mci'i normally produces mild or subclinical Trnmnowma 1/11!/leri Is not considered 10 be a patho-
disease in canlc. signs of cerebral tT)panosomosis arising gen.~'~ .th hough high parasltaemias can occur In cattle \\ith
from natural r. /Jrucei infecuons have been described.31 ' imcrcu.rrcm discase.2ll0 There are several reports associat·
One affected animal was de~cribed as a 'loner' - it would ing ·,: rllei/erf infection with ;.uch clinical signs as anaemia.

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270 " ""'" ""': Protozoa! diseases

diarrhoea or reduced milk yields. but conclusive e\'idence of cardium and ascites, and the carcass. in general, appea~
the role of this species of 1rypanosome In the pathogenesis oedematous. Residual fat around the heart and kidney<: i~ ge-
of these disorders is lacking. latinous. Lymph 11odes, haemul nodes and spleen are gener-
ally enlarged and have an appt>arance similar 10 that seen in
Pathol-Og)' acute 111'Panosomosis. but after prolonged disease there ma~·
The PC\/ begi ns 10 fall \\?tch the onset of parasi1aemia and be splenic atrophy. Haemorrhages are nol a feamre of
fever. From an Initial value of0,32 1/f il may be reduced to chronic uypanosomosis. The hean is enlarged and flabby.
0.20 ( I ( or less-in a maner of a few weeks. The early erythro- the li\'Cr ~wollen and pale. Whereas reel ma1TOW is present in
poietic response co anaemia 2<)9 is usually normochromic the long bones in acute trypai1osomosts. all the bone marro\\'
and normocyric. but an early macrocytic response may is yellow and gelatinous in the chronic disease.
occur. In chronic disease there is a microc~'Cic anaemia, GroSJ> changes are not always ob,ious, but microscopic
associated wi th hypoferraemia and dyshaemopoiesis due to lesions are extensive. These are found particularly in the
iron 1rapping. lnhial lellkopenia also_occursand maybe fol- cardio\'ascular :.ystcm and lymphoid tissues. and there is
lowed by leukocy1osis. Thrombocytopenla is a constam fea- widespread inllhration of many organs br mononuclear
ture of uypanosomosis and may accoum for the dotting inflammato~r cells. Dilation of the. microvasculature and
defect that occu~ in the r. 1•iiiax-haemorrhagic 5~'T1· oedema and stnictural changes in ,·essel walls are usually
drome. 1~0 - 20!'1 Hypocomplementaenua is a.lso a constant seen. especial!)' with T. co11go/e11se infections. Trypano-
finding in trypanosomosis.277 which is correlated wirb the somes are commonlI present in the lumens of blood ves-
level of parasitaemia:123 complemem levels are higher in sels. and, in the case of T. t•il·ax, parasites may be
cattle that have lower 1rypanosomal parasitaemias. The lev- associated with fibrinous thrombi in the larger blood ves-
els of other serum proteins also alter in the course of trypa- sels. In some acule cases. there is disseminated intra vascu-
nosomosis. Albumin is decreased and globulin levels are lar coagulation, when m1crothrombi may be associated
i,,
frequently raised.a,;· Other changes also occur in blood with haemorrhage and ischaemic necrosis. A constant fea-
chemistry- blood chloride levels rise and the calcium-to- ture of uypanosomosis is Initial hyperplasia of the l)·m-
phosphorus ratio in infected adult catde is significantly re- phoid tissues and of the mononuclear phagocytic system.
duced.fl~ More rccemly. auemio11 has been directed to the In the conical area of lymph nodes and white pulp of the
etrect of uypanosomosis on hormone levels. Results indi- spte·en. germinal centres form early in the disease, aJ.
cate rhat although plt1sma cortisol levels of infected caule though in chronic infections lymphoid rissue becomes de-
are reduced. parasitaemia appears not 10 alter le\'els of pleted.?05 The expansion of the mononuclear phagocytic
plasma adrenocorticotrophic hormone. 125 system leads to increased erythrophagoc)'IOsis and there is
The gros~ pathology of affected anima.ls varies with rhe widespread haemosiderosis.
duration and severity of the disease. and many organs and Monoqr1es and lymphocytes aggregate in the microvas·
tissues are affected. Acme infections may end \,ith the death cularnre together wiLh uypanosomes. ln\'asion of tissues
of an animal still in good bodily condition. In such case$ the by the parasites leads to infiltration by mononuclear cells,
carcass is weU-Oeshed. although the mucous membranes which is main!) perivascular. In the late stage of acute t~·-
may be pale. The overall appearance of the carcass ls one of panosomosls. severe anaemia and extensive vascular dam-
paleness. a nd petechial and ecch}'lllotic haemorrhages arc age produce degenerative changes in the tissues. but these
frequently presem. especially on serosal surfaces. In acUlc are less serious than ore the changes tha1 arise from ceUu-
T. vivax infections. there a.re extensive haemorrhages. re- lo:r i11filrra1ion. Since 7: brucei is more invasive, it provokes
sembling those seen in sepricaemic conditions: large, epi· considerable cellular infiltration. TI1e heart is commonly
and endocardial haemorrhages are presem, in addition to affected, and macrophages, lymphocrtes and plasma cells
those on serosal surfaces. infiltrarc 1he myocardium. Degenerarive changes and focal
Lymp h nodes are enlarged and. when incised, are necrosis of tn)•ocytcs and, in long-standi ng infections,
oedemawus and often have a dark pigmented medullary fibrosis are evidem in the hearc. Infiltrations also occur
area. Typically. Lhe spleen is greatly enlarged and dark red. frequently In other organs such as the pituirery, thyroids,
Excessive peritoneal fluid is present wh.ich is oft.e n blood- adrenals. kidne}'S and gonads. Congestion and interstitial
ringed. J\cule r. congoh•nse infections can produce exten- oedema also occur, a nd, when the subsramia propria is
sive oedema which is most noticeable in the perirenal and affected, corneal opacity then de\'elops. S,Nere degenera-
mesenteric fat, and the abomasa.l mucosa.1&1 Trypano- tive changes occur in the thyroids62 and a reduction in
soma brucei may cause kera1icis. although this is rare in erythroid progenitor cells of 1he bone marrow has been
cattle. rcponed.=
The gross pathology of chronic uypanosomosis is charac-
terized by cachc).ia and anaemia. The coat is dry and dull, the Diagnosis
skin is scaly and inelastic. the eyes arc sunken, and the skel- The specific diagnosis or rrypanosomosis is notoriously
etal muscles are atrophied. There is hydrothorax. hydroperi- difficult. :Sot onl)•are there no specific clinical signs. but the

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 African animal uypanosomoses 2il

intennittent and frequently low parasitaemias makt' mediately covered with a cover slip and examined micro-
detection of the parasites difficult. Furthermore. infection is scopically as a fresh. wut preparation \\ith a ><25 or x40
not synonymous with disease: many subclinically irifected objective lens. Approximately 50 to 100 fields are examined.
animals Jive in delicate balance \\~th potemially pathogenic Live. motile lrypanosomes may be seen as they bore their way
rrypanosomes. An element of clinical judgement rs therefore berween blood ceUs (the name 'trypanosome' is derived from
necessary when making a diagnosis of crypanosomosis. Sev- the Greek ·uypanon·. a borer. and ·soma', meaning body).
eral autl1ors6 1. 148· 223 have compared the different methods The wer blood smear method is simple, inexpensive and gives
for the parasitological diagnosis of animal trypanosomosis. immediate re.'nlhs. It can be used to exaniine a large number
lmporrant considerations are tharthe number of dececrable of animals. Depending on the trypanosome· s size and move-
parasites is not necessarily related to the severity of rhe dis- ments a presumptive identification can be made of the trypa-
ease and, in endemic areas, the disease is a herd problem. nosome species involved. 1-lowever. the diagnostic sensitivity
The detection of infection in a few clinically affected canle of the method is low and depends to a large extent on the ex-
warrams careful examination of the enrlre herd. aminer's e.>-'Perience and tho level of parasitaemia. Sensi1i\iry
Economic principles, the availability of expertise and the can be improved significantly by lysing lhe red blood cells be-
diagnostic requirements will guide the choice of a particular fore examination using a powerful haemolytic agent such as
diagnostic tesr. The diag11ostlc method will differ between sodium dodecyl sulphate (SDS),22*
situations. depending. for example, upon wherher species- More commonly, for routine diagnosis in veterinary
specific diagnosis is required or whether surveys are con- prnclice. thick and thin smears of blood are prepared. A
ducted simply to determine the presence or absence of the drop of blood taken directly from a punctured blood vessel
disease. Often a combination or diagnostic tests is needed ro in the ear or tail tip is placed on a glass slide; a thick and a
obtain the required results. rhin blood smear can rhen be made on a single slide. Lymph
Although in many tsetse-infested areas diagnostic facil- m1ears are prepared in a similar manner. The smears are
ities are not readily available, clinical signs of nypanosomosis kept dry and protected from dust, heat. flies and other in-
are we.U recognized. Farmers and veterinary personnel com- sects. Alternarively, under dusry field conditions. venous
monly resort 10 treacmem of sk.k animals, and use the re- bl<>od C-dll be taken i1110 tubes containing an a.nticoagulan1.
sponse co therapy for rctrospccrive diagnosis. In such areas, a such as heparin or BDTA, and blood $mears can be made
history of the presence of tsetse flies and the use of rrypano- subsequently. The dry smears are stained wirh Giemsa's
cidal drugs. when considered ,,irh presenting clinical signs, stain, thin smears after having been fixed, and examined
are sufficient to make a tentative diagnosis. Ho\\'ever, the under a xSO or ><100 oil- immersion objective lens. Although
presence of concurrem disease mar mask trypanosomosis it may be necessary to examine up to 700 fields to find a
and compllcate the clinical picture. ·nius, the only way 10 single trypanosome,51 the unfixed de-haemoglobinized
confirm diagnosis in clinically infected animals is ro demon- thi.cl: smear allows approximately 120 rimes more blood to
strate and identify the parasites in body Oulds. be scanned than a lhln smearl-1 8 and. thus. has higher diag-
The parasite detection methods are highly specific but nostit: sensirlvii:y than the thin smears. Trypanosomes are
their diagnostic sensitivity (the proportion of infections that easily recognized by their general morphology bm may be
the methods detect) is relatively low. This is especially the damaged during the staining process. Tbls makes irdifliculr
case when results are considered for an individual animal to make a species-specific idenrifica1ion on thick smears.
ratherthan in a herd. As a result, the apparent prevalence of The rhin smear, on the other hand, permilS accurate speci-
uypanosomosis determined by parasitological diagnostic ation of the parasites. However, due to the srainingprocess.
rests is an underestimate of the true parasi1ological preva- results are delayed.
lence. This is a problem in areas where the disease is presenr The probability of detecting 1rypanosomal infections in a
at low prevalence or is seasonal. or when attempting ro con- sample of infected animals can be improved by increasing
firm the absence of lhe disease in a panicular area. the volume of blood 10 be examined and concentrating tbe
The body fluid most commonly examined is blood, ei• trypanosomes. TI1is is achieved by collecring fresh blood
ther capillary blood from the, rip of the tail or venous blood (about 70 µl) in a hepariruzed capillary tube and separating
from an ear vein or from the jugular vein. Lymph, aspirated lhe different components of the blood by cemrifugation. ac-
(rom a punctured superficial lymph node (usually the pre- cording lO their specific gravity. Tr,'Panosomes are concen-
scapular), provides useful supplementary diagnosric mare.- trated in the plasma/white blood cell interface or buffy coat.
rial. Whereas cerebrospinal fluid is routinely examined for TI1e buffy coat is examinedmicroscopicallywhilst slowly ro-
lhe presence of trypanosomes in tbe diagnosis of human tating the capillary tube (microhaematocrit cemrifugarion
sleeping sickness, this has little practical use in veterinary technique or Woo-method).330 Trypanosome movemem
medicine.314 can first be detected using the x10 objective lens \,ith
Wet blood smears are prepared by placing a drop ofblood reduced condenser aperture: the trypanosomes can be seen
(about 2 µl) , taken directly from a punctured ear vein or the more clearly using a x40 objective lens. preferably of a long
tip of the rail, onto a clean. dry, grease-free slide. This is im- working- distance (allowing considerable depth of focus

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 ,_
2 -?

1.hrough the capillary rube). The microhaematocril centrifu- techniques are particularly useful in rhat the haematocrit
gation tech nique is more sensitive than the dlrect examina- or PC\' can be assessed after cemrifugation. b) placing the
tion techniques hut identification oftrypanosome species is microhaematocrit capillllI)· tube in a haema1ocrj1 reader.
difficuh. Alterna1ively, the huffy coat and 1.he uppennosl The PC\' of 1·cnous blood is a useful indicator of anaemia.
layer of red blood cells c:-,m be extruded onto a clean mjcrn· which is one of the most typical signs of rrypanosomosls in
scope slide and covered with a cover slip (huffy coat tech- dome~ric animals.a~9 The PC\/ of individual animals and
nique or;-, lurra~· method212 ). Approximately 200 fields of the the average PCV of herds can be detcrrruned. In individual
preparation are examined mic[Oscopically for the presence animals. haemawcrit readiJ1gssuppleme11t parasitological
ofmocile trypanosomes 11~th a uark-grotmd/phase-contTast diagnosis. In trypanosomo$is-endemic areas, parasitologi·
illumination and a ><40 objective Jens. 'fl")'Panosome species cally ne!fat!ve animals 1hat ha,·e low haematocrits are re-
can be identified by reference to the follO\\ing criteria: garded as having trypanosomal infections. In surveys to
• T. vh,ax: Lmge, eXtremely active, traversei. the whole field determine the presence or absence of lrypanosomosis.
very quickly, pausing occasionally. samples collected from animals with low haematocrirs can
• T. bruC/ei: Various sbies, rapid movemem in confined be investiga1ed more thoroughly to increase the probabil·
areas. ity of detecting a uypanosomal infection. In parasitologi-
• r. c1:mgole11se: Small, sluggish. adheres 10 red blood cells callr positive animals. the haematocric indicates the
by anterior end. severit)' of disease. The decision of the eattle owner road·
• T. r/leileri: More than twice the sit..e of pathogenic trypa- minister a trypanocidal drug is more oflen bM!!d on the ef.
tlosomes. feet of anaemia on 1he animal's performance rather than
1he confirmed p resence of infection. At the herd level. the
Since the specific grad1yofT. congo/ense is similar to that of haematocrit profile or the herd average PCV Is a useiul In·
red blood cell.s, these parasites are ofwn found below the dicator of Infection and herd health. However, other fac-
bully coat within the uppem10st layer ofred blood cells. The tors such as nmrition and fasciolosis may also cause
two concentration methods arc the most sensitive for de· anaemia on a herd basis. Therefore. it is imponam 10 es-
tecting T. co11golerzseand T. l'ir,axinfections.238 The sensltiY· tablish the haematocrl1 profile of ~'Panosomosis-negative
iry of the concentration methods can be further improved herds before relying on it as an indicator of uypauosomo,
by using the buffy coat double centrifugation technique. 15~ sis in a herd. Because of the lo·,, sensitivity of parasitologi-
:\ total amount of 1 500 to 2 000 µI of blood Is centrifltged cal diagnosric tests, baseline PCV-profiles are best
after which the buffy coai is aspirated into a microhaema1- obtained from herds of animals chat do not have anti-1ry-
ocrit capillary tube and centrifuged again. The buffy coat is pano$omal antibodies (Figure 12.9). In the absence of
then examined as previously described. ,\ modification of PCV-profiles of negative herds. comparisons of herd aver-
the microhaematocrit centrifugation technique is the Quan- age PC\'s at various distances irom the source of infection
1itati1·e Buffy Coat (QBCJ method. 1•1 This method is based on - along a transect - can give useful information on che
expanding the buffy coat by inserting a cylindrical noat in a variation in disease challenge (Figure 12.10). In trypanoso-
capillary mbe that is pre<:oated with acridine orange. The mosisendemic areas. herd average PCVis significamly cor-
method has been used to diagnose r. I>. g11mbieme infec- related \\ith parasitological prevalence (Figure 12.11 ). The
tions but is too expensive for routine use in the diagnosis of slope of this relll!lonship is determined by the effect of dis-
animal rrypanosomosis. ease prevalence on herd average PC\' and may be used as
The microhaematocrit centrifugation and buffycoat tech - an indicator of the impact of uypanosomosis on herd
niques give immediate results and can be used for screening health.
large numbers of animals. They require, however. specialized The subinoculntion of blood into rodems. ustral!y mice
equipment and an electricity supply which makes the tests or rats. allows a greater proportion or T. brucei infections to
more expensive compared 10 the examination of the wet be detected than by direct examination of the bu!Ty coat.
blood film. but this is compe11sared for by the increased diag· For prncrical reasons. subinoculation of blood into labora-
nostic senslrJviry: furthen11ore, haematocrit readings may be tory or other animals is not used as a routine diagnos1ic
obtained (sec below). Both parasite concentration tech- procedure. The mcrhod is expensive and diagnosis is not
niques rely on the detection of live, motile tl")'panosomes. immediate. Funhennore, ~ince rodents are refractory to T.
Since rrn,anosomes are heat sensitive. samples collected in vil'fl.t and nor all T. congoleuse and T. bmcei infections be-
ca pillar)• mbes should be kept cool: overhearing of the micro- come established in the new host. even this method has se-
haema1ocrit centrifuge should be avoided. Compared to the rious limhations. Mixed ~'Panosomal infections may also
mlcrohaematocrit cemdfugation technique. the buffy coat remain undetected.
technique has the added ad\'amage that preparations can be A procedure for the in virro cultivadon of 7: brucei from
fixed and s1ained for more accurnte species identification and the blood oi infected animals has been described but suc-
for retention as a permanent record. cess has been \'arlable. Moreover, the method needs sophis-
The microhaema1ocri1 centrifugation and buffy coat ticated equipment, yields results only after a considerable

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 African animal 11ypanoso111qscs 2i3

30 Herds In= 371 nega,M! for ant.·trypanosomal antibodies

25

~~
~~~~
~~~~
~~~~~
5
• ~~~~~~
Parasnolog1cal positive herds In: i58)

~
30

~ 25
~
~
"'
-0
a
.: 20

~~
0
5
,: 15

~~~~
8.
0
a:

~~~~~~
10

5
I~ ~~~~~~~ -~ ~ '. ~

a,

~
M
N
C"'>
N
\::; ~
-
7
M
<?
.,.,
M
,_ 0)

,.,, M~ 0..,.I
M -
~

'-
..... f:i ~ ~ N g ~

flan-;ie of packed cell volume (%1

~
I I
«>

Figure 12.9 Herd average haematocri, profiles of anti-uypanosomal antibody negauve herds ano parasitologicat positive
herds ffasiem Province, Zambia)
33

32 - •
_ 31
~

~., 30
-
~
• -----·
•
,.
:::0

i 29 - •
=
.;;
z
28

27
-
, I
I
I
26 . •
25
0 5 10 15 20 25 30
Oistar.ce from tsetse ctialleoge(in km)

Figure 12.10 Herd average pac,(eti cell volume at increasing distance {in km) from the tsetse front 1n Eastern Zimbabwe

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274 wa1c>., nrn; Proro1.oal dtsen,~'-l>

3i

30 •

29 •
~
?:
,.. 28 - •
~ 27
!
'C
;; 26 •
•
25 - •
• •
2,
•
23 ~

en .,, .n
~
0
~
.
'.r> e "T "'
I
R
7
.:;
,ec

~
...,
C

"
:::,
"' Paras,tological ;ireva!enc-e{lt) "
figure 12.11 Relal/011sh1p between helli average paO\ed cell volume and paraslto!og1tal 0te\'\li811ce ol 11ovme il)'panosomo~,& {%1 in a tT\IPan=mosis
enoem,c area in the Eastern Province of Zambia

de[a~ and is certainly not suilable for widespread use. A re· agm,sisof trypanosomosi~ has greatly benefited from the in-
cently described kit for 111 vitro isolation tKIVTJ of trypano- troduction of enzyme immunoassars. The enzyme-linked
somes has proved promisingforisolatingand amplifying T. b. imrnunosorbent assa) !ELIS.\) was first used to detect anti-
gm11bie11se in humans. domestic and 1,>ame animals.?3 ; The bodies again,t T. h. rhode.(/ensi> in human~. ' 01 It was further
rest"s value in isolating T. co11go/e11se and T. 11il'f1x is still un- dc,·elopcd for use in animal t~·panosomosisH•7 and ,,•as re-
known. Since it is based on the cultivation of procyclic forms cently modified for large-scale use in t~'J)anosomosb sur-
of Lrypanosomes. species differentiation is not possible. t;.z 1•ci-s.1 :; .·\ntigens can be prepared using bloodstream forms
A miniature anion-exchange technique has been de- or procyclic trypanosomes. 01 Purified ant igens arc used
scribed for field use in the diagnosls of human t~'l)anoso- prefcrabl~ m ensure that te~ts arr standardized and optical
mosis, 151· 171 but is mo cumbersome for routine use in density cut-oiT1'l!lucs do 1101 \'ary greatly between batchc, of
,·tuerinary practice. The use of a minicentrifuge \\ith huffy antigen.
coat technique has been ad, ocated. 1'' II is not satisfactory The !'LIS:\ com po.res fa\'ourabl~ \\ith the IFAT11''1 and has
0

in bovine practice where large numbers of ~amples often been found co give results that correlate wi th the local hbt<>~
have to be examined. of t~1>anocide usage. 1' However even if a uypnn<>somal in-
1n the diagnosis of h.rman trypanosomo,is the cere· fccrion has hcen cured. antl-uypanosomal amlbodic~ persist
brospinal fluid (CSF 1 is examined forihe pre,encc ofinflam- ror several monthsZ5 and antibody detection wsts do nor dis-
mato~· cells. raised protein levels and parasites. For tinguish between currcm and past lnfecrions. They can only
parasitology. the CSF is examined after centrifugation: the provide a presumptive diagnosi>. ,\I though the prevo.lc·nce of
5ediment obtained from 2 to 5 ml CSf is placed und11r a anti-t11pano~omal antlbodiesoften increaseq with increasing
cover slip and examined mkroscopicall~· for motile t~-pano- pre,-a.lence of 111•panosomal Infections in a herd, antibody
somes.31l detection methods arc not suitable ior monitoring disease
The non -specUic diagnostic te~ts to detect rabed protein chil.llenge in rrypanosomo;.i; endemic area;. Xevenheles~.
levels in the serum of animals suspected to have uypanoso· amlbocly detection te~ts, e<pedall~ the antibody-detection
mal infections 1 ' 8 have no role in diagnosis today. The ELISA. ha\'e been wry useftil t0ols for determining the distri-
de\'elopment or ami-t~'J)anosomal antibody detection bution of tl"}'P31lO\Omu~b.!92 ?% Thi'> i~ e.~pecially theca~e in
techniques has been a major improvement in the serndiag- rue.i~ where disense pre,11icnce is. low and \\'here trypano-
nosis of trypanosomosis. The indirect lnm1unofluorescem cidal drngs are used frequemly. Moreover. ihe prevalence oJ
amibod}· rest (lf:\T)·'~'" has been and still i,. u~ecl widely to anti·t~1>ano~omal antibodies h a sensitive indicator of the
dlagnose trypanosomosis. The tC$t has undergone several Impact of tset!le control operation$ on disease challenge and
modifications so that it can differemiate. co a limited extent, ror confirming the dbease-free status or animals in areas
between trypano~ome ~pecies iTI ruminam,. 14" The serodi- from which tsethe has been erndic~ ted.

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 4\frlCrul nnimaf iryp:mo~mno~i·!'- ,--
_,;')

Ano1heT ahernatil·e to lhe parasiiological diagnosis of largement of l)inph glands. Differentiation relies upon de·
nagam1 is 1he us!.' of ru.say, to deten ll')1>anosome-spl'cifk wc1ion of the parasiles in blond ~mear,. ',llhough lymph
anligen. or specic~-spceific or ~ui>,,pecics-specilk D\'.:\ :\n node enlargemem and fc\'er occur In l·.as1 Coast fc\·er,
antigen detection ELISA lor trypanosomosb ha~ been de- animal, oflen cough and haH: diarrhoea. The presence of
scribed~~s. ~~. ii, bu1 fit•lll e\'aluations of the 1e.~t have given \Chi,;onts in ,mined smear~ ol l~ mph node ma1erial and ol
inconsistent results. Addltionru work is needed to detvelop plropla,m,; in blood snHiars contirms East Coast fenir. At
the 1est for routine dia~no~b of uypanosomo~is. A poly- necropsy. 1he ahomasal ulceration and kidney lesion,
merase chain reac1ion <PCR1 method has bt:en den•loped charanerbtic or Ea,t C:oaSt fe\"cr are nm seen in tl')·pano·
for the diagnosis of infections \\'ilh African tl')'J),lllOsomt?s in ,omosb. Can:asse, of animals \\'hich ha\'C succumbed co
humans, animal!' and tsetse Illes.''-? Specific repeti1i\c trypanosomosis arc (>ftcn oedematous and anaemic
nuclear D~A sequence, can be amplified for T. 1·i1'rLr and Chronic ll')'Jl:lnusomo,i:, is an afebrile di5easc. in wh1c-h
each of the three T. co11gole11se subgroup,."' 10 A common anaemia. c:macin1iu11 aml !)'mph node rmlargemem ate promi-
primer set Is available for detection of the three T. /.>rued nent findings. It is imponant co tli!Jerentiate !his disease from
subspecie~. The test require~ 5pecialized equipmcm and malnutrition and hl!lmimho,is. In neither nf 1hcse two condl-
highly uaincd personnel ctlnsequemly II ls n<>t suitable for don, is hmph node enlargement found. The detection ofh~l-
use in most laboratories. Sample collect Ion has been simpli mimh cgi;s in iacce~ is a u,eful ad1unct to the diagnosl~ of
fled br adapting 1he rest using blood s1)01ted on fiht?r pa- hclmintho,i,. bm 1hc uhwnce of egg, in fael·al sampll', taken
pen;1~3 so that a large numht.'t of samples can b11 proces11ed from cattk \\ith rhronk fnsclnk>sis, an<l the occurrnncv of
at one time. ·n1is makes the tes1 potemially ~ui1ablc for anaemia and subcuia1w1lus 01-dema can make diffenmtial di-
large-scale surveys. Howe\·er, the cost or PCR analpe~ pro- agnusi~ di11kuh. In <:a$C:. of malnutriticm. tlw d1..-grcc of
hibits its rouiine use In veterinary lnvcstiga1ion. nw PCR a1l<1emia l, rarely n~ extensive ors('\ ere on a herd ba~is as h is
1echnique·s ability 10 detect lment or mixed 1rypanosomal with tr)l>llllOsom<>sis. ahhouih marked inchvidual variation
infec1ions in different hosts renders i1 suilable ro, rc,earch occur,,_ The lute ,1age c>f cn~oo1ic buvinc lcukosi~ is r:huracter-
into the comple.x rel;uionship1 be1\\·ce11 trypanosomc.>,, and ized by \\eight lo~ and lymph no<.11• cnlarg,•mcnt. hut. unlike
their vectors and hosts.2·•' animuls $ttffering from chronic crypanusontosis. appetite is di·
Despilc thedcvclopmem of mort•,cn\iti,1:, morc ,oplu~- rninblwd and ana1•mla is mrl'i} !>e\·t-n:.
tica1ed and expensive diagnostic methods, the clinician will. \.\1wn T. 1heill'ri is found In blood ~m1•ars it is casil}
for the foreseeable Future. ho,e to rely upon examination of di~tiniuished from ocher tl')·!J",-lnOSomcs by its large size. Al-
blood smear~. bulfy coat preparations and finding, at thouf!h r tl1eileri 1nay occur in caule infected with another
necropsy to confirm diagnose~ <>f trypano~omus.is. ,\ccum1e ~pcclcs of irypano,omC!, thc c.1rcful C'-aminatlon of thin
dia~osis is crucial to subsequent inrerventions 10 comrol Gi1•ms,1-stamcd blood smear~ is more Jlkely 10 reveal
nagana. and 10 monitor 1hrir succt'SS. At the herd h:vel. 1h11 the ptt-.;cnni or mi:xf.'d infection~ c,f h<'tW·transmitted
confidence placed in 1h11 rc·Mil1s will latgel1 depend on the t r,·pru,osomc,.
size of the sample. ln some cases. howl•vcr. repeated sam•
piing of an individual animal mar he nece~sal')· before a di-
Clinical signs, palholog) and diagnosis
agnosi> can b<- confirmed.
in goalS and sheep
Differenlial diagnosis Trypano,omosh of goat\ and ,h..,ep ha,, been ,tated 10 he of
In its various smges trypanosomosi~ re,c:mbles a number ol lillle significance becau~e natural infE'c1ions arc seldom de-
otlier di:;ease conditions. and 1l freqm•ndy occurs al 1he tected.~~• Howe\er. 1.rypanosomosis of ,mall ruminants is a
same time as ocher infection~. In the acute ll.'brile ~1age. major con~1raint on tlwir health and productMty, 1112 and
trypanosomosis must be dlfforenliatcd from redwaicr ti'} vanosomal infections in goil.l~ and sheep are !mown to be
(babe~fosisJ. anoplasrnosis and Ea~t Coast fcrnr The haem- pre\-alem in pans ol southern Africa. occurring in ~lo,.am-
r.
orrhagic syndrome In acute 1·it·ax inf..iction can be db1in· blque:!·1 and i'.ambiu.i1 Small nnninam, han• a much
gulshed from anthrax and haemorrhagic- septicaemia shoner gt•nera1ion 1imc 1han cattle. and it is likely that in
IPasteurellu multoddu infection by examination oi tsetse areas this factor has producl'd weater nmural selec-
Giem:.a-!>tained blood $mes~ when large numbers tion for II'} pano1olerancc in 1hesl' ~peck,,_ 1han ha~ occurred
of 1rypanosomes are found. with ca1tlc. Thi~ may explain the fact 1ha1 goats and ,hcep
A~u1e trypanosomosis rarely produces ic1erus and is are found apparemh· thri\"ing in tsclse-infcsted areas in
not accompanied by haemoglnbinuria. These fearnre; and which cattle cannm he kcpl without the use or drugs. Com-
parnsitological fmding.~ differentiate ll')'Pano,omo,is from pored with exotic breeds. 1he local breed~ of goats and sheep
redwater. The Lwo infections. however. may be concurrem. in the tsetse bells of r:.1>1 Arrica are more toleram of experi-
Similarly. anaplasmosis can occur in 1rypano~omt."· tnt'ntal tl')-panosomal infecrions. 10~ bu1 rrypanosomos1~
Infected cattle. On it'\ own, anaplasmosis often produces doe, nunetheless cause $lgnificam production los~es e\ en
icterns. as well as fever and anaemia: there may also be er,- amonglocalhreeds. 20 • 21 · 1~ 1

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276 ""'"'~ nm: Pro1ozoal diseases

Of the salivarian trypanosomes. 1: vi11(1.t Is apparently fast. Anaemia dcvclo1)$ rapidlyand the PC\' falJs from 0,35 f I l
the least pathogenic to these hosts, often producing mild to 0.18 £ ' ( or less in rwo 10 four weeks, when mucous mem-
infections. Trypanosoma congoleuse is an important branes become pale. Affected animals rapidly lose weight and
pathogen and usually produces suhacute or chronic dis- condition, and may die within a month of the onset of detect·
ease \\'hilst T. /Jmcei infections may folio\\' an acute. rapidly able parasitaemia. /\!though corneal opacity occun; in experl-
fatal course. especial!} in experimental infections. Fre- memal lntcctionswith T. vivll.~18 and T. brucei, ,zz this lesion is
quently, however, naturally acquired T. brucei Infections notc01nmon in narural cases.
are chronic. 273 Under na111ral condftions mixed trypanoso- The signs of subacutr tl')l)anosomosis are less marked.
mal infections are probably very common. although on and the course or the infection resembles that in cattle.
parasitological examination one species usually predomi- Anaemia de\·elops quite rapidly, lymph nodes become en-
nates. Tryprmosoma simiae also infects sheep and goats. larged (Figure l 2. I 2) artd. in goats. a marked jugular pulse
bm reports of clinical infections are rare. i 3 ;. 1• 2 can be seen. Although death occurs some 1O to 12 \,·eeks
after the onset of parasiraemia. the condition of many ani-
Clinical signs mals does stabilize and gradually improves. The impro\·e-
The general clinical signs of uypimosomosis of goatS and mem may be apparently complete: PCV, Jive weight and
sheep are similar 10 those in cattle. A.cute, subacu1e and general condition may return to nom1al, and fever and para-
chronic disease syndromes occur naturally in goats and sheep sitaemia abate. Other animals show slight improvemem,
in East Africa. 1Cl3 and have been observed in goats in Zambia.21 but they remain anaemic. emaciated, weak and chronically
Acute disease is characteri7.cd by intermittent and increas· affected. Chronic infection$ are \·ery common and may de-
or
ing dullness. During boucs fe\·er, sick animals walk sriffiy. or velop without marked diuical .signs. Para.sites are rare!y
stand with head held low. cars drooping and tail flaccid. Early- found in the blood and animals arc aJebrile. Clinically, it
morning rectal temperarurcs may exceed 41 •c in the febtilc is difficult to distillb'uish such animals from those \vith hel-
periods. when the pulse is rapid and the breathing shallow and minth()Sis. Funhermore, the immunosuppression induced

Figure 12.12 Agoat with a naturally acquired TfYpanasoms v1vax infection: presca:iwar lym~n nodes are greatly enlarged

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 African animal 1rypanosomose! 2ti

by trypanosomosis ln goats105 and sheep 173 permits the much more difficult because of the larger proponion of sub-
establishment of large helminth burdens. which chus clinical infections. In chese c~ses very low parasitaemias
exacerbate the clinical signs in affected animals. 1°" occur in the absence of ob,~ous clinical signs. It is undoubt·
As with caule, uypanosomosis of small ruminants is a edly ihese factors which are responsible for the serious un·
herd or flock disc11se which reduces productivity. Fenilily is deres1imarion of nypanosomosis in small ruminants.
markedly reduced. 121 The binh rate of goats In a <SCtse·in·
festedarea of Zambia has been Improved by regular chemo· Differential diagnosis
prophylaxis, even though few uypanosoma.l parasitaemias The main disease entity which resembles trypanosomosis in
\\'ere detected in untreated comrol goats.21 Resu lts from goats and sheep is helminthosis, especially haemonchosis.
eastern Africa also indicate the importance of subclinical Anaemia, ill-thrift, weight loss, submandibu.lar c,edema and
1rypanosomosis on goat productiviry. 109 high mortaliiy races are common co the 1wo disease com-
plexes. although diarrhoea sometime~ accompanies helm·
Pathology inthosis. Parasitological techniques ha"e to be used for
The pathology of trypanosomosis ia goacs and sheep does differential diagnosis. Frequently the two problems are con-
not differ greatly from that in cattle. current. and the final diagnosis and treatment must depend
Because oftheir smaller size and lower maintenance costs, upon clinical judgement.
small ruminanis are popular experimental animals. and chere
are many repons on the pathology ofeiqierimemal trypanoso-
Clinicalsigns, pathology and diagnosis in pigs
mal infections in these species. Recently, great attention has
been focused on the effects of uypanosomosis on endocrine In many parts of somhern Africa. indigenous breeds of do-
funcrions of these animals. Tse1se-1.ransmitted T. co11gole11se rneslic pigs suni,•e in trypanosomosis-endemic areas. Pigs
infections induce weight loss and Irregular oes1rou, cycles in are refractOr) to infection with T. 1,;,.nx. but are mild!~· af-
goa1s. rn 1 whilst che inoculation of gom.s with bloodstream uy- fected by T. brucei. T. congolense and T. suis. In contrast, T.
panosomes alters plasma le,·els or several honnones. Levels of simiae is highly pathogenic to Improved breeds of pigs. but
rhyroxine,? 19 1estos1erone.303 and plasma cortisol?15 are de- local breeds seem to escape 1he ravages of this let ha.I rrypa·
pressed while plasma pros1ag:landin F2 leYels rise during e.x- nosome species. Of zoono1ic significance ls che role of the
perimemallyinduced trypanosomosis in goats.218 domestic pig in harbouring the human parasites T. brucei
Functional abnormalities ha,·e also been demonstrated g<1mbie11se181 and probably T brucei rhode.~iense. 11 l -14
experin1emal.ly: semen quality of infected rams is reduced 1• c However. 1his laner association has noc been demonstrated
and ovarian dysfunction occun. in infected goats?16 Z-larked 'in southern Africa.
te,ticular lesions have been reported in experimental infec- Trypanosomosis of pigs in A[rica has been comprehen-
tions. In T. brucei-infected sheep, iaitial oedema of the scro- sively rcviewed. 12t. ~;4 and only an oLI!llnc of the problem is
tum was followed by a hatdening of the scrotal wall and presented here.
arrophy of the cestes. 120 1-listologically, che most severe in-
flammatory reaction affected the mnicn vaginalis and was Clinical signs
characterized by a thick fibrinous exudate on the surfaces Trypanosoma brucui and T. congole11se infections in pig;; gen-
which led 10 adhesions and organization. Diffuse atrophy or erdlly produce only mild d!-'lease.23; Occasionally anaemia O<:·
the seminiferous tubules was associated with intenubular curs. which is accompanied by loss of condition mid
oedema, mononuclear cell infiltration and fibrosis. Gross tes- progressive weakness leading to incoordinatlon. Trnxinosoma
ticular aLrophy also occurred in goais infected 'A'ith T. ccmgo· s11is Infections are of minor clinical imponance and have only
/e11.se. 1l9 while. histologlcally, degenerative lesions were rarely been recorded; infections run a chronic course. but may
e,1dent. Seminiferous tubules were atrophied and their lu- kill young pig:, in Jes$ than two months.
mina were devoid ofspem1atids. Whil;;t 1hes-e findings concur Tryp<mosom(I simiae infections in pigs eclipse in impor·
with field observations of reduced fenility. the same is cance al.I other uypanosomal infections in this hos1, lllis para-
1101 true for the severe central nervous system patholo1,'}' site may kill pigs after an incubation period of only four to sLx
of experimental infections of goats.3 19 The oedema and days. mid infected pigs may collapse and die within 24 hours or
inflammation of the choroid plexus observed in experimental the first signs of disease. ll1e introduction of the parasite to a
T. vii/ax infections may occur in aatural infections. but the piggery is usually by an infected !Seise fly, after which .:;pread is
under-diagnosis of rrypanosomosis and the common prac- belle\'ed 10 be by mechanical transmission involving biting
tice of slaughtering sick animals generally precludes full flies. often found in large numbers in pig pens.
evaluation of the pmhologyofnatural infections. The coul'$cOfinfection lsso short that pigs die before los-
ing condi1!on. although some animals may live for as long as
Diagnosis eight t0 ten days. Affected pigs are dull and inappemnt or
or
The diagnosis trypanosomosis in small ruminants is no completely anore)iic. The)' ha\·e a stiff, unsteady gait and
differem from that already discussed for cattle. but it is eventually become ptostrate. Dyspnoea is evident and

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cyano~i< may be seen. 'lnmc pigs rroth at the mouth and rectal temperatures may reach 40.5 •c and there b tachycar·
there may be d iarrhoea. Rectal temperatures rench ·l I •c but dia. Oedema 01 the lower limb, otcur~ early in the course of
the exiremitks are usually cold. <l1,caoc, a, alfoctcd horse$ arc le,, ac1ivc than normal. Sub·
The course of the disease within a herd j<; dramatic cutaneou~ Ot!duma then affect~ the venrral tlum1x and abdo·
After one or two early deaths. several more occur ,,·ithin men. Oedematous plaqu~ may form on the flanks. hm
a week. and after a short time there may be only a ft\\ depcndem ocdcnrn is more pronounced Wcakneo:. ~et, in
~UC'\ivor.,, rapidly. Initially, mucou" membrane~ ma} be conge;;ted
and ictcrlc but th!:~ become pale as th<: disea,u progres.se,.
Pathology Ophthalmitis b not a con,tam finding; iritb ma~ l1ccur and
TI1e necropsy findings \"ar\' considerably. and 1hcre are no up t(l 20 per cen1 of horse, clinkall~ alfened b~ T bru,·ei
pathognomon ic changes. the blood is u~uallv cyanotic: ma~ dc\·elop keratitis and conical op:icicy. It- \hhough
clot<' slowly, and ch icken fat clot~ ma)'bc found in the heart. l\mph node~ become enlarged, the~ are not ea~il> palpable.
Haemorrhages affloct the epl- and t•ndocardium. !;idnt'y~ The conditic>n of the animal deteriorates rapid!> and w,,ak-
and serosal su rfaces. The splC'Clll is usually enlarged and the ne~s may progre«s to parapl«gia. Oedemn bc,comes increas-
pulp soft with a ·s.trawberl") jam· appearance: d1ere is exces- ing!~ severe. Animals die within 1wo to four weeks from lhe
sh•e serous or sero•sanguincou~ Ouid in th1t pericardia! sac on,\?t of clinical ,igns.
and pieul'lll and peritoneal c;witie,: the trachea may he filled Chronic tl")·panosomo~is commonly occurs ln horses
\1ith froth when there Is also pttlmonary oedema. Thoracic.: untl donkeys. \nimab lose condition a nd weight (Figure
and abdom inal lymph nodes arc usuall~· enlarged anll 12.1 3) and the coat become,, har,h and dry. The dcvelop-
oedema1ous, and maf be haemorrhagic. me111 of .1nnemla follows a course similar 10 that in canle.
,\ nimal!, bc•comc extremely ,,eak and show signs of ataxia.
Diagnosis and differential diagnosis but lhllullr cominut! 10 1ta,. :.ubcmancouo. ocdl'ma. ini-
A history of s udden death in a number or pig~. comblnet! 1i;1lly nlfocting till' limbs and ,·.,mral abdomen. may iJxtcnd
with clinical signs as described. is suggesti\"e of T. simiae In to the sheu1h, scrotum, perin,·um und occa!>ionallr the
areas adjacent to 1,ccse infestations. '\hhough ~udden dca,h head I he condit ion is cx.1ccrbated b> ~trongylo,b and b
may aris(' from African swine fever, anthrni.., and other infec- fatal, often after several month, dur.ulon. In a ,urver in
tiou~and non-infectious diseases, 1hc presence of fulminat· the Sebungwe arcu of Zimbab,, e, a high proportion 01 don-
Ing parasitaemia,. readily dc1ectt'd in Giem'3-~tain!ld blood kl?~-s w;1, found to harbt:>ur T. /1rncei and ·1: ,·cmgole11,w in-
smears. co11firm, th<: dfa1,•tl0sis. fections. ~ Such mixed lnf~ctiom, in hor~es give rbt: to
Chronic uypanosomosb should be differentiatc:d from more severe diwase.
heimimhosi~ and malnutrition.
Pathology
The general pathol<>t.'Y of equine trypanosomosis resembles
Clinical signs, pathology and diagnosis in horses
that occurring in cattle.:\ \:ery full account of the histopa-
and donkeys
1hology of experimental 1. (Jrucl'i infections ill hor<e~ was
\\1th lhe advent of motoriicd transport and mechanized girnn by ::,.11:cully and '\e1tz 1"
cultivation. the role of the hor,e hab dimlni$hed in many The hbtological lc~lon, in the c:entra.1 nervous~~ ,u:m of
pans of the world, including southern .-\frka. 1:unhcrmorc. horw" \\ith experimental chronic /. //ruc,•1 Infection$ were
t.be peoples living in or near tsetse- infested habita,s in th is characutrt7.ed h~- ,1 severe pleocy10sb of the memuge~ and
pan of ·\frica have no tradition of keeping hor~c,. b111 in an cxten,lve ~ubpial glio,b. Tlwri• 1\·a~ abo segnwmed
some are.i~ donkey~ are 1mporian1. rlorH;s do not 'Survive demyellnation of uptir 1rac1, an\! ,onw other area,; of white
long in the presence of mfec[Cd t:;etse Oics. whereas donkey~ mailer a~ well as grey matter. Exum~t\·e perivascular cubmg
an: more tolerant of t~etse-transmitted trypanmome~ and with lymphol')'te~. pla~mo,~te,. large mononuclear and
frequently appear to thri\·e in lightly infested habitats. \iev- '.\Ion cl'lls also occurred. tr istological change, in the visceral
crtheless. ·1. bm.cei cau,es acure disease in donkc,s. and is and other organs were not uniform. but included re-<1cuve
abo a most :;erlou~ pathogen of horses. J"r)'f)(ll/oso,,u, con· lymph node,. imer~tltial myocardlth, haemoi,lderosi. and
golense produces a more chronic. was1Ing condition, ,,hll8t focal l'ndophlebitic ,~~ion, in \'l'im of th<> testc<.
T. 11i1•ax. although usually producing chronic and more mild
disease in equids. ,-.; has been known t0 cat1~e a<'ute infec· Diagnosis and differcn tia l d iagn osis
Lion in the horse ..;b. i:• ·1 he diagno~;ic: methods for equids are essentially the ~me
as those for cattle. It is often necessary to repeatedly test
Clinical signs those individual,. su~-p~ted 10 be ~ulfrring from 1rypanoso·
The general cllnical signs resemble those seen in ruminant~. mosb In order to confirm diagnosis.
l lorses with acme uypanosomosis are ,·ory dejected. Inter· ;\cute disease must be distin1tt11shod from African horse
miuent parasiw.cmia occurs and is accompanied b, fever: ~kkne!'S, amhmx and babesio,h,. in which high feH·r.

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 Mrican animal uypanosimwwi; 279

'
figure 12.13 Ahorse with a chron!. Tr;panosoma congo!er:se ·~!ect1on showing emac,at,on ano vent<a! oecema

sudden onset of signs. congeslion of mucous membranes sleeping sickness. bu1 such infections are subclinical in
and petechiation occur. Similarly. in chmnk trypanosomo- dogs.1 80 On the 01her hand. f. bmcei bruce/ is the most
sls the oedema has tO be differenriared frc)ln tha1 occurring p,Hhogenic 1rypru1osome in dog;. producing more acute
tn ·\frican horse sickness. and the .u,aemia from that of discose than T. co11golrmse. although strain difference,-
equine infectious anaemia. Chron ic bahesiosis may also orcur, some sin1ins bei ng highly Yirulent. Dogs a-re refrac-
produce signs similar to chronic trypanosomusb. tory to infection wi1h T. 1d1'(1x.
The signs of weight loss. emacia1ion mid oedema caused
by dourine (T. eq11iperdw11 infection), suongylosis, malnu- Clinical signs
trition and di·ntal disorders should be differentiated from The general clinical ~igns of trypanosomosis in dogs re-
those caused by sa!ivarian trypanosomes. semble those in li\"cstock, but the course and severity of the
disease are modified h~ Lht' virulence of the parasite and the
susceptibility of the hosL. In acute disease the dog suddenly
Clinical signs, pathology and diagnosis in dogs
become, dejccred and inappetant. Pulse rote, respiratory
Dogs are of greater cultural than economic imponance in rate and rectal temperature arc raised. Anaemia develops
•.\frica. The typical village mongrel sur.ivcs in man} 1~e1sc- l"Jpldty; mucous membrane1: become pale and the dog
infosted areas wirh little or no aitemion from !rs owner. show~ ,;igns of weakness and depression. Thl're is progres-
These are usually small animals. used for hunting and herd- sive weight lo:-s; as the condition deteriorates, the <"Oat be-
ing, and a;. watch-dogs. The exotic breeds, selected for come~ dt-;. and ~ubcutaneou, O<)dema. particularly of the
simil;tr purposes. quickly succumb ,o trypanosomosis when head and limbs. may develop.
they are e;,.-posed to tsetse challenge. E.xposurc of suscep- Ocular lesions characterize both T. brucei and 1: congo·
tible breeds may also occur as they are cransprJned through ll'nse infection~ but arc more ~e\'cre in the former case. Try-
rsetSe-infested areas, or as small numbers of infected tsecsc panosomn brucei in\'adc, all !issues of the eye, producing
are carried to otherwise t,ecse-free areas by vehicles. blcpharitb. coniunctivitis. kcrat!tis and uvcitis. There may
Dogs ha,·e been incriminated as reservoir ho~ts of be transient glaucoma, rn•, and excessive lachrymarion is
T. br11ce1 gnmbiense. the cause of West African human common. Additional!~·. T. bruct>i invades many other

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280 ,T.cno, , w,:,; Pro101.oal dlse._ises

tissues. and by entering the cerebrospinal fluid causes ataxia tlon losses. In these circumstances the need for supportl\•e
and partial paralysis. Death ensues \\~chin 4 to 10 weeks of therapy is greatly reduced. However. the cost of constant
the onset of signs. monitoring has to be borne in mind. There are numerous
Infections with T. co11gole11se generally follow a more well-documented examples of susceptible breeds of li\'e·
protracted course. In chronic cases. dogs become severely smC'k being successfully reared in many tsetse-infested
emaciated. and ulceration of the oral and gastrointestinal areas of Africa a~ a result of chemotherapyorchemoprophy-
mucosa may develop. leading to haemorrhage and la.xis.113· t6J. tllZ • .!116 Considerable success in maintaining
melaena. uo tnappetance may only be imemiinent. with caule. in the face of hea~· 1rypanosome challenge, has been
animal~ being periodically 'off colour·. ln exoric breeds achieved in Zimbabwe30 and in Mozambique.270 To control
death occurs after only a few months, but indigenous dog's u,•panosomosis successfully by chemoprophylaxis. it ls
may remain infected for long periods. necessary to treat a high proportion of cattlem risk at regular
intervals. This is not always possible, especially if o,mers
Pathology have 10 pay for treatment. and consequenlly a therapeutic
Lt?Sions of rrypanosomosis In dogs ' are broadly similar to approach 10 control is \\idely used. The.re is considerable
those found in livestock. ,\naemia is t11e most read Uy detect- evidence that such therapeutic control assists the acquisi-
able change earl>• In the course of infection. and the PCV tion of pro1ecth·e immunity rn locally prevalent trypano-
drops marked!>·· Whilst T. congolenseinfections produce ex- some serodemes: 13• 321 whereas with chemoprophyla>:is
tensive intersiitiRI oedema and ul<:era~ion of gastrointesti- immunity 10 1rypanosome:. does not seem 10 de,•elop. 11;
nal mucosa. tlssue 11.'sions induced by T. bru~i an> much
more severe. There is marked cellular infiltration and cellu- Treatment
lar degenera1ion and necrosis. 199 The heart. choroid plexu,, The satisfactory treuunent of uypano,omosis requires more
and eyes are consistently and severely affected. There are than a correctly administered rrypanocidal drug. and the
also marked changes in lymph nodes due 10 the initial pro- speed of reco\'cry is large!>· determined by the plane or nu-
liferation of lymphoid cells. although these may subse- trition, the amount of exercise during convalescen~'.e and
quently bedeplc1ed. the duration of the di~ease. Well-rested and well-fed ani-
mals recover more rapid!}' af,er trypanocidal therapy than
Diagnosis and differential diagnosis do undernourished animals which have to trek long dis-
Standard diagnostic methods are used co detect trypano- Lances to pa,;ture and water. Ho\\'ever, chronic uypanoso-
somes in the hlood. However. after 1he initial phase of high mosis or1en fails to respond to therapy; the ferrokinetic
parasltaemia, repeated sampling over several days may be disturbances and accompanying dyshaemopoiesis appear
necessary h~fore uypanosomes can be found. to be irrever,,ible. and ,1lfoc1ed animals may remain thin and
The anaemia caused by trypanosomosis must be differ- anaemic despite 1ryponocidal trea1menr.
entiated from 1.ha1 arising from infection \\id1 Ancylostoma
canhwm. Babesi<1 cani, or Ehrlichia ca11is. Hool..·worm infec- History of trypanocidal drugs
tion ma>' be concurrent with trypanosomosis and can be Not all compounds active against nypanosomes arc directly
confirmed by faecal examination. ln young dogs. babesiosis lcrhal to the pamsitcs, and consequently these agents
(or biliary fever) Is usuallv acute: rectal temperature may should be referred to as amitrypanosomal drugs. However
reach -1 I •c and imraery'lhrocytic picoplasms are usually it is usual to refer to this category of medicines as trypano-
readilr detectable in Giemsa-stained blood smears. The di- cidal drugs, or trypanocides. This latter umn is used in rhe
agnosis of ehrlichiosls (Nairobi bleeding disease. tropical followlng account.
canine pancytopenia or tick fever) in an endemic area relies The small but imporuuu group of chemo1herapeutic and
on t11e presence of such clinical signs as a moderate fever. chemoprophylactic compounds has been frequently
crusty mucopun1lent nasal and ocular <lischarges. foetid r~iewed. 83• 157 z:i~. :izs. 32,; Trypanocides have a h!s1ocy
breath, s1airted teeth and pale mucous membranes. which dares back 10 the early pan of the twentieth cenrury.
Bilateral corneal opacity produced b)' uypanosomosis when hundreds of substances were tested for antim.icrobial
must be distinguished from that seen in canine hepatitis acthity and for activity against the then recend) disco\'ltred
(adenovirus-2 Infection). In which vomitfon and acure ab- trypanosomes. the fiNit trypanocidal substances 10 be found
dominal pain are presenting signs. being several dyes. Since then arsenical and am!monial
compounds have been sho\,11 to be active against uypano-
somes, and sodium antimony tartrate (beuer known as
Control with trypanocides
rartar emetic) became widely used throughout Africa.
Underligh1 t,etsc challenge a thcrapeuricapproach 10 con- 1111>ano,ome-infected canle recei\ing an inrra,·enous
trol can be successful and economical. Vigilance is nece'S· course of the drug. J\hhough thi:. treatment was first used in
sary to ensure that infections are treated at an early stage. Zimbabwe (then Southern Rhodesia! with great success. It
before disease has progressed too far and caused produc- w.u. claimed that tartar emetic was used in Zambia (then

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 Arrican animal 1rypanosomosc~ 281

1'onhern Rhodesia) on a scale unrivalled elsewhere.' for up 10 12 days after administnuion a1 a dosage rate of7.0
1n 1921. suramin was markered ro creac T. ovmzsi infec- mg/kg,;iog The duration ofprotecrion conferred by isometa-
tions and human sleeping sickness. The drug is still widely midium is dose-dependent.242 a higher dose providing
used. nearly 80 years later, and is the drug of choice for the longer protection. E'.xperimentally. the prophylactic effect of
treatment of early cases of sleeping sickness. Research isometamidium has been increased significandy by incor-
which began in the 1930s on phenanthridinium compounds porating the drug in polymers in order to produce a sus-
led to che discovery of dimldium bromide. By che early 1950s tained release device which is implanted subcutaneously. 5~·
this compound was used extensively boch for cherapeu1ic 89• 90 However. results from experiments with laboratory-

and for prophylactic purpose:.. Dimidium is relati\'ely toxic: adapted strains of parasilcs in relatfrely stress-free experi-
it produces reactions at the site of inoculation, and treated mental animals. may bear liule relarion to events in lhe field.
animals may become phocosensirive. Despite this. hea\'Y re- TI1ere are numerous reports 1ha1 the higher the tsetse chal·
liance was placed on the drug. and in the l,ower River district lenge, the shoner the duration of prophylaxis. Addirionally.
of Malawi (then NyasalandJ cattle were successfully main- stressed animals on low planes of nu1rition or with incercur-
tained as long as they received regular irea1ments of dim- rem disease do not always ceceive the same level of protec-
idium. However, in Malawl. 188 as elsewhere, drug resistance tion as do bener managed animals. The most important
to dimidium bromide emerged in the mid-1950~. and this factor determining the duration of prophylaxis is the sus-
drug was superseded by quinap)Tamine, which became cep1ibili1y oi the trypanosome population to che drug.2~ 1
available in the early 1950s. Serious drug resistance also 'frypanocides are toxic compounds and have narrow
emerged to this compound and by 1977 it had been with- therapeutic indices. so therefore the dosage rates canno1 be
drawn from the market. In 1955, homidium was marketed. gready-increased LO obtain greater trypanocidal efficacy. Fur-
but once again resistant strains of trypanosomes emerged thermore, tl1ey are not equally effective in the crearment ofali
and became widespread. At almost the same rime, dimina- species of trypanosomes. A single injecrlon of diminazene
zene became available and was used to overcome quin- cures T. congolenseand T uiva.~infeccions at a dosage rate of
apyramine-resistance. Being babesicidal as well as 3,5 mg/k-g but a dosage rate of;,o mg/kg is necessary to treat
crypanocidal. diminazene has the advanrnge that Jr may also T. brucei infections. The efficacy of rhe phenamhridlnc com-
be used for trearing animals suffering from redwater. pc,unds. homidium and isometarnidium against T. congo-
P)'l'lthidium bromide then became available; it was mar- lense and T l'iva.r is morn marked thai:1 againM T. brtu-ei ar
keted in 1956, b111, again due 10 the emergence of dnig recommended dosage rates. Dosage rates ofisometamidium
resistance, it was withdrawn from the market in 1985. in excess of 2.0 mg/kg are toxic 10 canle, and even below this
\o\'hen isometamidium was marketed in 1961 the acLive level. the drug is irritant and has marked local effecc-s.
search for trypanocides diminished. This coincided 11~th the The prophylacticeffec1 of i$ometamidium depends upon
period when many African counLries gained independence the slow release of the drug from the depot created at the
from colonial rule and increasingly s1ringem requirements intramuscular injecrion sire. Diminazene is rapidly excreted
for the registration of drugs were demanded world\1ide. and has a negHgible prophylacric effect. Homidium is ab·
Funhemiore, comparable investments in pharmaceutical sorbedmoreslowly than diminazene from an intramuscular
areas other than crypanoc:ides are potemially much more injection sire. but more rapidly rhan isometamidium.
profitable, so at the end of a century in which grear scientific l$Omeramidium b lrrilam, and, ar the site of injection. It in-
advances have been made, Africa remains with only four duces an imense inflammatory response and causes necro-
compounds (Table 12.2) \~ith which to treat and prevent sis of muscle fibres. This lesion becomes enrapsulated and
one of rhe contine·nt's biggest ~co urges. is later organized by fibrous coimective tissue. From this
depot. the drug is slowly released. Anernpts to overcome
Principles oftrypanocide usage this irritant side-effect by reformulation of the drug have
The detailed pharmacokinetics and modes of attion of rhe been unsuccessful tc> date. Two important practical p()ints
trypanocides are incompletely understood. bur these com- are related to the injection site reaction. Firstly, the repeated
pounds arc regarded as ha\ing ei l.her 1herapeuric or prophy- inoculation of caule kept under rs.e1se challenge induces ex-
lactic properties. H~. 2 1-1, 2·10• 2az Diminazene and homidium tensive muscular scarring, and, to reduce carcass damage
ru:e therapeutic crypanocides, although homidium does pro- and subsequent losses from condemnation. the drug is usu-
\1de prophylaxis for several weeks.s9 lsometamidium, whilst ally administered into the less economically valuable neck
having a curative elfect, is \1idely regarded as being lhe only muscles, often with untoward side-effects. Secondly. from
trypanocide worthy of being used for propbrtactic pur- the viewpoint of human consumption of treated animals.
"°
poses. 1 The duration of chernoprophylal.'is is iufluenced the question of drug residues in the animals' tissues arise~. Jc
by se1·eral factorsr12 which largely relate to the race of excrc- is thought that because of rhe poor absorption of the drug
Lion of the drugs. 113 Diminazcne is more rapidly excreted Crom the gastroimesLinal uact, i;,o and che lack of untoward
than lsometamidium. After injection, initial high drug levels reports of toxici1y in humans. the risks posed co human
fall rapidly, although in cartle residual procec1ion may lasr health by residues in meat and milk are minimal.

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Table 12.2 Generic and t.-ade COMPOUND MANUFACTlJRER ACTION RANGE OF ROUTE OF AEMAP.'<S
names of trv!lanocides for the DOSAGE 40MIN1S·
treatment of Alncan amma GENERIC fRAOt RATES TllAT!ON
1rypanosomos1s l\l4ME NAME MG/KG
Oimmaiene Beren1I Hcechst A.G. T 35-7.0 11m Also
aceturate Germany babes,c,dal
slc Toxic to
horses.
donkeys. liogs
and camels
Homidium ~.n,dium CAMCO T(Pl 1/m
bromide Animal Health,
U.K
Homidll\111 Rhone·Me(~U)(. TIP) I 1.0 ,rm
chloride France
Jsometa· Samorin Rhone-MerieU',. P/T 0.25-1.0 1/11, Tax,c atove
m1d1um France uM··· 2.0 mg(~g
High!~ ,rritanr
<\void
soocuta1eous
aamm,s,rauor.
Tr,pam,:i1UP\ Rhone-Pouf911C P/T 0,25-1.0 11m.
Sante, France {1/vl"..
Ouinapyramine T:yc,mde Rh(JJ1j!·Meneux. T 3,0-5,0 s/c Rest .~1mals
sulphate· France before ano
ahe'.
treatmeni
Ou1napyramine TrlJ)3tide flhone-Mernrux. PIT 3,0-5.0 Oos~e
cai..salt D'.OSai;· • France calcularea 2s
sulpl'late

Aeimroduc!!j in 1935 to treat mainly T e,a11Simfections

Prosal; is a mixtvre of sulphate and cnlor,de sail$ 01 qu1napyram,ne
Source. reference 0
1/m-= imtamuscuiar
sic : sul;cutaneo~s
1/~ = intraver.oas
i : Therepeutrc act,011
P -= Prophylatt1c action
IP! -= Shorq1rophvlac11c actlvi!y

Trypanocide u se in cattle, goats and shee p The practice of using different tiypanocides alternately
TI1ese animals mlerace diminazene. homidlum and 10 overcome drug resistance warrants mention. Diminaz.ene
isometamidium ar recommended dosage rates (Table should be admin istered 1101 less than one month aher
12.2). bul quinapyramine Is no longer used In lhese spe- isomeramidium treatment,9 and should be follo\,·ed by an
cies. Trypanocides are dissolved ln warer immedialely be- i11terval oi at least one week before further isometamidimn
fore use. and normal aseptic precautions are necessary in treatment 10 avoid lethal synergistic toxic effects when nor·
order to reduce reactions at the injection sile. Work. oxen ma! dosage rates of the two dnigs are administered a short
and dairy a nimals should receive deep intramu:,cular in- time apart. Affected cattle lose condition rapidly. becoming
jections imo the gluteal muscles instead of the neck dehydrated and emaciated. Animals may recover after a
muscles. Calves, goat$ and sheep receive deep Intramuscu- long time. but up to 6 per cem die. :S:ecropsy findings usually
lar injections in the thigh, care being taken to avoid injury include severe fatty changes In the liver, with enteritis being
10 rhe sciatic ner.-e. The intravenous administration or more variable.
isometamidium as a 1 per cent w/v solution has been suc-
cessfully employed to n:eat bo\'i nc uypanosomosis/>3 but Trypanocide u se in pigs
careful injection is essential to avoid systemic and local Chemotherapy of 1: simiae has limited success because by
reactions.lij the time signs of disease are see11 it is usually too !are 10 save

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 Mrlcan animal 1rypunos-0mo,cs 283

the animal's life by treatment. c\'en when early cases are Trypanocide use in dogs
treated. few drugs arc effective. Sinutlmneous treatment Dimi11aze11e is toxic 10 dogs at doses higher than 3,5 mgfkg
\\ith quinapyramine sulphate at .5,0 mg/kg and dimina- and its use is he,l avoided in the treatment of rrypano~omo-
zene ac I 0.0 mg/kg can effect a cure. Other dosage rates or ~is. especiail} of T. brucei infection~ which require treat·
these dmgs have also been used, but relapses after treat- rncnt at 7.0 mg/kg. Jlowcver, simulrnncous administration
ment are common and ma) be due to the presence of para- of difluorome1hylornithrine (DF~lO) and dimlnazene ha.~
sites in the cerebrnspinal Ouid.334 Constant \'igilance and r~ulted in the successful 1reatmem of T. brucei infections.6
retreatmcnt arc therefore necessary. In the race of an out- Homidilllll. at a dosage rme of 1.0 mg/kg. or tsometa-
break it is usual to treat all pigs at risk prophylactically in midium, al a dosage,.tte of between 0,5 mg/kg and 1.0 mg/
order to halt the spread of infection, and for this purpose kg, may be used to treat T co11gole11se infections. However,
quinapyramine prosall has been used ar a dosage rate of the intramuscular injection of these drugs often produces
11,i mg.'kg (calculated as the sulphate) 1\'ilh variable suc- hot, painful inflammatoty lesions at the injection slte, which
cess. lsometamidium give,1 as a 4 per c~m w/v solution br re$ulcs in lameness of varying severity. Treated dogs should
deep intramuscular injection at dosage rates of between be well res1ed until the inflammation has subsided.
15.0 and 25,0 mg/kg are effective in halting the spread of Quinap)Tamine sulpha1e has been found 10 be suitable
disease. but large, crarer-like ulcers develop at I.he sites of for the procecrlon of dogs agains1 t.rypanosomosis. 10 and
injection.64 can nlso be u,;ed for treatment. Animals muse be cool and
Prophylaxis has been attempted with a surarnin- rested before and for o.t least two hours after subcutaneous
quinapytamine complc..\ at 40 mg/kg and with tJuinapy- injccrlon of qulnapyramine sulphate; a dosage rate of
ramu1e chloride at 50 mg/ kg, but neither of these 5.0 mg/kg i~ used. and may be repeated at monthly Intervals
formulations is commercially available. for prophylaxis.

Trypanocide use in equine infections Failure of trypanocides

Diminazene is poor!)' tolerated by donkeys~• and. on occa- When trypanocides fail to pro\~de the expected cure or
&ion. horses. The use of the higher dosage rate 7.0 mg,kg) pro1ec1ion there i., a 1endency to assume 1ha1 drug resis-
required to treat T. bmcei infections is often accompanied tance has arisen. Whilst this may be true, there are several
by nervous signs. lsometamidium may be used to treat and other reasons which can contribute to their fa ilure rrable
pre,·em equine 1: co11gole11se and T. uilr<Lr infections. bu1 12.3) Considerable logistical conslraints exist throughout
careful aseptic precautions and arrention to the intramus- Africa in 1he provisio11 ofvcwrinarv ~ervices to rural areas,
cular injection technique arc needed to reduce local tissue and farmers often reson to the trcanncm of their own ani-
reactions. Separate needles should be used for filling the sy- mals. A survey conduc1ed in the Eastern Province of7.am-
ringe and giYing the u1jC!crion. T() avoid depositing the itri- bia, for example, revealed that 6i per cent of 1he
lant solution subcutaneously. a sufficiently long needle rrypnnocides were administered by 1he cattle owners. 61
should be used: leak-back ~hould be reduced b~ using a l'rypanoddes are relatively inexpensil'e drugs. They are
small-gauge needle. applying firm pressure as 1he needle is sold in granular, 1ahle1ted or powder form and have to be
withdrawn and massaging the injection site well to disperse dissolved In \\'ater LO obtain a solurion for injection. fac-
the solution In 1he muscle. Quinapyraminc sulphate may be tors which c()ntribute 10 1heir l,idespread undcrdosagc are
u~cd to trea1 T. bmcei. T, co11gole11st> and T t•iu11x infections. incorrect dilution and the inaccurate estimiulon of 1he live
but animals must be well rested beforehand, since side-ef- wcigh1s of animals. The l1Se of un5te1 iii zed syringes.
fects usually occur. which usually pass within one to two needle, and the waler in which 1he drugs are administered
hours. These are transient and include restle$sness, sa!h·a- may Induce the form:uion of abscesses which, if they
tion. sweating. fasciculations, diarrhoea and abdominal dis· burst 10 che exterior. results in the loss of the drug. Con-
comfort. which may cau$e the animal 10 back away from the sequeml~. treatment quite often fails to produce the
handler or 10 become recumbem and roll. In weakened ani- c~pected results.
mals the dose should be divided in rwo; the second half The use of Lrypanocides at subtherapeutic levels is be-
should be administered b} subcutaneous injection four to lievl!d to promote drug resistance in trypanosomes- ll!I, ' 22
six hours after the first. Since diminazene is rapidly excre1ed. the risk of 1rypano-
Horses and donkeys can be protected by quinapyramlne somes being e~posed to suble1hal levels of the drug is less
prosalt glven at a doi.age rate of3.0 mg/kg. Since the pro~alt than L~ the ca~e \1ith i~ometamidium. Prophylactic levels of
1~ h1soluble. it is admlnii.tered as a su$pension by suhcu1a- isometamidium wane over much longer periods. Even when
ncous injec1ion. The injection sirn should be massaged well cattle receive isometamidium a1 Lhe correct dosage rate. it
w minimize the local reaction, bm a dcpo1 of drug b formed ma) be administered at the wrong time. Tsetse challenge
In the tissues at the injection sire. which induce~ an Inflam- rna) increase a1 a tlme when drug leveb have declined and
matory and fibroblastic response. The nodule may ulcerate are not protecti\·e. Furthermore. it appears that 1he duration
and lead to discharge of the drug to the exterior. of the protection is reduced in 1hose catlle which are in poor

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Table 12.3 Causes of epo.arer.t orug resmance in uypa~osomes: ia1lure
o; l"fpanoc1des withdrawal of many trypanocidcs io the past, and resist-
ance to the few rcrnairling current!) used trypru\ocides is
U!Y.leruw..ge documented.~· n 9 The degree of resisumcc in liypano·
underesuma1ea oocy •:,eight somcs i6 not clear cut: a spectrum of drug sensith'ity
ovsrdiluted solut,an cf t!'fl)i!not1tie occurs. and resistance to low h~,·el~ of a drug may be 01·er-
inccr111a1v caicu!ated dose \•olume come by increash1g the dosage rate. 11 ) Cross-re~istance
dellbe1ate unoeroosage 10 enable treatrneo: of ":'Ore .r.•mals between different trypanocidal compounds can also
fncorrect in1ec,1on tecMmc;ue occur."-2; '.\lultiplc resistance of a strain of r. uiL•tn to nor-
- shOrt neetlles mal therapeurlc dosage rates of isometamidlum. dimina-
- non-ste,1lity/ao-.csssat1on zene. quinapyrarnine and homidium has been con-
- early w1tho,a·:1al o' needle firmed.r... but fonunau:f~ such cross-resistance is rare.
- elastu: recoil or 1iss~e produces excess,ve ·1eak-cac~· Resistance to quinapyramine de,·eloped after onl~ ~ix
Incorrect suategy momhs in some areas of Ken)'a 321 - a sana1ive treatment
irregular treatment of either homldiurn or dlminaxene was administered to
prolonged 1n1erve1s bel\~een treaiments fcnalleo.ga when drug control this. Cattle thus treated and removed from the
levels have waned! challenge area wercltub~equeml}' found 10 be free of infec-
lack of knowledge of s~sonahty tion nnd also grew well. ' 21 :\evertheless. i nducrion of resist-
Stress-Induced metabolic changes ance 10 quinapyramine In T. congolense has been shown co
malnutntion rei.uh in cross-rcsisrancc 10 h<>midium. isometamidium
lactation and diminazene. Z.!9 Hence. quinapyramine should not be
intercurrent <Hsease used in canlc. Oiminazene and isometamid!um are
breed-assoc,ated stress regnrdcd as a sanative pair. since cross-rcsb1ance between
work and trekking the tw<, l~ very rarefy seen in the field 113 and, in fact, the
Increase in Cllallenge multiple resistance in '/. 11iM.t mentioned above258 \\'as
seasonal overcome \\~th dirninazene at 7.0 mgikg.
changed grazing area In southern ,\fric,1, the only W('ll-~ub~tuntiatcd rcp<>rt of
absolutg mc-eise in ts!tse nu!!lbe1s resis1an1;,• 10 isomctamidium was mad!.' from Zimbabwe il1
Re'aps,ng inre.:uons !9i:.i."111 Infection or cwo non-immune ca11le with the sw,-
T•~e drug 1es1stance pucted isometamldium-re~ismnt r. to11gQle11u strain was
not cured by isomctamidlum creaunent at 1.0 mg/kg. al-
th()ugh treatment at 2.0 mg/kg did effect a cure. The prob·
Jem in the field wai; overcome by herd sanative treatments
general health or are on a low plane of nutririon. and it is with diminazene. /-.lore recently. diminazene acewrate re·
thought likely that crypanocides require an acth·e Immune sistant ~tock, of ·r. co11gole11se have been i~olated from cattle
response by the host for them 10 be fully cffectiYe. 11 J in the ZambvJ \'alley of Zimbabwe1311 and diminazene and
A serious cause for the failure of trypanocides is the 1sometamidium resistance has been reported in Zambia r
phenomenon of relapsing infection~. 113 Trypano~omes in In case~ of serious drug resistance, cattle have to be with·
exrra,·ascular sites may not be e.~posed 10 lelhal concenrra- drawn from the tsetse area, but re~istant trypanosome~ are
tions of trypa.nocldes. and they may then emerge at a time cyclically transmHted by tsetse, and they may then enter the
when drug levels in the blood ha\'C declined Trypflnosom<~ wild hoM re~ervoir. rransmi~sion of the ,train 11ill then con·
brucei in the cerebrospinal fluid is not exposed to trypano- tinut', albeit much diluted. between wild hosts. 99• 100 ·" The
cidal levels of dlminazene. which does not reedit}· cross the reintroduction of livestock into such areas mny quickly lead
blood-brain barrier. 31 - Although uypanosomes are to the re-emei:gcnce of dmg reshtance.
cleared from the blood after treatment. the lnfcc1ion may .\lelhods 10 prove easily that certain strains of parasites
relapse as parasites reirwade the bloodstream fr()m drug- are resistant are not available. Currently. three techniques
inaccessible sites. A similar event occurs \\ilh r. 11i11ax178 are commonlyu~ed to detect drug resistance. Firstl~. tesb In
which is known to invade lhe aqueous humour of the eye ruminants pro~ide dlrtoct and accurate information on the
and the cerebr()spil1al fluid of goats. 318 The extent of the dose required 10 cure a trypanosomal infection.!o9 The fre-
occurrence of relapsing infection~ under natural condi- quem!y used mouse inoculation tests can, at be~t, only
tions is not known. a~ il is difficult to distingulsh between broadly indicate the sensilivity of a ~traln.269 Second!~·. at-
true drug resistance and a relapse in the field. and the tempts ro conduct i11 uitro bioassays \\ith culture-adapted
occurrence of a relapsed infection may often ~imply be uypanosomes are being made. but not all field strains \\ill
ascribed to ·resistance. ad~pt to culture system~ and. of those that do. some mar
True resistance to trypanocides is a spectre which acquire an altered pouern of drug sensilivhy. JJa Thirdly. as
looms large. The developmem of resistance has led 10 the an alternative to bioassays for determining the sensitll'ity of

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 African animal rrypanosomoses 285

uypanosome s trains, is the use of a trypanocidal drug-ELISA against trypanosomosis. The ultimate goal, and the basis oi
in conjunction with parasite detection tests. A competitive many control Strategies, wns the eradication of the tsecse
ELISA capable of detecting small amounts of isomela· fly. The Tsetse and Trypanosomiasis Control Branch of
midium in the serum of cattle has been developed' .;· ;s. 319 Zimbabwe's Depanment of \'etcrinary Services has
and has been validated under field conditions. ;'3 achieved notable success in controlling the disease by pro-
gressively remO\ing the vecror. Different strategies have
Strategic use oftrypanoddes been used. and wiLhin each strategy different methods
The single rainy season in southern Africa innuences tsetse have been applied as 1hey have bet'.orne available. /\ brief
dispersal and leads ro a seasonal peak in the incidence of I ry- consideration of trypanosomosis control in Zimbabwe is
panosomosis.'14 The detemtlnation of the precise nypano- rlius very instructive.
some challenge is a complicated rnaner. 255 but from the Vector control began in the 1930s when wild hosrs of
practical point of view it is more im ponant ro establish the rsetse were elimi11aced by large-scale hunting operations.
disease incidem;e or risk. This tends to be seasonal. with This me1hod of control was abandoned in 1960. Tsetse
many new infections being contracted soon after the begin- con trol then relied inilially on bush clearing and the appli-
ning of che rainy season. This is the Lime at which isomeca- cation of residual insecticides 10 Lsetse resring sites by
midium chemoprophyla.xis should be used. Repeated ground-based spraying teams. However, these methods on
treatments may be needed during the rainy season. and cattle their own did not result in satisfactory control ofnagana,
can be maintained by chemO[herap)' \\1th d iminazene during and in 1964 seleetive hunting began in fenced corridors
the lower risk period.fa~dence pointing to the potential ,·alue that delimited tsetSe-infested regions. To limit the extent
of this strategy has been obtained with sentinel cattle in of the disease, livestock were excluded from many tsetse
soutllem Tanzania. which also has unimodal rainfall ...; areas by well-maincained canle fences. As a result of pro-
To apply this or any other strategy it is necessarr to meet gressive operations, tsetse were well under control by the
all of the logistical requirements of drug administration and early 1970s. and a system of compulsory monthly inspec·
disease surveillance. h is frequently the logistical bottle- tion of all livestock. with the treaLment of sick animals. ef-
necks which reduce the efficacy of trypanocid~ rarher lhan fectively contained the disease. These gains were reversed
the inherent inac;rivity of the drugs.113 The cattle owner's at- during 1he liberation war years of the 1970s. In the
tirude rowards his animals and Lhe management of disease mid-19,0s an estima ted half a million cattle were at risk of
in his animals is an imponam factor contributing to the suc- lrypanosomosis. Every year some 1-10 000 blood smears
cess or failure of drug use. A survey of trypanocide use in were taken from caule in risk areas, and approximately 100
several countries of southern Africa411 showed that. despite 000 treatments were given annually. 4'; From 1980, large-
the m·ailabllicy of diminazene and isometamidium. caule scale aerial spraying of insecticides was used, which was
owners preferred to treat animals that were clinicaU) af- aimed at reclaiming areas in\'aded b)' tsetse during Lhe war.
fected using a curative rather than a prophylactic approach. By I 989. trypanosomosis was again well under control. In
Moreover, preference was given to the treatment of the the year to September 1989, two million cattle were in-
more productive animals, such as oxen and cows. Although spected and. of 67 563 blood samples examined, only 596
this treatment regime docs not improve the reproductive cases were detected. From 1991 onwards, odour-baited.
performance of cows, it significantly reduces monality due insecticide-treated cloth targets became the method of
to nagana and improves the condition of oxen. This may, choice of the Zimbab1\fean Tsetse and Trypanosomiasis
from a cattle owner's point ohiew. be Lhe most effecrive way Control Branch and almost completely replaced aerial and
of dealing ,~th trypanosomosls. It is not e,ident that other ground spraying. lly 1999, approximately 60 000 targets
strategies will be readily accepted. were deployed as an effective barrier against tsetse rei1wa-
The warning has been sounded: 'Cattle cannor be kept sion from neighbouring coumries and the csetse-infesred
alive indefinitely on the end of a needle'.' 76 and this must be areas in the north of the country. The barrier protects more
recogni7.ed at the outset of any control programme. The use than I 000 000 head of cattle, mainly in communal lands.
of rrypanocides can be highly effectlve when chere rs close from the threat ofrsetse.62
veterinarysupervlsion, but iris only palliative. t\trempts must Despite considerable gains in other countries. few
be made to reduce tsetse numbers and uypanosome chal- achievements have been sustained because of the prohibi ·
lenge. ln the shornenn, the benefits which accrue from the rive cost of preventing tsetse from reinvading previously
control of rrypanosomosis with 1.1y1>anocides amply justify cleared areas in rhe absence of natural barriers. 1':agana.
the costs. In tl1e long term this approach is nQt sustainable. therefore, remain$ a serious consrraint to development
throughout much of Africa and new approaches towards its
control have been devised.
Trypanosomosis control strategies
The new c.;onrro1 strategy aims at ioten•ening in areas
For more Lhan 60 years, governmenLs in southern Africa where achievements aye likely 10 be sustained by the benefi-
have made coocened eJions to comrol tsctse in their battle ciaries. Consequently, in most cases. rrypanosomosis will be

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286 ,i:rno, """ ProuT1.oal disease,

comrol!ed on a smaller localized :;cale in area, that haw to rc\i\\!d since tht· dt.'Ve!opm,mt oi ~11theric pymthroid~ ior fly
be selected carefully. Socio-economic: and 1cchnkal criteria and tick comrnl. Thi, me1hod ha, been med successfully and.
are the major determinants in the st•lection process. at first si~ht. its,implicity and its effect on ticband !Ji1ing nic;
Comrol imervemions are more lil:.el~ to be ,uMained hu\'t! prompted demands for ih more \\idespre:1d u~e \\ hl.'r·
where the bencf!ciarie:'\ perccivl' U)-panosomosi, a, a wriou\ P•,
e\'er cattle are l'xpo'ied to t.<.et~. lll. ,.n, I lowe1 er. caution is
constraint to their de\·elopment and where its comm! v.ill nec,;,.;sory ior several irnponam reason,. The first relates 10
result in significant economic gains. To idcmiJY ~11ch areru., cost slnct' p~Tethroid5 are rclariv<•I}· "~pen~lve. l'or a commcr·
accurate information Is required on the dh1rlb111ion and dal farm. changing from its u~ual acarit'ide to a p~'rethroid
socio-economic lmponanc.: oi rhe disease. nnd on farmers' could incrl•aSl' co~ts slgnifi«:antl\". In areas where traditional
perception~ uf the impact oi th<.> dist>ase and hs control. farming methods are pra~~tbed, tick conu·ol mea,urL>:, aro, not
The lmpac1 of nagana on niral devolopmem 15 often always routlnl'I~· usc,<I, and th(- ret,'lllar ui-c of ,,yrethrofd~ to
unknown and control opcrntions have rrequently heen treat canle I\OUld be ,1 high additional expcn~e. 111 \icw of the
planned and implemented ba~ed on.spurious or lncompleui high operational ,O.!,I~, !he dura1ion 01 trcntment should be
infomlation and i1walfd assumption,. The socio-economic kt:pt 10 a minimum. fhel'efon:. before embarking on a control
Impact of bovine nypanosomosis varies widely and is inilu· operation \,ith d1is method. thcfl! ,hould be 1Aidence 1.hat Jt
enced b) factors such as the disease challenge llu.• degre-e nf will br e/Tectivt'. Several Important que,;tions havf.' to bl" asked:
tolerance 10 infcctlon and the appiic.ation of control mea.\ures. Do t:.e1s.- in 1he area take a high proportion of f~d1, from
TI1e long-term succ:-ess o( an iruerve11tion to control na- cmlle? Car, tsetse reim-asion of the area be successfully con-
gana (and many other diseast>sl does not dC'pend entirely trolled?·'°' I low mnny animals $houkl be m"llted and a1 what
on its effect on livestock production. 1 he Uve,wck owner'i; trc,lllncnt iutcn·aJ? l'or ho\\' Jong must this measure be en -
perception of its impact is of unnos1 imponanc;c this per- forced? To continue 10 treat cattk with dchamethrin indefi-
ception will vary from area 10 area depending on the form- nitely 10 comhat constant ts<•tse relnvasion raises not only the
ing or Jivcscock management practices and may \'81) in qu~tion of it., cost. bur also of the elf<•e1ivcn~~ of it:. insecti·
tjme. depcndi11g on other iactors constraining develop- cidal and acarkidnl propenie, nle emerience of tick~ that are
ment. For el(ample. in areas where canle are mainly u&cd resis1ant 10 dt1ltnmrthrin \multi be serious, although thi~ may
for draught power, the effcc1 oft ry1)anoso1l1osis on r~ n ii it y take sc\·erul )'Cal', t<> develop.
is likely 10 be perceh·ed as a minor problem. In <uch art"as. The control of ticks in hidierio untn•ated herd~ may
cattle owners are uolikt'ly to ~us1.ain control measures that lead to in•tabilit~· in the rc-lationship, between ca11le and
greatly improve reproductive pcrrormanni The oppo~itc tick·borne disease agents. Thl' endemic smbility prc\·alem
may be true In areas where the stt1c or accumulation or in many traditionally owned herds (see Chapter 25: Bovine
cattle is of prime importance. Even if nagana ts perceived babeslosis) i, likely to be disturbed by rigorous p~Tclhroid
as a constraint, comrol imerventions will not ah\"il}:. rt•sult treatmem. ,,,. To prevent !o,-ses. dgilant \'eterinary ,talf
in financial and economic gains, This will normal!\" be th«: would need 10 he :wallable to treat ~ickanirnals. and 1here
case Whl•n either the potential for hcr<l expan~ion is limited arc implication~ for the vaccination or canle against 1ick·
or where there is limited opportunity 10 incre~e the area borne diseases. whirh ma} rel} upon modNate sustained
under cultivation. These considerations appl~·. for ex· challenge for continued immunlt\· rquallr. with the cessa-
ample. to many parts of '.\ialawi. Under ~uch conditions. tion of treatment and incrca.sing tick burdens. 1hc intensity
inter.cmions to imprm•e agricultural and li\·estock pro- nr 1ransmission or \11nµlr1~ma ,pp. R1d1e~ia ~J>p.. Theileria
duction mar ha\·e minimal impact anti are unlikely to be spp. and f./rrlidiia (Cowdria) ruminalllit1m might then in·
sustained by the li\·C\IO!:I:. owner. crease. leading 10 incrc:t~<!d mortality rates. l)t'~pitc the«?
Sew!ral methodlt are available to (ontrol trypano~o1110- negati\ e con~iderntion~. the method has much to com-
s1s. They may be directed against the vector or the para~ite, mend it It can have a rapid impact on tsetse number~. but
or towards the Iive~tock and modified management. In it should be us!.'d for the sh on est time po&sible. in conjunc-
choosing a meth<)d, technical and economic aspect, have to tion \\ith other control rncasur ...,. The management 01
be considered and the most viable op1io1\ selected such insecticide-treated cattle in nn active. offensive wa) h
Some of the tsetse comrol techniques that ha\"e bfcn deYel· also possible \nimab could be trcalcd and mo,·cd
oped with large-scale eradication in mind atl' not well 5Uited deliberately into a t,t>tsc focus in an a11emp1 10 reduce fly
for use in localized. small-scale c:omrol op<.'Tation!-. Currently. munbers quicklv. Protection of the,e caulc against tn·pa·
~etse control utilizes odour-baited. insecticide-trea{cd cloth nosomosis would be necc5sary. because tsetse could still
targets and insecticide·trtatments or cattle (see Chapter 2; foed and transmit 1rypanosomcs hefore being killed.,~:·
Vectors: Tse tse fUes) Odour-baited, insecticide-treated Ult· Despite tlrn proven cffcctivcnc,~ of insecticide-treated
gets have been very effeclive in Zimbab\\e and ha\"e been em· cattle and odour-baited 1argc1,, the ,ucccss of a tsc1>e con·
ployed in otJ1er Al'rican countries to co111rol vmiou~ ~pec.ii;>s of trol Opl•ration will depend largely on i1s planning and
tsetse under differem ecological conditions. The concept of implementation. Governmem-controlled imen•ention, an,
using insecticide-trea1ed canle to control tse1sc has b<.>en normally carefully planned and implemented by high!)

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 African animal 1.rypanosomost'~ 287

skilled lechnical per5onnel. Although <:Ommunitr·driqm memary feed. e5peclally during 1hedry 'l.'ason. is an important
lse1se control opennion~ may s111l benefit from ~klllcd per- factor in mitigating disease. L.acunlng animal~. draught oxen
sonnel in the plan ning phase. 1he responsibility for the long· and sell:'Cted breeding bulls benefit from a high plane of nutri-
term implemen tation is likely to be in the hand, of the tion: they are then b1mcr able to \1i1h,tand 1he effects of 11yp2-
beneficiaries. The degree orco-ordinn1ion and S}Tlchroni1.a- nosomo:;ls. 11ie gern:ral principle, or good management also
1ion of acthities between the communilv members will de· require the prevemion and treatmenc of other diseases. and
termine the effec1ivenes5 of ~uch tsetse control operatiom regular. tactical tick co111rol. amht'lmlntir treatment and \«Cd·
Synchronizing and t()·Ordlnatlng the tsetse control :ictivi· nation rue useful 11djunc1sto trypanosomosis control. Sadly. it b
lies of a heterogeneous group of community members. with often the ca~c that many livestock 0\\11ers prcfor a simple cure
dh·erse objec1iv1a:,. may be difficult. Therefore. where 1;.c~c 10 the hwe-;1mem or their rime. energy and resources in im-
eradication cannot be achieved it is unlikely that 1se-ts~ con· proved management. ·me role of good husbruldry in disease
trol measures will be the only too! to control nagana. and a comrol is frequently undcn.'Stimarro. Improved breeds oflive-
combinatit,n of method~ is likely 10 be used. Depending on stock not adnptt!d to tropical conditions gencnilly fare poorly in
the perception of the disease impact, preference will usually hot climates, where pa,cur~ are often of tow quality and
be given to a particular method. Where endemic human tick challenge is frequcmlv encountered. 111cy also n."3dil~'
sleeping sickness occurs, $UCh as in parts of Uganda. Lhere i~ succumb to rrypano"°mosis.
evidence t11a1 communities will sustain Lhe use of trap~ to
control tsetse numbers.
h1m1unologicaJ control
Trypanoc!dal drugs will continue to play an imponam role
in the integrated control of uypanosomosis. 111,•y appeal to The antigenic complexity of trypanosomcs has thwarted
communal cauJe O\mers because they pro1-ide a means of pro- :mcmprl; to develop a vacclnc:10• 222 Allhough potential
tecting private good;, and may achieve impre:.sive results at low immunological targets wilhio 1he paraslte have betn identi-
costs. Although the low-level usage of these drug<; may reduce fied, no 1-accine will be commcrciall}' avaUable in the near fu.
monality rates, such usage generaily has low impncc 011 animal ture, and thegreates1 hope for the immunological comrol of
producl.ion. This will bl' the case panic1darly in high challenge animal tf)1i.mosomosis lie, ln the exploitation of trypanotol-
areas and when farmers use 1rypanocldes thrrapcutlcally i,.,
oram breeds of live~tock. Trypanotoleran<:c is ii general
rather than prophylac1ically. The de1·elopmem of rel>istancc in breed charac-reristic and. as such, some individuals are more
trypanosom~ to uypanocide~ i, a continuous threat 10 tht-ir 10Jcrant than 01hers.2."1 Furthermore. the trair is complex: iris
susiainable u~e In the control of nagana. liven in areas where related to 1h11 abll!ty to resis1 anaemia. the abili!) to control
resistance to trypanocides h~ nm yet been demonstrated, the parasitaemia and. possihly. the ,1bility to mount a more effec-
probabilityofitsdeveiopmem 5hou!d influcnccthc ~election of tive immune rc,ponsc 10th<' u,;pa.nosomc.Z1l fl also appears
an appropriate control strategy. that while thc.e proces,e, are under genetic control. they are
~lillions of domestic animals are kept in csetse-infestcd no1 directly linkC'd to each 01her. ·n,c search for generic
areas, but the degree 10 which they are exposed 10 tsetse is tre- marke~ has concentrated on th~ major hbtocompatibility
quently detennined br management practices. The avoidance complex and on leukocyte antigen~.:?1<;
ofhea1ily infested watering poims or grazing reserves reduces In the mcamlm11. 1hc corrcl:iuon of packed cell volume and
challenge. and thus comrols crypanosomosis 10 some e>.1ent, producti\iry parameters ofN'Dama canle23' form, the basis of
but the availability of al1ematlve water ,ourccs and ~uppiiei. of a field tt-M 10 select young Lrypanotoleram animals.~·1 It i., in-
feed are then necessary. Prevemion oflivcstock movcmem into tended. throuih a process of selectke breeding of I ryp::inoml-
tselSe habitats also limits the disease. but lhe c-nforcemem of cranrnnimal,. to reduc:e 1hc problem of irypnno~omosis and to
livestock movemcm control in Africa is not eas1·. because of improve livcstoc;k productMry. Howe,"l!r, this appro.1ch offers
communal grw.ingpractices. seasonal migraLions and local po- little hope tor southern Africa tOda}~ inheritance is complex
litical pressures usurping the law. and the number of a;1>ano1oleran1 animals b relatively
If animals are exposed to tsetse fly, the se1·crity of the ~ub· small.zn On the other hand, the fact that some Zebu cattle ,ur-
sequent u:ypanosomal infecrions is modified by the animal~' vive in tserse.infos1ed areas prmides a basis for atrcmpring to
general well-being. Well-fed animals are benerable to resist the selt>ct the more tolerant of these anlmuh for breeding pur·
effects of trypru1osomal infections. and many animals at· pose$. To do so would hasten 1he proc,><;-. of na1uml ,;election
min an equilibrium with the parasiLes. The pro1~sion oi ~upple· that is alread\ occurring.

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288 ,r,rno, "'": Prow~o3l diseases

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a \.''0~'·1 1ss,.
·1mwnl Rt•J:K,n <Jfth< Ot.•pnrlmrnt o{Vtttnnnry S..'Tt'l<rt <tnd
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.1nlmnl HM/111. PI01crrn,1rit2buq:. D,•p,1nm<,n1 or AWirulrurnl $11,'\i.n Of Ttyt"1rtQ>(>tn(1 l'it"fll' tn flhod~ia. RJ:od~Uw \ri.-l(irftia:ty /Otlf'110f,
E.<."('lnnmlc:o. nnd ~1arl,inng. Rt:puhltr of South .Mrica.. l . .Si'-62
r.; 8'\U.IY. I w. &SMfMt. f>,H I~- The U$O'Oi the acrid.inc: orange QBC .14 nan. w ..P ,tAf>,,.l ~I'll', r•.>..t .~u~o.,. 11.D.1w. u 1.\\'l", n •• ~972- Th~
tccbmqu< in th~ diU$nos!., of Alncltn tr)'P""OSOrnosis. Trmwwons-uf 1mr10rt.am:e 01 1hL· donkt!)' lrr111usmmusi ns a source or (o()d and a
lit,• Ho,111 so«ttt) o[Tropu:al M1'(J1c1nt /1111/ TIJJgien,. 86, "30. rC"sN,·oir oftrypanosomcs for Gf9ssint1 mar!fiumS\\'ti~tw. Rlrodi.!tla
.~IMr:t','ir<N. 6, Ill.
lS BAU$,,.. 1977 ~otc.i-ur Id toxu:au: dt: l'JiQmctamrd.ium pM iniecuon
1ncro\\'cincu$e c11ez q~rl1.1ues man1iti'r1."S e1sp<-ci-aleme-m ch& le 3,5 f\RVCf. o .• US9S, Prit)Jlminary rcpon on the UK'bc 0)· di~3$t: or n,lgana in
clromcdnlrc. Ht1,11cd'/i/,i,ag11 ti d•· M>!dtdM V1crttratr, ,te,Pn>~ Zululand Ubomho, /.ululand, 0..'Ccmlw, 18.<lS, Outban: Sonnen and
1ropiro1,x. 30, 373-375 Unvld
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durinJ.; lhe <'"-1:'lbli ...hmcnt o( Tryp,mmom<1 btucei infection lr.u.1.~ITUuad t'~mgo/,!uS<, /.rif!oClmftfi,r Tropt"1>m•·di~i,1 rmd P1uuslr.olnffe• .2i,
by c.w1se Uy. J1ams1roto1-0·, 88. 67-64. 3;1)..371.
57 ll,\tt \tA!\, A,, M~(,lt, .\., \;HAL. T. ,-... SOWMQ:,,,, ,., 198~, IJu• 37 llumtpeulic~trett of 8tn·nll llnd
CIUTA ...UU). It. 6'Mt1\lt;J\\\',\, .\,. HHU,
h1unu11e-h}1>othol<unlc•pltw1l>r)'·ad~n•l a.'ds. Endo(rln, R,:1rw• 10, Samonn in mite lnlccitd w,th four trypuno:,omo populoll<>rn, ,,;o!,1,-d
~~-It;? from i'....unh\itn cunlc. Ve,~rlnnry Pnra..4-Uo/ogy. 39, 42-52.
18 IIA>u,, M.. 1~96, Flfoct nl dc(en<ivc bcha,·l~ur by rat de on the lc«f,ng 3fl t.H1.1,•,,...,". tt,r...ll- & au<~unu ~.M.t-..• ags.;. Dipping to c(1nuol ,·ectol") of
,ucc.,..and num1i011.~ '"''" <1( thr ~e1<,c lly, C:/o.'<Slr,a pallidif}4 CQUlf p:tra.,\ltt"\, P,mi.,itQ/QK,' Todaf. :?', l?.l
Oiptera· (.io,,imdaeJ. 811/~•1111 o/Ernoma/111,-ieal RtsJMrcll. 86, 3:?9-336. J.~ CUR'-!~~ . M.J. ~ ,:,.c.,fft. \,. .. 19'i'O. 1~inosomn tt/1.aax in rumln.1ncs.
19 RAYi i~, M."' 'l:,,Mtuno. c o .• 1\l93,: The cfrttt of amJc infection b~ '/'rmis,u:1,om of lht Hl1)'t1! 'Wri,'fl•o/Tropl(a/ ,\fi,d/.-lnr and Hyg,,·n,• ~
rr:.iJN'lmiMmn c(mgolDniaon the.1nrnc:tion, and feeding ~111:c~ of ,he H>+-165..

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 African animal uypanosomos<.>S 289

to CL\U~,. P.H. ,\fl£Yti)t1, i •• RAUER,"· liNUOt-1 M. M .• ,ucm,m•, L k ,r,\.U: , impres1,~11n1 ~·.a t:Jg..,, in lf')'{lo,nosomii1.,f., conirol. l'ropt,~,t Animal
c:•• ,,9d. Hun prdcrcrtce& of u.tb1,,· Olp:cnx: (;fo~1md.:c; ba~d on Hraltlr ,rnd l'mduamn. 20 it.7-2.GS.
b!oQdmeal tdentlncatlon,• \f,'.'d(ml
. rmd \ ir.-·rlnnry l:1uomofo~ , l!,
hi 0011.r\!\ ,1. 1~18. \odU•\'COI.Uffllc"iUC\'~·011~\·he.,,frt:i'~lJld
169-180.
ht\1-c·in!t,h.-d u.1..:-a, 1n 1ht1 t-.a:-.:ern Pnl\1n1.·t·, i'..arnl)fa, 19..C)i ReQ]on.11
:: ctou<.1-1. o .. 1929, Rcporz 011 lfrf(;m: .4/fnirs for tll1• 1~,.,,, J92!J, \ "al. I, T~t)<'at'ld lr.1-tano,utnf.)t,T"> t:c,111101 P'rug_r.:tmml." R ff(,P}.. H:.muc.
London: Harruon llnd ~on, L1tl, lJn1bi1bwc~
..t'.: ClOUGft. o .. 19;.1. U'!port /JtJ ~-tfrtron ."r.!Ji1it, for tTJ,. ~",'ilr 19-11 \'ul Ill r~ oor,A, ,1. ti;igl. Btm:llt: r.ou .m.1J~ ,i"' ofhorc:ftar 1.,.:1w cont ml
Guildford o.nd bher. ntlllng .ind Son, Lrd op1:ranon, 1n JJmb.1b.,,•r Rcgfon-41 T,ct,t- .and .. rfpiloo~omu:ii~Cunuo1
,lJ C.LO~ICiJI, n. 19.1::, Rt•JJfJTI ()It .vr"""
AJ}tur,, JOY ti,(' YMr J93Z, \'ol. tV. Pn>jtf,"H1lttw tftrTC.:.Pi, ll;11iU'c, 1.;mhKlm1.·.
Guildrord ond Ii.sher: llilbn_g.uid 5on,. Lid , 63 00\\U .,. ,,,,;., ~j1JtU\(,t,N. u. .., fOSSOM, Jll.1 .• l98!t, ~Qt(\'\ on ,h<" totnin.r
-4 COS'<OM •• ,. 1989, Finni ~~Wt,,, 1/:,· R,rglu11t1I rr,·11am,«>111fr1M F.t/Jl'fl. lntf\i\ l'.flOU-" u~· M 1,nmct.tmldhun in Jh!.' ctmtrol u( ho\"inc
Regional hc~t·and Tryp~no:,0mln-.. li. ControJ Pro,nunmt-. ~1a.J~,\l, 1i,i,a11usom1»I• un the l(onr;1 Coa,r rroplMl ,\11/m11/ l/r11/1/11111d
'louunhlqu~. l'.iUTib!n and l'.unb.,b,w l)ecember 19139. FGU,l\ronberi: Pr0tlu,1iq11. 2!. ~ 10.
c-,a,ulting<1nd f.ngmecrtng Gml>!I, kon,i;,1<1n. 1y.,_,, (;enn~ny 6J rwnt.L 1.1 .• 19-;;•.\nti~C!nit" v~ui;ui4>n tn .,r1lh:.u!.1n rrn>;rno~m1.-s. /u: s11so.
.a; co~ ,oi:c. k.t. ~ ttM UWHL It\,, _1g8;. 8o\1neo ll)ll:tm,11.umia~i~ in-.oi.nhc•rn J.• ,uut:R. t i. \JC.:.ltll ,·n. f • . (l--d.<') JJIIXXl•IHJrtt,t l'nrr~it/c Vlw11.,·1.•s. °\;l"\\ 1

lonr.anla: P.irJ,hulogl,:-:,1 •nd ,croloiirol Wl'H'Y bf pre, al,nct•, I rop/nil York! Plt1:1um Pr~s.
,\11111111/ l/ea//11 nm/l'rod11,tion, 1,. l~'\-172. Gs nu 1<n1. R.. 19:;..1 Tl)p3nOWmtuib In 7.u1u!aru.l a.11d tb.Lo control 0(1~\:~
.s6 c:os~Olt fl..J. &> \.lUKA~Ot. 0.1 A... 198&, Ccinamcm outbrt"a:kof hy chun1kal rnt.oans.. OudN'lt,!ptl(Jfl J()u11111/ uf \ 1.11,•ruu,n• Hvset11d1. :!ii..
p,;oudo-lumpy )'l..;n db,oas,o nnd ncutc Tr:.·1muw1.Juill 1-i,1tL1. infection in 317-38:"
cattle. Troplcnl ,\11/mn/ Hr,1/1/11111d Pm<t11nI11t1. 18, 12i-l :12. 6'i O\\'IN(;, R. JUf., l I 1nono, 0,1 .. A(;\'f.M,\~C• .:., Grm~1n·. G., C.AIC\'t·. ,\,5,
.a::- COXXOR, 1\.1,, )SU~\.SOJ. O,J,.\.~ KW lWlJ l n.,, .. t91:l9, Sotfoe k.OM. ~.. k 101.,,c:;.- 'MA .. l993, U1rnp11n,11,'\: .\tudit"!> cm N'Oimw ;md 1c.tbu
ttr'J)imo~miasis in .sou,hc.m f;i.nzn.nan: ln,es1igmWn into :hi: 1nc-ldc.-nc<· cautu fot1ow11,i r("r1c.t11:d iurt~11011,; wich Tr:,'P(mh.wmw ,vm1,.~lcm,t'.
or infecdon w,d dumti<>n of cht•1nopmphyla.'d" lwp,t"f A11tmnl llr"11h f<t~,('ll.fdl itl \"1°-lt'tJIUU)•Sc:{1•11tc•, S2:. :!92-298
nntl Prat11«tlOtt. 21. 1~1-10. 67 U\\'J:".filU, H,U. \(OH \\,\"l;Cj, "K,1 ~.'<:CJ\\', \\,I ,. I\AWU~(-~. P,o l.O'l URI:., 1• ,ft U,\JI,

18 COftft..,, J,J,F•.\1,, 1 fm IIUlJft.'\f.A..\.11.M- )IOOltllOU~I I l',IJ..!,. ~ U>. fC(.)()IJ, ~.C., M,L. m9;;. t•roduct1\.'i1yo1 ll)l);,JnOH)lcr;,nt 1,1111ic kept \HHh:r 1radllionnl
1968~ Prcv,1.t,mcr of 1r)·p;m0101m~ u, cn1de in Sou1h,wc ..1 /~mb1;i, mnn3~tml'l\t ,andh1on_, m 1h~ G~1mb,;,, \',•ttrlm11y(lw,m,t1,·. 16,
Tropical .•rr1111a/ ll('tlltl1111:tl PM/union, zo, 71Hl·I, 81··86,
49 OU)FT, \..L, tAbT; J.', &'MOf.'l"~J-U,O. D.H. l?)8:l. 1\mi•tf)1>:,utoll.t)n'mJ filt"lut ht oft D\\'IM'ar'A. ft.It. G.ROOTn:run11.. ,.().. \1UAk,,·. :\}., \IOI no,,,.,.. tr, (jf! n l:\nv~ , •••
th• h°'•molymph MOlo<J/1111, Arlfl l'ttlplc,r. 39. 293-302. 19Ut; ,ll!-ccp1Unthr of buffalovs. rrmk ,vid ~oat) 10 111(«11011 wiLh
50 ctiN:-.tSGlt.\'.\.I, M.v.. 1966.. ln\murtii'}' In hu\.itt(• 1ryp:inU"tnnfu...i" f.".p: dUturcm :m,c:k~ or Tf',,'JKlntJ,{Omn l'l('{l.\' 1r3r,-;:,niitc.·d by CIO.Ullf(J
.;frlam .\Mli'ro/ Jwrnal, -l:l, J94-397 mo1'111an1 l'1'1U1Ults.. &w.wrrlt ,,, \ ·e1,...rrnary::,cuu1,·~. ·11, J07-ll5.
~ OJNXIXQUA)t. \f,J' & \'A,:'\ JIOJ\'J, ~ .. t'JC1S Di:iRnOs~ or trypanosun111L~h, In 6'J O\'o'IMifl't, Jt.H, 1.UCJ,;tX:,, .\.(. , N\Jmt\.\" \f., RM P. & \JOlOO, ",t,;,1 1.98(,..
(.aafo. In: Jnic-rnalion«l Sc:Hmtltk Comnuth..,• fo 1 TryJ1,1nth,Dmi.l)i!'> lnwrkrcnec t>c:,,,l!en diffc,cnt ')troUcmc.~ of Tl"Jp,11,010""" t<>ngvfcms.· in
Re~an:h. Pt~;,.ffli11,:s11/th,• ·r,,ntlt .Hcf'lltt,:. 1,,:amp11fn 196/. lh.·riJord. th~· 0-5tobl1.,hnwm 01 "\IIH.!fln(ectum, m g,u.11, fol!ow1ng cydical
EnjlJnnd: Srephcrl ,\1J<Un &-Snn, l.td cran;rn1>'>lOn hy ,~·t.M,. f><traslri• lmmm1olog,·, 8. ?~;J..305,
$2. l'f ,\\\l(;O, I ., f'QIJS!1,IN'(1,\, J, :-.I i l )\,\5SOX, .\ 4li: t:W!'i,"O' n~l$-9:i- Pr.mqucs 70 U\\'1!\(~fl'M. N.lf ~ tU\,).J,-;o.. \.t,, ~IURR.n. \f A\t P , "- )101.00, ,;;;,J;,,. tgiS,g.
pa.stQral~ ~lboror c1 crypa11osomo)CS bo\1rws dons un ... ..co1w de lmerfcrtm:e in lhl' c-,.r"hh:,i.hmt-m ul hc1,1.·•mm;m1m.~ TrJ1xm~mm1
sa,;:nes l:umt<k:o de n.,nu;1Jriqui•. Rt1·wtd'El-t,~'!... r1 ,~ .\/rd,'t'inc rcm1,'0i,·nR, r iuut,•, or r 1'""'.A supt'nnrecuun.., m t:<>'-'l.' aln:~ld)·
:'-ltirir:air~d& P",\'f uopi;..'Oux. i8. 203-212 an!t."\:tcd ,-.,th 1~ co,11,0/~Ut.· nr r ,,11vM· hw·muuy Paras..110/0~·· :tlJ,
177 ·18~l.
S3 l)AU. c .. \\"'Et.BOK.,. ~.c.. MAUOUX, ,. '- \UUIG,x. ff,J,M .. 199S, Tht kini:-tlb-
of ma,urution ot nyp.1noSQme lntet11on11 ln LWhc. Pflmfirotr,g.·. 111. 7l nwr~c.ul.. n. u ,1uRR"·· ,, "1'11010<1. ),;.., 19gv. Pe&rn.,1:c •ancrl~ and
187--191 c\·llulor n.<t.,pon!'4.~ 111 J;o,ns mfeclcd u,1J ,11p,:nnfect<'d \\1th
s..i D.\Jttm: 1.n.. t982. ·Jni: lnfluL-nc~of g~n.:tlc:: fr.-ccor'" ,,n the tc-,L'l.ta1nct• of Tl)'PfllltHW>Ul rom:o!,•1r-wual)-s.m1tt.r:d by Glqt-stm1 mors.Jw,,., ,·,•111rnli1.
rumlnam,. ro gasttoinu:1rtir.al n~rnJtade .md lt)'P.UHhOmc lnt«t(on> .¾ta Iram<<• a;. 21--13
In· O\\i-.\, o.t•,\ (ed.) J\nlrnnl \t«lr/1 w l'urtJ.#tol~· London· '\1:,cmill.an. n o,,,:\(;at.. R.H.; Punrs. ,, .. \10toa., J.. ~ \\URJII.\\, ,1 .. 1988, t,c\·clqpmcm
~ 0.,\1~ox. c .. lgg('I. R\.-gfnntll T~ear 3Jtd Tr)-"?3.Uo<\Omta~~ Control ol Tr:.--f)tmQsomtt eoogo!N,.w~. ·r mNJ.\· t1nd r liru,c.•1 in the- ~kin n.:,~cuuo
Programme. Republkol ~l.'lla·,,i :\>nona1 Twtw on<ITrypnno<om,~,;• 1r.:i.luc1..-d in ~oQto.. by in(.,•ch."lf filmJHm m,,,_.fitrm., i"<"fllmlr~ ll<~m,h in
Sur,.ey Jgg,-1'1119. fon•I Rcp<>rt Cir.'< Ctm,ull Gml>I I l'lu•wluurf. t•e1mna~· ~-,r11re. ·l 1 15-' 16:.\
Fedoc.d n.,,ubll, aH,L'fffl.ln)", 73 tl~UiJt. ~,.c ,\ROWQU). n.n ..\,. ~HUI l L..\., \10l()('), !l>,1'.. IU)l.'\ll,;o,, ,. n. fl

56 Df.:tiOU~. J.P•• nu.w. l-t. !it suLot,r.x ..\. 1996. O:htn..ulon d"uno tl.lm~t" de M Rtr.RJ:,O.F "·"'-· 19-'J.1. honwramtd1um con<'l"ntnu1on~ ln 1hc s:er,1 of

lrfp?.nn~omosc •'quinc la T1:·ptutl'J1om:t a•iirtt en :t.onc: uthJm~ .iu Uoro11 \-.ante; ttmtJatm:i wllh rnophyiot.~ against t~l~c·1ran~m1111.-d
Sent'gol. TropittJ//1110, 14 35-26 h:r11a110$nma co,1J,.'<1l1•11~·..~rlil Tmpu·ll. 56, ,\q..3()

Si l'll:SQU.E:>S-ts. M.. 2997. ·1:valuat!on of:,; simple Pen tcchmqu~ tor the 7.; t l~U'IL ,1,( Ul.lQTI' t' -r-., I, JIOI \JI.,, 1•.11,. 1•)!)6. \ ,fmpft.•CQJllpt.'rith-c
diagnosis of Tl)'JHWOJoma , 1tia.r infection in lhe (l>naffl of cnulc in
1 enT,)'TllC ,mmuno:h:~'> for tht· detec-uon ot Uw tryp.1noc,dal drug
c-o.nparl.wn tea t>1:tra1o:1tologit&J a.-chniq,uf~ and nnti~e-n-t-ntymt.•· l'>'Htll't.1midium. nu•t(IJJ4'11ft1' ()mg .\toniror111R, iS, i'3-79.
l(nkcd,tmmunosorbt,ntob~.a~·. .Actn Trr,pftn. 63.139-148. 199i, 75 t l<s.lt'II. M,f.. (o\1HT, t .\,, '\111 It, Ii,\., J11'HH,Rl'.\I ,. .\.,.),. • Jt01,.\tll."'+ J",H 1-,,'3.
s& ou~. fl,. 01Au... o.. cun:;-. :-.. K.\t;;tRUi,;.\. r .. u::.,,~mutrn, , sru,n(r , , _ F.v.iJuatlr>n ,md impr0\'1,.'mcnt "' ;in cn~mc--llnkc:d lmmtino.,orbcm
aSL£.R. M..C.. s, 1101.r,.1u..11. isss. f1t"fd C'Valu:1unn or 1hc, pn:,phy~ctt,· ~"!-"l.l}' for chc l!NC'rtlon ur twmi."l.tm1dlum tn bo\.1n~ 1J('f'Um / l:v,n11t•r1tu·
effect c,f an Jromctamid1um su:st"med-rcl~~ dm1u~ :;gum~, n11,g .\/onlumng,. 15. !?:t~z.;2.
l~-panosomia..~,,,. in ~'1ulc. Antimtrroblal As.~nrtand Clu·mmlt,rt1('1)', '12, 7t) fl.Ct, at,, ag,;,, lht!' ir..1ni.mh,,fon of 1rypr,nn1r,111(f ronw1ln1S.'>thmugh
i01~101-1. fifMsiltlf mowtim1 omd the white mau.,.«:>. TflmwrriC11$nfrlrt" H,;_w1f
,&9 DOI.A,,, R,B., Oto:ECll, G., .\l,Us.HUL\, IL '-WT\J(a, M,.~U\.'l.~M),, P., >AHK, S«ldy,Jj tmpu<tl M\"dittn,·pnd lf.v-gt,m<. t,r,. 219.
P.O.... sJor..u. ,.H .. 1990. Ho:mdhtm bromide as a t'lnmmpmphylar.uc (or
n II :o,t ~. "·· "''1QUDI, :\1 •.\, &- tJt'.tJIRGO, \1., 19~0 l'rem,ertdc('(HJ\'Clt«!d~
cnuJe U)pat.1oson1ja.<is m t<:n,:·~ ·\r1n rn1pirtr,•l'i, lli-1-H dcu.>. <"Pi"<r:,. tit Cilfli.\:hU'.)c l!n Arabi~ Sol.<.tuJihr. rroh•il'mt Col'lft1rt•nc-e
60 OOIA~. fUl.. !l-,\\-UU, P,1)., AUJ\IIUL\. tl t, UL\1 H, 11.tt.. 1~U8. ~T,:thn:,Jd tnwrn.s\lr,11.il..- d~ fJllomolngi~tc~(J'rl\pr~ion hi,Jll~nJ,..c:. ,\.fcthQdC'"'i et

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290 s•r.no,nwo: Protozoa! diseases

r~,;ponsabilhcs d•• En1om9log!,1c,. d'o11jQurd'hul. G~mblou,. 9--14 99 ('IR."' •.\..n.,. 1w.rutR1"1 r..t .. 1r,1. rhc cyclf~I ;r.in:mif!<-..-lt,n Q( ~,r.iln, af
lulllet 1990. Tr,p,t1.nosomnccngol&ise('.nd T 1•f1•0.a-t<·.slstant m nomml lhempeudt
;a l'he appi!anut(f' or T. Vfrdx In
[\U'.Jt\', O,t-, IMHhl, J.D., \fOU)0, 1,,.1': .. l980,
dose, ohrypann<idal drug,. Pa1a,/1Q/Of;!', 63, bi'-89.
lymph (ollo\vlng chnllcng~ or go.m with lnlcctcd Glo>Jil:n mors11n11, 100 ""'"· A,R. • R<'IOIRT'i, c.1 .. 1971. rho $tabllltyof rc,si,tnncc 10 dlm!n.,r.cno
morsl11111s. Ar111 Trop/cr1 37. 3,3-37~. :rceturate- and c.1uln~p)'rnmine fn a ..mun of Tl)'fJ'mQ.~ma dt't2.t. during
:"9 ~'-»Jn·. o.L. a1 ,10UJO, s..x., 1980. The <cqucntial ct'llular cllang(~ ln dw cydh:·•I tm:1>Jnl<,,ion 1hr<)ugh anwlup,. Jlt1,mfm1ug)'. 63, 163-16'.l.
local ..,,kin r~ac:tion producit?-d in .,;oali by C(oJsina morslums 11wrsltcuu 101 r;nu~at.M .. >.:UMAR. ~,. ,c k'l'l:S\\'A, t .. t~g;. f,):alunrton .,nd comp~t'Uon or
infected "itlt Tryp/11:()S()ma I f"'!'/H11t1rJ>0111 /,rur,•l.kt<t Tmpkn. J7, ttnalbodyf:l$~s for w1>df,gno,r, ofb<,,1ne uypnnn;omu,!,. \·,urina,;·
137-149. Paras/10/ogy. 73. 197-205.
so u'SWN, 11.. 19'11· T11il Or/J/111 of1/f~ 01)m,ule ,111/muli of,Vrfcti. \"ols.1 &ll. 102 Gtu.Ftl~. L. 1978..J\trrr:nn rrypnno~,mrr,~;.,, in ~hL<cp and (,::03lS A rtvi~.,.·.
~cw York: Mric:1110 Publlshlnjl Corpor31ioi>. 1}1~ ,:f':tr/lUU')' nu.1/tUin, 42. 819-8~~-

81 l!VA"''"" 1680. Rcp<>rt on ·,um,·

dist<»C In tho t)ora brruul 1-.'hnn d1s1r!ot. 103 C.R.tr11s. L. &~UJ.osur. u.w•• 1919. Tf)-p:mcuolC'rJ.nt'f.: fn br\.'i..-d., or sheep
Pun1ab f"'ro,·crnmeni \1Ritar) O~panno.~m. ~o. 493-H&i and ,ctoa~ \..i1h an uxpcrimcn~J m.!CC1ion of 1'ryp<mowmn 1."'01'10(;:,:tt.
a, "'~""'· A,l<.T.•W.. 1570, l'olhngcnc<l~ ond palholo~· of anlm,\I \',•fitinmy P,uasil<>fJ>ll)'- ,;. 97-10:;.
~pan1»omlasis. /11: '1UWQ 11.w." ,s. =·
w.u.. led$). {he ,lfr/cn11 10~ CRIFFl~, l., .\UON6Y, f..\\·.-.. fl'~f,~'\, f.)lf,, 1981. 1'he inlcrnc:t!on (>(
T')7111no,omln.,e.;. t,,ndoo; Georg~ Allen ai,d Unwlnl~llnl$1f}' of TryponQ$oma "mgolrm~aod H(Jfln;qnclwJ ro11101Lu $ fnfecdons in t,,o
°'"''"'"" Devel<>prncnt. bret."d:- or 1,.-0:ii. T, Pam,ho1ogy. Jaum(l/ o[C,.,mµomric'C (>(11l:r1/0i::)•. 91
13 ta?3. Chimioth\!mpttH~t 1.·hhTiiopt~\Cntiun de
,..1;,;aJ.f, ,,., 85-~S.
n:ypan0$0minSe-3nimal~- Atqui!.ltion.s r~ntCl':11 Cl ihundon actuclJo. I.Lt( HJS c;ru.nL~. L, \\'AtUJY.t.,\, .,•• AUOMn', 1 .\, ... 1930. fhe lmmuno~uppressive
c,,111,~d• Mnlmn• \ ~,r,frmire. ,I?, 215-226. crfccQof cxpcmn&.tntal T. congolenSt: ln(t.>e:tlon.s lu g(l;us. V'«ttrrfnml
8,1 1-1.sos. 1 .. 1985- Vcrcrinary lnvest1g.:1rioJ\ Ccnuc. ~ltwurn. lan.t:inta. Pnmslwlol:)·, ;, 11-13-
Unpublish<'ll llntn. 100 C,KtXJn ,Hul~. ,.c;,. t)\\INOllff. -.u .. OOLA,'"· ft,1S .. MOt(K,), s.,,;. 6, MUKft;\\', :-.,..
&s rono. 1.. 19;1. The /IQ/~<1fllw Tr;,x,,,owmrast>S in 1\frlum &o/b~·. Qxro,d: 1990. ~u...,,p,ibflit)' or Afrfc,an bulfalo •nd 'R<Jron r:utl• to / ry•p,mowma
C.ren<!on Pres_<. congoldn~ tra®nhu:J hy Glos.d11ll mor,.i1amrcemtrnlfl'. \fl(c"~rh,nt)·
P(ltttsilQIUfrl, 35. 2 J9-~.1.
t6 nUo.:l.'CH, \1,11 .. 1935, -:!1rog,:n ond rnlncml 111ctJh11lbm In 'Tt;'/Hlttt;s,,mn
co11gofe1tA>dLseus;,. /,,,,1111w11/ R,porr oj'1he Dl!pnrrmem oj llwrl1111ry 10~ IL\t\U.\', J...\LL, l!JU6. fun.h er ,nu.ffcc;; on-~ge and trypanosontt in!t-ctlon
'><'lc11~ 1111d ,\11/nr(I/ H11JIJ,111dr;, T,mgm,ylkn, 1934 mtt> In G/o.<,/ttn /HI/I/di/ta Aust.• (;, JHlltx1/iJ.f11sr/1;ts NewM. ond G.
l,m·ipa/pi, 1'101-.1. in lJ~anda, IS:t1lr1in oj'E111omo/ogfcfll Hes..,1rrli. 57,
,- rn.t:l.'<.u. M.n.,. 1101<.~iv. 11.,.. 193G. Sn1dies conrcmlng the ell'l'<'s oft hr.
.159-47;
p!Jtnc ofnutritfon on the t1)ur« <n anlmnl rryp•n•lM>mi..,,1, JII. T.
<oni;t,follkdise;;!l(I from crlsi• to r0<0,~11·. l11:Amw<1/ /lep,,rt of<lle 1o8 t11:11.:u~os. \\',.193t. ·n,c reb.Hton,ohip ofpka lnc:aul~ to
0eJ)tlrtmrnt ofl'tttri.nnry Sdmct mut :\11/mnl H11$blmtlry Tart,&•tm,vfko. trypnno"'°miMis... r;,~ VrJNinniy /ounwl. 87, 518-523,
J93S. 10l) "'""· c.i,.r.•• 191!8. Th<lolfe«sof 11),ianosomill•ls propb~1:t.xis and
88 C41tt'.H~r:n. P.R., 1989. Rt?ccnr,tudies of1he biolO(O' of 1'f')1Jt1110.Jtwu1 ,,f,inx. 3.ntlh£'1mlntit' tr('atmcm Jn goa1s un<L.•r trlldidonat minngement in
Adt'011ces l11 Po!(,s/10/og:,. 2S. 219-317. somhcm Tnn:r..nnilL fr,: f.ilidlat:J: Pto(tutJifm m J"A!ut afftt1t1/ arvm oj
89 <.a,~1ns. ~~ .fl. UOL.\1t:~. r.H,. 1996. Drug ma.nttgcn1em1 and p.ir~~nc ..tcfrita. PtoMedillJ,tj u/a .\!rttlt1fl lt~!d !'3-27 ,\lm 1cmi>cr 198:'. NDtr.obi.
resiStancc in -nnimal 1t)'Pat101-omt~::- m Africa. PM r techn1c.l1 -:Jnd &11,m /l.C,VI/JIAD. Nlirobi: EngU,h P=~
SCie:ntiflc series. Jtom.e. 110 nt:.~~::.:o. M .w •• 19,p;ti. Anim.al Dit,vu~;. o, South,lftica. 3rd t:dn. Pr,.:tano:
90 GRR'TS, ~•• ~GllllUK.,. J'., 1>tIOEJ.:E~. tt., BMNOl, 1,lt,i\., )w\Z;Ut, J,)t., nt.\ftftA, Ct-ntraJ ~c~...-sAg<·n()' Ltd.
6 .. ESl..EJI, '.\1,C•• ~OlM;lrT, · - t) HOl\1F~. P,tf.. 1997. Proph}·l~ctk etrc,t~ c,f m 110.\RE. C'-\.. 19;~. 11re J'ry7,n110.tOmC$ of.Wamuu,J..f; I\ 7.()o/ogtcol
bom1:"1am1diurn-- :md t:1h1dium·)USt~1ted release <re'vi('(1:~ ti-.gainst .l/ni:ogra11/I, Oxford nnd F.tiinbtlrgh: Slnckwdl S<1entilic Pubhcattom;.
·1 ·,yp,mosnma t;Ol'J;(>ll!nS'! m canle••ilcco Trcpita. 6l-t z.3-31.
U2 HOt..\lb, fl.fl,, MAM'\fO, t., THU:\UO'.\, "-·• aii,n,tlT, 11..A .. I flt>,;f ,, K. "111MHA\'.
91 G1.11sos w.~-.
19$5. l\pplka1ion ofMw t~clmotogie!' 10 tp!den1folo~· P,K., '\NRR.\l\ '\I,. Jt..-,,..;r,t~ F,\\, j;- IJJl:QUIL\Jtt, C.,:.1,. 1$7,1,
Brlt#h .lletlle<1/ 8111/et/11 . .; l. 115-121. lmmurto~u,')j)f(!1'$JOII 111 bo,r1ne lJ1'J>0.110$0tnWis. Tire v~,,,rlnary R~rd,
92 cui:-0,. w .. 1g9.i tdentlficnrlon or tl)·panosomt~ l.n unhn:tls. hu.m3.tb and 9!>,g&..87.
G/O$SI1Ut. Bulle1/11 rlt• ln5"'/,t~de P111/10/ogie E.xatfqu,•. Ir.', 315-318. IIJ HOL\!H P,11 • SCOn', f.'I., 198:,. Chetnoth•rapy 3gnll\$1 :tnlmal
93 GlXGtUC.11, J,U., M.\C1'L.~, l..~1 .. JAO..':SON, P.fl~ & 1t0'6f,fft'S. 0,11., 1985 tr)'P•m=mlasl~. 1,c 6,,~•x.1.~. red.I P,rs,,«rl1v,i11 f?P<lll<W111lnsi;
r,,1,ano$0ma broct1: tnhance-mcn1 of infec~io1, t3.te,; m th~ L(t:lst Jly. RtEMfC'h. PTQ~fng.1; o/tJu.• Tu,~ur:, Fir,<l Tf')'P<Ul(l,wmlt.t#S5t!mlnm.
Gf(1,'f:,.i1w 11iot.sftansbyfeeding .irtificinf bloodmeal mlx(un:s. ,\mCri((m London .N September 1981, l.ctchwonh; R~om:h S1udi"'~ Pt.:».
/01mw1 of Ttopfral Mediral ff;'gi,·t1c. M. n-r.. ,l1\ JH)L\1f'~ 1•.u.. WHn'l'lA\\, n.o•• btLL t. )IOLDO, ........ tUKUMI, u .. ,.,u,ut.t.\',
9~ Gtl\1Dt., s•• 1989, Endocrine cff't:<:t)of 1ryp,mosomt~i.s: Rt:cent~tudl:t~ )1.li UftqtJlt\J\r-, o.M.. 1985. The ~1Ji.~qcf~1ton butwcun Sarnorin
D/$COJ~ry,m,t fmw1,•mitm, 1, 30,.33, cl10111oproph)·laxi> and Immune rn,pon,.,. ln ~attic under~rfmental
mct;!:t'}'Clfc Trypaiw.~omu .,v,n;;ol<um cl1nlle-ngt::. Th,• l'1rx't"l~ings CJ/rfo•
% r.o.""'· A.F.. 1990. O'A, SADCCworksbop on vector-1,Qmc di~ea,c>!
lmem<11fon<1l Scientfjk Council for Tryponosomiusfs R~,·arrh mr<l
Cou.ntl)· rq,on- .\J1gol• A pap<rprcsontcd t<J 1hc (..'T.\tSAl;)C:C
C;/11rrol. OAU :sntc. 18th \1ce1,ng. Hnrate, /jmbabwe 198.,. l'ublkotlon
workshop on \'CClOr·bome dbe=, Mb.i.baM, Swazlland. 30 lu.l)' l!t90-
3 August 1990.
'io. 113.
115 .HOP1'.I~~. f,!ot,,, CHl'rA\ftt(), U .. ,1:\~1111.,\, ~ .• t.UO.:tXS,"4,r. .. ML. f~F.. \';\~ Ol:~
96 uooow1N, 1.,<,... 1.970. The p.:ttholu~ uf AfriC"Jn tryp.;,no~rni;s~i.i,..
00..,....c,n·, 1• N 1:111.1 rit. M.c.. , 1998 ,\d3ptatlon nru:I \":.tlldatfon of the
·1~11.11,act/nr1s oj'tfle Hoyal ~oriei;· oj Trop/ml Mrdic/11d mu/ ff)-gir11v, 64.
ant11,ody ttopplng El IS,\ u~lng dri<'<l bluod ;poLS on r.lt<r papt'r, for
797-SH.
epldcmtoJogical ~un-ey.~ of tielse tr.1n~mlt1ed uyp.111. 0.somo~is fn cauu.~
9; CMY.A.11.. 1974, T')1Mnu,om1tl ant!1tc11s and th~ com:ol of Pret-enu,,~ Wrorlnnry .\1edlcitrt, 3';'. 91•99.
1rypanosomln,is. /11: C/J111ro/ Progrt1mt for 1'1)•1,n11ou1m,., and 1/irlr
116 HOR~m "·'-· 1!131. T11•panosom;is and tl)'Pa.nosoniia.,is of cunlc. /orminl
\•«1or.1.At1us du C.:olloquc Part,, 12-IS Morch 1974, ln<1ltu1 d'EJ,vage
01 de \•h!d~>cinc \1~11irin11lrt de, Pny, Troplcawc. rifv>mµ"rmit'f l'llrhQIOI:)', 34, 211-240,

98 C:-.R.,w.A..R. & .LUC.Kt~. ,\.CJ.. . 1$6. Antigenie v~rfatlon rn .i-olh-'.:lrhm 11; Ho1t,u, tu .. 195,3.,\nimnl Tl)'P(.Wruom!h.fis ftt Et.Mt Africa~ 19.19,
H"fJ)3.1\0Somes.. In: lU-"ilSOJl'...x, \,.'.H. R, & t,.'\·A~.:,. U.A.. rt."CB). B!OliJK)'Of1Ju:
u,n~on.11\\S<).
Kl11.:01>/1u1ltl,. Vol. 1. 1.ondon: Aeodom,c Pro<>.

www.ajlobby.com
 \fric:u, animal 11ypanosomose, 29 I

.\Jmr,n/ R1•1MJrt of r/Jr 0/1K1ttm1'1tl uf\·i-:rrirut'),' Sti~11u c1,ul :\nfmn/ l4l .:..\,Y.\111, 11 \\ , , ., 4l10'\:ll\. f-,\\' , WU ">0\, ,\.1, A- '1U,\'U \.. \,'•..:.,. t~i). S.0fflf!
Hu,1-.,mdr,·. ToJJ[lOll)'lk<t 193/. ,•C'()rtoinlC''l !t't1et... 01 rryp:mu."rllmb,i, jn ~r1.1l~. rm11/n1/ Auimal 11,"ttltJt
1

us llQt.\}(1.).1, C~\..ft.. lSCR..\~I. (';.,\, & \IOt\~l.UX. u .11 .• 198';. Hnem,1~~futhHJ\1t 11,td Prc,(111rllofl, 1S, !5.1-100.
~rtd parct.sltc aAA1,uti111ns In h:itmo:ymplt 0,11d gut ol GloJ.ti,:a. fro11/(1'1 M:t Ul'l•\H JU, S.... ,t.Vlt l. L\I, & Jt\(,!\'.\U , Jll., 1g98. RC\'ll"\\' of t!\N"t! 3J1d
.\ft'dfrJttr nnd Ptttlt.dto!"~ , 3:;, 1 ~I • 156. 1ryp.ino~mh'\i1r. it1 snu1h Afrk~. <Jn1/1•Tl:<•1H>t,n Jour,w/ of\ 'r1t1rlnflf)'
Flt"St"drt'lt. fi.5. 195•?03.
120 11(Wt.. ..n .. 1979. Cenhal I~sion> m t:~ptnmt'm-:sJ d1tomc TryPbJll)J(},,u:
brorti infci;non in rnms. R£'$¥1Jrtii m \',•trrit:111)' &it11c,• 26. 1.;s...1,.;1 H3 ~,u,,;-uR.,. h 1.US1'-<3\.. c., cmT:\\~fflJ. H. & lR..UH. t . l997, O,m.oeuon or

tZl tl>.'l:Df. a .o . 11 t{.~~\~. f ti". ., ,,1 .\\'TE. \,0,. H}d8. th!pm,hJLll\l' dl"-t1Cth:"" 1n
Tr,'f)(mtJ:.:onm i:.ort1tol,·n,-:,• «n\l T lm,c<r. suMpec1~ In r:.mJt, in .l:tmb1i1 by
.Afri(o.n trypano~umrn-.is:,\ h:\1e.., ...trrn TrupKt1, ,tS. i-tO. pc>lynu,·r.tk' cho\n retietton rrum blood t(lflc"('tcd <>n., llhcr pap~, .
Para!Jloloit.,· Hi."$4..wult. 83. 2,, 1.. 2,10.
C?> 1i;mr, n.o . ., w,o,, <..r.. 19;-:. Pa:holo1,') of 1he di>ea>e in ,h11ep pmclu<i,d
i-l l Ml 11,(.ut:..\•f\1, UJ~H \\'.\, L,., il)g&., J'btt pte'\tlJ~JICtJ of lr)'fJ;&IIO"-Q01U,i~ 1n
eq,enmtntn.U~· b~· TryJldm>l<Jnr.a lmu 1.,, V"11•rlm,r,• PmiJuftnv·, <J, 2i&-
:?89. '\lflu.U rummQnb and 511g,,-. ma 'i.lu~ping ~cknffl endomif.." aren or Bulkwr
Couruy \1Ui.:nnC\ dtsinrc. Uganda. H,·1·1Jttf 1:.'lt•:.r,g,·('l Jr \l&lrt:mr
l:.?'3 ILCA. ,m SmnlJ Humin(IIU Pf(klittlltm in 1he Humid Tropfcs. ll.C4 1'tlhint1irl' ((CS />11_;'1 lrOJ)i4'tlttX. ~9 Sf>.58.
~)~t,nu ,wdy !I. Addl,Abab;l: hnem.,uc.mal Livt:-.,lC)l;i. t('nH\' fr,r \frk.1
M5 ~\TU,(~m;.\·R\\ .U.bH \\. \, L, PATI)t:l~!l, t .J,, HSll\\10:, c;.,, MURR.\\',~( •• 6
1~ H RAI>. 198.3. II.RAO. l~a:? . .-lrzmull Ft~·por, <>I lltr ltJt1)flllll/rnu1t l.t1i1(Jf'(lt/U')' no1.\ff'!'o. P.n .. tm, llw ,onucncc af c-nt'I) mmkcon chC'
for R.-irth nu '\11111111/ r>f.kv:w \.tl1~b!: ILIIAll f1:llhoph-y~10JOJ,.''y or fryptm0~/1:tl (Qtt}t.'fJ/;•flSc" :n.ft"<.'tion m 5'."0ltt>h
1~1, n,.RAO, 198~. II.RAnt988: Ann110J Rtport of thtt lrt1t•nw.1lcmid l altfJra.tOf)'fo, Ulnrl:fat:"' \;it«p. \ ·nt·rtm,o· Pt1n13m1lo~.·. 59. 20;- 21s.
l1£tS£'OrtJi 011A11/11u.1l DfSl'D!,ttS. \!':tirobl. ltR:\O. 1.16 t,;.\Tt.\10£ l,\t ~U~l,;l.., .\.J,. '"'TIJl,H. , ,,,,. h ,.01mu:Rt~. tU,1 .• 198:- A
l26 l}IBUGA. ,1.0.. OSIR, c..o. ,e. I..\UOSM, \'.I... IY:92. lnMbflOf)' l'lftcl of nt"'i\ rm•thud for h'<3tltm rt1\d J>tC'-..ct1'·utfun Qf lr)lMnUMtni..il ,inltgcn,
1'()7NJIIO!:Oma brncirl brutd on Gfo.W,u, mn,;,/um.'{ l'ttid~t 1n·p~fn 111 for u,c in 1hc mdirci'.t hnmunollQUrbc-tn«' antlbod}· t~1 for di.i.gnoii~
,,uro. Poro1iwln,:y Ha.f<Ntrrlt. i8. 2i3-2Tb. of bovine 1ryp.111owmo>1~ rmpJcal Mtdrnnt· and Pnmsltoln1r1. :ttt
ll-N,
1:!7 IM,DUOA, M,O., OSIR, (..()., WU'>NOO. \ ·.1 .., O\lOI, ~ f,,()'flltS'O, t..H., t9~:..
Studitt on lSCt5,C mfd1,.."Ut rat10r-s th;u tndutc dHTcr(,1\tu1tlon of 111 t,;:t t.UY.,. -. •,.ftuu1:w,N,. n .. 1,tt3 lmpra\•t·d fide.I dl.:1gnc,stfr cuchnlquc-
bloodsue;,un Tryf)tmo.,·oma btuetl br,m,i iu ,,Jlm. Purn.•ftnlnJn Rtr$tYm:II. r.,, tf)-p.ano~omla~ts. b~· U"il! or a mtnic~rnrffuttl!', f'h~ Veter1,1,.1,y Umnd.
;s. 10-1s. Jll.219,

l28 J.\CKS.ON. P.ll. ic: omus.., C.L, t")H3 Ttj·pa>:tHanm rJi(){/i•siel'lH blood.~trl!'um 1.;8 ~1u1r.~.1,.1~nmr~. tt, 1~168, n,c dln~\t>"'i' oftni>~m,•t(HnlMh in
trypomnstiJ!tH~ ."Ind crutmra procycJk rcn S:irfac~ carbt>h~dr.ites. llve~tQd;. Afl'P\1('1.\' of currc:m l\e-chni,1uc."· l1t,• 1'.-,"rJ,mry lJul/ynu, ~e.
/c>:tr11t1/ ofPtrllQZOOl()/{f. 30. 6;;:?•668. 191-19;

l.cc1in nnat~~i'- of
1:$ (AC"'50:'<.. P,N.. IIO~l(,'IU1HC. IJ,\f. s, IIOtr 'S.C. 19;11:.
r U) >:1 ~.\tlo. LD.B. lq.')1'. Ph:atmarolog)' o( cxi!ltint drug, for unittml
T1)'Jl(ll1Q.i0mfl ro11go/4r11Jtl'bf1>od,trt.am rr·yp~rnn.">1ig,,te and c:ultun~
uypnno~ontlnr;.h.. ,ttm TmJ>fea. 5-L t&·:ii-18..'l.
y:OC>'ciic ~11rfucc sa,c~rid~ ~· •tAA!UtlnJ.1inn nncl cll"Ctron rnlC'n>sc.Op1c 1an 1(1,.,,m,. o,ts.'" ff()(;.,~. 1,1'.• t988 n~ pl\-,nru.u:olo~\· u! h.\)mcmmidhrm.
{tchniqucs_ Joumal n/PttJlt.1:ofJI~·. 25• .;71 ·181 ./oumal,~f\ t.•terJ,u,ry l'lutrmt,"CJlbtJ•mul Tlrt•fflJ~Wla. 11. 23J..2·15.
i:}U JUf'llJU . o .• uw Alf.I I . \J.P. -. \lOI 't''U:llX. n.u.• \937, (:iban.tl in(c,;Ucm~ i)f 1_~1 >,.~tl\\l,t~._• G., \lfl Ul, f;. +, 1'U..\<;t-;. \,J., H189 fll'I«UQll o( parn~lc pt:pt11;l:t,t"'
TTJ.'/J(lllOflJnw l'Wtl.( .>nd 'f. rqngo/,-,,s, rn (.ilo1s1trt1. Pt1tfl.ffl<JfogJ· R,~~nrth. in 1h~ pJa,m.J of hcircr,,. 11,f«1«t \\ ith 1,,·1,a11Q..Umm (t)UgQkn~
:3. 28~2!fZ.. \rf>li!C'llftlrtmd 8(Q(lt~W(lll PllfO.Sl/O(t>g.·. l·l , 25-3·1

131 Jt:\t\L. .,. a.·uu,m•JOS'f$, ,t.r ., 19,;13 .\),G<1i3r1on of t~IS\.' 1.ur'ltr<•t \\ith 1G'2 li.OMOl:'<,.·O~\, C • TRl.TC f". Kf.NO+\f.\. 7. FOk\1l'-T'I . J> L>U\',\ll.11.f>..
h~1h and prod11c1M1y o/ tu«lc m 1hc Oid= \'alloy, We$1em E1hiop10. t.AU,;f:\.1f, 1 k.\\nl, l,f\,, ·r,rPo, .\.r.. " Ul<>RH.\:\, .... , 1t1,£H l:wdc d~· In
l',e1•iJm1t.-e \ t11.m'mur .\1rdftmr 22 ~O. prr:vi1lem.:t< dN inf&uu11>:. 1r;'J).1no~mc.s chez dl.tiercm ~p«es
132 Jl!:\:\.f. L., :\lA!fU• .:,,, '\"0fWU1.U.. .• 8~11.\JIJ, £•,,. U P\C.\'., ft.\\'.f., \\W.:,
d 'anim:iux <JLU\\1g1...., du p:uc nuunn:d d,• In Comot tn Cb,~ d'l\'OIN
I,;\:., TAIT.,\,, l",\l~ll--\VQ1~f. p,, PAY>, L~ sm,'TRT. )I., 1986, Hybmt
rtsuha~ prflim.mwr~ f.-u1 bl comp.an.son dt.' &n1J:,. m...thodcs dl'
di;;i.gnn,.tfc, R(fl'lh' tl'Eln.11fJ!,t..,t1l ti,• .\Nlittm, Vi.•t,~rinoftrd,,s P,~y1
forma!;on bet\\ cen Aftic"..a.n tr:, pa.nusmtw\ durin~ cyd:00 tta.AAm~,>lon.
ltOJUCtltlJ.. li.18.q...19; .
.\'owr.-.322. 1-3,.-t,5,
153 ,.1tA1 n M, lf.t>. ~ o•ma>;~ t-.u .. t9\19, Thr burty coat doub!!! t't'Jttriiug-,mnn
133 rl::\,,. L., M01 \·;,,.tux. n.u., U\'l.:,.·n . 1.t..,.,. r. \I.us. R•• 1ufJo. feeding
u:chnique. ~\n 1mpron-d mc:lhod f,u dw dln-gn0i1,: <H Afric"n
be:ha\iour or tsetse me\ intcctcd ...,.;,h )tlivnnan tt)']>a.11<.•,om"", \'muN.
u11":iotism11MI'> n,e /7(JCi!tdlngsoJ tlit lmtnumunnJ ~J,mu/11: 011111<il
?113. 333-335,
for TryP,l/U>M/1/10.\1.1 R<'.<1'11rth find CQt1trn/, Q,\U I ISCTRC. 20th ~h..,uni:,
IS,: 10,f,·O•"•·'-" ,J,. 1966. Olmlru.t.111~ 4Ct.'IUrall! nnd hu111ldlum chlorlda \lomb;m.>. !,,em.1 l93'1, l'ubll<J11011 :\o. 11 ~.
reslstant'e In helM! fiy,1r11n .. mim."d uypa.nosomt.~ or cta.t.llc (n north<'m
is, L\Nll'-'l, "'· v r. f".ortnu,. o,c. 197u. The tt.-Ohttion of ~alh·.'lnnn
~1gt:rio, T11r \'rterln<1Q' llt•,-ord, 8J, ,,3l-t.37
lt)-pano'°mc:> from m,u, and mher inummah ui,:lng OhAE·cdlufow.
t3$, JQt1UA..,, .\.,,. 19:-6, TscL,t• Jllt"' 4\ v,·e1or$ Qf ;ryp.ano\Ome, \ 'it<rinmy f:XJ)fnmr-m,:l Jtam1uo/<>1,..'I'. :!8. )21-;J.,$ .
PUMJ'irll!~•, i. t.;3-15'?.
15~ l.\\fk'\.~. \,,. Mt~'-U., • .• I ~ (r:,7X't:C,fQfl:•~s ,md 'fry7HIII0$0fflllt{t~.
136 Jt')MO\:\:. "- ,r.• s986, Tryp,m<Ju,mfotO C.:<muo/ mzd Af11<m1 /{11r,1I f.n__glbh 1:-;m,-fo110n nnd enlar}i.t-menr :'\abarro. t) .. lCJO';" l.uodno
O.j1 il(1pmcmt. l.ondon and ~cw Yot'lc! Ltingmiln 8.)111 rrc, T111J3ll & Cc,,
1T, 1osuu.,. IL'- k ~\TJ, Y.'ll-., 1()8.;~ The «.ut~ qf fat.al Try?1'4ttOYJmn iimfllt' 15f'i r,,,,IU-\:(.f:, f.\ ... HXt<.n~. C. \I ... S"Off\11\L ft.\ I 1980. '11,r t:lTt"t1 U( \\,lf un
ln1t>ction In dmne'itlt- ,hc-.,p, Mm Tt<>p/CII, ,I 9l-!12. ,h~ romrol nf d1,<•'>·("' of Uu.-..tod: in Rh<u!t"°Mo ti'1mb.1l·Mel 11J..
l38 fO:-JIUA. JI.A•• onwm. .o. ,q .. 11\\ .t..S(:":tll
o . ._ '\\,\XO Et.WU~ l..i J99~ \ 'rt('Tlnnr.- R1·~·ord. 10;, 82-SS
R~t.Munce to dJmim1.wnc a<'~Utbr Tryf"J'10somt1 cons,:ofwue" from asr LL\CJI, t .,t k Rcun,Rr~. c ., .. 1qs,. P,,..._,,m1,:n1u\ nr ,·hr.mo1h1•rJp:,• :md
attth: in tht- Zn.mbO/J VaH1..1· uf7.in1~ab\,1('. v,,,nt,uu,• J>mu.wtrnl<J.B)'. 6U, ch,\mopruphyl.nh of .anirmi! trn>a1na ~omrll,.i!, m thr ca-,tcm
1-6. hcmt!\pht:~- PlmrmllN>lt>gJ·nml lhttmJ)l•rwN. l!i tl1-M7
139 ~\A. (,.P. &> OJ>UOH•OJ\.UO, !L l~)lio. The.- cffo(t, nf Fl)~IICJ$0UW 1(.8 u .,u.., ,c*..A. ,. RU\\1AW,, c;.J., 1%7- fhlt d\·namir-,; ol 1tyil:mownu:
tong,,ltn.utinrcetlon on I.he 1estis .md epidid) mi;,. of the ~o..it. Bulhm11 of lnfecllcm, In n.1111111 pupula1ln11 of,..,,,... l>lp!Otn. Clomnldao'.,.iudled
,\11/mal Health and ProduttlM In ,lfr/ea, 28. 1-5, u:.ing ,-.in~·ftJy .1ud uv.1rinn .t,;clng tcx.·hn1qu1., 8ullr1/u of
1:o li:A.).!t.S5l.'.V. n.. 1990. In ,·1tto ttchtuquesrarassessnrnnt of drug rc..(£smncc Emum.nlORrcnl llc•wan·h, IJJ, 27l·21l2
in u:ypnnosomt..~. AgBlot«lt X,1c1,w1/ lu/ommrfou. 2.. 203-2 lO. 15q u.u L\Ml. P.• 19is. Thl" pre1.k1min:mee ot r,,pmu,1.0,;ur vli·a.t: infc.:tion<

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292 <>:crID~ "''" Pro101.onl disc:1si,s

01 cattle ;u ~ d iMDnc;c from ~a,,annah tsetse ('Otu;emnuion. TroJnt't1/

.r\nimtil Ht.:altll mul P1od1,trt01', 7. 201-?CJ.l. 1';'1) '.\1\(,f):\.\, fN,, ~.\f \1\.\)01, \\,..OIA.110•~1lni,.\Xl. \\'., ._l~YJ-'.\'Ot-. J., V,4
160 t.fl\J), A,k, fl TIIOMS(),. ,,\\ 197.; Ob$tT\""1JlltU\~011 nn iW:nl"tamldlum ,,·M UMl·'.\'C.O, r.. t!Ul7, lfllemu,rrhajt:k f')pnuwomu 1th\'.ttUuth:e.a1 .in
r~ . . tanl;,mt:ln of TrJ,'JXl/fOJ.Onu, (Ortgu/1:n.•~ in Rhttde.-.fa. RhCJdoluu c.ittle- in Mbalt: an~ Torum th'-lrich in £lilW\•rn Uganda Jaumal 11;
\:ttcrlnmy Joumnl, .a, 6'2-67 Proto:oolc,z)1 H<.tl'(ltl!h, 7. -t8-5J
161 u_f\\f.Lr:, CJ.., 1uc..,,~~....c.•• ,m,no, c.o. "-c-s:.11n11-. 1.. 198i. l'Jt~ efiJ.>C'.( ot 180 ,1M,\1,1\',w1Nf, ,, \1,. 1~0. l ht.· cpic.fomLt>101tkAI ,-.aluo o( t.•:,cp..-rlml!"rU.ul hu,.1
T'J1llt.11,uumn r:onplrn.<, infection on 1hc t)("'SttOU< C)'ci~<if 1hc ~a.1t. and animal r(:s('n·oh' of 1,Ypuno,om" ,,,,,,..-1 gaml.,/4•m.i.• Rt'rto.•,t,•
Brl:uh VNt·rlnary )t;mnurl. 1-13. 423-;ll ,\f.'d~Ymrt ac'ri1111/r4• l.&l,8~.l-3'::'5..
1('2: U.£WFL\-;,.:. t;. \,. \tU~NU. C.D,. ltlC"Ji:I~~ A.e.-.• JOkltr. 't., \1UIUl", !tS & 11:SI .\tAKU,1Y.\\'1NI, .\., ~11111..11,. 0,, )..\(,[JtU);.,\. P .• !,,;.\l.\'U\111.\, G,l. I, \U)l 1.SltO,
tOAx.,"i:t..ts1, t.. 1&98. 1'ht" tifo(;sot J'r,1utn.oso11w (r}11,i:_oh11JS< infoc,inn on n .. 1939. L-t.· rL~rvoir anim;1l th• TryP1tnosom11 btur...·I gumb,,mu.,a.u Z~,ul':
the CX"SU'Otl). cyl'lr of 1hc- Rorn.n cut·. Br11L,t, l/t!lt•ri,w,j' /ounuJ/. \.1.: lo!ec1ion~ 1r,."J1~no~omfen11<:~ d,u\.. dt"U.'< fo~t·rt111 Bni·Z.."'tltL'. 1to:1frnl
379-387~ ,\Jtdicltir and 1'1Ua,:'tulos;,. ·10 258-2~
l63 l.QG..\ ..... LL. (,,OOOWN, J,1 ., TC~!t'Hl"~$. & C:RAI(~. T.M ... 198J ;\l:unt~im.og 1.82. \1,\10(), S.IC Cll.W.\.\.) , (:()S\OJt. A•• omo~I"( J l.'1\1CYTUO. v•• \fAEIII... ,.u.11 ••
Zebu Ml.\ure c:nttle inn i~t~c iDfL"Sted nr~ of ~lali. Tmpltal Arumaf '1U);• ~:>J. J '\lURH-\\. " · Mnn:,·.-\, J.).J, "'lflUI. ,., ,, .• 1968, Tlw u~,, u(
f1Mltl1t1t1tl Prod11ct(On, 16. l•l?. ch,•mopr<,pll~lui, ill t:a.1 Af11c,,n Zebu -,llai:c c;inl< <Xpo.<,"1 ,a
l~-p.JnasomWh. fn :,_Juhnka. Jv..•n~·;,. /,i, l.t1·t'ltot:k PruiluttUJtt 1r1 T~t:St
.i,
16' LOSOS. G.J., 1986, /nftct:ous Tropic«/ Disen. <>fDDm,·\I« .~nlmols.
t.ongm;an Sc:1cnllfi(':\nd TtcilniW in a.t;,s,0ci.,w,n wnh 1hr lmi:m;uir.,nal .1;f,.,·1.,v.i nt..-,,u of,·lfri,YJ. Pmc1.·1,.-dini,t... of .1 ~h.-.:tin~ar the.• Ahico1n
T1''P.lnotol<"rnm l.a\'l!~tctc-Ji.. ,~twork.1J ..,l:- SO\t.'mbt.r 19A7. X.iiruhi.
O.,.·•topm•rn and llesea,ch c~mro. Cinadt1
Kc.'nyn S:Jirob): F.'n~ltsh ~~-<-..
16,; LOSO).. c;,.&aiou1s.um, A.. 1,;-,
Pathoe,enlclt)·nf 1r)·p,mnwm~.
1113 ~,,,-..:t, ,,. , .1'f80. TIH., pathUt,:.t.. nbiS of iaf~tt>I\ with To·IJ(lnosr,mu
1
Procwdinr,> ofn \\'orksi10µ ll~t,/ m Sarro/Ii. Ke11)Y1, JCl-23 ,·~:·mllirr
1978, C>tra""' rmerru,tionnl U<'\'olopmcnt lle;.,,i.rch C~m,. l't('U,t m goa~ .ind ('0.llh\ n11 '"'ctcWl:tlf"/ R1."(f)rd. 10;. aSt-»-:-

tSB to~.s.c:;.J,. P,\Jtb. J•• \\''U.!l-0~ •.\.I. 6 V"-1\. l.~. 19;:i.. Po1hotog)' or the d),cJ.,c 1.8.t )1\,\EJ,Q;_:\ ,,,,.runA. ,. \l, ,').01 r.w.o +.MU!'OJ.:L,\.I J.9U.t Cercbr.d
In e:t:Ult! C":&u.srd h)· T1',·11twr,>011u1 C111t1;,oh•11,,,. Du1/4•tm ofE,,i:"°tlc uyp~no"'JmiA.."i~ m c.aulu ,uth m1.xcd Tf)·1Htr1QJ/Jmr1 roflJ.'0,1 'U1t' .ind /
/Ju.'lltc< of.'ifrlca. 21. 2J!l-2•l8. llrnul bn1C"~1 h1fcc1ion,.,_ •\,·rn Tmp:m, 41, !37-2,16.

16;1 LUOaxs.•,.e.. 1.911. Oe1ecu.on of ant!IJodics In h11'iln011Qmc-i1u·L"Ct1,.-d Serurn f.:11.:cor,

J8.; MAUlHJ,. 1.• j,,._,\f'!,\\'O, J.P.. t-l('Jf)f), '.'tU .1 , b.·l\",\~"'i. OA, t~i\
c~mlt by m<!'an.s of ,1 m1croplntC"cn~n1r,,lln._~J iinmunn'4.1tbcnt a~,ar.. .:,nJ the rrutlur.1uun of f~panowm,r rongo/n1.tf'"lnrt-ctfon., in <,!<J~um
Ttop:',al Am11w/ lftttltlJ nml Pmduc1i1.m. 9, 53-6".t. mqr1,uw~. l.i''IJS<hn.ft f"r Pnm.m,.....,~,md1.-. 70, 11-lq
l68 1.uc;~1s,. A.G .. i9;9. Trypwta>vmc, 111 th~ I~ mph nod~s o! ca ale .&nd 1Hr. M,,umJ!\, ,. ~ \\ 1t.HuK,, ,.r , t98i Lc.•<un medJai:t:d L->1:ibll,luncm c.•f
~he.ep mftoc1,-d ,,tth '1 r,•mmo,<omn ((mso/(!11$<', Rf!M'flf<h in \ ·rt1'tiwro· nudgut mr.....·,ton1. cir {r;·ptttt(IJOtnt1 Mngol,·tb,• Jnd Tryv~m().\i)lt:11 brur.-t
S<ienc;,. 27.129-131. in (;i'u~ma motsllfw.,. rn,11ft11l ,\ft"t.llrmt'nml 1;1m,sltolo&,,·, .sa.16;-r:o,
n;g lUCiia'\~. A.t.. " MFHtrr/.. u . 1978. E\-;slu.tdon t1r~10 indir~ct tluorrtet•nt 18';" \Huou,·.1 "WUJtmu,,:. ~.r. .. J ~ l'b.• rolt offl't11n" urut tl")-po.nol<IOrrtc
.u-alibudy te.1, ..:ney·m~·Un}..,;-d immu1\0'1'11'h~nt l't..,ur i11UJ qu:uudk;allfm ~l"nn;H'"' m thtl' matunu(on of mfd~tu tnfce1inn., in <ilo~'1irta munitans.
of immunoglobu1ins tn the dfa_gnoo;~ nr bovin~ 1rw11no"')ntl.J.-.t, IIOJ>lrnl \frdirUw 11,u/ Pt1ttit/UJl01;>·, ;i9, »-5tt.
TrQp/<11/,\mmal 1/ealclraml l'rod11Nlo11, I(>, M9-15!1. 188 MAhO~. IJ...\•• s9s~11 r\.n ln\.tsrJstat(on inm 1hc li,·cs1rn:k prub!\•m,,.uf the
1;6 LllO:.IS!o.•.\.t,.. :.un1on..\.\'D. 0, \I\\ ,\S\il. 0, &, IIIJPl:'.l'.'I::,., I, lgj.;, E.ul~ Mag,':'\ La\,'t.•r Shirl• , ·:1IIC')' nf N\'3s.tihmd with pr,rtfculM wf~rnc'-' to ch~ dhc:1SI."
o( infocdon ;,,·Ith TrypamJSQtnn wnsolt•n.w: p.at.1!->itC' kinc1ie'- Jt~d 1t'\·f\11no~1mio,i~. t)£,1>.trt1nent of\ ctl.!rin:,;n·Sc-n·lt·~ .inti Animal
or
cx-prcs...Jon mc.•1,acydic \':lriablt.: '1Jltlg;:n 1.~·pL·~ ,.\,111 Ttuplro. :,g, lndu~11) . ;\:\•ol"3laud.
195-206. 18Y )u.;cuu\•, H1\l. (,t;'llfllt.. •.,.o..
1971 CliniCOJ13thologJc~l smd~·on
t;1 tU:\bOg__"'., w.11.tt.. J.iJMB-~. (.U .. n· ,s:,., l),A. " 11( 11(.;., ..., •• 11\79- fl)1Nlfffl~mnu 1.--<perim~ntal Tl)'J>a,msomn lm1fet inrcccion.i in horses. ?an:?.
bruW. Mlnhuurc .anhm~cxc:homgc- ccmri(11t.,tttllan h.:chnlqu~ Jor t lblOJUlthuJ0gkal findlnAS :nth<" m.•l'\·oui.. ~},.tt.-m unJ (ICh.cr orgnn.. o!
dctecuon u(lo"' p,ar~it,1cmlas:: .\d.;sptadQn (or llckt use. Trot1,in,·rlrm.~of Lrl·lHIMS ;u1d untrc:ucd horM·_.;. ri:Jctlng to n.1~iln,1. Ondttstt/J(Jf)l't JoullkJI
rla• Ho_w1/ $(1(.:fol)•Qf Troµico/ ,\kdrtim•nml /JJpwr,, 73, 31:?-Jl" o{\l<'t,•ri,mr_t• Rl*:f,"tttt It, 38. I l I I i6.
r~ 'V\Cl:O\l1£. l',h.l .. U(J\"l, \\.V 4 [M',lJf., \',\\., 1~75, Twu Cll~l'>Uf Otltural 190 ,mcor,: s... OLU8,\\'0, Jl,fJ., u.,,.JI. ;\,. 1,, ZWF\'(.\1rm.11"' 1u93 Th-: mllui?m:c of
lnfc..\.CIJon of !ihCCl> ,.,•i1h Tr,·,,u11Mr,mn lN11111mmnnm: simi,w ltlru«"~11d ht:il1 biood on inff.'Clfun rtll(.\;1. in (;!~sin<, maromu:.< ipp. m1ccu:<S \•11th
other, 19121, 11/rr,,/,>sirlll \'eter/11tlr)' /1>11mrtl. b. U-14. TryJNmu,om,, t·1mpt,t,ms,•, T. llfuf?I .:.nd T 1uuia.tt Pura.si1ot,,so·. 107
l;"J \L~L!\'1..11., r.~,I., BOYT, \\'.P., 1''.\1Sut, ,. \\/ ~. 1.A.'Ol1t. .tvP. ,_. ~\\'\'.\t,J'01L It,. -11-18.
197:;., tmmuntbuppr1K-slon tn l'l\ine 1rypann"omit1ioi,.. l 1tiJ t·c...tt•nnal)' 191 ,nur<,.\'.\, r'.l,"'.'l.f. t, T1U..1'k. ctn. 1q8a, A model ohil~tw,Jron.. muu.•d animal
Fli.·Md 97. 452-l.'i:l. tl'ypill1QM1rl1Jl.1'is. / 1(1((1$110/(Jg)', ~6. ~! :-.23S
r:.: )IACU.\.Ni\.~, It.I.A., 1~;0.. The epu!oouolog)' ol uypnnosoml..1...1s 1n 192: \11\!11.\, u." f\'As-.., 11.<.. , 19,0 Z.im.inglo lnduttui.tl Corparo1tlon l.irnltcd.
Uvastoek in ,vcs, Africa. In: \tUUJ~""· H.W ~ (l()n!,, 'W,H. l~1, 11:1" lu,a~.,.. /,•nihiu. Unpul>ll,hed d,14.
.AfrU:on T')1>nnh,."cJmlll$l'..t. London: \n-01g.(.-Al~1~.md Unh"in l Minbln: of
193 MUtt>(), <...1,; , i'7iq3. Tht· d1,.:tribution of C/o,~J.nu ~t."t'ilr' in Africa :ind tht'ir
O\•c,~ca, nt!,•dop1nent.
nnwral ho~t~ ln>«t ~lrm·,1dud ii$ Ap,>Ttrm,t'm, 14, St !-S2'7.
1;5 ,v.cu x~Ax, ..:.J.k,. t9;.i 1hr- L-pi;,:omiolORr of tsets.r rra1umj11rd
19.: ~1010<1. ,.~ •. t98t Effoc:b uf mamt~Jiung (ifo11i,u, mot'SltllHS mMsfttJt:.,( on
tl')p~nosomi.ul> in relation to li<C>1ock dcvclopnwm :,nd ((>ntrol
Ulffor,•nc ho!->1' up<m the \'Mlor"·~ ~ubs('qut'm Jnfocthm 1ML--,,. wuh
measure$. /11: co,mol PMgrnmm~ fur TryfXJl1()$0mt>'tU:d tit,•,, \·«,ors.
p.tthu~1..·11.k uyp.:1no~rol.,.. :lrm Tropt,,1 38. 125-13«
.\ct~ du Colloque, P:.ns IZ-15 ~l;trch 1971, 111,"Tlull d !Je\.i,:e ,•t dr
)1EdL-cirte \'N,nnalrc d.!s P.\~ Troplnlux. l'::15 MOU><>. -s..,-..., >,;1rrn1.\. ,n. ~ UOJl'tn ,:.t.
,. r.1- ig&o .Siudics: an c;lI:Wma
pnt/Ulip..'>. G ftss<i,>t~ fuJ:t'lfh't an!J C. lmwfl)(.11,,,s in u:rm:)01 rh~
J:'6 '!. ,('.1.C\.""'\ ,s, ~.f.R., 197.;. ,\ m,M., of c.pcndon, n~m,t tfet,c- :la'"' '1nd
' trypano~mi.J.~1:, tn /.amb,.a. and ,ecoorn1cod,,t1on'\ (or lh,'. futon~. cp1dtmiol(:t\:\· and cpi1,0(11i<lroscy •1! U)pant1somiost1, In Sau1n Eu5tcm
U)!;iJ.Udoi, .J..n1111/st,f (u)pi((l/ .\li"d(c'hl<' IWtl PnM,,iWltJIQ,'. i•1. :?19-?37
R<•porJ to the l):roi-,o, of\ (J,,,rwao·nml TRJ..<t' (":tmtrvl Scn'rces. l.Jt,qika.
Zom/Jia. BU'llflgJ' (I/ l't)'/k'lrlOSOllta and
1gf, ,,U,!_\ "\.(\IX. O.tt ....\~llfUl)U. "\\' ig.8.3.. Tih·
1.-t.tlmu:ma Parfl.dtU">' tJ{,\/1111 mu! Dom1t,\lf(' .-l11111ml$, IAttdmi: T11~1ot
:7; ~cu."'~'-'•~,. R. 1.980. T'4:~c-tnsn..mmcd tryp:mo~ClmLl,b 111 ttdnuon
,md Francb.
to th~ 1t1r.tl. ecunom) in Afrka. I. r ><'l¾' !n!C\(;ltlon. \\'qr/rUn1m11I
R,11•/eu•. 36. 2-17 1,r ,1,.1u·."•.t. u'°'. o.u. • 11 11,vu1 -.. n., 1~&. h•t'<flr1l( bchJ.\iOur of
1:8 \IACLE..~~,. ~I.It. 4- M'I\.,\, P \o,, t!.t7U, f<vl:lp,tng rl)·;,a,;o,.ut11ft &·i1'1u
p,'l\ho1:.en•infoctc.'<f \'t'ttor-,. Pnrusur,letg:.~ 9?. -.?1-720.
lnft:Ction~ in ~lg:urfan ,t:hu cattl\• tn:Jtcd wilh dimlnatt.~nc.- ,.\Ct'IUratc. t~O ~101t111..,..J:>., \\ t.. \lutUt,,, ~1 • .,~oi.. c;,\,•,o., 19Hs lmmun,e u.°"'pnn~ of

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 i\friClln ~nimnl uypnno,omoses 293

c.1ntr to ,·\trit.Jn trypano:"1om1:",. /,,: nz.,nr,. 1., h•d,). Jmmunq/ogym1d r1.•:.l1'nn:\h1p \\1th cr.1rpu ... Juh.~im func.:tmn Ul the.• 1;{)3t. n1,rlOgi1110Io13·.
Ptttlto~m~i-. a/ T1)·1Hmt1M>ml,,.,1s. Boe:. Ra1on, Flo-rtda: CRC Prtt,, I cd -32 •• ;!',-S36--.
isg ,,o,ou~m•. \\,t,. \SUIOlA\·. ,,. ,. ,ui:.R, P.n .. t9"9, P:\1ho~<"nl!if!. afci~~U<' :i:19 ,,u·rA\OKA. "·"'·· o·uuuonu RI. It.ti.• c;u ... tltL !'>., ii,88, Tt)'J)at'IOSOl'l\e·
l~lou~ in 1: IJtu~I inh.-crion~ In: t ~)~. t,;. • u-1os1"'AHn. "· r.•dsl. in<.iu,cd dt:p1e11J:o.1on or pl:r,.1n41 th\lro-..:lnc lt"Vt•J,- In prepubo1'1nl and adu11
Putl:ogenlt:tt.1.·oj1,yptmfJ.,m,tl'1 Prc.1cct•tJln~,Qf a. Workshop held at female i:oais Mta E1ttl<J<·n110/uglw (Cup,nl1GS,·nJ. l l9. 21-26.
Xoirobl. K~n;., 20-2J ,ov,mber 19;"8 Ottawa: lntorruit!onnl ~ :i.nn>L-uu.,. t ,,. ~ l'L\JC'OS,, .,, •• 1981, Surface ca:rbohydr.Jt~s of proc:ydic
De"\~Jop:m..-nt R,¼.e,lr(h Ctnt,c. Conn, nf .-Unt'..m IJ1."J)3no,om~ -.tu.died usu-::R tlt1arts("(tt1c:r n.c1h,urd cell
:201) )IOR:Rl~OS, W,L wn LS• .,,\\ .. \IOI 04.'t, ...l: .. v.w~. J. t, MUM.RAX. ~f.. l9'8:l.. son~r tlru\1~·5-,~ and ~tu1 (nation or lc(titls. ,.\lol.xulllr andBfoch,w:ital
1nt~ricruncc In th,• ~1a.b1i,hme::t of >u?('d:1'1fl?clions ~,11.h T()1H'Jt:OS-Oma Plrrtditnfog\·,Z3. 16S 172 •
.:cmgoh•un~in c::UlJr./oumn/ of Poru.Sitof,w·. 68, 7SS-76-.. ~1 ,n,. ,:-iau • .,., ,i. ~ lOM'>~..1•.1.. 19;-9. lJJrro:,,1ruct\l.ro.l cho.nge,s ln blood
201 ,:os~ 1> k., cvoa:, c;.:\., ~Ul, O,f. • nurn. c.J>., 1989. Oetect,on ot T. \'\.~Cf~ or u,,1u,•..;,. of c.&tc!e exs,enmt•milJlr m!CClL'd With fryptmOSOU:(1
ro11:.,'<>k11,,• .an,i r '""<vi ...ub)p~,~ by ONA nmp)jfic::nioo ~ing the c.·ontof,·n~:md T. r•ltv,x .\ prrlln11n:u:y rerun. /11: v,so.s. C,-& cHuu1~ ., Ru,
1mt~mtrase chain reartion Pan~,rolOK)', 99, 57-66, "" llldsi. /"111t,oi~111<1ry of rryp,mo,om,1. Proce<ldingsor • Workshop
203 \iULJA~ A.L ~ JUCDL\~, t.H.. Ut81!. lio\\' do J-\frlc.tn gum1.· on!m.al~n:,mrol held •tt :\aicohl. Kenrn. 20-2;:s ~o,•ernber 1978, Ounwa; Jmernillional
tr)'µanoi..omr lnft.rcioru.1 Pllm~iwft>J.r.' fmlti,\'t 11, 1!l2-3!i·i DeVL•lupnumt Rt,~tarch Clm:rc~
203 ,1ut.L\ ..u. t,: JllCX.\S,\~. L,11,. 1988. E\.'i<lt•nce for tJfo pre,e11cc of a.n tnnnt~ :ta2 .~A~lUL\A, \·.,1 •. 19&6. lmn.1'111ofog1co.l .;pproach~ 10 the control of
tr~:p.1nt>'l,Qtnkidi;i1 !.-ictor In Lhi!~nJm of a nnn-hnmu.nL' African amm:d U)';>J1Mll~omm~,,.,, J>amiitulog,• TodaJ'. ?. 1*173.
\\'a,orbuck {~'"' ,lllpslpf)11111u,1. Tt11nS11<tloni oft/1,• Royal ~,,../cty 1>f ~ 1-..,,1un.,. ,·.~1 .. 19~u. ·rryp:,1rn'!Omlo.~i}m domt'!lth: :1nim3J.;.: thi.-
Tropf<nl M,'flirl11ea11tl Hygh•,..., ~. 4,-98. µrobltn~ ol ch:1gno·d"" R,.,,,,,..~·rf'nlifi(lf'~ ~, Ttchnit/u<t,lt I. 'OjJiu
20.l ;1,mwc.,s. tt.w. & J'Ul ,~. w.11. 19:n. The :\fnc4*fl ·rr)p:t.nosomia.IB~ lmITnnt(mutf tf~.f f.'pi:ot>U~ 9, 3S7-3E>7
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RQ\U., t... l97!1 .-\ te\1\t!W uf the prmpt•t,..,for ,·;u;.cina.tlun in African ,md r. i.mu·ri inf~ion.s In rnulr~ Tro,}!(ttl .\lrdlcm~mzd PnrfUirof"KJ', .io,
11')1)ano,omlu,ls. Pan I. Worlr/ ,\11/m(ll Hm<w, 32. 9-13. ?67-l'i':!..
207 ,u.11m.w. M. A- t11>if>:. ~.,., 198;..Afrirm t1)1mno,nmfa,.J< inca.ttlc: \\''orkJng :.i:26 '"~nu·., \•.,~·. u,'l>QUNT. li.,J. o St'l\r~,n;1,.. ,,.. 198g. rcm J1.rds improved 1

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1/zt• TroµiC'J. t'rutl'Cdlno" of the Sixth lnc~m\\tfonal Con(ettnc.c oo
208 '.\lURIUr• .\t., curroR.o, 0.1 .. GCTTl:-.-s\·. c.: •• ~"aw. w.F. s.,, '10:-..:n"KI. \\'.t,\1,
1981. S1J~c.,p1tbOlc\• to,\lric.in 11')-J),""tnosomin<iiit., of N'l},\ma.:ind Zebu I.J,·L'"il<K~ Protfut:1ion :tnd llMa!<.c in che Tropic~ twld in \V1;gc-ning1.•n,
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c.,1tdr- in :111 r.re{1 ,>f (i/0,.,.($iru,1 muri.i1011.( 1ul1mor.1/tn11t<'.h.~Jlen~~. Th~
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208 MURlt.\\'. l-1. &m:..xnn, T ..., •• 1986, .\n:um11a in t:.,cMncA(nam ,~a;. ~lonoclouul antlb<Jdl\!'. 01.1, dl>tonguJsh Tf)ponosQ111,, congolrnst.
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dettttlna uf mlcrofU:,rf.i :,nd rt)l),1nn~t>mh in thr- blood of t:mte..
211 \1lntR.\Y, ~! . \H.lJHtf:.c)~. \\",I." \\'Hl1'-l,\\\", u.n .. 1982. IJOJ,.t $.UM'i.'ptihlllt\' 10
A,uu,f~ tf~ la SwNrl 8tlgc,l,· Mt'l.licine Tropitnli."'. 75, I.t:5-148,
\(rican 1~··pnno'°mf;"l,: rrypnnn1oll•rnncc•.~/11t11ttt$ in Parasitology.
21. 1-"8. ~9 !"OOUT \'iL\.. c._ ~col.UC>.~• .:.. MUM.PU\', ,.a.6o J•EJU'CRl:\"t, ,\.5.. 199:,
Oer1,11tlon ond ch:u•ctoriroti<>n of 3 <(Wn>p)T>tnl111.. u,,;1>1nn< clorie t>I
21.2 '\tl1JOt.u. !'IL .\tu)i;Jc.\\', t•.l,;~ « \tCIXJYft!'., W.l.~L 19-;';'. An improved
h;/1umnJoma ci.1'1bifJfr.,m,r". 1\11111uirroblt1J. \g1."nU a1id Ch,,,,toth(rru(I:,'. 37
p.ira.~ito!o~lcnJ u..•thnlqur for tht" di:ittn~i~ of A(ri('An tf)'J1Jnor.omia~i"-
1163-1166.
rr,111t(Jtlion...<o/tlte RtJ.\·ul ~odt•l)· hf fro1>in,1 ,\letllC111,oml Hy;;:elJ!!. -1 .
.ill,-3.26. ;..'3(1 MOIJU, t .>,; .. O,iM, 1.0. rMUtJ<:\, ,1.0."" PU...'-\UJO, ._.. );., 1996. The-effee, ot
hn.,1 blood on lhl· in 1·11,0 1.r;rn:-;{o:mJtion of bJood$trea.'ll u:ypn.nosom~s:
::tl3 \1UAR.\\. M TM.\JL f,('_)1, 4 lf1l'JERf:O,. (.j,1),M., l.990, Tryp:!.:".OJolor.mc~in
b) t'<';;;c n'1tdRU: homo&Cn;HL~, Metiical nmf ~ 't'tmnnry Emam<1!Qg)•. 10,
anlt 3nd pro~p1.tcc-c: for 1hc C"onrrol <1f lt),-,.anMumia,i.s b>· s~1.-clh1?
31--:\22.
brei.--dio~ Ri:t'W! S<"it'ntl/i(Jut t•t Tttlmiqm• de L'Offecr fmtrnarlonnf M$
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S1r~1egic use oJ c;ho-n1opro1,hylnxss ,J1'..,,lll"t uyp,mosomia'iis i.Q di.lfcrem
2.\,1 ,111,ou.,, <··"·· \\IHt.m, 1l1 Phu.nnatoklnetiCS.
f. \oAR.\..,..,.,. w,M .• r99b,
~ypcs 01 Boran c-nule. K<'IIJ'tt \'l't?nnanm,. 6. i'
bioa\'W1ability and ll~we n:,,dues ot \4Clisom,·taf(lldmm in
no:·Hnfe-c,ed :tnd Try11n11f'J,mn:a <ungot,•rl.fl.!·lnCcctcd ~or~n catdt'.•-tcm ~ 0(,11:i...-n:,11. o ... ru~mtlAf.>f, t\.A., 19ij~. ProphyJa"t' or Afrk,1.n .u1lm.:d
Tropi,11, 61. 277-292, 1'1 pano"'l.om,Ase'- ,\ owlc" of ,Qmt- foc;tor-. t.htH may mfluence the
:,13 ,1u1.,YhU,\, a. ,1. & t<O'llllL :i,,, tg~9. 1:11\."C~ of ACr!cQ.n tcypar.o,omia-sSs on cturation of l>0ml't.m11tlh1111 c.·h1vrid~ pms>h:•h.\>.b. 'l,m:rint11)· nutteri11,
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pla\ma comi-01 and 1hyroxine conccmr.,t1on ln gom". Rt-tr:arch Ju
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2;;.. 33,
T~1JW\Ch,umc..induct.td ut..t.ria.n dy,tunctiQn. F"idcncc of higher rt""SlduJJ
(irmUrr fn tryJ>::.nott11t:r.tnt ,mall ~"\t A!rirrtn tt{;)at~.Att11 Tr11pil<1, ~s. ~3 i r;('I. j,..0\fJJU, O.t.l\ .. 1~.a6. Eff'cch of t:.'-J>Ct'fmenUtl
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?2!,-?37 '!?1~11101011,a 1•/1vu: lnfoctlon on hrs!·. "'C.Und·. and tlurd·tr!mesttr
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pn-gn~n11· ln h,-:ft". 1'11rrfo1J1·no/c,gy. 25. 383-39&
l~l)Jnotol~r;in<o or thu <m,ul E'1<t .-\iriran br<fl! or ~oats from di!!trent 235 oc."v, o.,. ""•u., • 1981. Tr.in,placont.>f trar.,ml.<s!o!I o( tl')']>>n"'omc
localith~, •o 1iy/Jc'WOJ()ma t:"011:gofr,uc infrction. V',um11n')· P,1r(Uito!~·. In ,1nim,1,J..- and m.1n. :\ rc\•it,"f\\ fl:~\ 't>iirlt:flf)' Br,IINm. 51 381-381,
JI. !lf>-10;. 23-11 o).:.tt:u, ""· w.u-..o~. 1.0. 1.uc:~1 \:,, \..11 .- \i.U.-\t\m, n.w. 1996. The ttit.."\'.t
::18 '\1UT\\()8-\ n.,,., \tf:\'f_ft 11.u.r>.• o~~"J· J. 6COMBL s... 1989. o( ".>.'J>t•nm1.m10J 1nit'C1ion o(Roran Ci1tt!e in ~:uly and mid ..pregnann•
r~,,a.nOM>mr.. inducl"d inri~.l"i~ in pro .. c-ij,llmdin·f·2·nlph.? nnd ~ \\'ilh Tl)p,W,vmn111ti1ru. RrlttJIJ Vt11rrilutr,/ourrm/. 1;;..'?. J ..,U-JSJ.

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.23: OMU:l. tt,c:.o., 1991, Pig trypano;.Qmo\i.,~ pu,°'al~nl't- ;uui .. i~nifit.,nc,• In Sctcn~-:-d arc.:.1r,uv.anio11 1941 U. Tht:'\4.•1.:wr roh• <1( c;tomn.a
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:.t38 P.,ru~. J•• ~nm.A.\\, M."' '.\:mrn,rnA. F.. 1981, A cou1p3n,ri\·l' t•,·.tlu:uion l>f 'fry.·1.vmoroma 1•it•tJX rn tJw Ton.a Ri,'<!r d{~1tlC't of ~{'nrn fl:< t '1.•tair.<Jf'}'
1he p~ra;i1ulogfcol icch.n!qur, currcnlli· .-ullobl,• for the d!Jgnu,,. of Ha:t1rd. i:r. 5-S~-S.58.
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m.c.u;urt~. Smitl:t•r,u, .,·o
I. 1-7.?..
m !'>.,.Le.,, I', 1 J l.., l'"J82.. '\l.·~1Mu>:&l in<•dcnc,• of hclnunths in sht..'C!p o.r:d ~oats
250 koorr1. 1. & PU,\.R:'ioOX, r.\\., 1990. The proc~·rlin co.it o( .\frkan
1n S0u1n '.\Jr.,1,wnb1(lue. l'et,•tinnn f'n,a,,rn.'·~·. I 1. 3ti-l28.
uyp.,nosom~ (Qr 1h.e ncn,-.:o,nakc.t.d lr)'J).,nfl,Qmcl, Ptu<liltll/QFJ,' TQ.tay,
6. ~'42. 2.:2 ~. fl'iH·'· t 1 .. 1!!,6(,. Pis lf)'JlilllO"iOIT\i.l',,l5 ln lropicn! ,\fnc-,t. Common·
\\t:ahh l:hm:.:u of Anunal ttca.h.h. RcHcw'i.·nC> 7':o. ~. COmn,Qnw<.•uhh
;lSt ROE.DOI, P.. ~<.on, f.M."' n1.~JlA.\I,
. 1u;.. t98.f. ,\nlh~ 1r;:1x,no.tomt'! 1•i1Y1.,·
\gncuhurul Hu,r\!'au. fomhnm Ho,u\c. f;41,rnhmn H,>r'11. tcut\."3 U.1'..
intcctton ot E1h1oplan c:ml~ fn tht! ,;1pp.11tt'nt ub\tm:.,,• of t:<tt~C!'. Tropfr:ot
..\niu:ot U~lth 11,ul Produr:ion. lb. 141-1 t7 l:'J ~u NO•.. , 1.1... i:f;o C.link:al mnnJf1.•.. :,uiotbof the trypanosomi.i~~ •n
li,·t."Mod, 1111d utht"r <lotn,:-.1J1. .inun.).~, !11: ,11111 u.hs, 11.w., 1'U~, \\',II~
'2s::: ROf.!A ,,s.
(i.L. 1.936. ~;:uuraT ~i\1-ance to .\fnc.an 1~p,1nowm1.1.\b
f'd.s Th,: ..~fn"'m fryvanrncntiniri. London: (ic1>rg4· i\Jlt.-n .1nd iJnwir.
J

Parasi1o:lmmu11,,l01,,·. 8. 1-10. \!!n,-.,try o(O.N-;,..n, Ot."'\'\1Jopmcm.

,:-33 J(OU.\:\~. G.t. 1',\.'\1KOURA, r.. sion.,, o.a.. flA..,q-:r-.;f,,'\, .\ .«, Pl~nfft. ,, .• 1983.
r.'J "-Ttrtll ~. L r .. 198f. nr1hmtMmnw.,t!-. \ t'tlt7'tml)' t•,1rs1!t'c·rilr-c ()\focrl •
Tryp.:anoioh:r.mcc: .\n ,odividuat nor a. bu:.-\.-d ch.;u.11.11:r ,r kw (1(Jpm1. .SO.
l'c.'rgnmon p~........
~l0-1.
.ti5 ~· U'IU, , Lt • \1 , .. ~-·'-·J'ft l .•.• 19-;-9, ~p~11mcnral 1'Jyf)t11tO:,.'(Jft;(I t'ii'/Ll.
,s., HO,..FR>, o.J., 191$0. l:pl,001lol~ Th• L<lll-W r.;111lc ln1e:t.i,,• 111 PAO lntt'C,h:1n Jn &.he l1orw. n,< \ ,·11·rlnur, ilt"ttJtJ. -1. 5li-511 .
.. Rn1'm Qflltt! F..qR~rt Cmuulu:n·o,, ,m llt..~11rrl, t>t1 Trypt111t):11Jmtful.\,
llomr, 1-.SOc'tol~·, 1979. MO-\C;,\;1'1)1"/ic:J25. Romo: PoodMd ::i6 ~UI I\U',';, tt,a.• n.u>M,\..\. J):.J.. 1q;gq P;uhQg~rH~lSand uctiulog:, af
Agnc-uhun, Org,nl7.aclon ;m:.i.emi,1 m lt}"l').rtrlnM>mui-c.J;"I , ...·uh 'Spt.'Clal rdnrcncc 10 1 brul1.'i a.nt1
T• .-l'.,mi. tt•1,·11m,,, 111111,•tm. :i9. 99-10;.
~'iS R'O(..tf.$. o,f,, a9a5. frypano ~mi:l\tt. ·fl,'k' or 'chalh:nitt·· \ tC',.iew. .-lt"fil
Trop/ca.42.~J.J. !!7' r.,111:1 .. H.., 197~. :,icnun pmttin chmir~ in ho,.·mi.• uyp;.umsonm1"*''' A
ft!\:h.'1\", Im J,OS(>"\ c;, ~ t lfOIJIS~RU. \•;\.~is~. l'ml:qg,f!nicll)'OJ
~6 ROGlJls. 0.1 .. 1908. A gcncralmodd ror the Mric.in u,p.r.no,omia,1,.. 1r"$.pa,:,>wmt•.t. Pr(x·t-.:ding.... of u \'iork.-.Jmp hdd ,U ?\~irobt KC'ny.l,
Pll.J'ru'ltologt, ~li, 193-212,
20.-2.i Xcnembt'r l97ti, 0C::lW3! lm~rnotion:sl fJ~·t1lop1nc.•nt R\.""4.:tnch
'l~ ROGJ;.RS.. o.r ... a.()RF:JL\,';1, v.r.L.1973. Slct•phlg.\'.ickn(~ "°"''C:Y In IIW 0.•ttUC'.

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 African animal ll~'Pano.somost•, 295

z,8 T.\fr, .\. &·l'URNl,-H, C'.M,M .. 1~9'). Gt'nctIC e.,xch:rngc in frypa1tQ>!Q(UO tJruet•J. !!~ V.\W. , .. 1-.a., Hl~B.ll}t(: r.. ,,. ;,,, \llt,1 .., ,,:-.:. t.9':tf, ?ethcl~' of T t."OUIJ4lt'IUt'
l'ar(JS:IO/Qgy Tc,;la)', 6, i0-75. In t,tl\c, In: w=. c. k cuou""""· , ...e,M Parh<>gfJ>ir/ryof
Zi!f T>.lt:.~L\., \\., r-AVLt>R·U \\ilS.. r.c:.ll wrmorono. !.t.n•. 1988. flC>ld!:nudleson TQ·panorome.<. Proceeding,, q( • Wooohop hcld ~t 'l:ulmbl. l(A.•n;'I,
anlrrul 1rypano;on1ia,i, In ~10'.:amhlquc. l. Eifocrf,'t,1\W of 1ho 2.0-23 Xo,.·embcr Ht78. On.1wa; tntcrn;,liorieJ l)evt•1oprnem R~enrch
proph)'l3etlcdrug; bomtta:'nldium thlorldo und 11yrithidium bmmld•. C:c.mut.
Ttoptcul .~n/nwl l·l,'illtlr fUtd l>tvd11c1io11. :!O, 2~3-255. :!.;1B VIC:Kl.'RM.\S, J.., lll~h On
1hc surfa~ cqatu.nd {tnge"llor- ad11{1:SiOn 1n
2Bo1n:umu.. A.R.1.. t9fl1, Tht! ste:corarian trypanosomes. In: SOULS!\\ 1:...1.L. t~·p~nnsomcs Jo,mu,l Qj Cf// !xirm:•. S. 163-193.
t l-d.~ Immum! Ri.'SµOn.$d ill Ptfrt,ufdc lnfi•ttlmt. lmmunol~·-
299 , 1cn11." ,s. ~.. 198;. l)<'l"lopincmol C)'<I"' 3nd biology orp,uhogm,c
lmmwwpmlwltJlJ' mtd lmmw:oprtJp/J)'ltu/$_ Vol. JI/: ProlO:,()(I Sota uypano,omt'>, Mrt,11 .\f,i/it<1/ 8'11/l'lirt. -ll. lOS-1 H.
R,uon. Florida: C:RC Press Inc.
3on,1crt:tt~1,,,,. ,.;, & r1tt.st0;.r.. T,"1 .. 197\i. Comp.:1rntivc,dl binlo~· u(1hc
281 mnu:i<. .,, 1903. A new Trypo11oso11111. nnd tlw dls~a,c c:aused b)· u.
kioeroplal'>ti4 fla:gc1h\tc-s. In: r.v'1snr,s, w.n.R. ~ E\'A~s. u ..,., (eds?. Biology
J(,z,r,111/ ofCom1Hm11;,,., Patho/r,~·. 16. 193. uftl1t Ki11etop/t.1.<t1tkt. Vol. 1. l.omlon, ,wadcmie l'ress.
28:nou~t!. "'"" 1,r,1, F.tutJc ultra.<tructur;d~d•; elJ'e~ prodults p•rd~<>
dro6'Ue, uypanocide_s sur des tr,'J'•nosomes. Rei,11c d'F.f;,·ag,.- ,1 d,•
:wi wn.l.l.R, ..... llt&>wu.u.. o.t:.& u ~•.., •• 1g,75, \ ~cro!ogl~nl ~1ud~ an
human r riuK(fs/e11,(t'1nf~lo~ u~ng m1cro~a!r Et.SSA.
,\/M11<111< vlrcrl11ml\'d~.1 Pros rrop,etm:i:. ?4, 3~1-39'i.
rn.,p,rnmrtli:iu mul P,m,ttw!ogie. ?6. 2,z;-151.
:!'U!! TftAtt.. J.c.:.t.. D'rli1U~s. G.O.~t' ' c:Ot...\RUfU.t.• r... :,.f-AII.-I 1-:. J.C C'UIU~lK, G ..
302\\'AUIMto T M,\\\'UW,?>. f'. & JE.\~Si~. P ... t'.)8$, T$et.it ttnd ll"jl),:tnosami~I~
• YA~eu1. c... 1990. Ev41u.ulon or"' t1e!4 lt'\t for
""U'-1'ROCIU + o
tn-panotol•ranc,, In young N'Danuicault.Acta Ttopfcn 18. ,J,-57 in caltle: The iD.1.frfo.cc of _grn7.in,g p.nn.ems i,l.Jld ii,. tmp<>rtnncr. /11;
Li>'<!S:<T('k P11x/11crio11 /11 Ts.-t.<4 .-lfl<!<·red .~r,.,rnf,lfnra. n11, Afrlco11
28.;. nlUL, J,C.~L. D ll'TI!RE!\', C...0...),1., n:ao:,:, .\.. 1'..WESli, o .• \lULUsc;o, \t, A:
0

Tryptmc>totemnc. /,ir'(fStOrk NelU.'(Jrk. Procc:edings of 3 ~k~dng. 23-2f

r-u.o, \t .. 1990. Effect or uyp:i,nos:omt infl.--ctlon, conuol or p3rns.trncmiu :\,1,..,mh~r 1~6; :\'llirabl, !,;ony11. 11.C\, II.RAO 1938. :-laicobi: Ens!ish
:t.nd control o!an~nilu d"\dopmcnt on productivit,·of X'Dam.:t\-:anft. Prc-s.....
..cia 1roplca. ~s. J7-15.
JOl w,uisoi. L'.\ .. C-..0,1sc.. S, AOOOUTI•OW.O, :>. )9':16. Pia,~ IC!,tOS:tCrone in
28S Tit.\lL 1,C,M.. U"U!l t:Rt~. c;,0.,1. & ff!A.t.f, A.,I,, 1989. Tt')-·pann10Jer~nC:f'.3nd
l'r:t'J}(l''lUOfllCI ccmgoltY1~.•-infcc1ed Togg1.?nburg go111s. ...\rch,,.~ cf
1he w1ue o! con.s-0n·l.ng,1i\'t!)tocl:. J.."Clletk. resources.. <"*1u>mr. JI. J1'1rlro!t>to'· J;, '9--lt.
805-al2.
30>1 \\'AR:O.F..,\,.\f.l \',\X or.~ 80S.'i;Clff, P., cmttlY,\, , .. ,:IJl>FX()f, I),. rmsr.v~...:.
2.86 TKAJl.. J,C.61.. MUIUI.\\", J.1., SON~~. X,, Jl811Q, Pl,t;., 1.>Vl(I'(~. J. t,, C,ttlff, O..
1'.f'. ~lliR.1---S;, w, 6 atuwxu.,. v.• 1~. Efficucy~(UJ.sl.-cuci<h.:·tre:tted ffluJt
1.985, llqrao canJe ll1'1ima.incd bychrmoprophylwo,; under U)'\>aTIO•
asn banil.'1' torc·lm'll.,tjon or,st'i1'C' fUc> 1cclt--ared ~ in north,easoom
somiasls ri>k.Jo11T11t1! ofngrlt11/111rttl S<ti''l<'WI, Cmnbrrdgi', 105, Hi-166.
7.hn""bw~. ,\lt1tiict1l (md vtrcrinoryl£11romo'!o$:>·. 13, 177...1&1.
281 TAUC:..... ,U..ftTS, t>.. \ICSt\,"oJ.\R.\, t.t.. c1.,n·" \,. AU -~(;HAM, k .. If- MA\, u. ~
coo..xrY. u,<;., 1ijg.2. Dlt,.-ct l..o!arinn i11 ,1ltr<J ur 1r,·11mw.1d11m bn(crl from 305 wcnsTat. 1•. 1939. Cndoq"losl$ b>' African 1rypano.so111os.. 1.
man and olh~r ,nimol,. and It, po1cmio.l ,·,luc for lh~ dlo~i,o,1. ~I Thrci.t·dlmcmstonal ,;ui.acm~ of lhl! cm!oc.1·tic: nrgnm•lll!~ in
:;ambrcn trypanosomos,s, Trm~q,((/Q'1.S ofthe flnJ'lll ,"«11'1;)' of Tmpicnl Trypm10.,omr 11111,01 and r. m11ga/c,"". Eum11,·a11 J<»11'11(1/ ofC,/18/ology,
,\!etlicm~,md flygic•"ct, 86. 62"'." ..-629.
-19, 29S-302.

2.881 UlC'-llk r_ \1,fl,, "Aun,. ,.u. /;,\'1ru.H,1AX. t .. 1988, t.o~" nr "·..Jii.ah!~ .antlg1..•.1\ JQG\\'ElJ.')TEJl. P. RVti'"tl• p,c;.w. & lll,\<.X~&.l,, .1 990. 1'hc: inwfn~tion of

during innsfonnation of Trypmiosomtr bru,·cl rhode;lmse from 1i).pm,ru·aum m'ucr!wflJ, nmibodJes m \'3tiam ~urfac:t" t1ycoprot~an~
bl.ood-.tn-.1m 10 procyc-lic form, in the hcl<c ny. Pmtr.{(tologj· Rht"Orci,. Joum~I ofCLII Srl,mn•. 96. 249-2S5.
7~.so--511. 3U7WE117,, 8.t,.f,, ,~w. H()'ll& or cur..~t,m In: ~IUHl<iA\, tl,W• .- PM'l'!t. ,,·.11 ..
189 UROUHA.R ,·. <,.M •• 19,h'>. Thi! pathogenesfa; amt lmnuuu">lt~· of.AJrk:!n {odsJ. 71,e .-\jnam T1)7><1J>o..<<1ml11.<fi. L<,nd<>n: G,·Otgfl Allon and U1m'inr
U')'J)illlOSoml:isis In dom.1dc anlm.1L~ 1'ra1uat1lo11SO[1lu:Roy11/ Soc1•1}' Mint~u1· o! Overw:.u Otwelopmenc
of (ropirnJ.\ft•d/c/11, lllld fi),giPJI~. ~~, i26-729. l.i!(lin ._ign.:dlins nr muturntion Uf
30g WCLDUP.'\, !'t•(;, 4.~t..'\UUl IS, I .• 19$9,
390 URQUU-.\f(r, G.:'1-S. & 110Qt"6S, p,u~, 198i, .·Un.can U)'t>anosom.iasis. /11: T. COligQlmH Tnfoctfon~ In ci,c~• •\11.vllc,II and\ •·t.:rlh<Jry llmomo((©', 3.
sou&SS,·~ t:.J.L. <ed.}. /mmu,rt Rt--sJH>rM,\t !n />(Uasirlr lnf«11ous: 141-1.t:\.
1,mmmotom·, hnm,mo11ntltoltJGJ' ond lmmwwpro11hylra.i$. Vol. n1. 309 wn.wc R.T. o. r.:uuMO, n ...,., 1.98.3. 1>el'$istemre ofRc.,enU in c~uic.. tropfro(
Ptvro:n,, Bot4 Raton. Florid:,; C:RC Press Inc. .4nflnd/ H,aJt11 tmd 1'rotfu1;1io11, 15, l4~iSO.
291 V,\...'\' Uf'- 80:S-'('IU-·, tt.. t9$7~ The conu-ul of Glu1,tl1111 m<1rJitmis mur,ilnm
311) Wl U.Ul a+ T., CHUMO, U,.\., An()yq, \\., l,;(W,\TCJI, 11,M., '\1\\0~Gf'I.\, f',,:\,;.
:l)iptera: Gtos.lnidac) In a settled area In Pe,aul:c O!Airlct iF.Mtcm
oPlro. £. , .. 19&,,, ffMniormAgfc ~)·ndromc in C<lulc ,.,,;ot1n1cd •,,illl
Pro,1nce. 7.nmblo) u<lng odour•bititt•d mrgc1>. ()r1d11rs.rvpoor1 /011mal of Tl)·f}(t1tf»Qm(1 ,,1,v,.T infotllun. J'roptaJI.-immnJ llehlth and Prorlm•tfon.
lcll:tmoryResearcll. 64, 251-25,.
15,9S-l()l.
~ v.,:,; n1, K0)$r.1U-. 1•... ~HU'-IU,\. \\'.,I ,,,,KtlJ\Mtlf.k,\, r .. ~ . The
311 \\'EU.tl"f.. Jl1l .. CHU:.tO, U.A .. ffhAJ-UMJ\'. \l.f .. \t\\·,,c1, J.~ .\$.LVlt,A.. 1989-
disiributlon and e1,ldemlolog)' ofbo\ino IJ)1>a1011omo!f; In \falew!.
l'tt~r/11111;• Pnrasllo/ol/.)', Bff. 163-176. Pre,mulng '"·""'"' of Rhude<lan -Jecpinptcine.., palfei1LS in the
t.untl',.., \'allc)·, '-•n)'>, A1111ttls ofTroplwl ,\/('((/rlnrmul Pnttt;/10"'1{;·,
~~ \'.A:\' or..~ Ui>lt'-CJlE. 1•. t. nuorATf.AU. L. tgg8. 1 he, clft•CJ of<fc1tl!methrin 83. ,J.-89,
pour,on opplicd to caulc on the 1mru;ma,ion of bovin~
31l \\'fl.UH--"· t .. UO~!>U-"YMC, \\,T~ lo.:OVAl( .H, Jt.\1., aux..u.. :\-1.S, ~ 01cc;.s.. ( .. L.
tr)1iano!.Omos1s. /l,tt•ut d'F.lt•i·l18'-' 11: cli.t .\l€1h"'Cln,• L-'eterinnir..•de, Pays
tropkm,x. 51, 12:1-12<1. 19St. Try.1Kmo:-,m1a conl!,'CJICttSi!. ~atuml :lndaC'quJred rr.sl>uu1ce in 1hr
1,o, me. fxp,:rimentnl Pnmsitology. ;z. ?IS-~2.
29..\ Y.\Nm-.., UOSSC:;K~ 1•. & MUOl!.'t';J'. t>.. J9:97. Pre,...J1en~ ot
tSe(<t;•transmluerl 1ry1>anosomosis nloDjt the casiern/nonh cas,~m Jl3 WLU..l)f..., H.T ut\flDUX, M.J., CHUM(), U ...\., J{0\',\1Cfl, ft.M .. \\'"tt--. M.\, 0.

bortk: of limbabwc. Zimbnbu\• \!tttfi1zt1')"]011r,wl, 28, 49-S9. \\'\'JUJU'. l).t•• \fWAl'oGt. J•• HO'\O:. \\~L.~ \\lUL\)t:>.. J.~.. 198g.. Ccn•brnJ
U)'P'Ul'"omlo:ds !n natumUy lnfec1cd cattle In thij l.ambw<! Volley. S.,1;1h
295 v.\."lo:,DW. !1"0$.)(:1-tf, st.. MUur..,cr;, o •• MVIL.\."I:(;,,, J. R ~0-ft.\',\L. , •• t99!1, Thc.1
:\)1107.4. .:cnya, ,\mw/10/ '/'mplra/ .\/e,/;,:m~a11d Prm.u/tology. 83,
p•rasltological onrl s<rplogic:ul pm;;Jcnce ort.,C1.SC-1mn,mhted bo,ine
151-160.
u;pan<a;.moos.i, in Lhe ~a,tem C:aprivl !Caprh i Dl~tnct, ~-amtb?a:
011d,wep<1or1 /our,w/ of\ ~ll·mwrr R,s.:at,·/,, 66. 103-111 ,\1.: w~. fi _\., 19'7::L 111c imponnnc.~ of mi.thanlt·il 1r.,.n~nd/li,.,..;-lun in the
~96VAN D£~ uo-.~J1E, r." "-'l.\.~. c .• 199;. Th~ impon:m« of caule-a-~ a food opldcm1olog) or nog\lnn: A rc,iC1,. froplcal .,nlmal 1/rnlth ortd
Pr<Nfortfon .;, 74-88.
source for Glosslna morsfwn, mor11wr1S ll'cHwood tr>lp<cra,
C::Uus.sinldae> In Ka;ctc Ob1rlcc. ~'ftcrn Pn.wim;u. /..umbta. .-itln TtTJJIIW. 3,;a \\'HL.", • ..,\ .. 1,16.. ,uhg,.mu.."' Mf'!,'lJl')tpc'WUm~ In: 1,U)tSDJ:..~. ,,.,ut. fo' £\'A.'"l'lo·
65. 105-109, n .., .. ~'d< ,. !liof~· Qftf:d.:111e1op/as1ida. \'ol. 1.1.ondon: ,\c.,d<infc Pt,•.s.<.

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296 5ECno:-. 1\\Y>: Protozoa! diseases

316 , ... t'.\lO;\". c ,1 .. 19:.tb. ProtQ:oo/,>g:?. ,t

\tam,nl for i\f(•d(,·111.\lt.~n clu:.rntiproph~·luc11c;- t1~l·1u-.. /11: >1uuJc;,\ -.. u.w, & J't>11s. \\',II , eti~ Ti:\•
i ru:-rinnrlrrm, and7.colog;sr$. \ol. I I on\l(m· U.1Uier-.-. t ,nd.111 o1nd Cox. AJr/((m Tr.,7>tmo,n,ni1),W¥-, t.ondorc G<-o:~· AftcnAnd Um,uli'-Unis.t~· or
J1-:- ,,un •.t...\\\'. o.o... 1~-- Lu«- ,tag.4! T lmu ,•I fnfccUu1u .,nd cffi("~t:1 Ch·~N!a, 01.'\'C'lopmt•ni
dlmilnuion nr 11)1)<ln(IClda! dr11~, ti.HAI> ,111111111/ ,<1,n11_rtc 11,·plJrt b[tlw 316" \\'ILu,,,sm•~•·· a9;6.. Chcmo1hc:rapyof \fri<'t\n :ryponoscunLl.Sl,... ltop:cal
Jm-.">f1:ntitJ1ml 1J.1lKJuut,r;·fur R,..~arch o" Aniuu.,t m~,,~. ,.urobt U1.<\"lt.k.• IJt1Ilt:i11. 73. .531-542.
c;em•ral Prin1crs Lid.
32; •,,nso!\ .,.:. 1909. V:tlu(> of lhl" indirect 1luor,.scc1tl{UHibody
~\.:.8 l>.ll,.. (..1\RDt'Xat. r.R ... ).JUJU<~\'. \l,. 1988, ~Mr4VO~dar foci of
\\.lHJ U,i,\\'1.',
d~tctllfJn tcu :IS ;isrrot~i(.11..1.ld tu dtagn(l... t, r,f Gf1J'5-lna,1r1.\Mtn1ttN1
T1)·panostJ1wi ,,fr,,u in go:it<· The Ct.".ntr.al nc,,·ou, ')'U?tn and ~QU\"(l\t'i.
bo\·ine u~·punn~nmO)i"· Tropftlll .!\11/mnl Jfrnlth 1111(/ /ltrNllll'llOn. I a
humor nf!he ere as po,.,,,ti.i ,aurw, of rclnps" in(e-1ion a lier 89-9.S.
chcmo1lter.tp} P11111>/rol<ig,,-. 97 Sl-61 .
li!tt. \,·1t..s0!\. ---1 ,·usM'\'.(Ut\M. ,1.l' , l~:'1. hnmunntot:1(~1 u~p,•ct~ f1rl,0\1nc
319 \\'HJTrL.\\\'. O..D•• G,\lllT, t .\. &MOl.)IES. ,.u.. 1991. Ut."\'tlopmPnt ofnn L

tl')-µ1t110'.\.(lmia,h .. I\ P.m,•mi In th,: ptodu,·tlott oi cornmon 31\tJbodit"',

e-mwnic,Hnkcd immunfhOrbent ac.!on,· tor rhc: d~rcctfrm and
Trc>J1f(·t1J .-lni111al /11.v1llh dutl /1mclur.titm. 3 , 133--139.
m<!la~uremcnl of the rn1>~nocldi'lJ drughomL•1.tmldnuu i ~.1.rnorinl in
,:-anlc. fft.(tftrch In \·'t•/t•rurar,,• Seft•ncf, Sb. 185-189. :n!f',\IL'\O'.\, ,.1 . l'.\Rt,. I., I 1Jt1't~. \.G.. 0\., t::.J..-4, C.RA\.. 1'..k..1976
Ob~tt\1'1Lioru, on a hctd or bt.'\•f cattle mntn1-nmi."d 1n .a 1gew arc:t.
320 wtt1n..uw. o.o.,., JC.,Rarr. 1 .. 1!,85. Cok·s.uo11t:uhf1.."T nt antlbodtt:) m
Troplt:ttf,.\11t111n/ lh•t1lll1mut PtndllttUm. 8. 1-12.
1,ypnnO'SOnm l)ruttf 10 tolli4Amlflb.$ tie /(I Sorlt!li: &rl~ tlv .,fl.•(11,'(/ll~
Trop,enlu, ss. 199-ZOS. no ww 1• t ~ .• l9:0, Th1,\ hac,n,all:)crit i:cntr1fuge h.'(hniqu'-' fur 1ht.' dt:1g:nru.is

;21 •,,·utn.s101 ....1-., ,,sa.

lbe con~ml of D.n1mn1 tl")'P-"noMimm"" 1n "'•ny,,. ,\ of..\fr1can tr')'J).1no11,()n1ia\iS. ,-\cut 1ro11Jw. 2- 33,i....300,
papc, propared for !ACFOS)'l11posium 19oll De1>.,nme111 or I •termary 331 \\'POUfOU~L M,I ,J., IUIAI It\, lo.,. \fC.:'.\',\\JA"--\, J,f. f.t SII..UlOSC;\\f I,, 199.:>,
'i,miccs, Kenya. Tr;11anosom1• mfooiorh t:1f cJw lbc.L,'-' fly Ci/0$SillC1 t'Nillitlipc$ in :be
~.a ,,·urrE.~mr.. a:i..r .. l96:t. lnttli)rnum, t.H.•1,\cr11 dnJ~. 11')1'•1no).(1mes anc1 Luani1wa Vrtll<fy 7.arnbla, lmmrat1011n/ 10,ir,111/Ji!t l¼mtsilo/o~·. 2~.
cattle ln the 11-eh.J. Im 1',oom,·;,, t..r. ik ~''"'°·~~Hut. " H .. te<b.• lJ,t,g,t, •tS'i-9!.tJ.
Pmwlt,'tmr<l NaStS. London: J. and I. o,urchnt. 3..u Wf>'()tJ1n11,1 ~ ,,.t
.1 & 11/'\RCn(W'f J.W ... 1996 on the in1\!'rprcuttion of
323 \\.1-10. 1979, l'h~ ,lfrlcw, Tl)'JWloJomfa.\dS. Hcport or u jnint. WHO t..x1x.on a{!.e-pr~afon(e curves ior trypan0$oml" Jnfecuon-.. oro.etsc m~
C,,rnmllll'i> uml FAO E.xpcn Consuha,rw,. 1',..:hnkol R~port ~,·rio< 63'\, Pm11utology. 116. J.19-156.
Gmuwu. \\'mid I ll':ll!h Ori:,1nlr.,linn. 333 \\T,OIHOU'-1 ~,.r.t .• llt\RORmT, I,\\ l 11(.!\ \M,\Jlo\. J,I, 1993. Epidt.1mi01os:y
32,; w1w:i-, x.c.. l970, tplzoouoiogy of crypanosoml~:,;li In lives;tork in Ej.)l of 1rypuno~o1n<.1 infcclions o(tM t~bl? nr GltMsiua JN1/lltliJlt!J' m the
ttnd Cenmtl t\friea. hr. .\JUW~·'-'"· II.\\.'. Ii l'Or rs. w.11.. ,,t.-d~t 1'h1•:1Jfirnn 7.ombc,J Yiultv. Plrm.<110/ogJ', l Oil. ~;9 ... s;.
TfJ'J)(11l~'omW..tc.~ l.ondoll! Gcor.,_~ AIIL·n and Unwin. ~h.n1 ..1ry •1r 334 t\\Hc;,,nn1. r.. _. 1t() nri1r-u, o., 19[$;. Thf o~currcnrc o( Tl)'JN.Jtl()J-Utufl.
0\'1!Dtas Develop.mun,. 1
\amwmoua>J simlO.t'ill thl.!.ccrcbro.•ipirnd fluid t>f tfamL~dc pl~.
32.ii \'tTl.11,\1.W~. 1.. 1910. Nt•\·iewof fhtmollu:n1pt!uu~ .:antl Auasuo!t>gJ.· R~·,·«rrh. i.t. H.9-480.

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 13
Dourine
Synonym: Slapslekte (Afrik.)

AG l.UCKINS, PR BARROvVMAN, \V H STOLTSZ AND JJ VAN DER LUGT

Introduction Schul1..f'° A shipment of horses from Germany to Namibia

may have brought the disease to the subcominem.:13 Schulz
Dourine is a chronic contagious d isease of horses. mules believed that dourine was brought into South Africa by
and donkeys. Ii is characterizt>d by swelling of the external donkey Jacks imported from Canada orb)' transport horses
genitalia and subcutaneous tissues of the vemral abdomen. returning from Namibia during the Herera wars.;a Dourine
cachexia and, b1 somt> cases, involvemem of the nervous has a S(:attcred distribution in -"'amibia, Botswana.
system. The disease is caused by the vcnercally transmitted Swaziland. Lesotho and South Africa.a. •~. io, 6!1
procozoa:n parasite Trypcmosoma equiperd11m. AILhough the disease is not of grear economic impor-
Tryprmo~om" equiperdum is essemiaJly a tissue parasite tance in South Africa. it$ continued presence nccessirate~
which has developed the ability to invade and sun~ve ln the strict surveillance of horses intended for breeding and
genital tract, rhus enabling direct transmission from one exporr purposes.
host to another. The disease consequently differs from othetr
t\frican rrypanosomoses in that lt does 1101 require an ar-
Aetiology
thropod vector for transmission.
Dourine was describes as early as '" 400 by Chiron. a Tryptmosoma equiperc/!1111 is a salivarlaa rrypanosome of
Byzantine veterinarian,"° and is probably one of the oldest the subgenus Trypanozoon. which includes Trypanosoma
recognized diseases of horses. The disease was known ro the eutmsi and the Tlypcmosoma brucei complcx. 17 The
Arabs and horsemen of :'fonh Africa for centuries before it mammalian blood forms of the three species have similar
was reported in Europe in the late eighteenth cemury. 30 The morphology and share common somatic antigens. which
dissemination of Arab horses for breeding purposes may pr!'vem their distinction by most serological tests.s2. ;o
have introduced <lourine to Europe. from where it spread to Trypanosoma eq11iperd11m is closely related to T. e11a11si. the
many parts of the world. Rouget. in 1894. was 1he first to tausanve organism of surra. which is endemic in :Xor1h
demons1rare a trypanosomc in 1he blood of an infected Africa.28 and it Is possible that they arc derived from a
horsc.3 5 while the first experimemal transmission of the common ancestor descended from T. bru~·ei.l 2 Evidence
parasite occurred five years later.56 The organism wns tha1 ther might belong co the same species comes l:'l'om
named T,ypa,wsoma eq11iperd11m by Dofiein in 1901. 18 isoenryme characterizarion that has shown isolares of
Control measures based on clinical identifica1ion,a. and '/'. equiperdum and T. e11c111si from different geographical
later by serological tes1ing."fi have eliminated the disease regions form a homogeneous group,3n and analysis of kine-
from many countries. Intensive campaigns led 10 irs eradi- toplast DKA, nuclear DNA and molecular karyotyping has
cation in Canada68 and the final pockets of infecrion in the confirmed the closeness of their phylogenetic relation-
US..\ were eliminated in 1949.3G By 1960 most European ships.39· 40 • 41· 71 A major difference between the two species
countries were free of the disease. 19 Ourside southern Africa is in the presence of maxicircles in the DNA of T. eq11iper-
dourine still occurs in certain pans of North Africa. Russia, d11111 and their absence in T. euansi,5 1 so ir is unlikely that
the YLiddle East. and Burma. 19 Recent outbreaks have oc- T. equiperdum could have originated from T. evall$i, as
curred in ltaly. 9 suggesied by Hoarc. 28 as this would have required the
Douri:ne was !mown to fanners in Griqualand \Vest in reacquisition of the maxicircles.
South Africa for several years before lt was officially re- T1ypa11osoma equipert/11111 is monomorphic and typ-
corded fonhe first time in 1914.64 During 1he same >'Car the ically occurs as slender forms 15.5 tO 36 µm long. with a
disease was reported from Namibia b>• Maag as cited by distinct free flagellum and a subterminal kineroplast.
29i

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298 '!cm<" ,wo: Pro102oal diseases

Occasional stumpy forms are seen.29 The organism is only breeding season when the parasite is tran~mh led from an
found nauu·aUy in the definirive host and multiplies asexu- infected stallion 10 a number of mares.5ij Trypanosomcs
ally by binary fission. Jryp,mosomn eqtdperdwn strains may be numerous ln the vaginal discharges of affected
ha,·e been maintained for experlmemal purposes in the mares or in the urethras of stallions, part icularly during the
laborarory In animals such as mice, rat$ and guinea pigs for early stages of the disease. Although transmission is erratic
many genera1ions:'9 · ,;,; These strains have enhanced viru- and wiU not occur at every macing. ii Is more likely to occur
lence and produce acute disease when used co infect when a recently in£ccted animal i~ involved.6 • ' 9 In Souch Af-
horses.49 They have a tendency to produce dyskinetoplas1ic rica, the disease Is seldom seen in Thoroughbred horses. as
s1rains. panicularly In rodents.30 breeding of these animals is usually done underscrict super-
Although T. equiperdum is es~emially a 1lssue parasite, vision. Great care is laken 10 cest breeding animals before or
paroslraemia does occur 21 to 23 days after natural infection during !he breeding season.
and may persist for three to four momhs. Thereatier iris dif- Occasionally foals become infected - possibly during
ficult to demonstrate the parasite in the blood of infected parturition (from vaginal d ischarges) orb) drinking the milk
horses.6• 7 of an infected dam.511 These foals remain serologically posi-
For many years, T. eq1tiperd11m could only be maimained tive and ma) transmit the disease when sexually marure.:;.1
by serial pa.,sage in laboracory animals <>r developing chick Non-infected foal~ may also become serologically positive
embryos:1'1 Recently an in t>itro cult\Jre system has been de- by ingesting colostrum of seropositive dams. but they
veloped in which a laboraco1ysnain of 7: eq11i/J6r-<i11m c-,in be lose the antibodies wid1in five 10 sL't months and are 1101
continuously maintained in a semi-defined medium, whilst infective.1s
still retaining its morphological and biochemical charncier-
istics.5·., These in 11irro systems have been utilized in stud-
Pathogenesis
ies on the efficacr of trypanocidal drugs 11 · ~ and could be
used also in the study of the physiology, biochemistry and Most pathological e!Tecis of T. l'(Juiper([um infection in the
antigenic variation of the parasite. hos1 are characteristic of tT)'J)anosome infections in generaL
particularly those of Lhe r. brucei complex (see Chapter J 2:
Epidemiology African animal uypanosomoses). :-cau1ml transmission of
the disease occurs when the parasite is deposited on che mu-
Dourine occurs in horse~. mules and donkey.s. Improved cous membranes of the genitalia during coitus. Like the other
breeds of horses such as Thornughbreds tend 10 be more members of the T. bruc~i complex. T. eq1/iperd11111 has the
suscep1iblc to the disease than native horses. donkeys and ability 10 invade intact mucous membrruws and lymphalic
mules. which are often subdinically or only mildl}' af. v~ls bu! also has a precillectlon for connective tissue. 22
fected.im lt is a true ,·enereal disease. Narural transmission Experlinemal infection oflaboracory animal~ has shown
or the organism occurs during genital comacc berween ani- char the major route taken br a labor.atory strain of
mals. and in£ection may occur from stallion to mare or vice T. l!(/t1iperdum from an intradermal inoculation site to other
tiersa. Biological transmission of 1: equiperd11m by tsetse locations wa..-; via the peripheral l)mphatic vessels. lt ap-
flies has not been reported but the disease can be uansrnit- pears that organisms reach Lhe regional lymph nc,-\e vi-a the
ted by subinoculating large volumes of blood from infected afferent lymphatic vessels. from \\'here they reach 1he blood-
horse.i 10 susceptible horses and donkeys.6 • ' 9 The disease is stream. Only after some time do they leave the node via the
not limited ro certain environmental conditions.Ii:! as is usu· etfere111 lymphatic vessels.63
ally the case wirh the tsetse fl)' ·transmined 1rypanosomoses. Trypa110$om(I equi11erd11m mulliplies predominantly in
The presence of dourine in a herd can usually be traced e>.,racellular tissue spaces. II i< seldom found ln peripheral
to an infected animal which showed no clinical signs at 1he blood and probably only uses the bloodstream as a means of
time of imroduction. Due to the in~idious nature oi the dis, transp()rt from one site to another. York, as cited by Good-
ease. it may take several months before clinical signs appear win.22 reported that the nutrient comem of extracellular
and by this time the disease may already be widespread in a fluid may favounhegrowth of the parasiles. To some ex,em
herd. Many infected animals are subclinical carriers26· ~,a this site ma~•also aid tl1eir evasion of the action of antibodies
and they can only be reliabl) detected by serological meth- present in circulating blood.
ods. Sexual acti,'ity is impeded only in cases showing ex- As in other uypanosomo$eS. T. eqrlil'erdum elicits a
treme emaciation or severe oedema of the genitalia. marked immune response in the host. 7 Antigen-antibody
The dlsease is more likely lo spre-ad in herds or areas reactions arc accompanied b~ the release of a number of
11·here indiscriminate breeding of horses and mules occurs. physiologically active substances including hisuunine and
Donke}'S used for the breeding pf mules ma-y be particularly kin in. The la1 cer may be responsible for the vascular changes
imponanl in this respect. Mule foals may become infected and oedema after the initial histamine release. The exact
in the same way as horse and donkey foals (see below). cause oflhe often observed anaemia has t1ot yet been dernr-
Transmission of the disease is usuall)' limited 10 the mined. A combination of the resultam local hypoxia and the

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 Dourlnc 299

pro1eol~1ic and lysosomal em:ymes released \in the host", Clinical s igns
bid co contain the infection! probably contribute largely to
che exiensi\'c damage caused to connective tfssues.:?2 Dourine in southern Africa1' :w ?,a. f,.; b similar co bur u~ually
·\lthough during the cour~e of infecclon rrypanosomes mlldenhan the disease in Europt>31 ~, and Xonh America..,.;
are constantly e.'tposed to che Immune responses of che host, Tho course of infection ,md clinical signs \'ary conslderabl~.
the} are able to evade the deleterious consequences of the depending on 1he \'imlence oft ht> strain and the susceptibil-
immune mechanism~ h} a process known as antigenic ity of the hust. After an incubation period ranging from one
variation. Trypanosonws are coverc-d by a surface coat con- week 10 a few mont hs.2"· .Jo. '"'8.;.:: the disease may assume one
sisting of a single protein kno,, 11 as the variable surface gly- of three forms. namely an asymptomatic, imerstitial or
coprotein NSG). This is a powerful immunogen and elicics oedema1ou~, or nl!rvons form. n,c ner\'ous form occurs a;:.a
the production of protective antibodies, but as the ho:sr re- sequel to rhe inrerstltial or oedematous form and is im·ari-
sponds co one VSG, trypanosomes ex-presslng another VSG ably fa cal
become dominam. The dltTcrent VSGs arc expressed In a South \frican strains of 1. l!(/11iperdu111 tend 10 induce
general order. with certain VSGs appearing • early in infeccion aS)mJ)tomaclc or laten1 infections. panicularly in donke~·s.~b
and some later (sec also Chapter 12: African animal trypa- These animals remain seropositi\"e for se,eral years and arc
nosomoses}. 4• 55 capable of trun~mitting the infection. Clinical sign) may
lmmunodepression is a characunlstic feature of infec- be precipitated In IMent cases by stre$S such as that caused
tions with pat.hogenic 1.0-panosomes in both domesdcated by unfa\'ouruble weather. physical ei.hau5tion and mal-
livestock and laboratory ho~ts,...• and it may well be an im· nutrition.~6- ·1' 1• ;a A moderate anaemia b often observed
ponant mechanism in the induction of disea:;e. lnfecLion re- in animals which show no apparent clinical signs. as well
sults in reduc<>d primary immune responses 10 non- as in tho~c with chronic oedemacous lesions or nervous
. 6,'-!.·-·~
tl)'Panosomal amig<>ns and diminished T cell proliferation signs.
and cy1okine proch1crion. 1 '- ;~ C..omplement components The imerstlrial or oedemntou~ form is usually a manifes-
are reduced, inappropriate antibody responses are made tation ol chronic infection, lasting sc,·eral months to se,·eral
and there are dt.'foCt$ in butype w,itching and impaired B year~. A common dinlcal sign in both ,tallioru. and mares i~
coll memorv responscs.2 · 67 In l'!~perlmcmal infections with progrcssi\'e wc-ight loss, with the animal eventually becom-
T. equiperdum, a rheumatoid factor-like subscance destroy· ing severely emaciated (Figure 13.1 ). These debilit.aced ani-
ing lgG antibodies and an unidentified soluble substance mals are prone 10 infection by other pathogen~ and
cau$ing functional hllpprcssion of r and B l}mpho- parasites, and man) die from secondary causes.53 TI1e incer-
cytes have been idemifled.~ 33 fhese factors probably stitial form of th.c disease mayor may not be associated with
predbpo~e the hoM 10 ~econda.ry infection~.za. 53 oedematou> ~wi.illings in tht! t.'arly ,tages. Stallions show
The onset of the ncr\'ou,; form of the disease appears co va111ng degrees of oedematous swelling of the sheath. glan:.
coincide \\ich the presence of parasites in the cerebrospinal penis and scrotum (Figure 13.2) and m sc,·ere cases. the
fluid CSFl. The CSF ma~ repre,ent a favourable emiron- oedema may extend to the perineum and , emrnl abc!omen
meot for the parasirc as the immunogobulin le\'els are much and thorax (Figu re 13.1 ). 5uch s,, 11llings arc neither hot nor
lower than in plasma, inflammatory cells are fewer and glu- painful and oflen disappear after a few days. only to recur
cose leveb ~imilar.Z The neuropatholoh'Y of dourine differ:. more severely within two to three \\eck& The oedema may
from thar described for other trypanosomoses. including persist for up 10 16 month$ after which the swellJngs often
T. bru<:ei infection in hor$es. 13 The primary changes in become permanent due co their Ofl,'81li7.ation by fibrous
T. eq11iperd11m infection$ com()rise cellular Infiltration~ of connective til-Suc.6 The penis ma) prot.r ude and occasion-
spinal nervl!'S and some major nlln:e trunks, but without ally there is prolapse of the urechra. 26 In mares, the ,·uka
much iiwolvement of the central nervous sys rem.,\ possible a nd perineum and occasionally the ventral abdomen and
t?Xplanation for the site of these lesions is the drainage of uc!der are ~·wollen. In some case~. a ,iscous vaginal db-
CSF which moves along che sheaths of the spinal nerves. i.e. charge ls ~een which m;iy be copious. Hyperaemia of the
antigens of l0'Panosomcs presem in CSF may elicil a hose mucous mcmbrm,es of the vulva. vagina and clitoris is
response at these sites where nerve tissue is no longer sometime~ accompanied by a few nodule;, and ulcers which
protected by the blood-brain barrier."· 7 heal h:a,~ng whire scars. Some mare~ show frequent mictu-
Passive proteccion against T. eq11iperdu111 iniectior1 by rition and behave as if c:ominuall\' in oestrus. ln other coun-
1ransfur of immune serum 10 ,usceptible animals has been tries. signs such a~ ulceration of the penis. frequent
demonsrrated.'11 and passive transfer of immunity from in- erection$, and ocular manifcsratiOn\ such a~ photophobin.
feCTcd dams 10 their offsprinii may thus occur via colostrum. keratitis and corneal opacicy have been reponed, but these
The efficacy of thb. procection is. however. limited by the have not been rei.·oroecl in southi;rn Africa. Raised unicarial
ability of the parasite to rapidly change anrigeniCall} during skin plaques. 40 10 50 mm in diameter. referred to as ·sih11r
rhe course of infection. Resistance co reinfecuon is limited dollar' plaques, have only rarely been recorded in animals
only to homologous sirains." 1 suffering from dourine in southern Africa.6 · 58• i;.i Early

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300 s..:nox J\\O: l'rotowal diseases

fig·ure 13.1 Horse showing

emaciation, ventral oe!lema
and swollen p1epuce

Figure 13.2 Horse showing

scrotal and J)'eputial Delle ma.
and phimosit

reports from Europe, North ,\frica.50 and India described are often set wide apart and the feet are frequently dragged
these plaques as being of common occurrence. is Although along the ground with knuckling over of the fedocks. Ani-
no1 frequently seen in Canada, \\lmsonliti consfdered chem mals ma) have difficulty in rising and lying down. ~love-
to be parhognomonic. ment becomes increasingly difficult and affected animals
:'-len1ous disorders occur in some cases and may set in are therefore reluctant to do so. Eventually they go down in
soon after the emaciation or the oedema appear or. al rerna- sternal or lateral rccumbenq . Although some cases do
tively. some rime may elapse before their onset. Inftially show e\idence of apparent improvemem, the condition of
there is acme generalized or localized hyperaesthesia of the animals suffering from the nen•ous fom, deterlormcs
.skin.31 Lacer, sensitivity decreases and i~ followed by paraly- progressively and they invariably die.26
sis of motor nen•es. The motor involvement originates fn the Early repons from Canada and Europe desc1ihe facial or re-
perineum and hindlegs and may eventually extend to affect current nerve paralysis affecting the lips, nostrils, ear;: and
the forelegs. Affected animals become restless and move un- tllroat. 31 • MiThe,e findings are far Jess frequem in southern Af-
easily from one hindleg to the orher. There may be ~tiffness. rica but they have been reported from the Cape pro,'lnces.1,11
weakness or lameness in one or both hindlegs and the Pregnant mares may very rarely abortG-1 but usually carry
animal may have a swaying or straddling gait. The himllcgs their foals to full term. i;_ ~8

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 Dourine 301

Pathology The clinical signs of dourinc are not specific and, I\S
many cases in southern Africa are as~mptomatic,6• ~. ,q se-
Animals tha1 die from dourine are usunlly severely emaciated rology remains the only rellable and practical me1hod of di·
and anaemic, and changes similar to those found in 01her agnosis. The c:omplcm<.'nt fixation (CF test has been usccl
wasting diseases are seen. There is subcutaneous oedema. for the diagnosi., of dourine since 1915 and is still the most
particularly of the perineum, male and Female genitalia, and commonly u,ed serological technique.27 However, it is only
ventral parts of 1.he abdomen and chest. In longstanding appropriate for making a specific diagnosis in coumries in
chronic cases the oedema Isof1.en accompanied b) fibrosis of which other members of 1hc 1: brur:ei complex ore nor
the scrorum, prepuce and tunica \'aginalis. In mares ,.iginitis. present, since antigenic cross-rcac1ions within 1he group
merritis and cystitis may be presem. 53 =-:umerous cys,s occur occur. The sera of infected animals gi\'e poslth·e CF tests
in the uterus of some mares. and the ovaries !lte enlarged and from 20 to 30 days after infection" and remain positive. "11h
sho" cystic degeneration.Sit :'-lost of the internal lymph nucrnating tiue. for up to 10 years."° Sera from donkeys and
nodes. but especial!) those draining the ipguinal and pel,ic mule:. are more frcquemly amicomplernentary than are
regions. are enlarged. oedema10U$ and firm on palpa11011. On those ofhor~es. which are rarely so.611
cu1 surfaces r.hc affected nodes are pale and there is li1de dif- Other serological tests forthe diagnosis of <lourine have
ferentiarion between cortex and medulla.' been applied t?xperimcmally. n These are the emiyme·
In horses that e.xhiblt neurologlc.tl disturbances. the linked immunosorbent assay LELISAJ. and the radio immu-
perineu.ral connecti\'e tissues of the larger nerve trunks. par- noa~say. lmmuno-elecrro-osmophore~is. and fluorescent
ticularly those leading to 1he t!Xtremirles. are oedema1ou,. •· aml body 1c~1s. n,e results obtained shm, that rhe CF test is
31
• :;a or the neITes are transformed into fibrous cords fused stlll th!' most rt:'liable although 1he ELISA has a satisfactory
with the surrounding fascia.3 1 There may be a serous infil· concordance ratio whh the CF t~t. Di~crepancies encoun·
rration around the lumbar an<l sacral portions of the spinal rcrcd wi1h th!! 01her tests are attribmed to many cause,,
cord~· ;:; 5; ancl the cord ma~, be sofiened.26• Gu Including the subjectivity of rest inrerprcta1ion and differ·
:-licroscopicall}'· variable numbers of l)mphocy1cs. ence~ in antibod} populations detcc1ed by the differem
plasma celb and macrophage~ infiltrate the dorsal and test,. Caporale t>r t1/., 11 howe\'er, consider tha1 the ELISA is
11cmral spinal nerve rotlts, spinal nerve~. ganglia and the me1hod of choice as th<' amigen ls used in minute
meninges of the lumbar and sacral region~. and in more quantifies and appear~ to be more stable than the CF test
advanced cases. the larger peripheral nerves. notably 1he anrigcn. This \,·as ~upponcd b)' the work c,fWassall and his
sciatic and obwrator nerves. 6 · • ~• -'•· ' 6 • 6'l In cases of co,worker,,b, who concluded that 1he CF test was less sen·
longer dura1ion. spinal nerve roots in the choracic region sirlve rhan either the indirect nuorescem antibody test or
and the brachial nerve, arc also affected, nlthough 10 a the ELISA. They considered tha1 ELISA was 1hc 1est moM
lesser extent.r,·' Earlier workers referred to ·ctcgenerative !ikl.'ly to lend Itself to standardization, allow considerable
changes·. 'atrophy' and demyelination of nerve fibre, in automation and could be adopted eventually as an inter·
the spinal and peripheral nerves and the spinal Cllrd.i1 31 • nationally appro\'Cd 111st ror equine health ccrtifica1ion for
15 06 6
· · • sr 61 but Barrowman slated that the lesion~ In dt)U· freedom from dourine. ELISAs based on Trypano::0011
rine in southern Africa are largely confined to the penph· group-specific antigen have also shown promise for dc-
eral ner\'ous ::ystem. Marek42 d~cribed innammatory tec1ing ontibodie~ to parhllf:lenic trypa11Qson11:,~, including
changes in thr facial and trigeminol nerves. ·c eq11iperd11111, 10· ~n and tests based on defined antigeus
:'-licroscopic changes in affe<:ted lymph nodes mdudc l·oukl more readily enable standardization rhan tests bused
exren&ive lymphoid hypl.'rplasia. reticuloendothelial prolif- on the ust> of crude. sonicated antigen preparations. En-
eration and 1r:irying degree:- of lymph stasi$. ru; well as area~ zyme-linked immunosorbenr :t,;says for antigen de1eclion
of fibrosis in the more chronic casesJ (Ag-ELISA) have also shown promi!,e experimemally10 and
both .'\b· and ·\g-F.I.IS/\ tests have been used to in,·estigate
the pre,·alence of T. ,,q11i11ertf11111 ln horses in Elhiopia. :\
Diagnosis
commercial ca rd agglutinauon test for trypanosomosis,
The diagnosi!> of dourine is usually made indirectly. due to using formalin-fixed T. t'V<111si. b a,·ailable3 and might be
difficulty in demonstrating organisms in either 1he blood or useful in 1he diagnosis of equine trypanosomosis. fhc test
tissues or in discharges from the genitalia. It is generally fu- has not been evaluated for u,e in the diagnosis of
tile ta search for trypanosome, in blood ~mears because T. e,1t1i/)C'rd11m, but a ~imilar te~t. based on T. brucel gam-
their presence here is transient. altl1ough organisrn~ ma~ be /Jit•11se amlgen. has been sho\\ n to detect anribodles
found in recently infected animals. If an attempt to demon- in horses that show dinicat signs ofdourinc.53
strate them is made. a large quantity of citrated blood Diagnosb by subinoculation of blo()<l imo experimen-
should be collected and either be centrifuged or allowed to tal animals is not prac1ical as tl1e southern Afrfca.n s1rain~
sediment out. The lowest pan of 1he plrtsma layer is then ex- of r. eq11iperd11111 do not transfer readily 10 dogs. rabbits,
amined for parasites.6 -1 9, fi.l ra1s or other laboraWf)' animab.r. 26 6'' Infection of these

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302 "' ""'· ""'': l't'OlOZOttl d[sca~c><-

laborawry a nimals would occur more readily after passage blood-brain barrier. The use of chemotherapy for the con-
in <!quinc animals. bill this method is rarely used for the trol of dourine i.;; thus 1101 recommended as treated animals
diagnosis of the disease. 2"· 19 mn~ show clinical impro\·emem but remain carriers of the
parasite.
Differential diagnQsis In most C()t11,1ries dourine is a scheduled disease and
control measures are enforced through legislation. In coun-
ln sourhcm 1\ frica. oedema of the genitalia in hon:.es may tries free of 1he d isease. the issua of import pem,i1s for
occur in o ther conditions, e.g. nagana (caused b~ T. brutei): domesticequids is subject 10 certificarion by an official labo-
equine \~rnl arteritis. which was recentlr diagnosed in South ra1ory of negath·e i;emlogical results (usually the CF tesn for
Africa: coital examhema: purpnrn haemorrhagica. where. in dourinc.
addition, exten~ive pe1echial and ecchymo1ic haemor- TI1ere is no legislation restricting 1he movcmem of
rhages may occur on the skin and mucous membranes: and horses within the borders of South Africa and :,:amibm as far
lrauma oft he genitalia. . as dourlne is concerned. In these coumrics all 11utbreaks of
Conditions leading to chronic weight loss and emaciation, the disease are no1ifiahle h) law. but compulsory testing of
such as malnutrition, venninosis, dcnt:11 pathology, other equine animals tnll ~tallion$ and maiden and hurren mares
chronic infections (e.g. crypanosomosis) and chronic liver under the jurisdiction of the lockey Club of South Africa
disease (e.g. that occurring in scnecic>sis) can be confused m,tst be 1e1-1cd for dourine prior to the commencemem or
with, or mask. the signs of dourinc. The nervo11s signs asso- each breeding se;1sonl 10 control or eradicate 1J1e disease is
ciated with dourine should be difft•rcntiated from slmilar n01 enforced. Serological testing of animols is performed at
signs rec()gnized in a \'lll'iecy ofiufcccious and non-infectious tl1e request of the owners and is subject 10 them signing an
causes.such as equld herpe..~~ruS· l enccphalomyeli!ls, cervl· indemnit) form which requires them 10 ha\'e all positin~
cal stenotic myclopathy ·wobblers)11dromc'J, injuries to the animals destroyed. castrated or ovariectomizcd. :\emering
back and pelvis, and iliac rhrombosls. 1,; ma) lead to clinical impron•mem bu t animal~ remain in-
fected.58 Casmnion of infected stallions is not alwO)'S effec-
Control tive in pr(wenring them from nansmitling the diseast'. as
rhe~ ma) still serve mares for some time.
Several chemotherapeutic drugs, particularly homidium. The presence of fences may help 10 control the spread of
sunm1in. quinapyramine and some melaminyl amimony the disease, although stallions have been reported to serve
compounds may ha\•e some beneficial effect In clinically af- mares O\'er fences. even 1hose constructed of barbed \,ire.
fected animals, but none ha\'e been proven effective in the Due 10 the presence of \ariable surface amigern. on
sterilization of 1he infec1ion.- ;o The efficac) of most try- T. e.qulperdwn. the developmem of an etrecth·e vaccine has
panocidal drugs is limited by their inability 10 cross the not been achieved.24

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under fi<'ld condl<lun, in Thulland. !1111111.., If• /11 Sm·i,·t,· IMgNfe JO BlS)toP, s .. l'Jlll'l!S, I' '° I uc1uxs. ,\.(i.,. 199.;. Fl')1'J'/(,/lt(Mf>'UJ1 r(/lli/Ntdum
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T f."IWP"rllmn, T tc·m~· T~ r1JOtfai1.m.w and T gamb1('ti.1t•. 11t,· c,,ropran T r<1u11wrdum trlbmion. bioloQ·, Uitatmcnt '1ml ph)1ogcnctlc
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il!\Nltl\\")tA:'\, 1J c'..U'QK,\U ,·.P .. l\l,\\t.·ot()M1, 1.. t-ttt,_'C'.uR,,. t., m 'tAmo, .,., s,~v1,1 o.r..
clfnlc<ll ,lgn, and (h~mothernpyr,f douri~~ /Tnpu,10.,011111 ,,111/pMl/11m1 Uft.K,,,. r.... 19th. C:<>mv:irhnn .,r vunotH, tt.·,b for the ~crologu:aJ

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 Oourino 303

d!.agno11ls of Try/1antJ.,;<,ma equip,wlurn lnfocti011 In the hnt"\C'. 36 uvtxF. x .n.. t96! Prow~11 P"ro.~lt& oflJoniltJtfrA11imnlsnnd Alm,.
Qlm/l(lrlllit'<.' lmnwm)(otJ•. ~\1irmlliul<,gymul J1,fi."Ctfm1.~ l>ln>fl,f(',,. 4, Mlnnc-Jpolb; Surge».
?43-Z·l6. 3; u..,r.:,an. ,., 1904. l11e1t)p;tnc}\nn1cn(dou.'incand 1Nlifoh11oton,.
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doutine./011mnl Oft//c S0111l1A[r/rt111 l',·terinmy Medical ,\$$1Jdmio,,. lO. Prot:«dlr:g; ofa11 /11tl!Il:u1l01wl ,ympq,lum on dlt>g1101ri, a11d torurol
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IUS', l'.. R., WJ\:l"lU.\M. H,,~
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18
London:Ac:ademic Press.
.
um,u I:\. f., 19.01. Oio Pro:owi•n 1tf.l, Pnrasftt11 an.d Krt1nkheir1.,,uglr.
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,II MASI(",.\, cu, ... C'U.&SOS-. w.. 1990. Spcdfic probes rorTt)'f}llll(J$()"1(t
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.13 Mccuuv. n.~r.... Slitr7.. w.o.. 1971, Cliniropatholog,cnl >1udy on
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rm11sao1ioru oftlte RO)YII S0t:ltlynf7'ropi/:III Ml!dil'l11f an,1 ff.~"'"· &.I. )ollmal of\iwrl1111ry R""'""'''· 33. 141-Ho.
7l!Hll7 +.; ~1rra1w. ('...\., \\',\Ur...'""FR. '4,\ 1,L, HMTII, ... ,:. '-' \~(:).1MCHl1.,., O,<, •• l9:::l,ft
2:_'4 C.000\Vl~. L.G., GREEN", O.G .• Ctn', \1,\', Ii \'0-t.Ll!K, ,\,, 19?2 Prellminlll)' note on 1be wowth or frypm:ou,rr:n ;qu/;wr,/urr, in the
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f.J.11,rlme,1wl Pnr/rology. 53. ~o. ;1. n'i-22•
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immuni,arlon ng~ins1 African tl)'Jl~nosomlasfs. Trn11s,11·1/011S tJftlre ea.;c ofchmnicdnurine r,r mhl d!,? tl')it, ttruJ ccunp.,riso» of lhl!' !lame "1rh
Ro.111/ Stx:l,r;•ofTw;,ic11/ M.:dfc111nmd f~1gt,11v. 70, 118-f:!I. thos.< found In sl•"Jllr.g sicl<m,;s. P1«&tll11iv-(Jft/1<• Ruy,11 .'w<lrr,·. s,,rlc•
B. 78. 1-12..
:t.-i JL\tG, D.A. & w.so. A•.s~. 19,18. Transmb~fon of the South Afric:m "imlin of
dourin• to !nhomtOt')' antntnls.. 01:d~m•1""'" /t>1m111/ iJf\l,i,ri1111r;· ,16 MOTT, P.w.• 1906. Cll'lltgcs In the nef'\'>U»Y"~m produ~ In chronic
Scltll.cy><,md Animt1I lnrluirry. 23., 59-61. ll)'Panosontc !nfocuo,is. Br/rlsli Mtd/,:ol Jo1m:nl. 300. 1772-liT<
2.6 tf1l~NtSC, ~1. u.t_ 1956. /11timal Dl~fL.({!J. flJ Sourh A/rfca.3td cdn.. 3:outh ,f7 NIL H.Y.. f.A.SG. Y., \'E. w.x.. HUA..XG, r i &W.\XO, Y.l-. 1991. Studies on the
Mrk.i: C-..entrol Ne\\~.Agenc). ltd. 111 vitro culture or Trypnr:osomll ('(/IIIJJC!llum. Chl11t'ta/011tfl(I/ of
vcu1ri1:rtry MMfcinc.'. L7. 2-1.
2'j llti1tR, s.. UUCU!f.AA.Ml:-\'l:1'. I t.P,tl..A.... i:E SRO(;GL I...A•• \\11u.t\M$0~. C,L, r<oo:,.
J.A. S: .scmur;, G.J .. 198,5. 111~ use ufa $:inih.· e0-mple.mtn1 fix.a1lun ce.:<1 .,u ort1m1 ,.,. \IOUL1ox, 1,b., •!13:1. Splcnic cell 1mmunosuppre5'1ion In
1echnlquc in bov!M bn,tcllosil.. John~·s cfueose, dourlno. equln~ cxpcruncma.1 trypan(l~Omiasts. Tropir:<J.I Vi:urri,mrinn. 1. 92...S7.
p!tor,la.;nio,;i, onJ Q•fewrsorology. 01:di•rsiepoor1 /011mt1/ of l'et,rlnnry 19 PAtu;1~. 11-,.
19-iS. The dcmonstNtfon and lr,;msmiss~on of 1he South
f/es,w"11. S2, 2.79-~2. African strain of lrypa,m$Qn>a •·qtup('T<ium ofhnr!'o~5,. 01ulert(('f,QOrr
28 HO:\nlf. Lt\~ 1967. Evolutio1to1ry trends lnmnmmn1ian 1ryp.1r.o~umL-s, /01,rual of \'tten'nat),$.cif'liel! rilld A11i11rnl ludoslfJ'• 23. ·11-57,
AilM1trt'1. In Pnrmi1ofogy. 5 . ..;';"--85,. so f'2t\S~. u.T. l904- Sur~ and dourine. 1J:,, ~ ·e;mnn,;i1Jnunml. 9. i87-20D.
29 t?OA:R:n. C.A•. ;.970.. S)>":r-lc:n.tliCdc..script1()n o!the m..tmmalian m tttov. n.F.,.. '-AJJCtE1'., l,M .. 19;s. Charuc1cd1,uion or 1hu motucultu
tl)')>lltl0501Ue5 orN11oa. /11: MULLIG.\S, 11.\\'.. (<"<I.). 771• ,lfrlr:rttl componcm• tn IJnotopl•n-inl1ochondrialDXA ofTryp1l110$<l111a
Tryponcm,1111,JS(!S. 1.tmdun: George ,\111."n.& U11win. N(U/p~r(111111.Ju11rt10luf~II Biology, 8? 243-263,
30 """Ke, c:., .. 19,-1. The Trypa1tosom,·soj.\lnmmnls. ;1Zor,/(1[!.i,'<1I $,:!' fWtu~sos, E.~1 .. 1927. Serofogic.al Jn\'~st.JgJ.tlon~ Into somedise~e.~ of
MoMgrop//. Oxford, Bl<l<'kweU Sci«mtific Public.nion<. dti!l10,ricat,'d animnls In S0111h Arncn rou«d by T~1>n11osot11"'-
31 IIUTYkA, r. ~ MAHV:. , ..
igz&, Sr>t'Clt1l Pml1ologymtd Tlwrapeuntsufrhe £1t•,1,m1h mu/ Tu"<1!fth RrJH'Jrt~ 0/1/zr f')i*rW'tnrofYc1u~n11ry £tlucmfon and
D"'""'" of Dmnestrc Ammals. 3rd odn. l~indon: llall!lbre. Tlndall & Cox. /1-rrll Vnion ofSoulh.Afrfca. 9-2.,.

3,3 KU.iN 1 F., MI\Tr1!1t.~, r. ,& );Okt'oi \fA °'1:•\'A,'\' 1>1;~ Pf>S4.:JI, lt,f., 197'0. S.'l Aotn:\.sm.:. F,.M., 19,.18. :'\oll:s on ~roJogfc•I ttM.S c:.o.rricd out o:l cqulnc
F'1)erimental induction of rheum:uoid.J°nctor•llke $Ub5tance ln unim2tl •Pe<:le• lnfocted with doutine. 01ut11rst.·poofl JoumJJI of\ltMlrw.r;·
,rypanosomi:><ls. Cllnlc«I a"d Exp,irimrnrol Jmm11110/ogy, 1. ~l-$63. Scfom:e «11tlA•w1111/ buhtstry. 23, 33-36.

~ 1o..-xw1·n. J'. & uu torr. v.f,, 1921. llmulhut'I: tltr TrotJNl }(mn4·1:firtn. s..i .1\os1ssos. t.)1,. 19.:.e. Oourlne infc.cdon an young equines, o,uletM.e11oor1
Lt!p1.4!1 Banh. Jc>urnr,l ofF,uertnary·Sclttm. •and :\Jilmal /11d1cstry. 23. 39-40.

34 KUH'<. Ln•. 19s8. The effect or splenectOm) ;1nd bl<>cluid~ on the !",S RO I ll;, c:. r,\(,."'QUE.V.OT' C., GIH(Jt,ID, (:.. BRISG.'\UD, F., U$C:-., JI." U,-\lTl r .•
pro:ccti\c ti1r~ ot .ami ..crum .1gnlns1 Trypt,11QSIJ1mt cqt1iJ)f)rdt1m.J<mrn11I 1991. Anlf~"enic: vnriatil)n In 1'ryr}t11J'1it>rntl tqUff)('rrfum. fi<'Stt1tch f11
ofln/«tlous Dlsed:k"$, 63. 2 l 7-224 J\tfcrolJlolo~. H2, 72&-730.
56 scu~F10m. G, 11 Ot.>fPMU'l, ).1., 1900.. t..a dourinc et son pn~irc, R1.'C'l.fil dP
35 UI\U...~. A. h M(.<~IL ·- 1~04 Tr;ptltl0$Clltl;,$<11ld TfYP(/IIOSOIIIWlS.
Englfsh translation and enla,iren\ent b)· :'\:,harm, D.. 1!)07. London, .WMfcmc- V4rtrwairr, 7, 81-lO.S.
Balllerc. Tindall & Co~. S7 ...,Hotw<c;. "·" ~ ,on.,,.o. KJ .. 1923. 1>nidroithcscrology, 1he

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30-l »'tno, '"": Pro1ot011l diseases

ccrebro,splnal fluid, ;ir.d rh• pa1h<1lnglc;,I chnn1te< in the spinal cord in 6G \\';\TSO..'.;, IL..\ .. 19:!0. {)()11rin,•tlt
CnJuufn. H/Jtory. R/!SM1t'/J nm(
Jourlnc /ournnl of.-lgrlculwral R<Sttlf<il, 26, 497-505. $11pprr1#011. Dominion ofCan4da, D1•portmcnt of i\grkuhur<>. He•lth of
58 sci1u12. ~ .. 1935, Dour11w or slapSiek<e.Journa/ ofthe Somlt ,.\frlC/li1 lnlmal !lmnch.
t'DtcrintU)' Mlull~al AJ$(J('((llion. 6. .a-1 S. (i7 \\11.U.\~i,s:, t,.J.L, TA\'LOR, )t,\., X[\\'j().\;'. l, ~·GlCUUKI. 8 .. 1996. Th~ role.' o!
S9 S1UGH01', M., J-l.Y~:,... ,.,,., P.\ltJf, ,\,. ne IJM.fS8U{M, p_.j. ~.,Es~:;-c,s. ).. \9'9;1. an!l,,'tlrlable surl'4ce&lrcoprotein antibody responses In hn,in•
Afri+.:wl 111i,,.mosomias.~ fo; Kll;~S.Zt;...~OAUM, 1-.. (ed.) Parasillc lnf~€fi<>ns 1ryp.,no1ofcrancc. /lt(((c<lt~ lm1111111ology. ltl. 200-218,
niul tire lt111111111t $;41<111. San Diogo, AC!ttkmtc Press. pp. 1-51 68 \\tUJ.\.\\~OS-1 c.c. ~ tum.A, s.. 1986. Dourine fn Southe.rnAfrka.198! - 198-t
60 >MITH, f .. 1919, TheF.11tlyHlstor:,·ofV,r&fr1aryl./tem111r,. l.ondon: Serological Rndlns;,; rwm 1he VatMn~· RCS«lrch lns1hu1c. t)nd~n<l<pcl<IM.
llallier~. 1lndoll .ind Cox. /Q11mtti oftiw S/1111/r ,lfrirr111 Vttt"1'/tu,r,· .:UWC/nl/011, 57, 163-1115.
61 soLn~. M.A.. &- \\'00. P.1'.~.. l~':'7- T~pano~ome.s prududng disease In &g ,,,u..i.,Mk>~. c.c.. srotTSY.. w.n.. M.Antrr.u'-, "- &,cun:.u::. C.J.. 1988. An
ll<t<t~~ in ,ltrka, /11: KRI rnn. 1.,.. <t'dJ. Pr,rnsiif< Proto::ot,. ~tw \'ork, in\:f'.c!tl[tntinn h:iiu fthc!rnn.tc- methP<h, fur tht> semd.J;1~1'l~hfs o! dourint!_
s.an Fr.,.nd)to. London: \cadcmic Pr~. 011a4rst.'JJ(l(1tJ /11uri1t1/ ()f V1m•r/J11,r;· llhtt1tr!1, Sa, 117-119.

62 t-n:PIIEX. LfJo-, 1966~ rr:,,panosomfasis: ..\ V~u,·i,uuy p,,rs1Jec.1li,e, Oxford, ;o \\'OO. ,.n:~ 1,r.,. SnUvarian uyponoiomes producing dlscru<e !n u,..,stoc~
NcwYork:Pe-rgamot\ P~. outside nhub-..,,hornn Africa. br. •Hmft. 1.r.. C<-d.). Parl1:tltlc Pioto:on.
63 THE:L'i, p1. & 001.ro~. v.. 1980.
Trvpano$flm(I qq1,ip,:rdun~: ,-1o\~~em ~c,n\ York. San Fmndt.eo, London: Acad..,~mic Pr'CSS.
from 1h~dcnnls. ExJ)Orrmental Prtms110/ogy, 50. 31 i•33P. 11 zuA.._;c. z~q. ~ tiM.l"/., T.• 199.i. ld~nlllk.:t1lon of T')'f)tm010nut t1t 11m1l.
,;4 \\'AU:tR, 1.. 19184 Th<' oc-current~ ot dourine (sl:ip,.ie~1.c· ln South Africa. Trypnno1omt1 equlpcrtlum :ind Tl)"J.'IWOWlil4 lintcL,Jusing repttitl\'~
"i
Fifth mu/ Sixth Re11<1rtl tilt D1ttc1t1r of V1tttrinm, R,wuclr Sen,fc,s. ONA probe, v,,,n-im,ry Pt,m,(itQ!b.f:31, ~l. t9f,-208.
Umon of South Afrlco !89-206. 12 Lil \.\;C.. ~Q., r.utouo. c.. & t1ALI% T,1 :99L '!be fo
vil 10 3.nd 111 virro
63 \\'Ass.AU.. O.;\.. GntGORY. R.J.f- & PlOPPS. L.P., 1991. Compantti\'e en\(Ua:titln scn.shi,;it)' of Trypattosoma ~,·a,isf Md r. equlperdu11, to d.iminazcnr.
nf M1.)""~-Unkcd lmmunos<1rben1 Oil$8)' (F.I.IS>I) ic,r the di11$J10,is of •mramh'I.•\tel Cy. quinap>·rnm1nt.• and i"OtnCtamidium. At:rll Tn,p,lll ..iO.
Oouru,e. v.,1nri11t11)• P11ra.<it()/og;·, 39, 23:1-299. lOI-JIO.

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 14
Trichomonosis

PC IRONS, AP SCH.lITTE, SHERR~ A~D J P KITCHING

Introduction ease prevalence or 40 per cem. 110 Economic losses in an in-

fec1ed dai~ herd were estlma1ed at USS 665 per infec1ed
Trichornonosis is a venereal disease of canle caused by ilie CO\V.a~

protozoan Trlrriclto111011asfoe111s which parasitiz.es the geni-

tal tract. Clinicrtlly, it is characterized by aberrant oestrus
Aetiology
crcles. infertility. a low percentage of earl>· abonions. and
pyomerra. The causative organism. Tr/1rfcl1omo11as Joerus. has be,en
According m Morgan 85 T. foews was first identified by variously named, the more common names being Trichomo-
Kunstles in Fra11ce in 1888 bu, .Mazami is generally credited 1ias foews. Trichomonas ho11is. Triclwmo,u,s genitalis and Tri-
with the discovery of the parasite in Italy in 1900.~3 chomo1u:1s l/011i1111s.94 • 110 It is an obligatory inhabitant of the
Trichomonosi~ occurs in virtually e,·ery country of bovine reproductive uact and has only a trophozoite fom1.
the \\'Orld where cattle arc raised. ii was first recorded in TI1e similarities berween Trirrichomonas suis and T. foews
Pennsylvania in 1932 a11d ha~ since been diagnosed have prompted suggestions of reclassil')ing these two species.
throughout the USA'· "· lJ. 119• M; The disease is also "'ith recent studies of the haetnagglutination and adherence
reported to be present in Europe.63• 149 Ausnalia,32• 33 propenies. neuraminidasc acti\·i1y and sequencing of the
Africa. 23 · 103· 123 lndia,88 · ~ij South and Central America, 10• 5.8S rRNA gene supponing this view. 16• '>6 ~lolecular analysis
i;.;, 194 and Canada.?~. 40· 120 and should be considered has gone one ~1ep funher by indicating 1hat Tritn'c/1omo11os
endemic in extensive cattle-raising regions. 130 mobilensis may also belong 10 Lhe same species.96
The disease in cattle was fir$, recorded in South Africa by The pata$ite can be identified by the presence of three
Robinson 122 in 1937 and has since been diagnosed in herds anterior Oagellae and a characteristic undulating membrane
throughout the coumry.1 5· 78. 84 • 1~· io:1. ui. M 3 Allirs1 ir was (Figure 14.1) which is visible because 01 its wave-like mo-
not considered to be of imponance in caule breeding. Later tion on one side of the organism when viewed underpha~e-
reports, however, !denrified it as a major cause of infeniJiry contras1 or dark-field mlcroscopy. 16• 115 The undularlng
in beef caule it1 soulhern Africa.711 · 113, 137 membrane is typically somewhat shorrer than the body. and
Trichomonosis occurs in beef and dairy breeds. At present its external margin is supponed by a recurrent flagellum
It seems 10 be more fmquem in beefherds probably due ,o the which continue~ beyond d1e posterior end as a free poste·
extensive use of artificial insemination in dai11• cattle. which rior Oagellum. 119
effectively reduces the chances of 1ransmisslon. H 3 , 121 The flagellated protor.oan tends 10 be pyrifonn and elon·
Economic losses associa1ed with the disease can be as- gated when a fresh specimen obtained from an infected ai1i-
cribed to the increased number of days that cows are open. mal is examined (Figure 14.2), bu1 on culrure it does show
decreased calving percemage. decreased growing time of pleomorphism and a tendency 10 become spherical.94 In
calves born later In the season. cost of culling and replacing fixed and stained preparations Its approx:imate size is 9--25 K
animals. and tl1e cos1s of treaimem.55• 9u. 109• 110 l\lodeUing 3-t5 pm. and ls thus slightly longer and slimmer than poly-
of the.extent of economic losses indicates that the losses in- morphonuclear cells. 1 :4 In wet prepara1ions its jerky. rolling
crease el\ponentially as the prevalence of infec1ion in a herd motion a1 magnifications of 100:,; or 250X under phase-con-
increases. with an expected loss ofrevenue of35 per cem for rrast and dark-field microscopy is characteristic. However,
each cow confined \\ith an infected bull in a herd ,.,ithadis- ancillaJ) strucrures should be confirmed at higher rnagnilica-

305

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306 ,r.cnll.~ """ Protozoa! diseases

,,-L-- - - 3 amerror
flagellae

Unc!ulat,ng
n,em0tane
Nucleus

Axosryfe

Figure 14.2 Tvpital morpholOQ',' 0 1 rtirni:homonas foetus in a sp.ei;i~n

from an ,n;ected animal
per cem.2. J, t:t. 34, • ~. •r.. s1. Bl<. a9. 9o. io-1. M S Preval-
ence data for various regions in southern Africa concur with
these figures. with various laboratories reporting infecrion
Figure 14.1 line rates ranging from Oro 25 percent of samples and Oco 46 per
drawing oi
cem of herds. 15• 1•1• 711• 84 • 102• 103• 118 The population dynam-
Tr,11,chomonas
loerus. The undulaung ics of herd bulls generally keeps the prevalence within a herd
membrane ts an to below 50 per cent. 110
important Specific risk factors associated with an Increased pre-
.oemifving fea11Jre valence In bulls are:
• increased age. 18• oo. 103 • ID-l. 110• 13 l Bulls of up to three
tions. Protozoon parasites \\~th similar motion, morphologi- years of age are relatively resisrant to infection. 16· 31 · 3"
cal features and size can be recovered from the preputial Howe\'er. some young bulls do harbour the organism
cavity of bulls. 20 and are significant in the perperuation of infection in a
The existence of different serotypes was first recorded by herd, 18. 49 , 1J J
Kerr and Robenson:'4 and 1,;assubsequentl~· confirmed and • beef herds,
elaborated on byothers.4• 22.. 4; . sa. 72, 107• 11 9 Threeserotypes,
• sexual resE. 104
namely Brisbane, Belfast and Manley. are recognize.d. 42 The • commingled grazing, 54 and
pathogenicity of the serotypes i11 villo appears to be simi· • exposure to a greater number of other herds. 54
lar.4 • The antigenic characteristics of T. foews have been rc-
vjewed.H A.n alysisofthc total protein and antigen profil~ of Bulls of most breeds are susceptible to infection. 1,ith some
different field isolates showed strong similarities of the pro- l'llriations In actual infection rates which may reflect
tein profiles and strong antigenic cross-reactivity, inillcaring differences in breed susceptibility. breeding acti~icy, 104 or
its relatlve antigenic homogeneity:11 • 65 Polymerase chain managemem.11. 18. ,14, 110. 1 13.130
reaction (PCR) studies have shown both conserved genetic Once infected. bulls remain life-long carriers and per-
loci as well as polymorphic loci in different isolates. 120 No petuaie the infection from season 10 soason, 17• 93 although
attempt has been made to establish the antigenic relation· spontaneous recovery can occur.39 Concurrent herd
ship between Sourh African isolates and those from other infection with Campylobacter foems subsp. 1•e11erea/is is
pans of the world. Using an Indirect immunofluorescence common. 91 103• 104 although there is no e1idenee 10 suggest
1echnique. all the Soulh African Isolates tested were shown that Infection 1,itb one agem affects the chances of being
10 have common antigens.
126
infected with 1he other.26 • 90
Trichomonosis Is u~ually contracted during coirus. How-
Epidemiology ever. T. foiJI us can also be disseminated by anificial breeding
as it can sunive the standard processing mc1ilods for semen
Persistently infected bulls are the primary source of infec- used in anificial insemination. u. ;o, sz Despite the low
tion. Infection rates in bulls are typically between I and 40 [requency of infection. bulls in artificial insemination units in

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 ·tnchornonosis 307

the USA continue to be screened.92 ~lechanical rransferofin- ln heifers and cows. the ,·agina is initially colonized by 7.
fection by young bulls from infected 10 susceptible females foetus. From there organisms migrate through 1he cen'ix lO
has only been demonstrated in one case in which two Lhe ucerus and Fallopian rnbes and during the process they
services occurred \\~thin a short period of less than 30 min- produce a cerncitis and, in some cases, endomecri1is and
Utes.3'' saipinglr!s. Endometrllis is only seen from approximately 60
Transmission of T. foetus between bulls when a suscep- days after infection. 101 Tritricltomouas foe111s may be iso·
tible bull mounts another which has recently been contami- lated from exudates collected from these sites, depending
nated by an infected bull is possible but documented on the stage of infection. 13· r.o. 121 H I While clin ical e,·idente
evidence is lacking. 1 15· 121. 130 Iatrogenic transfer may occur of vaginitis. cervicitis or endometritis may be noted, gener-
if basic sterile techniqu~ are overlooked when collecting ally there are no observable signs at thi~ s1age of the
prepucial material for diagnostic purposes or when per· dtsease. 1 17
forming ,·aginal examinations on cows.55 Transmission of Ir salpingitis develops. the nonnal process of fertili7.ation
the disease between females by comact or mechanically by m,w be disturbed. However, conception is generally not in·
flies is unlikely. 34 cernipted.101 Endomet ritis may cause death of the embryo
Sexually active females of all ages are equally susceptible, or foetus due to interference \\~lh Its nutrition or nidation.
\\~th spontaneous clearance or infection wirhin seven The occurrence of foetal d(?ath is closeli• associated "ith the
months. pro,~dl ng that normal oestrus cycles are occur- appearance of histological evidence of endometrirls. 101 The
ring.1· ~,. 55· 121· 111 If pyometra develops after infection. the exact mechanism offoetal death is not known. although se,··
oestnis cycle is interrupted and animal5 remain infected era I hypotheses have been formulated. 8• 17 Embryonal :ind
and capable of disseminating che organisms for much foetal dealh have been reported from M 10 18 days up until
longer periods. Shedding of large numbers of organisms five months of ges1ation. with occasional lacer abonions. 1~·
33
occur,; following abortion. Individua l females may also re- lij. \\'here embryonal death occurs af1er recognition of
tain the i.nfecrion throughout a normal ges1atio11 and only pregnancy (day 17). Irregular oestrus cycles occur. Gener·
become free of the infection when cwo or three oestrus ally, up to S per cent or more of the foemses that die during
cycles have occurred after panurition. 1. I:?.\#. 1 n ,\ panial the l!arly stage of pregnancy are retained. 91 These cows sub·
lmmw1icy develops in infected females, which does not sequemly develop pyome,ra which may no1 resol\'c for
prevem reinfection but enables the animal 10 overcome the many monrhs. 11> With the exception of animals which de-
infection in a shorter period or tlme.35 • 111 No immunity has velop p~·ometra or abort la1er. 1he majority of infections will
been recorded in bulls at any age. spontaneously resolve, reported clearance rimes being 95
The number of cows in a herd that become infected by a days and 1wo to fourmomhs. t. i-. 33 ~. &.~. IO I Clearance irom
carrier bull varies. In one survey 42 per cent of the ;3 cows the vagina. cen~ a.nd uten1~ is simultaneous.53· 131 Infec-
served during the firs1 year became infected, while all those tion does. however. persis1 for longer In some cows, result-
which showed resistance to the init.ial exposure became in· ing in a so-called carrier status which can perpetuate the
fec1ed during the second breeding season.J 5 Banlen 13 re- infection in a herd despite commonly applied control mea-
corded that 28 natural services produced infection in 24 sure$. 55 · s;. 131 Rarely, cows may carry the infection through-
susceptible females. In another smdy. 23 of 26 heifers he- out pregnancy and calve at ierm.55 Thb was demonstra1ed
came ilifected after naruralsen~ce which resulted in the low in two out of 40 infected cows in one study, lhe two cow"
pregnancy race of 45 per cent. ui remaining lnfec1cd for six and nine week$ postpartum, re·
specrively. 129 Repe:1ted challenge leads to 1ransien1 infcc·
rion in some animals, which is cleared within three
Pathogenesis and clinical signs
wccks. 131 This partial immunity is of ~uffidem duration 10
lfa bull is closely observed following infection. a swelling of carry over froni one breeding season to another, reducing
the prepuce with accumulation of a mucopurulent e:..-udate the rate of pregnancy loss in the subsequent season.3; How-
within the preputial cavit) may be seen; 1his does no1. how- ever, cows appear to be fully susceptible again after 15
ever. lase beyond cwo weeks. 62 · 119 In older bulls. T. /Ol!IUS months:35 Jl:ormal fertility usually is restored two to six
readily colon ize.~ the preputial and penile muco~a and, less months after the Initial infeccion. 1 lnfenlllty follo1A1ng the
commonly, the urethral orifice. 100 The organisms are con- development ofa chronicendometriris or pyomerra is rarely
fined mainly to the preputial cavi1y and Its secretions. and seen. 119
do not Invade the mucosa.I~· 100· 117 It has. howe\•er. also Host-parasite illlentction is the subject of recent re-
been iSolated from Other pans of the male genital trace. such vic\vs.•1• 130 Evasive properties of the organism include il$
as che seminal vesicles. 6 1. 62 • IC>O. 121· 141 Older bulls remain non-specific immunoglobulin-binding properties and its
chron ically infected although spontaneous recovery has production of extracellular proteinases.139 Tritrichomonas
been recorded."-q The development of deeper epithelial foems organisms bind to and cause extensive damage to bo-
crypts in the mucosa of che prepuce in older animals has vine vaginal epithelial cells in 11irro in a species-specific
been put forward as a reason for increased suscepiibilit)·. 16 fashion suggC!'tive of a receptor-ligand intemction. 128 This

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308 , Ktro~ ' "'" ' Protozoa! di.cases

binding Is en hanced by eJ1.posure of the vaginal cells to several litres of exudate which per•lsted in the uterus for
progesterone. Only JgG and complemem-mediated de- eight momhs after infoction. 13:
fences are ~nown to kill trichomonads.~ 1 Btills show only a Placentas from aborted foe,mses are ordematous but
weak circulating amlbody response. but antibodies are se- othemise unremarkable. 115 Aborted foemses may be fresh
creted into the preputial s.ecreiions following challenge, or autolysed. 1 1· 11• Gross changes were absem in most foe-
suggesting local processing of anrigens?'· 117 135 In comrasi, tuses.1 15 Foetuses :,boned in lace gestation show pyogranu-
heifers mount a significa111 systemic antibody response by lomatous bronchopneumonia and necrotlzing enteritis
11 weeks after infection. 131 u 5 lg.A. and lgGl are secreted characterized by thrombotic lesions with tissue im·asion by
into rnginal mucus from the fifth week post-Infection. trichomonads. 1 H Placental lesions consist of ;uomal
remaining h igh up to at leas1 week 24 post-infection, and oedema. mixed inOammato~ cell infiltratlon, and focal ne·
responding co challenge,.53 · <>'l, Ill, 135 lgG2 and lgM crosis of the chorionic epithelium. 115 Organisms may be
responsl:'S in the genital tract are tTansicm and incon- found randomly scattered in chorionic stroma and in
sistem.131 Uterine responses were found u) be slower and or necrotic exudate adherent to the 1illi.115· 116
shorter duration but qualitativelys!milar m those of the ceni.~
and vagina. 131 The local immune response in the uterus and
Diagnosis
uterine tubes Is thought 10 be a• a result of amigenic stimula-
tion acting on lrmphoid follicles which act as inducuve siles,8 The clinical picture of an infected herd is usually one of
Anti -T. Joems antibodies have been demonsrrated in some moderate calving percentage:.. prolonged intercahing peri-
unei.'Posed heifers in serum and genital tract secretions. the ods. heifers failing to conceiYc. sporadic abortion,. aberrant
occurrence of which is thought 10 be namraJ. l 31 oe~trus cycles. and the prcsenc:e of post-coital pyomerra in
Typical calving percentages in chronically infet:ted herds some animals. 113· 130
are between 70 and 85 per cent, while much lower percent· A positive diagnosis of trichomonosis depends on the
ages can be expected in susc.eptlble herds. 11 · 3s Staggering of demonstration of !i\·e T. /oew.; organisms in specimens ob·
che gestational ages oi pregnancies follo,,'ing the breeding tained from the genital tract of females. preputial material of
$eason is often the first indication of the disease. i; The dis· bulls. or aborted foeral and placemal tissues.
ease tends to be more Insidious in dairy than In beef Bulls are the best animals 10 select for testing. unless
herds. 130 While pyomerra is generally ciled a~ an uncorn- females suffering from pyomemi or recen t abortion. or
mon occurrence. it often prcsentS in 5 to 10 per cent of cows aborted material arl.' available. The number of organisms in
in a recently Infected herd and was reported In 21.3 percent 1he prepu ua I secretions of infected bulls fluctuates. reach-
of exposed animals in one herd. 17 91 ing a peak e,·e~ 5 to 10 day:..•0 Samples should preferably
not be collected from bulls during the breeding season. as
active breeding appears 10 diminish the numher of organ-
Pathology
isms in preputial secretions. 35 Suitable times to collect
Gross lesions may be e\'ident in bulls infected ,,1th T. foews preputial material from bulls is therefore at least two weeks
during the fi rst two weeks of infection and are manifested by after the encl of 1he breeding season and shortly before the
a mild swelling of the prepuce and a sllght mucopurulent onset of a breeding season. The moderate sensithity of cur·
prepurtal dischargc. 33 Histologically. a mixed cellular infil- rently available tests and the fluctuation in numbers of or-
tration characterizes the inflammatory reaction. Debris ac- ganisms necessitates che coUecLion of samples on three
cumulates in the crypts of the preputial mucosa. but the occasions a week or more apart in order to reliably confinn
causative organism is not demonsirablo. 71 · 100 that a bull is free bf infection."' lo,; 133
In rhc female. few lesions a re observed within 30 days of Preputial material may be coUe~'ted by sheath wash or
infection.to1 Varying degrees of vaginitis. ccniC'iris. en- sheath ~craping, the 1wo method,; not differing significantly in
dometritis and salpingids occur in most case~ between 50 sensithit)·.125 Both methods ha\·e been weU described. 11· 2 ' ir
and JOO days ofinfection.8• 101 Tri1richomonns/oe1usmay be l'.l8 Rinses taken from artificial vagina liners immediately
dernons1rmed on the surface and in the lumen of euidome- after use hal'e proved to be almost as reliable as sheath \\-ash
trial glands using a silventain. A sligh t opacity of the normal samples for the detection of infection.;& Whi.chcver method
clear oestral mucus in infected females has been ascribed to is used, measures musr be implemented to e.xdude the poten-
the infeccion. 71 but this phenomenon is often a normal tial of pathogen transmission bet1,·een animals.
occurrence brought about b)' the presence or migrating Sheath washes are collected by mean$ of Infusing 50 ml
leukocytes at the end of oestrus.~" of sterile phosphate-buffered saline imo the prepmial t-avity
The uterine content in pyometra following T. foews ,ia a funnel and flexible tube. massaging the preputium
infection is usually voluminous. fluid. odourlt>.s~. gre;ish - ,igorously. and retrieving the fluid into a clear glass bonle
white and contains trichomonacls in great numbers. 16. 11 ~ forinspecrion. Asuitable sample is opaque and contains vis-
followi ng experimental infection. a single heifer in a group ible flecks of cel lular debris. Urine comamination of the
of I 7 manifested pyometra characterized by the presence of sample does not ki ll the organisms. 100

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 Trn:homono~ls 309

Sheath scraping is performed with a rigid artificial insemi ·

na1ion pipette conneeted 10 a hypodem1ic syringe. This is in-
troduced into the caudal reaches oft.he preputlal ca,iry and
then, 1mderg1.1idance or a hand massagingthepreputial skin,
is manipulated vigorou~ly. scraping agains11he preputial lin-
ing and glans penis whlle negative presslll:e ls placed on I.he
S)Tinge plunger. On \\ithdrawal or 1he apparatus a small
amount of thick. opaque material should be present in the
p,pcue. The presence of blood does not constln1re a problem
and is considered by some 10 indicate adequate depth or
sampllng.11 The material is then flushed Imo a
fluid medium in .i comainer by repea1edl) aspirating the me-
dium up into the pipeuu and then ejecting ic into the con-
llliner. Placing a drinking Straw ifllo the preputial openillg
facilitates repeared passage of I.be pipe11c In an effort to ob-
tain a good sample. while roughening the tip of fhe
pipeue may also enhance the quality of the sample. Custom-
rnade in~o-umems for lht' collection of prepurial material
have not shown any advamage O\·er 1lle foregoing method.99
Co\\'S selected for collection of genital tract fluids should
include I.hose which have aborted recemly and those wich
pyometra,:;o The detection of infecrion in clinicallr normal
cows which have no1 been recently infected is less reliable.
Five successive negative cultures a re suggested to confirm
the negative statu~ offemale animal$. although two sampie,
may be collecu:cl at each sampling opponunit) .55 \'aginal,
cervical or uterine samples may be <>btt1ined from females
u~ing a pipeue ,lllachoit.l lo a $)Ti1\ge or bra ~wab, douche or
scraping methods. 1·1• 2 1 '' 2 · fl". 11:1 Pipettes or $Wahs may be
protected with a sanitary sleeve to reduce comamioauon
of the ~ample. The pipette is passed 10 the relevant section of
the genital trac1 and suction applied 10 rhe plunger. Infusion
of small volume of Rir1ger's solution facilitates sample
collection in cow~ with \·ery thick secretions.50 Samples
colleccerl in plas1ic plpenes may be chilled and cransponed
in the pipene with the ends sealed.
Figure 14.3 ·noculat,on of a comll'21c;a,1t available trar.spcrt alld cuit~'.s
Suitable sample5 from aborted material include foetal kit with crep,,.a1 mate·ial collected oy prcp.r.,a! scraping
abomnsal con1cms and foetal fluids. 1:oetal and placental
tissues should also be collected for histopathological exami-
nation. bright field microscopy with the correct condemer settings
\/lats or pouches of transporr or culrure media may be in· can also be us1:d. Care must be taken 10 posi1ively identil)
oculated at the crushside immediately afrer collection 1he parasites as 01her protozoa closely resembling T. foerus
(figure I 4.3).8z llel'rigerated buffered saline containing have been isohn11t.l from prepu1ial samples from \'irgin
foetal bovine serum, lac1a1ed Ringer's sol.ution. Kupferberg bulls. 20 Direct cxamimu ion or sheath wash samples is re-
medium, and milk-based media maintain viability of the or- ported co have a sensitivity of20 per cent. 118 while direct ex-
ganisms ror 48 10 96 hours.:?S. 11 ~· 130• IJ<> Alternatively, amination of vaginal and cenical secretions from recently
samples should be kept cool and transported to the labora- infected heifers was found to be 36 and -10 per cem sensitive
tory as soon as possible. While organismssun~ve in buffered respectil'ely. 132 in comrast. it has been found that direct
5nlioe for 24 hours, :.amples t..ep1 for this time in transpnrr examination of vaginal nmcw; i$ 73,9 per cem sensicive.' i 9
n'H?dia dn suffer from a decreased viability.'''· 13-: and mu)s- Direct examination and culture of u1ertne secreuoos from
pon co the laboratory should noc be delayed. clinically normal animals is less sensitive. and the cen'ix ap·
Samples may be subjected to direct examination or cul- pear~ 10 he the hes1 area for the isolation of organisms. 13•
tured and then examined. Dark field, phase comrast or dif- Centrifuga1ion should be applied 10 concentrate samples
feremial interference contrast microscopy are favoured for when necessary.
demonsmuion of 1he motlle organisms. However. normal \'ariou~ nmriem media have been used for the culture of

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310 ",:nos n,<>: Protozoa! diseases

T. foe1us. Most consist of pepmnes, ;•east extract, serum. This test does not identify infected bulls. A haemolytic test
maltose or dextrose. buffers and a111ibiotics. Diamond"s me- on blood samples has also b<?cn used.2ll An inrradermal rest
diwn, \\'hich consists ofuypticase-yeas1 ex1rac1-mahose or has been proposed as a screening test in female animals but
TY~!. is the most commonly used in the USA. and has a re- funher research is rcquired.'·15
poned sensi1h/4iy of 8\.6 to 93.2 per ceni. 76· 99 · I'.!$. l3l The While an early DNi\ probe was found 10 lack sensith~ry
~en,iti\ity of a l'Ommen:iallr a\'ailable modified Diamond·s and specificit}',Y the u~e of PCR ampllfication technology
medium for diagnosis of Trichomonas 1111gi1111/is In humans ha.~ enabled the development of highly sensitive and spe-
was found to be poor in isolating field st rains of T. foetus. 9 cific tesrs for diagnostic and research purposes.46• G-1, 120
Organism, are present in greaie,t concemra1ions at rhe bot· While these techniques are 1101 yet commercial!)' available
10111 of culture vessels. Positive cultures are usually identi· in somhem Africa, they are under de\'elopment.
lied by 18 hours. bu1 examination should cominue umfl
,even days.
Differential diagnosis
Various Lmnsport media are commercially available. The
moSL widely used consists of a protease peptone medium in a Early abortions and infertility may be caused by a number of
dear pouch (In-Pouch TF. Biomed Diagnostics). 21 This prod- factors. 1101 all of which are irtfectious. The syndrome ofinfer-
uct has \,arious advanwges facilitating sample rransport. in - tility. irregular oes1ms cycles and occasional abonion result·
cubation and examination in situations where sophistica1ed ing from T. foews infection is indistlnt,'1.rishable from that
laboratory facilities aw not a,-allable.21 · 138 Howe,·er.recovery caused by C foet11s subsp. renetl!alis. Therefore, laboratory
of trkhomonads other than T. f(J(!t11s bas been demon- 1estS are essential for di~tinguishing the one from the other.
strated.20 Overnight shipping of samples prior to Incubation In beef herds where vaccination <\gains1 camp)1obacte·
reduces lhe sensitil'ity of this 1est by I (l per cem. ;g Another riosis has been practised for~orne years ii may be possible to
combined 1tansport and cuhure system (Trichrube, Vrede make a diagnosis of 1richomonosis based on the clinical
Veterinary Laboratory) as well as a transpon medium suitable manifestations alone.
for bo1h trichomonads and C. foellls (Ste,·e·s T:-1. Vrede
\'eterinary Laboratory) are commercially available in South
Control
·\frica. but Lhese have yet to be extensively tested. 102
:-.•10,t investigations have found Diamond's, Claussen's There Is no universally successful and easy 1rea1rne111 for
and tht' ln-P01JCh media 10 have comparable sensith~ties T. ji,e111s-infected bulls al1hough curative treatment has
for preputial samples. generally in 1he i'O to 95 per ctint been achic\'ed by a number of workers. Early methods in-
range."· '"'· 1~1· IC~:. IOG. 113· 138 Some investigators slightly cluded administration of potassium iodide orally or sodium
favour the lr1-Pouch te('hnique, which also has the advan- iodide imra,·cnously, and topical application of nilrofura-
tage of identifying positive cultures mote rapidly than is the zone, iodine. hydroge,, peroxide. acriOavine. dimlnazene. or
case with f)lamond's medium.21 • 79• 136 Culture of cenico- an oin1rne111 comaining 0.5 per cent diaminomethyl-
, '3ginal mucus was 56 per cem sensitive in naturally infected acridine and 0,5 per cent bisme1hyl-aminochi11olylcarbam-
cows, 5 "while greater sensitivity has been found after experi- ide hydrochlorida (BovofJa\inJ.~8• 62. 63• 83. 121· 148· 141 A topi-
mental infection. 10 1. 132 cal high-pressure treatment of3 percent hydrogen peroxide
Trichomonads are often \·isible in haema1oxylin and \\'ilh a wening age111 has also been described. 63 Topical
eosin-stained !issue smenrs. but special staining may be rreatmem methods have Lhc disadvantage ofbeing cumber-
necessary to differentiate them from leukocytes. and some and time-consuming. 1
idemification <Jf the parasite i1. often 1101 possible.'~· 116 The above methods have been largely superseded by the
lmmunohis1ochemistry has proven successful in demon- use of modem ontipro1ozoal agents. Metronidazole, lp-
strating organi$mS using poly- and monoclonal ami- roniduzole and dimetridazole have all been used to good ef.
bodie~.28· 1 H. 1 h,. 1t, This technique can be used for the ex- fect. 3• 12• 51 - 16-~i. 133• 134· 1•· Successful use of Lhese agents in
amination of foetal lung, gastrointestinal rract and placen- bulls requires simuhancous intrapreputial or S}'SLemic anti-
tal tissues, as wcli as vaginal mucus .smears.28 · 114 Marked biotic therapy to reduce chc resident bacterial flora in lhe
au1olysis of tissues makes identifica1ion oforganisms more preputium which metabolize 1he imidazole compounds. 113
difficult. 1Iii Due to the unavailabillry of ipronida2ole and che cost and
Vaginal mucus agglutination tests have been weU docu- residue concerns of metronldazole, dimetridazole is the most
meme<l,62· n. n. ~.I. 107 1113• 1rn but iacl< sufficiem specificiiy commonly used compound although iis use in canle consci-
for accuraw diagnosis. 111 119 An ELISA assay co the protec- tutes extralabel use. It is available in South Africa in a premix
tive surface a mi gen TF 1.1 i' has been shown to detect long- and a soluble form, whereas the parenteral route is used clse-
lasting lg/\ responses in ,-aginal mucous starting six weeks whert>.~6 Practical problems associated with the use of these
af1er infcction.M Preliminary reports of a serological assay compounds Include Intractability of bulls due 10 discomfort
with an apparent sensirivitr and specificity of 85 and 95 per caused by the uea1men1, indigestion. inappetance. absc1?sses
cent respcc1ivcly for cows and heifers are encouraging. 17 m 1hc injeclion sites when injecrab!e formulations are used,

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 Trichomonosis 311

and I.he rapid development of resistance by I.he organism. 11· nation of infection from pregnant carrier cows. 129
, i;. 134· H, In addition. not all bulls are cured and testing must One strategy for elimination of the infec1ion from a herd
be repeated following a course of treatment. I\ delay of eight involves I.he es1ablishmenc ofa clean herd, consisting of vir-
weeks berween treatmem and retesting is recommended due gin heifers and 01her previously unexposed animals. on
to I.he greater number of false negative tests which occur dur· which only young, unexposed bulls are used. 11 This herd is
ing I.his period.133 The negath·c status of bulls must be con- 1hen built up as the previously exposed herd is gradually re-
firmed before they are used for service. duced by culling. This strategy requires absolutely stric1 seg-
A commonly used regime for the use of dimetridazole is regation of the clean and pre\~o,1sly exposed herds. making
oral dosing with 50 mg/ kg once daily for five dars, either by excellent fencing and managemem mandatory.
drenching or using capsules. 11 This is preferred ro feed Bulls can be ensured 10 be free of infec1ion by various
medication due to the difficulry of ensuring adequate intake means. Young bulls which have never bred are assumed to
using I.hat method and the likelihood of treatment failure or be safe. Unforiunately, even young bulls can act as sources
the development of resistance if the fu)l dose is not con- of infection and when 1heir negative status canno1 be guar-
sumed.130 Oimetridazolc is also effective in the treatment of anteed. testing should be performed. 18 Older bulls should
campylobac1eriosis in bulls.21> be sourced from a cerrified negative herd and should be
Due to the practi113I problems outlined above. treatment tested three times before use. Previously exposed animals
of infected bulls is seldom applied: they are generally culled. must test negative in threesucce~si\·e tes1s at least sLx weeks
Treatment of cows by the systemic adminisrration of ip· after withdrawal from breeding. Annual replacement of all
ronidazole has been suggested. IG The efficacy of such treat· bulls over four years of age ,,1th young bulls has been ap-
ment is difficult to evaluate due to the poor sensitivity of plied with success.38 but may be ineffective or impractical
diagnostic methods In the chronically infected cows. Trear- due to s ignificant numbers of infected bulls remaining and
ment of cows suffering from pyome1ra with pros1aglandin F~ the marked shonenlng of the productive lifespan of the
leads to evacuation of the uterine content and return to ey- bulls.90 The inability 10 detect and eliminalf.' clinically nor-
elid!}· but fnfection can persist. leading 10 recurrence of the mal carrier cows may also render a control programme fo.
pyometra or reinfection of I.he bulls in the herd.•9 Funher- cused solely on ensuring safe bulls ineffective.59
more, cows treated in this way have a high incidence or per, Wherever possible, the maimenan<'e of a dosed herd is
manent infertility. 7 fundamental ro the effective prevention of the introduction
Artificial insemination is I.he best means of preventing ofT. focws into a herd. This entails good maintenance ofpe-
transmission of the disease. However. the semen must origi- riineter fences. limilecl introduction of scoclt from other
nate from disease-free bulls and suitable hygiene measures sources, and tbe tes1ing and quarantine of all animals which
mus1 be applied during insemination 10 avoid perpetuating are brought inro the herd. Virgin heifers and bulls, as pre-
the infection.~9 Unfortunately the management, logistical scribed above, mar be introduced into the herd. Older cow,
and physical requiremems of artificial insemination pre- can only be assumed nega1ive if they are from a known
cltide its use in many herds. negatl\·e source or if Lhey have calved nom1ally and have un-
Many managemental s1rategies have been used 10 con- dergone regular cycles for three months withou1 breeding.
trol 1richomonosis. All arc based on a sound understanding Where avoidance of commingling is no1 possible. limiting
of the epidemiology. These srraiegies have been reviewed. 11 the nwuber of herds in comact with each other during I.he
Control measures are often dicta1ed by the practicalities and breeding season has been recommended as a control mea·
economics of 1he situation and 1vill differ from herd to herd. sure although It is not expected to eliminate the disease
Comrol of 1he disease in an infected herd depends on the completely.5-•
removal of infected bulls by treaanent or culling and retest- The reduction of the dura1ion of the breeding season has
ing prior 10 the next breeding season, reduction or the infec· been advocated rorthecomrol of this disease. 1• rn, 43 Shorter.
1ion ra1e or elimination of the infection from the female more intense breeding is likely co reduce the trnnsmission
herd, increasing herd immunity byvaccinotlon, and preven- rates by infected bulls.' 1 In addition, a short breeding sea-
tion ofreinrroduc1ion of the infec,ion. son allows for accurate identification of the po1emially in·
Se)(ual rest before rebreeding allows for spontaneous fected females and segregation or culling of these animals.
cure to occur in most clinically normal heifers and cows and These high-risk animals include any which are open fh·e
a rernrn to acceptable levels of fertility. a.s infertility may months after the end of the breuding season. as most preg-
persis1 for two ro sb: mon1hs following i11fection. 1·8 ·• 2A,, 55 i\ nanc)' loss occurs before this time. 11· 32 Cows \\1th pyomerra
period of 1hree monlhs is generally considered to be ad- should be culled. Some authors advocate the culling of all
equate, providing that the animals are cycling regularly dur· females which are not pregnant five months after the breed-
ing this period. However. some clinically nom1al animals do ing season and those which fail to cal\'e.59 The disadvantage
remain infected for up to IO months. necessitating the use of of short breeding seasons is the likely reduction of ihe preg-
additional con trol measures.87• 131 A similar period of sexual nancy percentage in infec1ed herds. 11
rest following normal calving is necessary to allow for elimi· A membrane protein vaccine administered to bulls gave

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312 >«:110~ twn; Protozoa! diseas~1,

some protection against infection and a significant rise In TF 1.1511.li' amigen also enhanced the antibody response.
antibody titre of serum and semlnal plasma but not of induced significant proteclion against challenge and reduced
prcputial washings.r. 3 6. 3 ~ 134 In comra1>t, the current com- the duration of infection and lesions found in infected ani-
1
mercial whole cell vaccine (TrichGuard. Fon Dodge) has no ma1s.& ~· '1 Fun I1errnore. th'ts anugen
. . weLI conserve d b e-
1s
proiective effect for bulls and vaccination of these animals is twcen different strains of T.foenis.~
not recommended. The immune response follo1,ing both whole cell and
Vaccines containing killed T. foetus organism~ or rnem- subunll l'accinatlon b chamcteriZed by predominanrly lgG
brane proteins do not offer complete pro1ec1ion but do re- production systemically and a significam increase in lgA in
duce the percentage of animal:. which become infected and cerl'icornginal mucus.4 1. .;3
the duration of the infection.so. 53• 66• i;;-.si 1~4 Furthermore, a Vaccination of heifers and CO\\'S, in combination with
significant impro\'ement in first service conception rate and other control measures. is recommended for control of the
services per conception and a redut:tion in pregnancy loss disease in infected or high-risk herds.41 105 Two doses of
has also been sho1111 (62,1 per cent calved compared to 31.S inactl\·ated \\'hole cell \'llccine lTrichGuard, Fort Dodge·
per cent for comrols). 59 · 66• 81 It is suggested that the more should be administered to heifers a11d cows witl1 a two• to
rapid elimination ofinfecrion avoids the occurrence of foetal four-wi.!ek interval. the second dose being given four weeks
death, which usually occurs between 60 and 120 days of ges- before the on~et of the breeding season. Annual revacclna-
tation.9· 53 · 81 Vaccination has the added benefit of reducing or tion with a ~Ingle dose is required. The high cost of the vac-
eliminating the carrier stare from the cow herd.!IO cine has limited adoption in southern 1\rrica. In Xel'3da.
Some progress toward the production of sub:mh vaccines USA it was found that 62 per cent of ranchers were poren-
has been made. Surface proteins which elicit a local immune cial users of 1he vaccine. these being mainl} larger herds
respon~e in infected animals have been Identified. and ha\·e which made use of I he sef\ices of extension officers and
been shown to be well conserved.$.l Vaccination with purified veterinarians. 1-1

References
"Rn1n. n., ,gflo, Trlt:l:omon& in cante. Tn: MOJtRO\\'. v,,, .• ed.}. c,,m,,u Dfagn~1s. pt11h~ene,i.s. treatment and cf>ntrcil. Pron"l.'fllngs n/ 1/1"
i1lf'{(IIJ)' 111 l11aiogcmOIOg_\·. Phi111d1..·lphi-.\, 1.utldc.>n, roromo: \\-,8. A.oh·n(au \tjOCJ'ruioo of Ixwin,· J>r,1(1ittoners. 16.
:,.,,under.; Compon)'• t3 aunun. u.r:,, 1~t;8. ll<,\ine vemm:al uichomonlu~I" and buvinc-,
~ M\Rrrr. 8.,., '\ll'Y~t-10l..Z1 c;;..\\ . 19;.g. Trft.·homofltA')l>i•t11, infection nf ,angt> .wonlon /,:, >Wl<M.l<- t.c.. (t..S.1.,tbar1hm D/Si'Omof/Ji>-s:ock. tlllnoi,
bull; tn South Florl\la l'c,,mi:m)· ,\lttlldn,• I, :,,,u,/1 -l11i111,1/ Cl/tti(/mt, Clt:ulhC..& Thomtt, Publi:J,or,
-.:. l3J9-13,ll l.l OIJ.\.ff.\.Ul\11.\'\' ,. \ . 11.uu,1.s. r.R . .l,.'\".,i.~,,Ch,, \\".(,•• \"I ~TA..U, G.~t .• 199';.

3 i\.l.1:-.unAm. ''·"·· t980. lnddent~ af bo\'fnc lrkhomono'li, In ~lgeri:i. Fac.t0r~lntlucndng nttes of ,1doption ofTrichomonlasli \·Jtt.i.ml' b~
Rttrm• ,r£t••1"tger: de .\lidt•tlrw \ t:lrinnir1•dt$ PnJ, Trc,plmu.,. 33, '.'\t:\'J.dil. ran~1.· r;i11fc produc\!t:,;.. /otlmat of.-ts,rit.ultuml rmd Rt"Jdurt~
381-:llH tl'(IH(Jtnl£'$. !:?, 1711-190

., .u»utn. ,.r..
198.1. Antigc,n analyst, of ,01era.l pa1hogenfc maln,of 1-5 llOF.\1ro,1U\; \'a tfCl~\ll\ I.ABORI\TOJ\Y,. 1.999 U11pul)Ji... hcd d.tta
l'ncho,nomu r-agllJld/.J. lflf«:lt111 at:d Jnm111t1il)· :l9. ti).; l-!04'" 16 BO:SDURA'.'.T, 11.n.. 198.s, rnaino~t~. tre-a,m\!nt .ond ~nttl'JI ol b<)\1nC'
# ,\U)S"SU, "·· Mfr..n. A.. ,\t..fOSS.O. u.;..• BR.WU.>$ •• ~llO!l'<;I. 0 •• \'JOAUX, 1• .tt mchomno1a~1.-.. Tiu· (:nm~dlum of(omirming l:ducnnq;:, 7
t:\\t.,~. ft,, 198a. Tr.uantlcnto dr Jn trlcomtmra.iisdd bQ\ino<on $t;&-S188.
Mecronld:11:0. Rtr1$fl1 C.Uba,m dr Cit11rf11:1 VttcrimttinJ, 14, 2eit-lft4 r. no'.DURA~f. u,n., itttri. Pt1thQgcne'ils.df,sgr.tosi"$, ,md managc:ntcnl o!
6 ,\Lll1AO, ,\,0,. t.:JWGJI. D.. HSCHE1l. )t,. cti~"l,\l~IJN. ~ •• e,~... r1 n .• Ri.:JC..t.rtRJ, l,
uiehomnn1~1~ tn c:tutlc. \ 'tJh,•rlmu')' Cl/ma ofNur1h 1\Jn11rlra. F00<I
41 l!A-V.t-llt, t .. lgt:I~. Tric-h.omonias.is In a bee..•£ hl!td. \1t•writU1')· .\fi'Clirln~ &
,!11111111/ l'rlltlf<·t•. IJ. l•t:i-:lu I
'u111/I .-\nimnl Clmk:t,w. 79. "708-709. t8 UO~D1J1t..\.., t. ,tn ...\..~1)1:Jl~,. '\1,L bU.~c:J:L\.AD. t•.. tutm, 0. l>\S,\,,·. ( c:..
PAl!tilf.fl, C- ""1M" HO. \\'."I., SUTIU'R, :,., IJTl'EA&\CI>,., \\' £ \\'UGUt'I.. D.J .. tS90,
» COSRAO. P••, •• 1gq,1. Prnto1.o.,1 t:u.: ... ~ot
\'SDt.M~~. \t.l .. ftARA. IJ,C.
Pr~l!h!'nc~ i:,f trichUmQl1i.tlit. amons CaJiforn?:t bet( ht:1rd\ Jour1tal of
tepruducth·~ rnuurc in domtso'c ruminMIS.. t'e-Nrlnory· C/initsof.V<1rtt1
tit, A111,•'1tt1n t ,i,..-:nnr:i· H.-,//,,11 .wocfa,iot:, I ~ti. I S-'l<l-t r,q3_
,\Jnttrita. FoodAt1ll>wl Pm~riu 10.439-161
19 1:l(l',;0U1\.\:-.!, 11,H,. CORtlt.11, II.It.. CORI.LIL. L.B... 1993, lmmunr,;auon of
8 A'I.Of.M~S. \I.I. .. no~V-8.\.,,. R.H .. {.08.Sl.ll. Ft.A., O.'>nBUL L.8... 1%6
1,i~n cows. \\'hh ,urfu('.t' ann~ TF 1.17 of rn:riclwmrmt'JJ fo...•ms.
Immune ftnd inllammtnory ~pons~ 10 reproduc-th c tract inrcction
lnftoeritm (l,U/ /lmmmlt)' 61. 138'"""~1'39-l
\\ith Tritrlrltomonos.~ms ln immunllec.l ,md c:on1rol ht:lfer~. /01,maf of
Po.rmtw/1,g;·. 82. 59,l-600 20 80'\0UJl.\'\*f, Jf.H ... G,\J,\r)lf,\fl ,\,, ("_\:\tPLRO. c.:-.i •• JQHS"S();\. J, u,~. 'l,ft,
NOA:t)HAl,,I~+-', 'A.W. VAS' tlOO~f.AR, J:~ \. \'lUASUt.VA. M.Jl \\'.\U.'f:J\, JU
9 \11,.•t,l.l, t..11 .. )UQi.r,IJQ?~. W,O,. THOM,\S, ~l.\'\,.,& 11,.\A\IOX, \\', ~t-, 1g9l .,:\.
1999. Prclinunary clmnactctri7.a.1ion or a Trftrichomo,m.< fo,!nL.~·lik
comp;,,rison of 1cchnlqL1~ u~ !t>r lh~ diagnu~i~ of Trltrfch(1mo11a.(
p,o:ozo.'IJ, !solArcd from pr¢puual &m"l!ma or virgin bull,. 8ot•ir:t
/or1us 1nrcc1io;,$ In beeibull.s, .f"\grf Practic1:. 14, 30,..3.;.
Pttl(((tll>nt•t.:-13.12·1-127.
10 K\ 17, KOH'!Y, A,H,. ai Jr.on~ •.A.lLB- 1g8J. Tr1chomonin-"'s ~· \ibrio.~,Js genital !U 801\CH,\RDi• .:.,\ ,, '\"QflJ.l"-"· ts,!\", TIIO)L\.:). '\f,\\ • H,\K!:110('., \'\ \f,. 19~2.
bovin:,; ~ltuacion o.ctuaJ en con11mtes ~,i.on~ proxinm.s de p10\i11da....,
F.vJluauon o! n Ill"\\ cv1tuu.• me1hod tor dia~m1s"lllg 'I rltrrci,o,no,;as
,·ecin::l, \ ·1.•Jt•rlunrfa .-\rge.mJrut. I. l..&2-1 l;I.
J<k'llls inf{-cuon. \ 'ittrfnflf)' .\t«Ucwr. 8-:" 10-1.
11 • .\11 L, OARC.\TZ. D,,\. C:tU:!liH", ,.,•. j;, \jOltll}ltR, ft,(#. 1!)8; Control nr n aum-;r»- n.c •• H•*"· rrirrld11,mo,:(li.fnr:ms: Pn-panufon uf immuclonaJ
ecnereal diwa<e in infoctc'<i hffli<. I ·,1,ri1111ry· Cli11(,s11f.\'or11, •.;,11,-ri,a. at1tlbodl~ h11h dfoctar runctJon. 8.·p.~nnu,ntal t1,1rmltalol!J'· o2.
Food Animul Prc1ttlc"'e, 3. 361-574, 2fm-2:"4,
lJ uw\~co,101. o .. 19;9 Oi*a..~t..., c:au:.ed br pro1010:t h1: ,,uc,t.R., c.)1'.,

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 Trichomouosis 313

(ed.; m,.,,,,,,.
o/C-,a/1/t In Twpfwl ,lfricn. '-•lmbl, Kcn)'o.: Kcnr• urn MFR~n . .u,t~. 198,. so, ine trlchomnnlatih tn th,• ~n,rth \.\'~tcm
laerarure 8urc3u C,ipc Pto\1ncL!. \\'~>h:m Trnns,\·11:d .md tht.~On,nge- Fret St.m t. /Q1m1at of
2.: Ci\..\\M'IHL. J., St.HUMA' '\, f,, f,\'\:l..l'-. I. .. \•, \C.,hR. D. 1.-POI.U'li'. L. l9Sl.
,1r~:w,,1/r ..\jrlrn11 lrtmnm)' u,o.-lnuon. EO. ~1-S2
Tnc1Jo,rum11J.(~ttJJ tn bulb in ~.J,;4tchcwan. Omudlm: \'°ttMnal)· "" rrtUU!JI. O.. fA8RtUO. , .. >lt.\NCO, M., I.O\'l' F, < \t£.,llL 0.JL, 1935,
Jo1m10/. :I.. 5rl-572. Pre.enc• de trkh1>monla.,l< en el non• de lo Pto,1ncia du Corr!~nte<
2; C.\..\lPJ:."o· c "·. t98.l. ).cfl-dl~ di: ~r.u~~porte p:trJ frl1richomo,u:sfo1,.w~
l'c.1ct'1lnnrl(, _Vf;,,·nrtna. 2. 636-638.
JkiithJll dLt Jli"tlfrlun \ L•teru:nrta. -lrgont1,w, 66. 200-202. :?0,;-209 ,16 rau1s1:.~. ft s..,., 1tt97 ComparJ.ti\'l· "cqut-nca~ an~tl~ ,;s of s.ss rRXA
26 C.\.,iP'f'11Ct, C,M 8NIA-t'tSI. S,C CIPOUJ\, ,\.~ &.: 7A'10M, .\..S, 19.8;-, Dual g.:n~, AnJ inttrnal tri1n1tnbcd ,pac!'t (ITS. ~OflS. of trirhomonildld
in!et."1.iOn o(bulh \\'ich C'.afflpylobatm;riolls- a;,d trkhomonlasls: protniaa. Pa"1</m!nio 115, 111-119
:.rt."!;itm('nt, bith d1metridv.o?e chlorh\·dr.ut?. Att.'itJalian Vt-Mm«')' &i Fn 1.t;nt"-Ul. P 1\., 1986. eovtnc trh:homonia~. Vi11winnrr CUnico/Y()r1'1
Joumol, 64. 3:!0-321 •.\m,•ru.n· Fadrl .·h1lmttl Pl'tu.'licc. :?. Z:{-28'2.
2; <A,1PF.11,(), c.,1 •• 1i1RST. n,c,;., i~,ob:,. r \\ .. ,·.,uou~s, J,.\,. !'~tut,-. D,L. "' 48 l'lTlGt HAU>, ''·"· 1963, freounen1 of g«nml trlthnmonin$ls ln bulls.
w.,-rso~. 0,1 _. ·1990. Meawrcmunt of :mu body In s<"rum and gt."nlta! Joumnt nfrh~· Amrn«m n,,rrmor)' Mi!dlcaJ Au.ociotton, 143. 259-262.
Uu!ds of bulµ by m.lS. \ alter rnccmat1on 1111d c:hallc11gc w1Ul
•9 ~...,xu. 1.1>.. 1g88. Tridmmonlu.sls In~ datl)' herd, control b)' anllldal
Tri1rft•l1()111ouas/Mlt1S, Alatrnllnn \'r;,nnary Jour,ml, a;. l ;5-11l:l. in~mi~tion and pro>ti.1Wundin F:?:l!phu tR:llmt:nt. nu:rlo~fnOlogi•, 29,
23 c.,~u 11tto, t
1 M., tAUO~ P.\.\ .. HUl~a. R,r.... V'\U(Ut.i\S, I.A kL\IEJtY. 1>.E. t,SS. 1367-lJ:'4.
n.iectlun of Trlsrlcl,v11101tM fv,>tu,< ~ntlget\J> ln fonnnlln·fi~•d. par.,tfin s,, G,\1'f-<, f.D.. 1!tl,o. E.~plorlng tho tJt.1>0,• of subfenilil)· ln nntur.il«:"'ice
embedded ,cc1loa, b~ the peroxidase ,,m!pcroxidlL'<! t..:hnkiue.
dal1}' hcrch l',•1..n,111n· .\liv//,•/11, RS. 29&-302.
.~r1srroUm1 Vtti"nnary /ormwl. 61>. 264-:?0l;l
29 c,..,,~,.._M>, c.,1 .• Ml£D1"-:.\. o.• »o»i:n,. o.• ,, ,atr.onco.uo., ,, co!-E..,-nso. n,, s1 c..,w.w,,. G.. \'.,om. ,1. &, l Alm \.'11J .. "'· 1~63,. Tre:iunent of b0t'tne
1rieho1nl'>nitbh. ,."ith n1ctranlduolc-. ,,,... \'t•teritwry· Rc:cotd.15.
,w,cos1, 1.. CARIIA<"O· ,1,. 1998, Subc111att,-011> and intr:tl'.l$lttol
940-94:i,
,-actlnatlon ,1gllln$t tritrkhomoni.1.,1, In holfer,. H1•:·l,m,d~ .1/1d/t/11n
V,u~rlnarln Bm:nos,\,r1,.,. -9. 3..;;"-JS2.. .;2 G•ULT. I\.\,. IIAU. ',I.II., ~'VA$"<"', W.o • IL\XSi. 0.X., 1999,
Chnrnctc-ri7~lfon ofanngcmc prottln> from Tntri,homonM f<ttms
;JO CA..\1.VUtO. (:.j,1,., ttOSSl:'Tfl, o .. \:(t'IJ~\. 0 ....Mt£n;OfSf1nrM. CL ROPl•tL \I.J.:.,
rl!'C'Olt111Z~d by ru1tibodi~ in rnbt;lit ~trum. bo\int!' scrum and bo\ine
1999. Jmmunh.:lrton of h(''1ft•n \.,ith a: Trirrlr/umu.mas /oorus nlcm.brane
rt-nicO'\·j,~,l.J mucus fourmll of Ptittuitot(Jf.J•, 85 244-251..
\'2tc.im,'- \t('{/rl1:nrla •"-'8~,nrtm,. lS.L ?.i0-1~
:i3 U:\.Ul.f. ,t.\, ~\ .\5'1Clrl:..\. \\·,(i ,, IIASS.S. 0,1 tl,\:\J,;.,'--M. &JL\IJ., Y.,t\., 1~,ii.
Ji c1uu.STI..,'.St'."", 11 n.1,;CL.AkJ,;.,. H.L.. tr,g.~pread ufTritrichomon(lsfoew.1
:\pl.'Cll<' antthodlos io wrum and ,·•gi~ mucu• ot hci1cr~ lnoculaie<l
In beef bulls 1n ,,n lnf,-:1.<'d Mrd. ~usrr,1/11111 Wlfri11ary /otrrnal. ss.
\\'Ith il \'JCC"ln~ contnh,u1g Trltriclwmur:tli fU.:nts.:\ml'Tican Journal OJ
205.
1·eun1tt11')' Rtst'ltfth. 56, 1J5'l~S9.
3l C:lfHl~H-.;~1 ~. II (\.. CL\ftf;, 6,\, k PAlt"-Q~)OS, t. \1,, 197;, lnddenc~ o!
54 r.,.\'I'. J.:\1,. f.au..... o," u.""1.1 ·•· w. "·· 199&. Commlngl4..-d gr;v.1ng Cl'S n ris-k
fr,rnd,01m11111$/o,·111s 111 young replQccm,111 bulls follD\\ing
facror ior tnchumonosis in b<x.!c hcrds./ouma/o/the•.tm-.•r,con
m,mduccion mm mi in(ec1ed ht:rc..t .-lf~truiim, \'ou·ri111J()' /(}uri:tzl, 33.
\ \wrf11at;· .11<'<1/cnl ,l,<s'1riWlorr. 209, 6-l:Hi-16.
13?-134,
55 r.Q01>C:rff, \\ .1. ~ ..;;,;tRnow~ +.:,J , 1986 Ep\dcmlolOJCY and cronOmiC'
33 cu••· s.1-,. 1>1111v. 1.11 .. 1980. The frequeMr<>flnfcnilil) and abontnl\ in
nn.11)-Sefi-o( iln un1J ..uj1ll)' long cp,wot1, of trh:hotnaniD.sb m Q lnrge
cQ\,:ii in!t-c-tcd wnh '/'rltlrcm1mw:1fortrc.1 var. JJrislHmt!, .-\.us:tatfmJ
Callfurma dan)' hc,:d. Joun1(J/ nfth,• ·\Jni:ncfllJ \'nrrlnnry Mcdff(I/
~ ·ru.•1inury/fJunwl. 63. 31-32
A.uocimton. is<J, 1r2:-17r-,.
.,4 CL\.R);, s.L., mu 1\-, ,.tt.
~ .,.,n-;n,"SOx, u,1 .• 19;;. S1Udh.•> pn roe
3b t.,tu<.OR\, ,1 w.. a .. h. o." ,ouwoou, h w.• t990. Comparl>on o( .,~mpling
1rn01,mfi;.~lon ()( Tritrlthomo1rc1j{Ot1tus.. ·l11ssr11linn v..u'1'inr.1'' Jnurnnt. 53..
170-1·2. method> lor th~dot«:1lon <11 Trtrrlcl1omon1U/0<ntstnfec1lon in buU~
Tl:.-l<'t<t/na,y 11,rord. 12; lb
ls <tAHt,.. o L. burn'. ,.u. &. PA1bo,-ros.1.,1 .. 1983. rh~ L•ffact oi
Ttitrl<ltomo,r41 fc,r11~ infocrlon on C\tl\in~ rotes tn bee I cnrtl~. s; f.ROllLU~••t.., D,\.l,, CJll-Sl;'\, I,, UIIU~~. n.n... ~m'J'n'l.l:H. O.J ..
.11a1,ulu111 \'i1erm11.,,Joumt11.F.O. 71_;,;. $;1'\JtilU,G. '- \ f \ ru~. \.(,,, '\:IU,r~. )i..\ ,.. \1.\R~H. ,,.,,. 1994- Bo\ini:
trkhornoni~1,: toulh Ol 3 ;-lau~tum )m,ey in Colorado and ~ebras~..i.
36, Ct>JU. fU.. nm-rv. t.tl. & r.\i\'60S-'0', 1. ,, .. Jg.SJ. flnmunt.zation o1 bulls.
ThrTJOf;'l'Ol"ll:•'• 42, 16:.-171
~gain,, trichomonias:s. "tusrralwn \ f1L•ri1t(u)• /qttn,a;. 60, l "'8-1 ;-9.
58 HAIJ.. M.ft I~. Char,,c1cti7::ulan(Ji7rurirhomonns/ottustmtigcn~.
3; CtAR.A. a.t... nu.nY. u.1-11 uu.-n·, r.n .. 19(1:.a. 11uuap,eutic immunit...uton of
u.\in~ b<nint• anti1ocrum. ~m('r/rnu Joumrilo[Vitrrlnnry Rcs.v,rch . .;;.
bu.lb,. with mtrubro1nl""f. .u1d glycoprote-m&- of Tr[flramunns fr,~w,s \.V.
2~i9-?.;5..t
8ruban~. Austmlfnu V.•tcrnnmy lflumal. 61. 6:t-66,
S9 11,\U, ,uc.. ):\',\... '\If ),'.A, w.r... 1tA.,'1o.~. P .. Ot.\\'l:t. L 6' ~4, ~DUt.O)t, lJ .. t~~l
38, CL\1l.~ . 11,L, P,\~$C'JN~~. 1,M .. \\Hirt, M,.B,, &.\~f-11!.LD, J,C. &\'OU~"G, f•"·,
lmpm\'td C'Ontml o(trkhmm,n1.1~i<t \\1th Tri,bomn,)asf~w.J. \'aCdth.•.
1971 Control o( rrkhl)monla-is In~ 1;,rge htrd o( b<'<.'foml<,..~u.,irnlian
>lb'ri Pmtrlcc. i.i. 2!>-14.
\•t.WrJ1wt')' Jouninl, ZiO. 424-l.26.
6o H\\\\lO.,t>. t).\I ... u.\llT'l..crr. o ..E-1!?.al, PJ.n~m of ttue-m:uion in nurnbt.'h
39 CLAM. "B.L. wHrn', ,\t.l\. 6 M'IP1nu. 1.r. 19;1~ Dlngl'1osis or Trit1:om11,m..t
of r,1,ho,no11m /()(}ttd occurrin~ In u,o bo,·IM ,,og1na during lnll!ul
/o..-m:t1nfoctlon in bull.s.Austral:n11 \:~wrintuy Journal. ,p. 181-183
inf~ttlQtK, t1m'1ril"llrt /m,mnt oJ'l.t!ti!rlnttry Rtstt,nlt. 6. 94--95
.:() CO?l!L\XJl, $., C.:U'fUC£, S., KROH:-;., C ... HUMtnUl'CY. '1'•• \\;Aua:ft, E 'H()U$'TO~.
61 1tA~1~10~0. 0.,1.. n1M1tw. \·, A.. JU·~ t. . .& DJ'.':,~ wH 1950.Aquantiuil.i\'~
L.. 1994
Tric'hornonlns1$ in bull~ from S3Ska1chc\~n :md Manitoba.
:,,lUd\' o( Trirltumom,.tfv1J/ttj in prDpudAJ sampf~ fron, infected bull-...
Cmuuifrm Veterrm,ry Jnurm,r. 35. J88..J89.
A1mmr,m/nun,ltl o[Vtw•rlnnry llf!.,1-arth. 11 30ff...3 l ,f.
41 co~UL. LU., 199-1 Vaccmauonbtr:st<-gi~ ug;lill.',I tr;rrid:omonasf~rtl$.
li1 rn:s~1:,.:c:-. M.\\., ;956. Arumnl JJ1S('rY..<tt (IJ 5ottrh Afrtc.'"(l. 3rd edn South
Pnm111a/Q..:,•Tcid1t)', JO, 10.'1-106.
Alric>: Ccntr3I ~ow,.'\jtenry L1d.
-i:t u·.-,,..,~. q .• t!lllG l)logno,i> ;11wcntlo1> and conuol or ijJ Ill>> ~.19;1. Diagno"' uml 'Jnemple derTnchomooodrn,eucho bc!m
QJ11P)'h:>b,tttcriush. and rrkhumnnia,tj~ T11r 8'.ii,in~ Prruutron,n.11,
7ucht.:.uc.r. T!,•mr;Jllchc Umsc:iuw, \>.191·1~0.
18!H8J.
<4J ot;~!l.n. D fl .. Rf:.'fcr. R.L. D..\R.U.\, J.o . .. JOH:\"SO:'\, Jt.H., 19:.; Obrie:,:.1tions 04 110, \I.~,•. , f.c,,ff;,\11 P.,\., CO:':RAO, P.I., U-J•Ut\·JU:. R.U. ftfJtE.7..,. t
..
on the incidence and dl.icibuuon or ,e<Ol')'P•• of Trfrncl:omono.s foetu.< ua,UUk.\ \'T, ,i.u .. l!l!U, Dt.·1cc1ioti of buvtne Irlchomonf;1~ ,,i(h ;i
LQ bt."C.f ,auic.• m north·cau~m A1umaha. .-\ustm(imr \lN,mrwryJo:"not. ,pecllic 0:S~ probe and !'CR ompllftc.1tlon ~'>t<m./oumol of<:Ji,ua,/
SO 4?~-UI .111rmo1olo,;,. 32. 91H0,1
.;.; t"ftA.."-,)JU~, J.;\., W.tT, ,.u.• M[A\\f Ii.IC .. ru ...,AK. G,(,I .. 01;•\\'?'T, J ..\.,t.. Ai\'.\,,

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314 sfcr•~~ 1,m: Protozoa) diseases

Anrigenk relntlon,hlp> am Mg field ISOiate,; or Tritrfclrnmunlll /0<11u Persl~tenc:c af Trltrldromo11as /onus in naturall)• lnfooed cows and
from <an1t.tlmtrfcanfo,1mt1I o/Vetor/111,ry RescarcJ,, 50. 1064-1068. heifers lnAzgtnrina. 11,1,rfr,ary Pnzm/10/ogJ·. 59. 7-11.
66 HU0$0N, o.e., 13:AU.. S- CML"-:l:V, J,M...MOll'flMl!R,, l\,G,. !l(.l\'\'l;~. tt •.\. ~L\RSIJ, 88 \IAXO,\I., P., Clt..\TfnRfl:Ji. .\•• Cl"'l{.M.\.&OltrY, M.., HIC)YA~Y.\, 1·• DI!. IL~. A. Sf;"+,
D.J." PEl!T'~. R.H.. 1993- O~·clopmenr nnd testing of n bo1inc c;.r,., 1963, (:ultural l~olatlon a.nd serological 1yp1ng ofTriclromonM
1richornonlasls 1s,ccine. n,crlogtno!ogy, 39, 929-~~. foetus frorn brcedlnghuib. ht West B,ngal. /ml/n11 l'~t~rlt:nry /otmral, 60.
6: llUOSO!<ri, o ..u... UAU.. t.. on:sn. l~t•• MOR:t'JM.LB, lt.C ... W\\'E."i, ft..,\ ~u. '19l-7ll!
1>,1-" PEUZ. R.IL. 19g3- Tc"1ng ofukhomonla<is \<a<t'hto In hei(U< 89 ~1.i\1m~l.J... LH.. MJWO'-', ,.c., NOSEDA. R.P. It CORl)h\ 10LA. J.o.<1., Jg.85-
1

mat<-d lO infee,ed ~ulls.- 71wrlllSMOlogy. :19, 93;-~H. Dfag_nOSficO dt' trlcht)moni;t""t< tn ~oros. Pmpu~Ul dt! un ~uerna d\"
68 ino,\, J.S... 80SOURi\..''T· R.H.. (A\tl'ERO, CM .. COR.BBU.... 1-&, 1993, diagn<»l:co. v.,..,11111rfnNRJ!nt/1111. 2.9%. %11-!17;.
Constt\"at,on of a prott-ctn"t surface antigen or Tri1til'l:omcmt1s fo..•w.J.. 90 )~CCOOL. C:.). fOWSSf.,..'.O, !ol.l'.. Wbl.FE. S:.-G.. S!MPS<>N. M ..\., m.M. Y-.C.
Journal o/Cli11iet1/ Mirmblology. :n. 3289~1295. IM .\WARnttA~A. t).A• ..,cAR..?o:EY. 1. ,-.. 1988. Prt"1.·aJent-t• ofbo\1ne \'t!'ntre3l

69 , ...'l:DA. ,,,:,... BONOURA.,~T* R.ll r.OHIH]L. l ...t;;.. 1ij95, Bo\1ne \',tgma.f

a.nul,c;dy dlsc3~ fn the Viclot!o Rh-er Dlmkt of tile Nanhtm i·errltQry: like!\·
responses rn 1mmunoaffinity•purtricd surf:ice a:migcn of Tnm'chonumas c,ronOmic: efTectS and practfoablc tontrol ml'.l1,u(C:S..J\cut10iimt
f~w,.Joumal ofC/f11ia1/ Microbio/010·, 33. 1IS&-11~. Veu•rtnnry Jotmwl. 6S, IS~l-56.
':'O 1ot~"1;tt, a...r. ~ )IUJ.u P.c: .. 1g~ lhc 1rururntsslon of Trfrlromomu /04,'llls 91 ,)IICK[LS~. W,I) .. PAlSL.n. LG.&A.'\'.[)Ensos. P,R•• 19tl5, Ptevatenceof

during the ~olloctlon o! semen from infested hulls, n,, l'.•trrinnry postservlce pyon\ctro 1n o herd of bm~r cows inft.-c1ed i.vnh
R~ord.&1.1-4 1richomoninsis.! A ezas{\ r~port. n,erio;;Nu>lngy. 25. 1•11-74•1.

PntltologJ•oft>o,m11f{r.
i'l Jun,,. ~.v.a:•• ~!\'EDY. r.e.1- l'Al..,\lfk.. x,. 1::,85. 92 ,so~n. 1M1. 1998. Ris~ analysi.S! CSS te$Wlg protocol for tricbomom~.&:s.
.'\Jrlmals. 3rd <'<In. Vol. 3. Orlando, San Diego, New Yotk, London Pr0<·et<d111gs of tlieSer:r111,.,111h T«lzmcal Co11f~ret1ct on ,lmfic,al
Academic Press. Inc. lnsmmna,rion ond Ht·prt>d«ctton. Middlrion, Wisconsin. 2S-~6
SeprombM 19SS. 3i-42.
12 \."CMH. w,H, ,., 1. AM0;\.'1', H,G•• 19.:9, Tht.• lrnmunc r~pon.se in
ukh.ornoniasis. Pmct.~11£ng:s oftht Four1,¥mh lmema1£onal Vt1"rl11ilf)' 93 \ fQRG.,,:\, &.l\., ~g.i:b. Bo1ri11e trfchcmir,mns-l-s. Mlnncapoli~ Burgess

Co11grsss. London. z. 16-20 Publishing Company.

73 1r,:E1m, w.R, &> ROnurrsos. )1 .. 1.9.u.An Inves1iga1ion imo the infoa.!on of 9,1 MU~Ol. o. 195,1, Bcrtragzum Problem clnc\-F.nn,·ichlungzyklu'f cfcr
cows \\ith 1"rl(homo,ws{t><lltS by mc.u,~ of the aggtutlrunion rcoctlon. 7rtrlt0mQ11t11 /Of'IIIS untcr besond'1er Berotc~,ithtlgur,g ,akuoliger
Vcurfnary/oumal, 97, 351-365.
Rundfomrnn. l}nP1t1ktuclre Tiemr.t. 6. i3S-137.

74 1rCE:Rlt, w-.R.,11 ROB~~. M., 1945.A note on thenppea.ranceof sero\Of:tea.1 9.i 1'A1Sl.lX. L(i., Mlrt:l!.L..,;FS, w.n. $- A,~?>F.B!sO", "'·"'· te)86, Ml"t'ht\ni,im~ and
v:.ricr!es among Tri(Jromot1as /0<:rus ttr.uns isolated rrom infe<:1cd anle. thempy for rc,•lnl'd frtal rnwibr.ml!S and uterine infe.;!lons or rows, A
rc,~w 77reriog,11oloio•. 25, 353-JSI.
The l'tu,rlt1ary R=td. 19. 221-222.
15 Jm.t.SbY. P.a., 1997. Bovinr tnchomoniasls.. /n: YDUX<,qtrlST u .. (ed.). ~6 1~.uu•.•o,;01- M.• GRukunr.nrR. t .. l99.J. Som, pr<>pertil'S orstaflc,ndd
O ,rttllf Tluuapy lu IArgeAnimal 11umogtmol(®•. Phi!3dl'.tphi:a: W,8. bindlng !.)'.Stem:; in Trhrlclwmont# suts 11.nd Tritrlt:lwmo,ms/«ws.
C,,,nparatl,..- Bl<J<:Jwmlstrya111l P/ryslolog.,, 1088, 529-536.
S;u.mdors Company. pp. 275-279.
76 1ii1),l~EY, ....... DARIE."+, 8.1.. "f'.NOMICJ;, J.\\'. & TR..\.'\"Tl. Cf,, 19-8,(). Ba\'mf!
97 t'ALL\Dl~O • .)1,1\.. (ZA\11'fk0. (,M~&.AC.UX.\. c .. 1983,. Uti1i1.:adon de

uichomonfa,ts.: Dia.gno!is and ucatmtnt, l<>umal oft1J~Amtrlcon dl\'ersasdrogas lr1cornoulcidas ttn ,oros.. Cttut.n V111~rl110.1:o. .is. t239-
l"rc,r/nol')' .W,d/ca/~tlon, I;;, bl6-l,18. l 295.
98 PAK£EJ:. 1).).,. $1L\ltMA, li.n.• \"\'AS. u •.Jr.., 1:ATMR. P,n BllATNA(&!\R. G.P• .&
~ >1tcKl,<G, J.1•., ,~. Allmon R~onal Vet•rina11· i.alx>rotol),
Pictcrmarlo:burg, South r\!nca. Unpubii!hod data. t.0D1l• ~.M~ t96u. :s;o,c on Trichomo11otf~iu,, fnfoction ,n natt\'t can!~
ofRiljaS~irL hulton /1>urn<1f o/AnmurlSckr:uJ. 50, 282-284
7fl l<l'TClll"r,, ,.,.. 199.9. Trkhomonl~$i< In Xoml Qecfhcrd>c imponauc;.
cradit'tltion and ron1rol. :>:ewslcrccr of tht Uvc.<roek Health and -95 P.\AJ:ER, ~. C,A'.\IPROl 1.. ,., fd68LE, c .. O.\J \DHAR. A 1999, Comparison ot
Production Group of the Sou1h African VctcriMI)' As«>l'lation. October two ,nmplrng tor>ls for rho dragno;,. of Trl1rrclromnnri, ['1twsrn bull,
1999. and clinsoal inte111re:n1icm of t:.ulture resu.hs Jounmt of 1hc American
\'tlftill<U)' ,\frtlkal J\$$0Ctatlo11, z 15. 23 l~-35.
79 l.rnu., D.R., 1998- Compa.ri~on or di.:agnostic m,e-thotfs (or detttuon of
arth~ lnftttion with Trirriri1omo11asfo,,111sln bcc,fhelfc,:,; /011mal of 1/ru 100 PA:a....o~.s-0~. ,.111 .• CLARS. u.t., &- uum\ J•• 1&7.i. The p;1.thos~n~!s of

Am,•r:ca11 l'•tcrft:111yM,1/lc111Asso<'i111io11, 213, s19-,;22. Tntriciiorno,:a.t/oerus in(ecdonJn the bull. A1,.urolla1t Vi'ttrinmy
Jo11mal. SO. ,21-123.
8o Ji:\'AS...... IC~, ._,,,G•• HAU. ~,1 •• IL\SJ.~, IJ•. !81:J.. L Kf.ARl.£\', D. k \'.1"nF.. .s.E.,
1996. Currtnt conctp1s-in tht- con1rol oftx.wfne t:!chomoni3'i~ 101 P\MO~s<I~. , ..... Cl.AK~."·'·., DUFTY 111 .• 19;6. Emiy pathogent<!> and
CDmpentlium on Continuing EdnuuJon for th~ Prac#sin8 Vett1rlt:1Vinn, pathology or Tritrl{lromon,,s ftwus lnf..,tion in l'irgin helfen,. Journal of
18. 103-111. O,mparatlt~ l'ilr/1~/ogy, 86. ~

f;ll k"\'/LSNJCK.\, \\',G.. HA:-:xs. o .• HUANG, 1.c.• llAU.. i\l.K,. SANDllLO:M. o .. CHU,
102 f'fFAMS. ~.,, •• 1999, Vtcde Vctcrin•I)· Lnbor.nory. Vrcd~. South Africa

H.J .. CICAV[!J.. L & ACNFiE.. \\ ,!-1,. 1992, Cllntf".at l"VO.luo.rion o! the-effit.:tcy of PcrwnaJ c(lmmunh:-n.titm.
lnoeuhHing c:nu.le \,~th ~ vncclnr contrunins; Trirrtc11omQm'I$ f('ttu.c. 103 ~~PA.NI~ ~.M., H\::JUt. ~., \'T?<,.'Tl:ll. C.C., l:KU(.;.EM; W',, QUl•JR.OCL\, C..C:. &
.~.m~riew, Journal o/Vi11cri.JU1')' Rtsl'Otclr. 53.1023-202:" A~AMt. L , 198ll. Trfohom~nla$1$ ~d <i!mp,lnb,ttte,losf< In bulls In the
8-:t usns,u-, o.s.• ousm·. f.P. &, ITLAGWR.);', s.L., 1996. Findfo.g 1he t:ause R~ublic ofT.run~l«i. tm,mnl of tlr~ Sot11ltA{rfran V,•1r'rinnry
of panisitc·indu(ed abortions In t3ct1e. \'otrrinory ,Htdtei,ie. 91. Assoclotio,,. S9, 1:19-140.
&1..;1. 104 P"ERE2, £., GO:,,.·AA.O, f>.A,,. lltft.D, O., 09TUX0,A., Cll\C:OS. t.. 801\'t>UJl.\~T R..&.

83 ur,u., F..w., 1951. The uea1ment of tri~homoniasis in bulls. n" Irish sooROHUlll!'i'. J., 199,2, Pre,·aJenec: ..ind rlsk: fartors (or f'richomon.as
l'rteri,rnry Jaumol. 5. IS~-1 57• fotiru.s fnfect1un in rattle rn nnnh·eastt'm ~m Rlc:a. Pt,•v,mrfrtt
• Vutrn·m11)1 Ml!dirin,, 14.155-16.~
84 LOUIS TRlt.:IIA~l)T Vt.TUltl/,R\' IA80RATOR¥, t!lU9, Unpublished d.110
103, P.E-RISO. LJ, & Ru1•~. C.P,.199.J, lmmuniz;\tfon orthe hccfcow tmd lh
8.; LOIi'. c;.u .• 1978, Tlr, Tricl10111UllbdJ. Public H,-ahh Laboratory Sc,rvite, inOucncc on fCtoJ nnd ncano,.i calih<ahh W1erim1ryCll11/f.<o/NoTtlr
\lonogroph Serie;. 9. Londo11: Hot M•J<';ty's Srat!oncl)' Office Amtrlca. Food,\Jzlr•u1I Pf/let/co. 10. 1;;.s,;.
BS )tA....:tuo. O-"- ~Mo:-."tO. c.M,, 198..,i;. Trkhamonfasl~bo\in~ 106 PFTtiJl,. o .., .• f'AtIS, \\',II,. ),UU.(K.- lt..lt .. \'0Ui\"GQ1,11S1, tt.-S., RA~nu. a.F••
Tritrlrl1amo1101foerru'I, I. Di•gnostico )' prw..Jcncfa en la Prcmnd• d• GA~JA~L u:.1.- 1,\'bYER, J,L. 1995. Trttricl1omona1/o~m, infccrlon Jn a ht:rd
fonno'wl !Argentina). l'<1eri1lllrfn Mg,111t/11a. 2, 43H39. of MissOuri C;l llft! . /fJUTttfJl oft ·.ueritiary DitJ·w wslit l11wnfgmfo11. i.
Si \to\SCIJ10. U.A ... .RUSSO. A,M·.• (AAAJl-4.!At.. 1..,1 ~ Mf>S'Z.O~. CM... t~9!.- t?8-?80.

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 Trichomonosis 315

,~; l'lli!ICL •.t.. 1949, ·n,t muc\ls ~gglu1lno1lon 1c.,1 for thu di>S,rlosk or I.ti sonrrrr., A,P.. flout, s.. "~t'!'<:H1N(,, J,$1, , 1994, TrkhontC)fliMi~. /h: C()liT?JlR,
bo,in,,irlchomonln.is, 77re WU'rinnr; R«ord, 61.341-349. J,A.w•• THOM~~. G,k. ~- ,u~TIS, A.,(;., cc.'d1. l,1fe<tiousViStth:S.eSO/IJ1'tSI«~·
108 Prucc••,. e.. 1950, The host·p:uas1te relatto!l,hip bc1we;,n the female 111ith S1>«fnl Rtfanmtt 10 Su111//rrr1 .4frit:t1 Cape Tuwn: Oxford Un\VC1$lty
bovine and trfthommrm footus. Tht Vt'1eritU11)' Ri"Ct>rtl. 62. ~I, l'n><>. pp. 213-219

109 AA£. o.o .. 1989. lmp>ct of ulchomoni>11> un 1hc cow,calf produc~r·, 128 ~ISGlt, 8- s .. IUCAS... r.J .. UMCII, D.111, ~m~. s.1 .. GU.BDO'. rt.o•• 1999.

profitability. Joumnl <1/ th~ ,:\ou•ric.an V~uri1u1ry ,\1rdkal Assoc1mron, Adhesion or Tr/11/el1omo111«f(Jl!111s 1., bo\'ln• ,·~ginol epilhcllul "1'11,.
IS-.. 771-i7S 1,1f..:1/o,, nM lmm1111/1y. 67, 3&17-38:;.l
HO ML o.o.. autSO\\ fi'tlt, "·'·· cE.a,'1HJ, ...c. ~IC1ST0:)tf, .,.n.. OtOWW..._ • Ilg CiKIRAOW, s.. iq&-:. lcfrmific;mon u(1richomonf1d,carrier CO\..~ /mtrnnl of
MOOJU'. S-A .. 1999. Pr1.'Villenct or Trltrlclwmonns /n11s an a: bull tltt .·\,,itrlco-1: l 4trrfrib.1)' AledlttJ/ Mt«inlitJn, 191.SS3-S54.
populallon and dl'l-c1 on 1>rodµ~lon in u l3rg,; t~w-r,,lf tmcrpri~. 1:iO SI.IKROW, SY.• .& MSOURA.N"t, M..11,~ 1983. iJ0\10C tnchomomasis. \Wtnll4t)'
/oumal oftlw Amwlcan \leterlnmy \/,r/fcol ,..,socfn/10,1, 2H 103l-lOS5. 1J111/i1in. 08. 591-603.
m o,. i,) 1011,~():.:. A,1-r., 1981. Tri,ricl:omt>uaJ {MIU$
1U.t:.Ct. LL, Ol~..;r.rr. 131 "''""""' ,,z "lt<>Nl>URM,r, • .u .. 19sa, lmmunoglobulin ,,;orype of
agglutination 11!<1$ upon :.amplC$coll,>ct<d rro,n caul~, C1o~Huc1ion~ ~p«itic u.n11bodie-s m rnpruduc-u,·~tr.1ctse-cretio1u and.M.'ra m
4~socittted with vaccin:uion :tgainci Qm1p?1'oltt1&tl'T/Dt'Ws>ubsp. frlmclzomonai /ocm.r,lnft-cted h,.'ife~ :lm1•rrmn Joumal o/Vetrrmn.ry
&.'c!1ttr..wUs~Aus1r,,lltw Vt1t1rlnm; Joumnl. 57. 352-353 H~tnrc/J. SI. l>ls-653.
U2 RE.Ca.. LL. t'l(,.\;:",,'E1'T. DJ>. ,a, JOl!XSOS. Jl.[L. l983. Sbmc obset'\'utiOlh) on 132 ~J..'lRJtow. :,,1.• .t .BOS't>l.lRA!\'T. ,t.u .• 1 ~ . Induced Trir-nrhomorms fo,m-1$
cultural and lrailSpon condJrions fot Trirrlcltomouas /otlllS\";U. infection an t>cclhrift=~. /<um,nl of11:,, Amr,rir..,-11 \'eierwary· ,\ frdlc:al
Btubattt.Aus1roUn11 V,•1<trilltlfi,'/Ournu.l. 60, 62-63. As.ioriotw11. 196. 835-8$.
113 Me:rtEF.. G.P, £-.s<:uunr.. ,.P•• ag;s. The diagnc>sk ond 1reaunent or 133 t,,t:IRR()\\, ~. 8():-.UORl\~"T. R,. MlilUiV. ,. 4'- f.()ftkl.A. 1.• 1985. lffi<?.tc)' of
trichomonlasl,; in bo,ioes.Jn: Symporium 011 Reprod,mlw ()is<Jrd,,rs lpronida.wlc •s•in,11richornonla.l, 111 b«>f bull~ Jo11rmd ofth~
and Ferdtit}· Rwninanrs. S-Ourh Afr'lcan \ ..tum nary .,-\S$0Cf..'lcion. Juh' Amf'rlcori l'-.,111rlr:my ,\ledfrol A,sso<fntto11. 187, 405-407.
1Si5.
134 '\OTO, P ~ U1tcttf11. 1•,r.. 19ff6. Tra,~mtomo lmmunolugico 1k 1.,
11'1 lUfYA..°", J,C S.1..'\.'-:CUARO, r ,c;.. ~ 'ASSla:A. W.G.• lift\.L, )Ut. &HA.\;'lU). t>.
1rh:homnnla.,n· gcnhal en 1oro.) dt: n>tfo.O'- C'Un qwnuo1e«,.£s1encia..
199> 1'issue-11wasl\·e Tri1ri&l:omonasfootw; m four aboncd bo,1nc ,~eurin,1rio ,_.,8l'nti11a. 3. 980-98j
feruses.. /()umai of\ft1i.'"r111ary Vtngnos.1:c J,,1.,,,rigm/011, ;, .;09-; 12
,.as ~OTO."·'- .-.,o,~u . ..\,f. .. 1989. l'hc.: immune ie~-pons<• in c3.ttltt 1nJec1cd \\ilh
IIS llHY.\S. ,.c.. sr.,c~Hou.•. LL ..,()UIS!>, w., .. 1988 Foetal 3nd plnctmal fritr/J:lmmo,u>.r /oer,., V~wir1nl)' t•11rlll/10/o].<)'. 33, 343-3<13.
lesforu, In bovine abonlon due- ro Trirrk#tamonn.> {~tu,$. \ltifflll(ll)
Pmholog:.•. ZS, lS0-355. 136 sro&~TL i=-.n_"' ''"'"' Rk6MS. s:.ri...1.• 19-;8. l,1,"C.bu dc!t."ft:tn:ida como mcdto
de cultl,o para frid:omo,uu {0i11U.s. CatNn V~ururor{a. 40. 2,l-32
u& nu,·A.,. ,.c::.. WILSON, ,;,t.., lltJk(,tSS. O.E. ~ACtlUOUSL l..,l.. ,. Q\H\; ', , \\',I ..
199,. lmmunohl<1ocl11:mlc>I dttee11on of (rltrlc/u,r>10111Uf0<t11.1 in 137 n: l)RUGG! L\., 1.985, V~wrimuy Rt.sc;irch ln~titutt'I. Oncler$(epoon
fonnnlln-/l,ced. paraffin-embedded 1cctlons ofbo-,nc placent• and fe1al Unpubllshed d.11>.
lung./011maJ nf Ve1t•r/m1ry D(ag,l(lsrlc ,,,,..,,,1g11tln11, ;, 911-101. 138 IHl)MAS. M.W,, 1(A.K,\10S, w.,\J . .-wmn.. c .. 1991.An imp,o\-i.-dnwlhod fo,
11'; RH\:,\S. J.(!•• v.11.sn~. LL... , \\'J.C:SfR. 6.,AS-0!:llSOS, ~l.t.,.. kO~OUM.,'1. R.11 •• the dolcc,ion qr Tritrlcl,omonas/<M•111~· mrcc1ion b)·cuhure In bulls. ..\Kn
a-u~cess. n..r,~ Mun,1t1u, o..t. coMEtl. u .. 1999, Demon.ttto.tlon of Prriul,e, 1 l 13-17,
{ri1rlcltumo1ws/0t·111s fn Lhtt t'XC~mnl gcnh.:dla. anti of spedfk· ~1uibod.1t!I> J...'i.q THOMFOJtU, f.W.. iAUlot, J.(1 .. 11'.U,J,\, r.>., COk.UUL t~n,, 1996.
In prepu1Jol st'ctNi<>IIS of 1\an,1'111!)' 1nftt11'd hulls. Wtirrl,11;0•Pm//o/oJ._'l'. C:ha{att~riz,nlon of cxtrac~·llular prc>telnn~ of 1'ritrfcJ1omo11u.sfo~w.s.
36,•l<llkll. Jorirual of t•nrosrralof(J', 82. 112-111
118 mu1I110., t ..M,.,s,. 1&,)9. •,n efficicnL mtdium for 1hc isofat1on of r.to VMIOt1•0,;._~rii>. >1 • 1946, Her 1><,~1rljdcn ,-an de tnchomonaslnf<!Ctla bij
Trkhomonas facw.t. Omlenr,r,oort Jourrml ofVtwrinhry Rl·Jen,clr. S7. ~tloren. V/aamscl, /)/,rgem'tSlmrul,g fift/$<·hrift, 15. 1-IS.
209~10.
1,U VANl'>EJ'l.,\S~fE, \t., fLOft~T. A•• PA.JU Db. I . 4 kRb~J..1 .. 19311. PQtt>g<'.'n~t.•
1111 nu.o>JOlJ.(Jt. L, 19;8. Tr/1rld1omom>$[0,,ws. 111, •1w11.,1. ;.1•.. (('<LI
PatMlt!c Prour..oa. New York! Acadt:mtc Prts~
dingnose en behandclinl( vnn TrichQntona.,-infcl<.Je hr nmd~r~n.
Vt1'W8'f" Oi.•, .\'111'0rsingm. No. 13, December 19M. Unl\'ct>ltyof
12() RILEY. f),R.1 \\'Af:Sr~. 8,. voux,· L.. )."RIE(,[fl, 1,11'.. 1995. PCft.•bascd l,:lUU~· of Ghent
eonsorved '111d "•ri~blo ONA sequ~nccs of 'trllricho111onasfm.•111s
isolates from Snsbtchewan. Can•da.Joirmnl ofCl/11/cal \flcr11bio!l//O·,
142. v,. ,. A.(l:,-.'$UIJJtC.. s w.J.. 1953. l{tS,e3.Jt.h into lnfortlllty in t":tltlltt and Sl'll-ep.
33. 1308-1313. /oumal of//11• So111/J Afrlrtm v,,wr111ary.1J,ttiralA<socu11/011. 24. 6~79.

121 •oBI!IU'S. s.,.. 1971. V,rarinmy 0/Jliretrfcsand C'.c11/1a/ l)lsea.,,s. Mkblgon: 1.i3 VAX ac.oorrs. c .. 1998. Venereal dise:i.ses ean impact r3rmcrs hnnnciolly,
Edw:uds tlrorhcrs l11corµof'l!tlon /.11ndbou11.,.kblr.d. 16-17.

w ROa,~sos. E.M .. 1937. Triohomonll< lnfoc:1ion in a heifer •nd a brief 1.;4 VAS SOMl'.IU:."., v.o.. 19-,1,u. A Mmple cechmque for the ti.xanon and suumng
rov!e1< ofahorl!ori In DO\'ln<H due w 1richonro11a,- infe..·tll>n /oumnl of o(Trlchomonas fn cullurc and ":igtnnJ mucus.. Vru,.riruuyJo:,mo!, 102
1//eSo11r/: Afrl<n11 \fe1~rl11nry M<!dica/ tlssoc/11111111, 8. 13'1-140. 73-78.

L23 ~ctotmr •DUMO!'.I _A...\k.\..,, 198.: The, it1dd1mct" of Cnmp)1lobaeur/MttlJ US \'A$Ql;[1.,. JI:,, ,\\'II.'\, J,I},. HU~M;\JrA), (.f;,. k >.\OF.It. 1t.l., t984,
and Trlrrtd101110111tSfO<'tlJSIII c:itt!~ in Sou1h•WCM AfricalN;imlbi:L rnchornonia.~t, y carnpylo\,(l.tC.crioMs ~n la u1g1on ~enuaridll C1mm1J
PrQ<dd,ng1 of1/1, l'eml, lntlmwlonlll 0,1,grl,,so11 A11im11/ Htprud11c1/n11 \'e,~rinom, A~"'"u1111. I. !1-10. 942-9-17.
Argcnti.oa.
and ,1m11n·n1 /11~111/na1wn, u,i>ana-Champai,;:n. 10-1, June 198'.. 1.;6 ' ''ALSJI, r .r.• ,:c,""UTT. s.H. ll'>IURRAr, c;,, 1934. Trl,l1omo11as bo1risinfc-ction
f2.t SCH:ri.ACltl.. J.A.. WA.Li.ACT. S.L, ~-VAS):fcx.\. W,G .. llAN..:S. tJ,R, & HALL ~f.R•• of cattle. <Amfl/ WtNin1rrlo11, 24, 60-74.
1990. Tn'rhomm,,u /OPttl.J. \'tlcclne immunogenicJty uial. lgri Pwr1ict', o., .. '10.WAN, r,., 111:R'lMM,) u., GJU'U>OCR. ... as-a;. Tre-atmem
14; \\'IU1A.:,t,..,,

10.11-H. of mchomonia1ls. AIISlf(llum l'PNt'fnnryJo:,r,ml. 64. lSS-160.

l.25 SCJ(OXM.,\.~~. M f .. BO~l)URA.,T• .RiH., (;.\R.D.~U:) I..A,. tu)OS.I!.\~ )r. \",. 148 \\1UO:-O. ~.K., XQC"A'o;, A,!,... (,.\UDY, t:. T ., (';O()UWI:\, n.. 1!17'9, Thi: prcval1mce
BALTZFR. w.• J;AC.,101.IS, C. ·$' \'AX•lf005~R. ~. J99,I, Componson o, ofmchomonla<>s in Oklahomo IK:efbula. 8'mn, Proctitloner. 14.
S4mpUng and oultur~ m~thl)ds for !he diagnosB of Trltricl1omo110J 109-110.
fM1w infection in bulls. n.,, Ve1,r/nary R,'<ord. I 3·1, o~2. 1--14> ,,"'l'i::,-.u.~. M.., OL..D'l''iEJt. r. RJ«>NI!. t.. ~ 80~"Tf. P., ~ . Ecm gt.~""'31 "an
126 >C1111Tn .•,.,•• 1sn Tauru, Coopcn11iv,.1tcme.Soulh:\!rtca Tricl:omo,uujol!WS infcctfo in \Vbl•\llaandrren. \'l(rmns
Unpubllsh~ dn1a. Dn-rg,•n.ofs~imdig Tifd."hrlft, 56, 328

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 15
Amoebic infections

J J VAN DER I..UGT

fntroduction cal and uninucleated. They contain a smooth or wrin.l:Jed

puter enocyst wall and a polyhedral or stellate inner en-
lnfec1ions caused b~ the para~ilic amoeba Enmmoeba his- docyst wall. The trophor.oites of Naegleria spp. are sllghtly
rolyrica and free-living amoebae of lhe genera Xaeyleru1, smaller ClO 10 35 µml but the cystic stage has no1 been
Acam//amoe[)a and Balam11t/1ia occur principally in hu- observed in tissue sections.s. • Recently. a new species of
mans and are rare in domestic animals. In humans, 5. his- free-living amoeba, Balamurhia mandrillaris, has been
ro(vtict1 rs the cause of amoeb iosis. which is chiefly described. but has yet lO be Isolated from nature. Both
characterized by the developmem of ulcers in the colon and. trophic stages and cysts are formed in tissues. The cropho-
In some cases, abscesses in organs such as 1he liver, while 1.oites measure 12 to60 µm and are usually uninuclearewith
the brain is specifically involved in disease caused b}' a large vesicular nucleus and central nucleolus. Cysts are
Nattgleria, Aamulwmoebaand Bc1lamwhic1 spp. Fe\\' cases of generally row1d, sornetimes oval, 6 10 30 µm, uninucleate.
1he diseases have been diagnosed in lives1ock in southern \\1th a layer of granule.~ below the cyst wall. 19
Africa.

Pathogenesis, clinical signs and pathology

Aetiology, life cycle and epidemiology
In most species, the clinical signsofamoebiosis vary accord-
The amoebae are prornzoa belonging ro the subphylnm Sar- ing to the location and severity ofrhe lesions. In me ime~ti-
codina, class Lobosea. and they characleristically movewilh nal disease a cat.a rrhal or haemorrhagic diarrhoea or
the aid of pseudopodia. 11 15 dysentery may be presem.s. 10
Emamoeba histolytica. although usually a commensal Humans infected \\1th E. his1ol,vtica are often the source of
inhabicanr of the crypts of rhe large imestine. is a pathogen infection for -animals. After lhe ingestion of infoctive cysts.
of humans and a variety of animal species including dogs, trophozoites invade lhe mucosa of the large intestine where
pigs and caule.i." Emomoe/Ja hinoly1ic;a occurs worldwide; they may be responsible for 1he de\·elopmem of numerous.
infections in humans are more generallr associated with pinpoim ro large, confluent ulcers which are often flask-
poor h~rgiene in iropical and subtropical countries. 10 shaped aud covered wilh a greyish exudate.:i. ;, 9 The tropho-
The rrophozoites have an irregolar shape, are 15 to SO µm zoites of£. histolytia, commonly occur in smaU clusters in
in diameter. and ha,,e a vesicular nucleus containing a necrotic debris or in the adjacent \~able tissuc.4· 9 Amoebic
small, central ka1yosome, and peripheral!~ arranged clU'O· abscesses. especially in the liver and lu.ngs, occur relach'ely
mat in granu les. The fine granular endoplasm often contai ns commonly in humans. but are rare in domestic animals, 1. ,;. 10
glycogen and food vacuoles harbouring red blood cells and Naeg!erfafowleTi is the cause of primary amoebic menin-
cellular debris. Cystic forms of the organism occur in the in- goencephalitis in humans and is almost invariably associated
testinal lumen but not in solid tissues.2 · 5 \\~rh a history of recent swimming, diving or engaging In
,Vaegleria and Acanthamoeba spp. are free-lh~ng. ubi- other water sports in fresh water lakes and ponds. h Is be-
quitous protozoa of soil and water.s. 9• 11 In tissue sections, lieved that the amoebae invade the cranial ca,~cy ,ia the nasal
the trophozoites of Aca.111hamoeba spp .. \\'hich are 15 to mucosa and cribiform plate, :--amral infection resulting In
50 µm in diameter. have a granular cytoplasm and a single meningoencephaliris has been reponed in a South <\merican
nucleus whfoh contains a large, cemrally located karyosome tapir (Tapiris sp.J.' 2 while sheep are susceptible to eKperi-
surrounded by a halo. Cystic fonns of the organ ism, which mental infcC'1ion 1,1th .\', /0111/eri following intrana$al
occur in animal tissues. are 10 to 25 µmin diameter, spheri· inoculation of lhe o rganism. I"
316

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 Amoebic infection~ 31 i

-·
figure 15.1 Cerebrum of a sheep. ence;inalomalac,a show ng G,tte:
cells. and 11ophozo11es and cysts resembling those of ~mnamoe/Ja soc Figure 15.3 li«;her magn1ficat1on of a cys1

dogs.1. 16 A multifocal necr0granulomatous meningoen-

•
cephaliti~ and cerebral vasculitis associated \,ith 1rophozoi-
les and cy~1s morphologically resembling Acawliamoeba
•
•• spp. have been reported in a sheep in South Africa (Figures
15.l to 15.3 ,.' 8
Ba/amurhia infecrions ha,·e been diagnosed in a sh~cp,
lowland gorilla (Gorilla gorilla,. orang-utan (Po11go pyg·
maeus). and a mandrill (Mandril/us sphinx) as a cause of

. ~
.
menmgoencep

Diagnosis
h a1·II Is.6. 19. e

-
The diagnosis of amoebiosis is based on demonstration of
the parasite in faeces. or in biopsy and necropsy specimen•.
and on positive serological tests. for example agar gel diffu-
Figure 152 Highe: magnification o' a :1eoho2011e sion. cellulose acetate diffusion. and indirect Ouorescem
amibodyiests. 5· 10
In humans, Aca111hamoeba spp .. in addition to causing a
gr,mulomatous amoebic encephalitis in immuno~up- Control
pressed or chronically ill people. can give rise to keratitb,
uveitis andgranulomatous lesions in various organs.~· 6 7 13 Melronldazole is lhe drug of c:hoice in amoebic dyscnterv.
Acanthamoeba spp. rarely cause pneumo11ia in canle1 14 Control mea~ure$ in humans cnnslst of improving environ-
and necrohaemorrhagic lesions in variou~ organs of mental and food sanitation. 5 · 10

References
.,em. K -.; .. l9U3, Hnnmn.nrK"llosts in caln.w; , ·.:1etinlll)' ,\lc-dklnei:,uu,/1 0 C..\NJ,U;.UJ. 1•.J,,. \'f'J4CU.~UT. I,. CUXM~GK\\t, \U,., .:C\'~')\'0\'Mtt,\ G..~. h197.
Amnwl Ch,,iehM. 78. i'·i.9-7'"50. Amoebic men,ngoen«phwltls c•u.ed by Balo1>111tl1i<1 mn11cfr/1111ri, ,nan
2 "1MCJI, K. ,. ,., 8001>1~. T., 1:ti'6, Amoeht,~. /u~ >,;RlJr.Jt. t,r.. (('d.J. Pt,rtu:ttir tJtn~·llf:\n . .At1.11mlinn \ 't..•ti:rlull()· JtmmaJ, 75. .97-100.
Pr~to:on. Vol. II NcwYnrk:Academic l'r<S>. 7 DUMA, ft.I., 1SJ8. :\mrtt.•blc infccHons of the nel'\'<IU~~'Ystcm /,,: V1;\Kl :,.. P,J
J .\\1..!t'>. .: ..,1 .. oru.1ws.. LH, &G.\~'""6:R. t .,s.. 197:!. .,.\canlbm1\oeb?osf> in~ "BllU\'S. ~-'' .. 1e<1,,. llc1111/Mok a/ C/i,1icnl sn,ro/Qg;', P;u-1 Ill
doi: 1·,1rr111or,• Pmliotl)f(). 9, 221-226 Am~1crtlan,: EIScvwr Konh Hollnnd Bibn1edlcal Press.
ii n,1ou 11 , .).; " v.,x JJR£cUMt L "4A. i98S. Tlu: alicmm!ary ,r~u·m 111; 1un&, 8 f-Vl'"Tl.Al&,. u:.,. \\11;~. ~L. RL\O, W,Jr... tD\\ARDS, J ( .. VI-S\'h\'A~. G.,
~.\'.f, Kt.'-~1.D'i',. Jt,C. "rA1.,1F..R. s ..feds.. /'(ltholog;,· of()Qme,m: inuncils. 1g9-1.Aml.'b1~ mrnmgoonct•pho.htb, in a s:h~p./or,rnu/ oflht'.o\mt.•ffCQ11
Vol. II. 31d odn. Orlando: Madffllic P,..,,., \'ftt"rlltO,j .\!C'(/{ml .4.#0('(lttiOII. 200. 363-365
s Jrt.l\n.R, •.c ... ,o,t,., u.e,. ,..cu-PP, 1.,.... 1~4. Cllnii:nl Pt,ttuitt1l(l((,•. 9th \•dn. g OR1fH' , .• )978 ra.1hog~nfc fo..'\:•ll\"ir.~ .}:fflOtba~. /ti~ ):$tUEM. J~I".... h.·d J.
Phllad•lphl• Lea & l'ci>fgc,. l~tl,,u(:tc l'roro:..on \'ot. JI. ~t•w York; .'ic.."ad:1.1mlc Press..

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318 SliCllru- rwo: Pro1oi9al diseases

ro M...uuur:s, J•• 198~. AmoebtoS't(: A l't."-'ie\,_ Journol ofth.-Roy,tl s«rcryof Ameble mening~nce1>halill$e:1used b)' .l,:amlrnmo;,bil tn.ttaltnm ln a
,\ftd/f/nc. 7S. l90-l97 dog. Jtn,mat o/1Ju•Amtrfcan l'cttTUUJI)' ,\ltdita.l Assotf(11fon. t8i.
11 u.,•1s:1,. :,,;,o., i.985. Vt:tnnary Proro:oology. Arru.•i, tow.a: to,,.-n Sune- !151-952.
Unlwrsil)• Press. lj Sl,\IPS()S, (.1'.1 \\1l.J,..\1Jfr, F.., :-:-ML, J:.C.. STIJ\'[..,"f., A.It. k YOUN<:, )LD., 19$2.

12 t..01.A.NO•ALi.\R.CON, f., 8AAUtrY, G.A., HOUSeH, HS. k \1.$\'li:S\'.AftA, (';,$.. 1997, Experlnienull Nn•glerwfowwrl mcnlngocnceph:illtls In sheep, Liglit and
Prunary •mcblc mcnfngocnccpholltfJ. duo ,u .Va,,glcr/i,/01•/cr/ in• elt!'ctron m.icro.scc,pic sludies.Alttt'r/can Juur,u,t ()/ \lett!rl.nory Resea.n:IJ.
Soulh Amcrlam tapir V,torloa,y Porholol()', 34, :?39-2~3. 43. t5.;-15'i'.

13 """TIM,,. A,J., 1980. l<Aranthmnotbtl "nccph;illds an opportunl,tlc 18 \"A."\ l)i:Jt l.UC.T. J.f. t,, VA,'\t DIR ~tlWE, Jl.f•• t990- Amoebic
lnl'e<tlonl ,W11roll,~·. 30. 567-~74 m•nlnll""nccphalit!,, In a sheop./oumal ofthe South Afric,111 Vet~rlt:ai,·
JW«(at/011. 61. 33-36.
1" :.it:cossiu. £.E.. CAJcsER, •·.,t.41 ..,,-,.i;:, p1 •• 1968. Hortmannellcsls in a
bull. PothOIOflia VtUrirrnrla. 5. I-& 19 \'IS\'t..,V..\AA. t, ...... ~<:11u~'TER. Jt.t_JJ, M,\RTINf</., .q.. 1993. &tftmwthia

l5 ,1EHU10H~. u,. • w\l.LDORF, v.. 1988, Ufe cycles, It,; ,,m1utoRl'l>.11 .. (cd,), numdrlll11ris. ~.C.. N. Sp .. ogcnt ofamoblc mclllngocnccphilllti• in
/><1msl10/ogy III Fae,,,,. Berlin: :.vnngor,Verlas, humons and 01hcr nnfmols. Joumlll ofl:11knryot£c Mlm,blol~. 40.
504-514.
IG Pl.!.-UlC(. pt,, W>Wl!LL. lf,.io.,, C:H,,SOU"R, F.\\',,., \'1$\-f'SVAfci\. G.~, 1985,

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 16
Coccidiosis
Synonym: Koksidiosc (;Vrik.)

CG STEWART A ND B L PE!':ZHORN

Introduction tests have also been used in cartle. sheep and rodents.38 The
developmental cycle of the parasite occurs intracellularly
Coccidiosis is a prolozoal disease whJcb occurs in many within the host (see below), while the final infective fonn.
mammalian and all domestic Liveslock species. It is caused the rnarure oocysr. is passed out to 1he external e1wiron-
by infection with species of the genera Eimeria or lsospora ment. Oocys1s of sheep and caule can be distinguished by
and is characterized clinically in li\·eslock by enteritis. fol· their shape and size, the appearance of their surface (par-
lowing lhe invasion of mucosal cells of lhe gascroincestinal ticularly the colour and texture), and the tllickness of the
tract. Subclinical infections are frequent. The disease is rela- wall. 37 Variation of oocyst morphology occurs within spe-
tively common in sheep, goats and caule. cies. however, ru1d there is a tendency for 'bi7.arre forms· to
Coccidfosis occurs universall)'. but is most prevalem occur in sevcrcl}' infected sbeep.7 In a typical spomlared
where animals are subjected to overcrowding, unhygienic oocyst of the genus Eimeria (Figure 16.l ), the oocyst wall is
em'ironments, or where other stress factors are present. composed of one or L\\'O layers and mar have a micropyle
Eimeria spp. are important causes of lhe disease in cattle, whicb in turn may be covered by a micropyle cap. An oocyst
sheep and goa1s. Coccidiosis is rare in horses34 and in Eur- or sporocyst residium, consisting of material remaining
ope and North America is an important disease of pigs. 68 AJ. after the iom1ation of these stmcLures. may be present. A
though a number of £imeria spp. are commonly found in sporocyst may have a knob, the Stied a body, at one end. 30 •34
pig faeces. clinical disease is caused only by infection with At spomlation oocysts of the genus Eimeria develop int0
lsospom suis. Coccldlosis due to /. suis has not yet been de- four sporoc}'Sts, each containing two sporozoites, while
scribed in southern African countries, but it is associated those of lhe genus /sospora have two sporocysts, each of
with clinlcal disease of very young piglets in many other which contains four sporozoites.
pans or the \\•orld. Coccidia of livestock are primarily intracellular parasites
Coccidia are generally host-specific, and, in the gut, spe- of the gastrointestinal tract, but in some animal species tl1ey
cific to a panicular location. Host specificify is more strin- may be round in other organs such as the liver or kidneys,34
gent in Eimeria than in lsospora infections. or even the utems.41 Development in the coccidial species
Economic losses from coccidiosis due to monaliry, poor mentioned in this chapter occurs by means of an initial
performance, a11d the cost oftreatmemand prevention may asexual cycle, followed by a sexual one. B01h 1ake place in
be considerable, especially on stud fanns and in intensive the same host.
calf or lamb rearing systems. There are three stages in the life cycle: merogony (lhe
asexual cycle), gametogony {the sexual cycle) and
sporogony (Figure I 6.2). The first two occur within host
Aetiology and life cycle
cells, while sporogony occurs outside the host.34 Following
Coccidia are classified as protozoa in the class Sporozo- ingestion of infective sporulated oocysts by a susceptible
asida, subclass Coccidiasina. and suborder Eimeriorina. host animal, the sporozoites are released through the ac1ion
Both morphological and biological characteristics are of the digestive juices. and. depending on the parasite spe-
used to identif)• the species of coccidia.3'' These criteria re- cies, enter epithelial or other cells of lhe mucosa of the gut.
quire subjective assessmenc and are therefore not always Here ther become meronts and multiply by merogony
completely reliable. For more accurate identification com- (schizogony) to produce merozoites, the number depending
puter-assisted image analyses have been used in cattle62 on the species. These enter nearby epilhelial cells in the gut
and pigs. 5~ Random amplified polymetase DNA (RAPD) and again mulliply by merogony. The number of asexual
319

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320 s,:a,o:,; m-o; Protozoa! diseases

S0crocys1 resldium

Sporocyst
Sporozone

Figure 16.1 Structure of a sporuta,ea E1me11a

oocysL (Adapted from Levine 198:l"l

generations which occur remains consranc for each species. E. weybridgensls are more elongate and lie lengthwise in the
Merozoites of the !,1st generation infect new host cells and sporocyst. while those of E. cmndaltisare more ovoid and lie
become gamonts. thus initiating the sexual stage of the life ar either end of the sporocysi. 37
cycle. Some of the gamonts develop into macrogameres The meroms ofthenon-pathogenicE. gilruthi are \·isible
without further multiplication, while others divide b) mul- to the naked eye during necropsy as whitish nodules (300 to
tiple fission (asexual) to form a large number of flagellated 900 µm in diameter) in the mucosa of the abomasum of
microgameres. These microgameres are released and pen- sheep and goats.34
etrate the macrogametes ro form zygotes. A thick encom-
passing wail is formed around each zygote, which is then Goats
known as an oocyst. \.\1hen the cell is ruprured, the oocysts ).,lost of the coccidla species which inf«ct gears are. in spite
are released into the lumen of rhe gm, and are excreted in of being morphologi,cally similar. distinct from those of
the faeces in an unsporulated state. If environmental condi- sheep.~9 The more important species to cause dlsease in
tions of remperature, oxygen supply and humidity are suit- goats can be identified bytheirclose similarity ro cheircoun,
able, oocysts sporulate and sporocysts containing infective rerparts in sheep {Figure 16.3). These. with their ovine
sporozoites are formed. This process takes berween two and counterparts given in btackers. are: E. a.rloingi (E. bakue11-
five days under optimal external conditions, which include a si.s). E. chrlscen$eni (E. nhsmn). E. hirci CE. cra11dt11/is), E.
temperature of24 co 30 •c. 10 jolchijei.•i (E. gmnu/o,m). E. alij e11i (E. parva}, E. kocharli CE.
Coccidiaare haploid throughout their life cycle, except in imricma). £. apshero11ica (£ faurei), E. ninakohl_vakimovae
the zygote stage. The firsr sporogonous division is meiotic, (E. ovinoidalis), E. capri11a {no counterpart), and E. <:a.·
re-establishing the haploid state. Oocysts administered to prol'ina. which occurs in both sheep and goars.25 Eimeria
aberrant hos1S will excyst and beginmerogany, but the) dis- capri11a oocysts are 27-40 x 19-26 µm in size. have a micro-
appear after 48 hours5° pyle cap and can he distinguished from che other caprinc
coccidia which possess a micropyle cap. by their very much
Sheep larger size. Eimeria apsheronica differs from £ capri11.a in
The most important species that cause clinical disease in that it is ovoid and has a small micropyle.:l6 The most patho-
sheep are Eimeria ahsata, E. ouinoidalis and £. /)(lki;ensis.33 genic species are E. 11i11akohlyakimo1,ae. E. chrisce11se11i and
Other species (Table 16.1) are less pathogenic but may play E. arloingi.29
an augmencative role as mixed coccidial infections are rhe
rule i.n field outbreaks.55 Cattle
The characteristic feanires of the sporulated oocysts of At presem there are between l9 and 22 species of Eimer ia
the most common coccidia occurring in sheep are shown in which have been identified in cattle. Most of che$e are rela-
Figure 16.3 and Table 16.l. Oocysts of E. weybridgensis tively non-pathogenic. and field outbreaks ofcoccidiosis are
are similar in size ro £. crandallis, but che sporowires of generally caused by mii,:e.d lnfections of E. zriernii, pla}~ng

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0

Figure 16.2 • .te cycle of a

cocc,a,urc sileeo
· ~sooicroite
2 ~mero14
l = merozo,te
4 = mitrtigamete
S = 'N!trogamate
6 • zv;;o:~
7 = urs~rulatei! occyst
8 = s~o·ula:e-:1 oocyst

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322 ,, 1111,s ,-,·o: Protozoa! di~ea.~

E pa/Iida E ah1ev1(G) E nmakoh1Yok1mov~e !GJ

(S&G) E parva (S) E. ovinoidalis 1SJ

E apshe1011ica (GI E. puncta1e E.caprovma

E faurei(SI (S&GJ !S&G)

E h1rclfG) E weyl111dgc?11s1s !SI E. Jolcni1evi1G) E roµinalGI

E c,andeflis (SI E granulosa tS,

E arfo,ngdG
E .:hos1ensem !GI
E bc~ue~Sls1St
E a11sa1a1s·
Figure 16.3 The sporu!ateo
oocvs,s oi cocc1dra from gca,s tGl E kachatlilGI
and sheep IS) (Adapted 'rem lel'ine. E inrrice1e ISi I I II I I I I
i9Si"I 20µm

the major pathogenic role . and £. /1ovis pln)'ing a lesser one. mero:wites, 12 occur in the last pan of the small intestine of
.Eimeria nlabamensis has been reported 11s a cause of coc- the lamina propria near the muscularls mucosae. The sec-
cidiosis in northern Europe. Eimeria t111/.J11me11sis and £. el- ond generation of meroms ( 13- 18 x 13-18 pm in size. each
l/p.(()idoli$ have produced diarrhoea In experiments. 12 containing II mean of 32 merozoitesJ and gamon1s de\·elop
The oocysts o[E. zuemii <Figu re 16.4a) are subspherical in epithelial cells of the caecum and colon.05
to O\'oid. measure 12- 29 )( 10-2 l µm and have no micropyle Oocysts of E. lxwis (Figure 16.4b) are ovoid (23 -'.!4 x
or residium34 (Figure 16.3). Large. first-generation meronts 17-23 µml and h:we an inconspicuous mkropyle but
(l 15-225 x 78- 202 µ111 in size), containing about 120 000 no polar granu le or residium. The sporocysts have an

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 Coccidiosls 323

Table 16.1 CharattetiStlCS EIMER/A SPP OOCYSi SIZE PRESENCE MICROPYlAR OOCYST POLAR Si1EOA SP0ROC¥ST
of sporulateil OOcYSts OF CAP RESIOIUM GRANULE/S SOOY RESIOIUM
lµml
occurring •n sheep. MICROPYLE
(Adapted trom
Levme, 19853'1 Ea~u1 23-48 X 17-30 •
E i;randallls 18-28 X 1&-20 't
f rauro: 25--37 X 18-28 •
£ granutase 22-37 X • 7-26 ._
f inrricaia 39-53 X 27-47 •
f /Ja.<116nsis 23-36 ~ 15-2; -t •
£ 01mo,rJa/is 15-30 X 13-22 'T +
tf]8lftda l2-20x 8-15 t
E par,a 12-23 X 10-i9 •
E punc1ata 18-28 X i&-21 + T + 't t
E wev!Jr1dgens1s li-31 ~ !J-19 +

inconspicuous Stieda body and a residium. There are rwo uansponation or when animals are maintained under un-
ase.xual generaLions. First-generation meroms develop in sanitary conditions. In well-managed animals there is a bal-
endothelial cells of the lactcals of the vii.Ii in rhc caudal pan ance between the hos1 and the parasite.
or the small intestine. They reach a size of 207-435 " Young animals become infected after the ingestion oi
134-367 µrn. contain 55 000 to I iO 000 merozoires and are sporulated oocysls and they in turn excrete oocys1:s, often in
macroscopicaJlr visible in the mucosa as whitish spots. Sec- large numbers, in their faeces. Hot. humid, u11hygienic con-
ond-generation meroms measure about 10 x 9 pm and con- ditions, such as those that frequently pre\'ltil in overstocked
1ain 30 to 36 rnerozoires. They. and the gamonts. occur in feedlors. pens containing srraw bedding for raisillg lambs.
epithelial cells of the caecum and colon.31 kraals and irrigated pastures. favour the survival of oocysts
and result in higher infection rates than is Lhe case u11der ex·
Pigs tensive fanning systems. lnfection may spread 10 animals
lsospora !tlis occurs in the small i111es1ine and sometimes in housed in separate pens by the airborne 1ransfer of' oocysts.
the colon of pigs and is the causacive ageru of porcine neo- Most animals are subcli11ically infected and as Lheir immu-
natal coc;cidiosis.35 Oocysts are spheric;.)! 10 subspherical. nity is not absolute and continual reinfection occurs, they
li-25 x !6-21 ;1m in size, and lack a micropyle. Stied a bod~. shed small numbers of oocysrs in their faeces for long per!·
polar granule and residium;i.; (Figure l6.5). Excretlon of ods. Spomlated oocysts are very resistam to adverse em~-
ooc~'StS in the faeces starts four 10 five days after infection ronmental conditions and may 'O\'envinter' on pastures
and continues for one to three weeks.3S which then act as a source of infection when favourable
Eimeria spp. whi'ch commonly occur in pigs are E. tie· climatic conditions remrn. 16
l1lie.ki. f.. 11eorlebliecki, £. pQrci and £. s11is (which are all
smooth-walled). and £ scrabra. E. polim. E. perminura and Sheep and goats
E. spi11osa {\\'hich are rough-\\'alled).68 All Eimeria spp. of The disease is most severe in lambs rwo 10 eight weeks of
pigs are <;onsider.ed to be either non- or only mildly patho· age24 and in lambs two to three weeks after weaning. Sheep
genie and are no1 significant causes of disease i11 pigs.3 ; entering feedlots, or after experiencing severe stress Such as
that caused b}' uansport or inclement weather are prone to
Horses developingse\'ere disease. 16 The most important specie; are
Eimeri(1 /euckarli. \\1hicb is occasionally encountered in 1;·. 01,inoidalis. E. cmmlallis and£. ahsnra.J
horses. ma~•be pathogenic 2 but is usually an incidental find· ;\,lost sheep and goats pick up oocysts as young animals
ing. The rnlcrogamoms are 300 " IiO µrn in size and so either from comamina1ed pastures or from Faecal contami-
are visible to the naked ei,-e. They occur in the small intes· nation of the reats and udder of their dams. The parasite
rine.:i.. population is increased enormously during the firsr passage
in susceptible lambs putting later-born lambs at greater
Epidemiology risk.9 Sheep and goats usually become infected ,,tjth a num-
ber of Eimeria spp. and the paiencr of oocyst production ex-
Coccidiosis is essentially a disease that is prevalent in young tends o\·e.r several months. Mixed infection; prolong the
animals under intensive managemem systems. Older :mi· patency and increase the number of oocysts produced.6
mals are occasionally clinically affected. Disease usually oc· Faecal oocyst counts may become high in apparently
curs when the resistance of the hos1 is lowered after stress healthy lambs.r
such as that re..~ulting from o,·ercrowding. weaning and The presence of the more pathogenic species, such as

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 a

figure 16.4 Efmeria ,uemiilal and :imeria bovis (b}OOC'1m lr~m canle Figure 16.5 lsgspara suisoocys:s frcm a plQ

E. ahsa1t1 and E. 011i11oid<1/it, plays an imponanc role in the b) weaning or cold weather. may be imporrant in initiating
production of o,•ert disease in shecp.33 outbreaks.5? The disease may occur in young canle kepi
Coccidiosis is often a serious problem in feedlots. Spe- under extensive farming conditions wh,.m high oocyst con·
cies of coccidia co which animal:, in the feedlots have not 1amina1io11 occurs ar()und warcring poims.
pre,iously been exposed mar be introduced 1oge1her ,,ith Eimeria tllabamensis is reponed to be a i>roblem in
newly purchased animals. Conversely, non-immune ani- calves in Sweden and German).G9. ' 1 four to seven da~-s after
mals may be placed in a pen containing i11fec1ed animals ol' being placed on contaminated pastur~. The source of In-
which has been contaminated with ooc~·sts. Clinical signs fection is from oocysts that have overwintered on pas1ure70
may develop 10 to 20 days after a ne,~ lot oflambs are intro- or comaminated hay.72
duced. Mortalities In feedlots may be high and arc often The presence of subdinical coccldiosis may interfere
nssoci,.uc<l wirh sudden outbreaks of the disease. Once a pen wl!h rhe response 10 ,·accinarion against other diseases.~ 1
has become infected. the oocysts may remain there for a
considerable period, creating a h07,ard for any ucw intro· Pigs
ductions as well as for animals al ready in the pens. Feed Unsporuta1ed oocysts of 1- suts sporulate very rapidly
trough, mar become contaminated wich faecal pellets (\\ithin I 6 hours at 30 •q under suitable moismr1: and tem-
which .ire lnadvenentlrconsumed and. in this way. the pro- peramre conditions, whereas £/merit1 hl)Jl. take from 8 to 13
duction of massive infections or reinfections is facilitated. days to sporulatc_r.a Day-old piglets are more susceptible
High con.ccntrations of oocysts in the pens may resuh than month-old pigle1smaincained free of/. s11is infection. 68
when animals are penned only at night. Good immunity develops after infection. /\dult sows do 1101
Very 1,•c1 summers are kno1,11 co result in a bigb preval- appear 10 shed/. suis, so the source of infection for piglels is
ence of coccldiosfs because of the survival of larger num- unkno\\ 11.68 ·me most li kely source of inrection is contam-
ber~ of oorysts and siressing or che animals caused by the inated farrowing crares.:'•
we1 condirions.

Cati.le
Pathogenesis
Clinical ct)ccidiosis is primarily a disease of cal"es between Factors which play an importam role in the pathogenesis of
three \\'eeks and six months orage. although older animals may the disease include the pathogeniciry of the coccidia species
~how ,igns of infection. Oven disease occucs when large nmn- concerned. environmcmal conditions. size of the infective
bers of oocysts are ingested while !he ingestion of smaller dose. age of the host. number of host cells destroyed. loca-
uumbers of oocybrs leads to subclinical infection followed by tion of the pa.asile in !he tissues and the presence or ab·
the development of Immunity. In 1hesummer rainfall areas or sence of acquired immunity 14
·South ,\frica the praclice of removing cakes from d1eir dams Some species ofcoccidJa are parhogcnic whcr-eas others are
duriJ1g !.he winter momhs ofJune and July and placiJ1g them in not. The most important pathogenic Eimeria ,md lsospom spp.
pens 10 l'aciliraie weaning commonly results in outbreaks of of livestock infect epllhellal and other ceUs of the mucosa of1he
coccldlo"'is.o.; Ourbreaks occur despite the fact that under the gastrointestinal tract. the site along the length of the gut 111ry-
dry \\inter conditions oocyst contamination is likely to be fairly ing \\i.th the species. As the number of ase.'<ual generations oc-
low. Outbreaks of coccidio,ls in winter have also beenreponed cuning is constant for each species. infectfon is seif-limitfng
in Canada:•R h would appear that stress, $UCh as that produced pro\:ided lhai reinfection does not occur.34 Many of the

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 Cocddlnsis 325

aifecLed cells are des1royed or 1heir runction impaired.31 Clinical signs

meront~ probably being rcspon~ible for most or theceUularde-
stnicrion. In SC\'C.'rt' infections there is an acute inflammarory Sheep and goats
reaction. Capillarie~ in the denuded lamina propria may The ons<?t of diarrhoea is usuali> the nrst sign of Illness.
haemorrhage. leading to hypoprotcinaemia and anaemia. Sec- Affected animnh pass sofl. unpelle1ed faeces and later <1
ondary bacterial inrec::1ions may exacerbate the lesion. All these watery diarrhoea. Blot1d may be present in the faec<.<s but
changes ~timulnte ,lJ1 increased rate of peristalsis and result 01 it is less common in sheep and goa~ than in cattle. The
diarrhOE'a. The reduc-ed absorption capacity of the parasltizcd perineum and hind limbs become ~oiled and Lhis m.1y pre-
muco~a results in h)5,: offluidi; \1a the imesrine and decreased cipitate 11~ ,trike ·\norexia and dehydrafion develop a few
absorption of electrol}'tcs such as sodium and potassium.. dars after the commencement of the diarrhoea . \\ith result-
CQmpensatory increase of abwrption in nonnal ponions of ant los;, of bod) mass. Tlw,.e arc followed in St'\'erc ca-.~ by
the gut may redeem this lo~. 11 Dehydnuion and acidosis due depression. recurnbenc~· and dea1h.
to the loss ofbicarbonn Le ions in the faeces, as well as anaemia, The morbidityrangPs from 10 ro50 pen:en r and 1he mor-
hypoproteinaemia, and shock combined \\¼th secondary bac- 1ality may be up Ill Hl per cent. In \nl{ora goat !locks in the
terial infec1ion. are lhe main causes of death. Easwrn Cape ?rm·ince of South \Erica morbidity rnn reach
The effects of coccidiosis due 10 £. zuemii :ind E. bo11is 100 per cent \\Ith mortnlltv as high as 50 per cent.= In the
are mainly the resu lt of damage to rhe muco~a of the large sandy. a rid eastern parts or Namibia a disease known as
intestine caused by the second-generation meroms and 'bok,iek1e· (goat disease) i~ due 10 a combined i,ffect of
SC.'1.'Ual smges:•~ Second-generation mcro nts and gamoncs coccidiO$i< and infestauon \\1th the worm S1ro11gsloirles
of I:.. ~uemii arc simultaneous!) present in the caecum. pa11lllos11$."' The cour,e varil') from one 10 two weeks and
colon and rectum. and it is 1herefore difficult 10 determine 1he disease is manifested hy the cliniC"',d signs or ataxia,
which is 1Jie more pathogenic. ;>.ta.ximal de,aructlon of the opi~thotonos and convulsions. in t-onjunction with \'a.nous
epithelium of the large intestine occurs in cattle just before degrees 01 emeritii,.
the peak output of oocys1s.61> Thi:, resu lts In a reduction in
thl' absorption of water and sodium and chlorine Ions Cau.le
,,~th a consequent rapid lo&> in bodrweight. Animals dying The incubation period is usual!} about three weeks. but may
at this stage probably do so from dehydration. There are vary between a wee!.. and more than a momh.i"
marked decreases in packed cell volume, haemoglobin The predominant dinical sign is diarrhoea. which mny
levels and erythrocyte cou nlS in severe cases off.. ;:uemii be poorly developed at first, bm becomes more watery and
infection. This is in co111ras1 10 £. lm11is Infection where linally hal!morrhagic. As the disease progrt':.ses there is an
there is an increase in packed cell volume and increase in the nmoum of fresh blood in the racces. and
haemoglobin.ll<i blood clot~ may be present. ,\fter a few day~ the amount of
rmmunity develops after infection. but the determinant~ blood in the face-es may dimini~h. being repla~d hy mucus
of protective immunlt1 have yet 10 be elucidated.00 The cel- and muco-flbrinou~ l'Xudatc. Later the dischruge>s become
lular immunity which usually follows the development of blackish-red 10 black.ish. or gree111sh-block and foul-smell-
thl' merom stage is mo$t important as far as prote<:rion of ing: shreds of mucosa may be present. The diarrhoea.
the host is concerned. while 1he sexual stage< are poorly which 1113}' be proiuse .ind foetid, is often accompanied b\
lmmunogenic.611 Subclinical irtfoction a1 a youiig age resuhs tenesmus and pronounced borbor}!{ITli. The animal mar
in the development of sptc:ific immunil) early in life. Thb constantly grind it~ teeth. :\bdominal pain I~ usually
immunity may. however; break down under condition$ of present and, if the te!lcsmus is intense, prolapse of the rl!'l'·
stress such as during Lransponation. esposure to climatic 11,1m may occur. There is often a fc\'er an increase in the
e.,1reme-,. changes of diet and overcrowding. or when respirator} rate and loss of appetite.:r,
massive infections occur. 1:1 Drugs mny also lncel'fere with During th<' course of the di~ease. scvert>ly alfoctcd an-
the developmem of immuniry. Lambs exposed 10 an imals bec:ome extremely emaciated and anaemic. ,\fier
aureomyci11-sulphameznthine supplemel1l did not develop about 8 10 12 days. they eilher sho1\ ~igns of impl'O\'ement.
immunity when fed 500 mg/iamb/day for 35 days, but were or deteriorate and die.~"
able 10 develop resistance when fed a lower dose of 100 mg/
lamb/day. 1 lmern1i11em or continual reinfection enhances Pigs
the level of imm unity. The immunity 1ha1 de\-elops in Cocddiosis usually occurs in 5- to l 0-llay-old piglet,. and
response, 10 an Fimeria mfection is species-speci.fic.60 is charac1cri1.ed b} the development of diarrhoea, dehy·
?rotection against iniection has been obtained by drat ion and loss of condition. l'ael·e~ are yellO\\' 10 gre) and
immunization of lambs \,ith £ cmndallis and £ 011i11oida- do not contain vislble biood.n \'omltlng may occur. On
li,.!I ii and in calves \1~1h E. alabamensis.71 A 'tricklc- some farm, all liners may be infected. wher('as in Olher;
infection' appear~ 10 be the most effecli\'e mean~ or on ly a few liuer~ become infected . The mor1ality is u-.ualiy
producing immun ity. less than 20 per ccnt.',s

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326 s.cno" 11,•o: Pro1ozoal disease$

Figure 16.6 Flat to dome-shaped. white focr(consisting of coccidial Figure 16.8 Nacrotizing colitis in a calf
parasites) in the mucosa of the small ,ntestine ol a shu p

Figure 16.7 Various stages of &o::cidia in epithelial cells of tile small Figure 16.9 Haemorrhagic proctitis in a ta!f
intestine

Pathology Sloughing of the. intestinal mucosa may occur, sometimes

\\ith ulceration. Haemorrhagic enceritisofthecaudai part of
Sheep and goats the small intestine occurs in the case of E. 011inoidalis in
Decreases in the levels of serum albumin and plasma alka- sheep and£. 11inakohlyakino11ae in goats, and blood may be
line phosphatase coincide with the onset of dlarrhoea.5 The prese!lt iri the faeces.29
,mimal may be cachectic or even anaemic. Microscopically there is disorientation of the villous
The location of rhe lesions in tbe gut, and to some e>;1e111 archicecmre. a reduction in epithelial cell height, and a
their appearance. varies With the Eimeria spp. concerned. decrease in the width of the brush border.57 The villi may
but most infections are mi.xed.11 be shon or absem and may contain proliferative epithelia,
In masc natural cocciclial infections small. flat to dome- with various scages of coccidia being present (Figure 16.7).
shaped, whice foci, l 10 2 mm in diameter, each consisting of Malabsorption may occur if these lesions are widespread.29
an area where there is severe infection of epithelial ceUs,
occur In the mucosa of the small intesri1,e (Figure I 6.6). Cattle
These foci, which have a diameter of up to 300 µm. may also The sodium and chloride ion levels In the blood decrease
be cliscernible from the serosal surface of the gut20 and are about 18 days after experimental infection and reach their
due to the schizont stage of E. ovinoidalis an.d E. ahsaw in lowest le\·els seven days later. 66
sheep and Ii, 1ri11akohlyaki11ovae and E. c/lristense11i in The meroms of'£. bo11is mar be seen as small white spars
goars. 29 Macroscopically visible, discrete opaque patches In the mucosa of the ileum, but those of E. z11emii are not
may also be see.n In the mucosa in£. bakuunsis infections be- \iSible rnacroscopically.12
cause of the presence of groups of heavily parasitized intesii- In severe cases, fibrino-haemorrhagic 1yphlocolitis is
nal dlli. These may appear as raised patches or even develop typically prese111 as Is. in many cases, procritis (Figures 16.8
into polyps.22 Affected areas of the intestine of severely af- and 16.9). The terminal ileum may be aJfected.w The con-
fected animals are oedematous and thickened. and there may tent of the caecum. colon and recrum becomes semi-fluid
be focal or diffuse congestion or haemorrhage in the mucosa. and bloody in lhe early srages of infection. During the later

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 Coccidlosis 32i

stages sloughed mucosa and copious amounts of blood and a glass microscope :.lide. covered v.;tb a cover slip. and eX-·
fibrin may form fibrino-necrotic pseudomembranes. The amined under a microscope.
content of the large imestines has an unpleasam smell. The Species idcmification of the coccldia requires some ex-
mucosa may contain erosions. and both it and die submu- perienct'. bu.t by comparison wilh tex,book dra\\ings (Fig-
cosa are oedematous and congested and contain petechiae. 11res 16.3 10 16.5), some of the more common ooc,·scs of
The mesenteric lymph nodes are often enlarged and sheep and cattle can be distingui~hed:1 Oocysrs may be
oedematous. more readily differemiated after :,porulation has occurred.
Microscopically. the mucosa of the caudal part oi the Sporulation can be induced h> mi-ting a small sample of
small intestine is invaded by first-generation meroms and faeces with a 2 per cent aqueous potassium dichromate so-
may be infiltrated by a few lymphocytes and eosinophils. lution and allowing it 10 stand at room temperature fonhree
but. in general, the le..~ions are mild. 12 The lesions caused by to four days..;• Accurate identification is a task for a parasi-
second-genera!lon meronts and gamoms. on the ocher tologist.
hand. a re characterized by extensive changes. 12 In the case Estimates of the number of oocysts in faeces can be car-
of£. zuemii infectinn~ these are present in the large il11es- ried out by the modified McMaster method.•·1 However, it
rlne. In severe infections almost all the epithelial cells of the may be difficult to interpret tl1e results as high counts can
alrected pan contain parasites in various stages of develop- occ11r in apparently healthy animals. Low counts may be
mem. l\lany afl'ccted cells lose their integriry and become presem in diseased animals as the majorfry of the parasites
detached. while the crypts become distended by oocysts may not have progressed to the oocyst stage at the time of
and cellular debris and the lamina propria is congested and tl1e examination. Sb In these cases. smears of faeces should
infiltrated with plasma cells, lympho<:}1es. neutrophils and be made and Mained with Giemsa to demonsrrace the pres-
eosinophils. E.xtensive haemorrhage occurs in the tissues. ence of merozoire.s.3"' Sheep tend ro ha\·e higher oocysr
Capillaries in the lamina propria denuded of epithelium counts than goats.30 At necropsy. either scrapings from le-
may be exposed 10 the intestinal content t?ecause of the sions or rissue sections of the Intestine may be examined mi-
sloughing of the epithelial cells. croscopically for che presence of meronts, gamoms or
oocysts:l-1 :Slixed infections usually occur.
Pigs Healthy lambs between four and eight weeks of age may
Fibrinous or fibrino-necroric enteritis in the caudal pan of have vei, high faecal oocyst counts (of the order of
rhe small intestine is present in some faral cases. Micro- 500 000/g or higher). but after chis age there is a progressive
scopically. severe destmction of the villous epithelium of reduction in oocyst numbers so that by five months. OOC)"St
the jejunum and ileum. and marked atrophy of the \~lli are numbers are similar ro those in adult sheep. This makes the
evident. Endogenous stages of/. s11is are apparent in the vil- imerprctation or oocysr counrs in relation to their clinical
lou~ epithelium.68 slgnificance vei,1 difficutt.S!>
Coccidiosis should be suspected in yow1g cattle whose
major presenting sign is one of a haemorrhagic diarrhoea.
Diagnosis
As most field outbreaks of coccidiosis are due 10 £. =ttemii, 12
The hi~tory of the case. clinical signs and necrops) findings. the oocyst ~houid be identified (Figure 16.4). fn some cases
as well as the demonstratlon of the parasite. are all relevant diarrhoea may begin a da)' or so before oocysts can be de-
when diagnosing c:oc:cidiosis. tected in the faeces. Microscopic examination of the faeces
In order 10 visualize oocysts. a small quantity of faeces of these animals for oocys1s should be repeated. It ls some-
from an alrected animal is mixed with water and examined times possiblt' to detect merozoites in Giemsa-staincd
microscopically. but. in some cases. concentration of the smears or faeces in these and other cases.
oocys1s is requ1red before they can be d.cmonstraced. This is In piglets the demonstration of oocys1s of I. suis in the
achieved by noration of the oocysts in a nuid v.'il'h a speclfic faeces (Figure 16.5) may be difficult as the}' may not be
gravity bet\,'een I, l and 1,3. The most commonly used is ei- present until about three days af1er the onset of diarrhoea,''1
ther a 40 per cent sugar or a saturated sodium chlorideso!u- Death may occur before this srage. in wll ich case it is neces-
rion. the fom1er bei ng preferred as it is less detrimental to sary 10 perfom1 histological examinations of sections of the
the oocyscs. Approximmelr 2 to 3 g of faeces are first mi.,ed intestinal tract in vrdeno detect the organisms. In some ani-
with about 10 ml of water. The mixture is Lhen passed mals. mJcroscopic examination of smears made from the
through a sieve to remove gross debris. placed in a resr rube gut contents 013) revea l merozoires and meroms.
and centrifuged. The supernatant fluid is decanted and the
sediment mixed with about LO ml of the flotarion solution.
Differential dia gnosis
and again centrifuged. The oocysts ·noat' to the surface and
many of them can be removed in a droplet of fluid obtained A n11mber of conditions of sheep. including colibacillosis.
by couching the surface of the supernatant nuid with a na1 lamb dysentei,·. salmonellosis, C1,'Ptosporidiosis and ro-
object such ns the end of a glass rod. The droplet is placed on tavirus infection. which usually occur in young lambs under

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four weeks of age. 1" should be considered in 1he differential ments. \\l1enever possible. clinically affected animals
diagnosis of ovine coccidiosi,. }.,Ilxed infeciions may how- should be isolated.
e\'er. occur. I lelminth infosiations, either on 1heir own or in ln a heavily Infected environment, ~terilization of bed-
conju11c1io11 with. coccidia infec1io11s. may result In dia- ding and soil "~th aqueoussolurions of l.25 per cent sodium
rrhoea, but the eggs of helminths are much larger than hypochlorite. 0,5 per cent cresol or phenol. or by fumigation
ooC)'ltS and can readily be differcntiawd. \,·ith formaldehydc. 6~ ma>• be achie\·ed but this is not a prac-
Clinically. bovine coccidiosis should be differentiated tical conrrol measure. Exposure co sunl!ght (for at leasr eigh1
from condiiions in \\'hich diarrhoea is a major presenting hotm,1, and de~iccation (humidity les'S than 25 per cent, are
sign. Foremost among these are infectious diseases, such as the most co5t-effective me1hods of killing oocysts.2:i
salmonellosis, colibac:illosis. rotavims and coroi1avirus infec·
tion&, bo,ine viral diarrhoea-mucosa! disease. and crypw- Chemotherapy
,;poridiosis: lntoxicmioru. caused b>• the inge~rion of plants. Clinical coccidiosi~ i~ difficult to 1reat. and success I~ nor-
such as the chincherinchee Ornithogalum spp.J, tulp (cer- mally limited. By rhe rime clinical signs arc evident. the in-
tain oflhe Homerir1 and Momeaspp.)'and slangkop (Omitho- fection has usually progressed 10 the sexual stage of t.hc life
glossum and Vrgi11et1 spp.), 31 and helminth lnf'e~tatlons. cycle and ooc:ysts are appearing in the faeces. E.xtensive dc-
Jsospom suis Infection should be differentiated from srruction ofiniestinal epithelium will have taken place and
other infectious causes of emeritis and diarrhoea in young recovery \\i.!I be slow. In addition. tl1e parasi1es ma~ have
piglets. such as colibndllo~is. and rola\'inh and corona\'iru~ pas~ed through the stages again,t which the ancic:occldial
Infections. Concurrent infections mny occur. drugs are effective. Treatment or clinical!)' affected animals
~hould rherefort- include supponh·e therapy to aid in the re-
pair of damaged gut and 10 limit <econdary infection.n
Control
Supportive treatmem includes the adm inistration ofan-
,\pplication of the principles of managemem. hrglene and 1jmicrobials. antidiarrhoeal compoun~ and fluids. and .
chemotherapy play a role in the conlrol of coccidiosb. •·1 when necessary. haematinics and blood transfusions. 2
Once clinical signs are apparent. the disease is ad,·anced Specific treatment of concurrent Infestations with helm·
and considerable comamination oflheenvironmcnr ha~ oc- inths or other infections may be indicated.8
curred. Although clinical cas~ mny he successfully trea1ed. Where animals arc penned. those showing clinical ~igns
ireatmem may be expensive and the subsequent prodoc1h·- should be removed and not returned until at least two week.~
i1y of the recovered animal is of1en poor.3' For 1he,e reason~ after the cessation of clinical signs. a~ oocyst shedding muy
prophylaxis is th11 most effective approach t<> control persist for some time. Other animal~ In the pen should be
treated prophyloctically. as the~ may harbour early develop·
Management and hygiene mental stages that arc susceptible to tre-dtlllenL
If the Intake of sporu!111ed oocysrs by young animals is kept The devclopmem of anticocc:idial agents for use In ru ·
at a low level through good management practices. infec- minnnts and pigs has largely rcsulled from an extension or
tion~ may become established a1 levels which s1imulme development of the,~e compounds for use in the poultry
immunity without cau~ing clinical disease.? 1 Young. ~us- industry. Relatively few of the compounds registered for u~e
cep tibte animals should be kept clean and dry. and feed and in poultry have been tested and found co be effecti\'e in
\\'ater troughs should be constructed and maintained in ruminants and pigs.
\\tCh a manner that ns liule feed as possible falls to the Amicoccidial compounds are used either prophylacti-
grou11d, leakages are preYemed. and faecal contanlitlation is cally or therapeutically. The most imponan.t anticoccidial
minimized. Properdrainnge of paddocks. kraals and feedlot drugs for use in cattle, sheep. goars and pig$ are lis-tcd In
pens i~ also important in order ro pre\•ent the accumulation Table 16.2.
of hirge numbers of oocvsts. and 10 create an adverse envi- Amprolium administered in feed on'/3.ter for four days or
ronment for their development, The faeces of animals longer results in clinical recon!l)' and reduction of oocyst
,.
should, if possible, be remol'ed from pem before oocysts output.·'
ha\'e sponilatcd. Su·es~ associated with such factors as Sulphonamides are common!~· used for mlaunent of
weaning, sudden changes of die1 and transportation should coccidiosis in ruminants but are only partially effecti\'e,
be minimized . Their concurrent antibacterial acthity ma1• contribute to
• Older animals. although partially immune. may continu- their efficacy by controlling secondary bacterial invaders.
ously shed OOC}'St5 in their faeces and so contaminaie Potentiated sulphonamide mixtures have been found to
pastures. Susceptible animals. when placed on these havesyncrgisticacth<icragainst coccidia in poultry. but have
apparent!)• safe pastures. may suffer severe consequences. not been e\'aluatcd in ruminants_ Gut-active sulphonam-
This occurs especially \\'hen animals are transponed from ides (e.g. succinylsulphathiazole and phthal}'lsulphathia-
the dry e:-aensi\•e famiing areas of southern Africa and arc zole) should not he used in treating coccldiosis ~ the~
placed In intensive production systems or 1,·et ell\iron- become active 01\ly in the large intestine.

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 Co~cicl,o~l> 329

Table 16.2 Recommended

DRJG USE ANIMAL DOSAGE
i.se ar.11 dosage or anticoccidia! Amll'Oilum ilieraceuir Calli~ IO mg/'<g oa,ty 1or ~ oa-,s
drugs um in ca111e, sheep, Sheep 50 mg/kg da•lv for : days
gca:s a~t:f pigs Goats 100 mg/kg <1a,1v ior:; cays
Pigs 25-65 mg/kg Ol'te or tw,ce dally for 3-4 davs
Prcohylac1,c Cante 5 mg/kg da11v in feed for 21 days
Sutohooamides Tberaneuuc Cattle, sheep an<I gaa:s 50-100 mgfl(g daily for~ cays
S~lphadimldine Ttwaceutic Cante. sheep and goats 50-100 mg/kg dailv fO!, davs
Sulphaquinolt81ine Prophylawe Sne8l) 0.5-3 g/lamb/day for 20 days
Su1p1>agua0tdine Pigs I gt IS kg oa,lv fer 7-•Ooavs
lanoohares Pronnv,acuc Cattle. sheeo and goa;s I mg/kg daily ,r. 'esrl for 30 days
Monensm
tasa1Cl(id ?ropnyiact,c Canle. sheep anc ;oats 0.5-1 mg/kg pe, !lay ,n feea tor up to 6 weeks
N•irofurazone lhe.rape!Jtit Canle sheep. goa,s and H}-20 mg/kg da ,v :o; $ days
•
Pi!IS
Decoqumate Proµhylacuc Cattle and goats 0.5 mg/kg in ieed ior at le.isl 28 davs
i oltrazoril iherapao11c Sheep 20 mg/kg (smg1e treatment)
Dclaz~ril Therapeutic Sheep and goats 1 mg/kg orally

The carboxylic ionophore~ have broad-spectrum acth·· relapse after withdrawal of medicarion.43
ity. are coccidiocidal in action and ha,·e a narrow margin or Toltrazuril and diclazmil are symmetric tria7Jnetriones.
safety.'12 They act by facilitating the movement if inorganic a novel chemical class of anrlcoccidials. :\ single dose given
cations across cell membranes. thus upscuing the balance to lambs T to 10 days after being put to pasture was highly
of physiologically imponant ions such as sodium and potas· efficacious in pre,·enting clinical coccidiosis. It reduced oo-
sium. Ac11\1ty is most marked against the sporoz.oites, but cysl output to low le,•els, prevented the development of di·
merozoites are also all'ccted. They are therefore best used arrboea and improved wcigh1 gain during the four ro five
prophylactically. weeks af1er treatment. 19
Oral treatment with nirrofurazone (10 mg/kg per day The level of drug required 10 be mixed In feed varit"> ac-
for fh•e days) is e[ective against cl inical coctidiosis.~7 cording 10 the amoum of feed consumed by rhe animal each
but nitrofurazone is no longer a,·ailable for use in food- day. Distincti.oa must also be made between full ration,,
producing animals in many countries due to persis1em creep feed and ~upplememary feed when calculating the
rcs.idues. inclusion rate. The ionophores have a relatively narrow mar-
Dccoquinate completely conrrolled /:, bo11is in cnh·es gin or safety. and errors in dosage or feed mixing procedure,
when gi\'en in daily doses for three 10 four weeks. 11ith no may result in toxicity. H. 61

References
''"', 1..\. <'i. TOon..,.c.. 191. Relation>hiJ» bC"n,·tcu cwn le\!eh, of Einwria 1<·,,·brldg,n.:'1>umJ other cocc1di• m b:m~ in Engl:and .and
3ureom)'Cm•,u1ph,;1mcthaz.tnc supp?e.tnenmt1nn amJ ",CQUlsition \\'all.'$ Brirtlh l~:mnnty/011rtwl. 131 329-101
ol re,.IMance to O\·uu.- coct1di(t,;;fs. Rrt1ult t'tt~urmy /011,m-J/, 133,
8 <"-"TCH..OU , .• ,ok'IO~. ct;.." 10\''SUt. L.P.. 1976. [~rimtm, \\ith
1~17,;
defined ,nulti-,P""ilk tocddlal lntcruon, tn l:llnbs. Partr.<ITQ/"1,f, il.
~ n.,11t1u. v.·.1. 0·1i..,nv.1 1• "auc.:o,.h. r>..s.t ~ 1979, Elmtrln lruktlrriin fkc ISi-lr
\1'nnc,oto1 hon.\·~. \ltt,•rilt111)' \Ji¥iirt,ic1Smnlf A11lmtl/ Clinlrinn, i.;,
~ J. '\UktU~. c C.. .. (Ofllt(;.()lt\', ~,. w.• l-g93. lmmuniY.a.litm n(
C",Alt711PULI
77-80
Jnrnbs ~a1n,t cnc:c-fdfm,1!-, n:e \'"r"ri11tzrJ Httord, 16. ;&-59,
] HfHKLHU~. ~ (,1lMt.'i:, L. L. t. ~i()k.<P,.'I:, lo:. I. 199.;, t\ dt.scripU\'c,\
i,p,dcmlologu:,i! s, ud)' or coccldlosts in "3:iy lurnbtng hou,,..i noc1,s. 1n c-.ouvAJtMt. .... \A' "u ..,, ,H,., u.P~,.. n1o~t.,,i;..s..2 ., r~aa. F.trcc:t of dlt-t;:r~:
Vttcrt,:al) i',wuilOli>X)•. .3.t. 337-35 l. lo,nlncld on cocc{df.~ 1/<lC)'>I number,, fttdlo, !""formon,~ anti \\U<t!
groMh of lnmb>. Jottmnl o/ ,,,. Smtih Afr/con \ ·,11•rlrtn')' ,lsstJclol/1111. SI
.s C.\1.10\\• 1.. 1~4. Anlmnt Htalth
/,: ~.lr.t1urt1/lo Vol ,t;, Ptottn()(I{ tmd
111-113
Rll'kemlnl D/s,vi>e~ ~le!bnurne, Ausuallwi G,M,rnmenc l'\tbll,hlng
•,,,nice. 1l UAU<..~cun,,. \.. DUtlCF,R. IM & AKl,\t.\kU. \!., 1qg8. Appatcnt dlgl'"!lotlbHiry c,t

5 <; \TC.HPOU. I,,_ GR.lC.OA.\", ,, \\., ~5- P:tthOgt'niclt) af th(• CO.:cldlum

nutrie:nh ~r.d nlttoip..'11 biJltnu·c \lurinA C:\--perlmPntaJ htft."("licm nr c.ih.~i-
E:ml'rill trmutallls m lnbora1oty ta.mb>. Parn.s,rotoro•, 91 • .i;>-52+ whh Fmrtrir. l1'>Z'U \'1.~1l'fl1wryPar,~J1nfr,KJ, 77. 'll-102.

6 c.,-rc.itPOU 1. .._ H-\..,JU\, T.J .. 198,. lnt~raction betwi.:i.:n~occlthn .md 1~ t!R~~- 1., & an :\l. ,,.\,., 1986. lntt.."'dn.1.I C'O(:'C'id!os-is or catth,•. Vi•t'-•nnan·
\1-'mntodfou l,r,rmi fn lamb> on pasture Thr l ·,ur,muy R,'nlrd 12.: CJb,fD-:ufYonh .·ht:t•fidl: l"QfNI .\nillud Prr.c1lc.t1 . 2 .?83-291
60-5 13 r.wrn. a.. 1980. fp,d~rn,ol<>g)· o( pro10,o~n infectlom.: The cna:i<ll>.
; f~\TocPOu. 1.. M1~rn.s, t.c.-&. ;o-r~ER, LP.. t9'i;,.1lic ocnirrtncr or Vt1,~1inn-,, Para.utaln,rJ·, ti, 75-103.

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330 ,rrnl'>~ "'": Protozoa! dl~cases

M R.,k.'1,..\Nl)Of M.A..• ln$2_ f>:2thntogy:intl p:-tlho gcnidl\. Ill: lOSf.:. P.l_ ted.l. 38 \tAr"PJtr.kSOs_ I :-.,. 11 (;,\J!iUIL\lt, ,\ ..\.. · ~ Dlffcrentlf.tt!tm of ).C\.'l'l"l
Th,· Blofo/;y<,{//rc <:occldfn Lo11do11, £cl" a,d Arnold £/ntrrln ,peel es b)' ,..,,dom 3mpllti,...S pulyn1orphit l)"IU. l't'll'ri,w?'
J!i tn?'<;-tx\.U>, r.n. & ,t,,:.,;1-n:w. M .r•• 1979, f.01t"',.sf)' of l4..~1tond ag-aSrt,1 PtUtJJlla/'1~'. 4..5~ 2S7-20fi
CO<cidi3 u, c:aul,. /oumol ofParMitQ/O;;y. 65. 82-1-825. 39 \f.\kt nw, c 1110 .• 1968. A,nproHum :as ii coc:cldios-t.a.1 tor ,\ns-:or.i goo:,,:

16 roi:t6\T, w., ..
1')86. Epidcnuology and ..::01Hml of cocc1d.i.l m roh\•1,.•p.
/rJttrn(II af rlu.• Sour!, .-1/ritan V1•r,,•rltrttt)' .\1cftfi('t1f ,tsg,r/mirlli. 39, 93.
t't"lennnl)' Chni~, tJ/Xortlt Amrria,: fOO(/ Aninwl Pmrm:w. 2. 383-388. 10 \\.\RQu.,1t:or \\,,,., 1981. J lo:.t and -.itr i-p,·dficnr ,n 1hu coccid1n A
ptn-!><Clt\"< Jo,mu,r ,if!',om~o[:)· Z8. 213-24·1.
i- TCJJ\t!\T, \\',f., ~"Th, , ..t;... "ffl(;H. J t 1981, E\.".dU;ttiOtl ofl,s.,afocfd in ""fl
Dgain ..1 ovinc coc<idlo.. .i\me-rinm Jm11nnl of \lffrrin(l1)' H,~,·nrrl,. ~2. ,H \Jt".<;UU\, Jt.\f,, M,.~O~. 11. \ Of \'~ \', 4 Ul' H)'-, A.I .. 19:u. Utc.-:im.·
.54-57. c:nrrtdk~"''\ of 1hr unpali;1 c~u.1.\lod b) famt·rln 1:rlt:-1 .. pc.lC. UO\',
IR HhnlL A,(),, OUU.. rl!,.~('.\, ,., • tL.\.DrJ\'.\1,\8~ n. L IIN;'. ltc:atmt•JU o!
Ontl,m,7"""' Juumnl o/V.rerinnrr R,"Sr1nrh. 37, 4S-58.
c0<cldl~l fn(<e1ioM In larnb.,,i,h ><1phagu•nldlnc Pro<e,•1/mg;oftli~ 4:!' ).~Cl>OUl,,\U'). L., 19CJO. t'()JHtoJ of (Occhilt1~l~ rht·lUtithcmp~· In LO"l;C.. 11 .L
Htlmmthu!ogltYJI S,,,ifl)'U/ m,i1/11gw11, 8. 33,18. '('cf J Cn<i'uUtu-ri 11/Mrm mu/ /'kM1r.,,t1c .1.,11111/tls... lw~ Raton: CRC Pfi'i-,.
pp.3~7-~D.
19 GJEllne. ff, /.I HLLU.11, U,, 19&1 Chc:mopre>ph)!JU\i....;, (Jr (Otl':idtD0\ in f.unb..
1,ith a~h,ilc oral do,c .,r1ollra:curil. I ·,•r,r/1111?' Pt,m.\lr~u,ir., 31i!. 97-107 J3 ,11sut !\l.l.1< JL'\~1.5:--. J. 11•• 1976. DcC'oquin,1tc in th~ control of
r~rl.mcnQIJ,• induced ("C)(Cidln."'t,. of cah'~..4.mrriMn Journul o{
:20 csw:olli'. M,\\ 4,c,.rcurou.1 .. 19S~ o,fuorocctdJoi!~, P.uholo~· uf
\-'c:...•rhm,:.· Ri..'St'mr:lt. 37, l~l3-l045
F.iml'rin oviuolduti, ini-eccJon. /mtmatio,wlJour,utl t.Jf P,.muitO!IJ&'\', I7,
1~1111. :.& \HM)nn·o,, .\GP?CULTUru1.11JMlt:.Rlt~ " ' , HlQO. 1!>lJ';. ~(tmuu.l CJI
\'~erln~ry l'llratj1ol0jt1Clll J.abnmo11 r,.,,hnJqij.,._ Techn1c.1i Bulie1111
~ Gl\£C".O.fl\". M,w. &C\1'U<POU. 1.. 198g, <hind cocdtllos:1,: I lea\·\ tnfer1lon
No. 18 IIN ~1.111.'oi1y',;Sfat1uri~· Officc. l.l>ndon
In young Jambs inrtl!'.l.'\o~ tt\l"--umtt! \\ithou1 causin,.: dilllt:3~. TI1t1
Vimtrltta')' R('C()rd. 124, ,158-461. .&S "ltrTCITf U., C.. ~ U~.,_LJ,11 fl,).., lg.3,l. i)\tTCrt:nlinl d1agoo.,.ic;. of 'l('OUnng 1n
fomb~. In Pra,:hrc". 5. ~ 12..
22. GSECOJt\'• .)J,W,, C:ATClk>lL , .. Pm1LO. n.,1.-. N'OATO'., c.c. 1967, Chin"
0

cnccid:h;,sjs: ol>SCn'Ution~ on ·1>0C)·stp:1trbl!S and pol)'lb fn n;tlUrallf .at'i SMCRO\\, \1.1\.. ~Jlt.)IU~O.U It. ~·l-R.."'-~1-11, C..'\1, 197.,,·,\n\(trnllum for
3cqWred in!tctJons. Jnr...,1:tuitmt1/Juun111lfer P(irtwtof41*'· li. l I 13- coccidio>!S ~n cunt\', Vc1,vi11nr:r \lt1lk,,:l·t11:d smnll Aoimdl C:tum:inu.
112·1. 69,4b!-l~'\
1.1 HMOU,.~ \\.,., 1~;,t.. Oinirnl \lt;n~ nml
..&i' :\'o\no~. P.\\.~c..,iow,:, .,,,,,.
23 (;REGOR'i, .:,1,w.. ll'J'iSllR,. LP. ll C.\lOlPUlL t98;:, Mcdii.:ation :sglnn.st 0\1ni!
cocetdJosls-/\ rt\icw \ 'rtmmuy Rtfi·flrclt CQmmtmfrm:onJ. 5. pa1ht>lt>ll)· ur acddcn1.1l mon,••1>1n p<>i...,n1n~ in ,h,~. umndlntt
307-3~. h•t,·n,1111)· /011nml, ?.'l, 3?..1-326

~ GJlfGORV, '1,\\' .• JO\.~. LP., C.\Tt;11f'Ol,L ,. "~OUtOS', c.c. .• 19$0. Onne .aR :\111.0. L.~ 19":'0. 80,•btc roctidio~fs n Can.tdo.. Cb11,u1.i1m \ '(•it•rl11nry
coccidiosi> 10 F.ni;land ond \\'ale,, 19""3-19711, n.~ t'«11•mm1y R«w1/, 1(16, /01rmal. 11, 91-.98.
l6l-16Z. .19 ,(>1rro.,. cc., 1986. C'Pre,dia of thf domll.Stiqoa1 r.aprn /,rr,.-,,s, with
25, (;KYcour. ~u\. ,, suNtt>"'· c.c:•• 1u8G. Ctpnn ..• corctdlo:;,i,-. ct0t,1 V,·rmn"'J' notti on Ei111.:.ri11 ol'inoidalls:,.nd E. Ll(t/wt11.,,fiJ ($\n, Ii. ovinal from the
$o.cu!~r/ournal, ;.~i~-31. )http Oi•ls t1t-fll~. Pnm.~lwlt,w·. 92. 2r.9-:!89.

50 :i.:ORlO~ ' .c 1or.o,.ut. LP. lo Ci\ TC..UP<>U.. 1, 1.9-; J, F.lm&:11r, t('t"fl>rifl~~,Jsis ,p.
.26 Hf:\'.:\.:sn, ,1,\\,, 1956.i\mmo/ Dlsn,s,ri in ~utlt ,\/ri"'· Pn..•1om,.- Ccnu31
Ne\v6- .\s;~nq·. OD\' 'Ind tunerw m•itur (rom th<' donw~11c "'ht.-cp. P,m,siWl"'fP'· f,g,
a~-~s.
27 HOft.W, i.e.. kA\MO~u. 'LM, t,, umw, 1,1••• 1~69. Th+: us.e ofamproliurn 111
~· <lt11u, R.D. • ou L,r_ , .. l!J9'J. -~1'ma~mcn1 of cotcldi"<i; in d:lliy tat_,,
the trc.nlntcn1 at· coccidi()~ in dcmt~tfc rumlnrmr:,./tJzmmJ q/1h~ S-0111h
-~fr/m11 V~i,r/11n,y .lh'fflr11/ ,\$sCH'lt11/011, ~O. 293-299, .-nd t('pl,,cemM\t hCl!e~. C.omfh.·trdnun ota C<Jtttim1111g &lucmlrm fur tltt'
Pm('tklng \·ct."'rill1ma11, I~. AAl-8~.;t
26 uoRTo~. c;.>-1,1. Ars-roc~tHU:. P.JuJ.. 19~. EtTccts or :unproliwn :md
ii2 P.\RKU'I, Jll., .. Jl;)~I ~. C'i.W., l lJ.l.,, l,;.J.. ltMtllt ""'"" S"Cll"CTO·I\, ,-..1 T\'I t.ff, JI..
1no.u.C!1\'sln oa OOt')'SI d!schargt>... fol.'d udli?:~lllon !ind rumln:il
ml.'1rtbtilh,nt
oil:lmbs wirh r.ncddh)~i$. .•lmtri,•tut Jm111>11f of\'i'"t<ri,UJry.· Rrs,v,rcfi, ,io, • Holfttwn. 1u; .. 19K6, Pf>)t•wcanmgc:occldio-11, in beef c~,h('), h1 the dt)
9fl6-9C0.
troplC:b; F.!11:1>crim<·nt,d rontml wilh (Onlin\lou.s m<>ncn~ln
,upplern,,m,nion vlo ltmanimina1 dt•,1cc,. Tro11lrt1l /111/11111/ Hro/1/1 and
29 fUBO. ,::,v.t-•• ~(NMU>\'. P.C.« ,,,'1..MUt. ~•• 1985. P<ttlwlDgJ.'0/Domc.-c,it: Pro.11mto11, 18, 1911-208.
l\nimals.. 3rd edn. Orht.ado: :\c."td~ni~ Pr~
S.l Pt ~,11011,, Jt. t..,., ,wA~. c~. t:.. 198f,-, Cuccit!iv!l-1~. Ju: Jtt',W.\RtJ. J,L. '.\i.d.J.
30 AA.'O'MU, w. ~ .. 1993-The rd::stioniJ1ip Dl.f1w~n COC"ciduJ a.nd htJminth c,,,,.,m t ,•1,r/11(1,,. I 1,,,,.,,,,,.
.3 t'oodA11/mnl /'t(l,J/<l'. Phih1d,~phia: I\', fl.
Intections in ~h<!ep nnd ~o:.·us in Kenri1. \ 'Nt:rinory P11ro.(ft0I~•.5l. S.:'lundu:s
137-l~l.
s.. run "·· l'1Ako,s ...... JO\r11t\f, .\." :,.,ucwm-s.., .• 1999. lntornctiv~
31 1aU.l..fl.)~~. T.S.. COE'tttff, f,A \\, t... S.\UOl.. T.w .. 19U8. J>l(mf Po:1Q1m1XJ C'l.l.'-..Jtlc:Juon of µordne--J;'im~rl11 c.pp. B~ computc."N1 ..""Sh,11..-d lm.Jgo
and ,\/J'('QTo.tic.o.q~- of I 11,'$1Qd: Ju :-.omhrm Afrir(I, Copt• l't.1\\'I\'. Ox-!qrd zma.l}~fL \ i.•tc.•rimir,· PtlfllJi1d!o~·· 86.. 10.5-l 12
Uni,·ersuy Pr= Sau1"1tm Afric11.
S$ ,,001, o.u 19Gg. C'O<"i:id 1osis of ~?H.l\'p. A critt<'31 rc,,'ie\\ of lhe di~oo..se.
p uu:. Jt.G•• Mnk, f,. * ,ir.,our.uu, . "-"'·· t91fi. Effect ot moncn~in on \ 1!1¥1/ut,rJ• Bulltw:. 39. 60'J-6llS.
expenmenrnl .nrecuo1\.) of f:.imrr"' 11makoh/y(lk11uo:-n... in limb~
.~ l'OITT D.o. 1973, C-..xcid,osl, oi lamb> I. 01>><'"'41lon, c,n 1h,• narurally
Amrrica11 /011m11I oJ l,1on11or)' R,f.Cf1rch, 3:1, J3~-~~3
acqum.--d 111(~c1jon. lJriti.Sh \ ~,rrmmy Jaumal. 129.SS.~'iuli.
33 LE\'1:"'C. ~.o.. 198:, Tuonom)' and Jlfo C)'clt.~ of cocrldta. ltJ. toM;, P.t...
bd,J. 71,~ Biof{)//J·ttftliaCor<itlia. Umdun: ~ilw,ud AnJuld '17 '"'" D.D ·~~~- (:OCcldl<>,b of1,1ml>$ Ill Th<t IC'JCIJOII ol 1he •mall
U1l1t.."'Stin.tl ntUCo-.JI U> cxpt!nm~ntal lnf«-1rqnq1.i1h F nrluintt ·tr and
34 1.1"\T\l,:... .u.. ,985, V~tcrhuuj· Ptotc>:.oo/r,;;,· Anur,: low~ ~,.rn.- Unl\·eKII) F.. cm11d"/II.,. /Jrlri</1 l"rt<rirurrr /~hlllfll, 130, 4>-53,
Pres.s. ~ J"CJlll , o.u .. nS-nu,o,i..... 10\',ut. w, ,. ,01110,,, c..<-. 1s6ti rh1.~ cncodfal
33 U"l>S.". D. <.., OL\OOJIUP..'<, D L ~ PO\\"t, T. ,\. • 199t. F.n1<rlc coccldl:11 popula1ion In dlnlcally nonn.il ,h••P· 1i11· 1·,,,m,u,ry R,~·ord. ;"8.
lnfocclon~and cocdd.io•fr" ln M\in~·. <:1m:1N.·mU:111, t1,1 O,.m,m,ins .;!'i:,-460
F.Llut:i1t/()1t for tit,• Prn"tfdtJB \•'t!t~rfr:.nr1i111, 1-S, $!:lll-702.. S9 kAU<,~ o.,:. & ~tOC'):1'):\U. 1,.1ui., 198b. A brief tc\it-v.• ofbo,in.:
36 u,1\, J,u.. t9;9. Eimt•riil aJp,bto ,rt.
u. from the domhdc s.;oaL Coµra t:ocddiO'lf~ in \\'t."',tcm Canada. Cmu1dftm \t-,t~f,wl)· Journal. 21.
/t11,iu. from 1hr USA. /011mnl t,fParrulto/ogy, 65. 902-903. :ir.-:uo.
37 t.OSG. P.L & J()\'Sf R, LP,. I~. Prob1em9 In lh!'.1' idfntittc~cton Ot lopt!'Clth.Ot 60 ROSL \l.l ,. 198~. lmrrtuntty to l:Jm~rla tnfttthm-t. Vni•tlna,y tmmw:olos;y
EJ111rrih.}Olll'llh/ ofl'tatO=(>(JIOg)', 31, 53S..54 I and lmmuunr>mlwluJO'· 1;, 33.3-3.J:t

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 Cocddiosis 331

61 ..;cH\\'Un:ER~ o •• KIM.£11::kLtS'G. c.. "'SPR..u.'11', 'l ., t98+- :\cddt:nlal mom:Mtn of.\'ur1i1 .J,t1ll'rlca; fl><JI/Aninwl Pwttlu. ?, ~SS-168.
"odium lntox'itadon offoedlot caufo. /011n1nJ <>flht•Am,.•rkcm \:4"rcrinnl')' 69 •WL~~ns. c •• uno~n,L\"-IHlAt>. 1>•• l'U1RSO~. n •• roRSQut'lf M, &. U(.GI.\,
,\l•vltcat Assoctmln11. 1a.1. 1z;3-1z-6. ,\,., 1~ utre-s:ion 01 Efml'tfttoot:y~r5, in e:,,lv~ during 1hcir first three
62 so,,u. c.. 1998. Qu,uult31lw ch.1rae1erlzo.tion. t'lti">ilir:,Uon ond \\'e~b ~tt'r nam,out to pas1ure..lrtn ,·l·unarid S<a,1dbwvla. lit 175-
or
rcconscruc:rfon ooq~t shap~ o( Elnu.1ria spttles from t:mlt.•. 182.
Pamstu,Togy_.116.21-28. 10 ~\'L'~ox. r ., U<NLA. ,., "vurnso:,. n.. 1~94, EJ,m·ru, atnlmmtn,f~
63 souL!B\·, r,.u. • 198:L H.tlm111d1.-:. .-\nl1raJ}O(ls amt Proro:oo of 1nfl!C11on m cakes 3t pns1u.re t 'ttermal)' Parosuology. 53. 33-43.
Dommlwwl Animals. ;,h edn. Phlladelphoa, l.i.':l & P,•biger. ';1 ~\'L~~-;C>N. G.• OLOl-~'i(>l\, 11 6, UCGLA . ., .• IS~b. lmmum~tion or c.1h·e,.
64 snwun c.G .• t91jl, ;\ledlcal Unh,,.1~· ot S0u1h~mAfrka.:.ledun"1 agnln~1 f.im<na alalmm,ns/$ corn~lo,~, .,pp/led l'nr(ISimlogy. 37
Somh Africa. rero:onal ob<erwitlon<, :ZOS..216
65 HOO:D,Lf. P,fl.(.., 19r,. Proposed life C)'ciC of Eimtr/Q :wrn/(, Br/ru/1 n <S.\'t:.~"-SO~. c.. 1g:r,. l'he survtv:11 :and tran.fimi'S$!On or ooq'Sisor llin:1.trltt
VNrrJnaryJourunl. l33, 4;1,_;73, n/11/J,111w11s/J In h,I)'· l'Nrr/11111y Pnrn,<lrolpgy. <;9. 211-2111.
66 5T0(..1'0A.i..t, P.11.(0"# 8.\IS·BO-ROUGtc. C:11 •• B.\11,.N, c.u... Nllto.L. 1981. Snme TJ TAkl.A'IZl', c , l•A~l-,.WS, A. Iii UltA(;()~AS P•• :9.5;. FunhL·tcxt,cricnc()~ \\ilh
pathoph)"Sio!ogiC':ll cho.ng~• n.s.oclmed \\ith lnfoctlon of E./,;a•rln :u1,r11/1 fur.tcln Jn ttc.i1mL•m of (,Wint· and ca print" coccidto-..t'-, Jt>utuaf oftht.•
in cak~~ C.tmmlfan Journal o[Compara1htt.• ,\Jerilrint•, 4..5. 3-t--3:-. Amt1rlM1l \'t'u-rinnl'}1 .\!c-d/Ct11 ,,1.JJ·oclt1rio11. }31. 47-1.
'6; ST'OC).:OAt..:E. v.•r.o .. SHCAR.Q, ~ 1.:mr1s-.G.fl.•• tg82.tftcs.ist:inct ro Eimtrur 10 rosss. n.c , 19ll6. A rc,1ew of porctne nooMtlli co,:ctdiMls..llodun
b<n•isproduced nfter chcmotheraP} of ~~rim~n1nl infcc:tton.s in ca1vel'. l'e:cri11n')· Pran«<', 67. 899-903.
Vc1erl11al)' Pamslrology. 9. ii I-Iii. ;t, , A~ msrn.Jt, 1-.,1 .. l!t7'6, ~<He,. on i;omc dl.,c-.\~problcmi 1n Anbt0rn Y9'3l$
68 SJV,\.R'T, 1J P. " 1.1~0S1\Y. o.s., 1986, Coccidl<,$!S ofswine.. V.uttrmaryCfirrics ,n South Mncu. Vnerlnmy Mmlfclll //11dew. 112, 109-138.

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 17
Cryptosporidiosis
Synonym: Kripro~poridiose ,.Afril-.. 1

CG STE\VART AND B L PE;>.ZHORN

Introduction and is associated whh disease in mammals, while C. muris is

found in the cells of the peptic glands of the stomach or abo·
Cryptosporidioshi, caused by the gcllus C1)•p1ospnrirli11m, i~ masum and it has no1 been a~sociatrd wirh disea.~e.2
usually an inapparenl lnfe1.-iion of a variety of mammals, C,yprosporidi11111 meleagride.< and C. lJaih,yi infec1 birds.3~
birds, fishes. and reptile,, but it mily assume clinical propor· and C. na.<omm and C. serpemi;: fish and reptile$, respec-
rions characterized by lesions ln 1hc gastrointe~tinal trac;t rlvely:111 Crn,ro.(porirlium !)fl/'1'11111 generally parasiti:t.e.~ in-
followed by the development of diarrhoea. This is particu- testinal epithelial cells, 2 and its oocysts can be differentiated
larly so in neonatal animals or irnmunocompromised host~. from those of C. murls by their smaller siie; C.. pcm111m
Cryptosporidia arc very ~mall cocc(dia which were fin.t oocysts rneasure -1,5-5.4 x .J.2-5 pm and those of C. 11111r/$
described by Ty?.zer in 1907°1 from the irit~tine of normal fi.6 -,.9 "5.3~6.5 µrn. 17
mice. Umll 1955, when the)' were Incriminated as a L~du~c or Foll<m;ng ingestion of oocyst~ b) the host, sporoz<>ices
disease in turkeys, they were thought 10 be c;ommensal~. 3• are released into the intestine ll'herc they pcmo.>trate the
bm since then. diarrhoea associated with Cryp1osporidiw11 brush borders ot 1he epithelial cells to develop into mii.'ronts.
spp. has been reported in lambs.' kids,:$ piglets. 16 immuno· Although organism~ appear to be attached superficially to
logically deficient foals ..llj and human~.~:, Bovine lntii.'~tinal cells, all ~tages are in1raccllul.1r 3nd are contained within a
cryprosporidiosis was first described in 19,l in 1he USA?" parasi1ophorous vacuole which is situated between the cy-
and subsequent rnport:; from the UK.41 Canada.31 -\ustra· toplasm and outer cell rncmbrane. 15 -n1e pard$ite is sur-
Ila. 22 and Gcrmany1a indicate a wide geographi<'al distribu- rounded by the microvillou~ membrane which fuses 111th
tion of the parasite. In South Africa, C1yp1osporidiu111 1he epithelial plnsmu mcmbram.• to fonn a thickened adhe·
infection has been associated 11ith diarrhoea in calves.2 1 !>ion zone at the point of auachment 10 the host cell (Figure
and a high prevalence of the parasite has been demon- 17. 1). l here ls also an adjact.'nt comb-lll.e strucwre formed
strated in the faeces of healthy calvcsw and in children hith by a raiding of the parasite pellicle:~1 These specialized
diarrhoea. 9• 17 stn.1cture" ;ire thought 10 facilitate the intake of nutril'nt5 by
the parasite from the hos1 cl'll. The meroms are 2 to S µm in
diameter and coma.in eight merozoites which arc re!eased
Aetiology and life cycle
into the lumen of the gut without destroying the host cell.
Oypros1;oridi11111 spp. are protoxou ascribed to the cla51; Second-genermion mero1H:. may occur, and these will con-
Sporo1.oasida, subclass Coccidiasina. and suborder Eime• cain four merozoitos when marure. These merozoires invade
riorina of the family Cryptosporidiidac.2" The parasites new host cells where they undergo gamogony b) forming
complete their life cycle in gas1roimustinal epithelial cells.~ 3 macro- and microgamoms. The mlcrogamom has sperm-
The life C) cle of Cryprosporidi11m spp. resembles 1ha1 of like microgamems which. when released. fertilize the mac-
other intestinal coccidia but differs in that the parasites de- rogamonts which then develop into oocysts. Oocysts arc
velop just under the surface plasmalemma of1he ho,1 ccll.2:' round or ellipsoidal. and comain four sporo7.Clites. It h gen-
In young animals the life cycle is completed in three to four erally acc:ep1e<l thal sporocysts clo not occur.l4 Sporulation
days. which is shorter than the life cycle of most other coc- lakes place 11ithin the host cell and is followed by the devel-
cidia. opment of a resistant thick wall around ,ome of the oocysts.
Cryptosporidium spp. infec1 a broad range oi venebrnw which are shed In the faet·es. ,s 01h11r oocysts. howevei'. do
hosrs, including fish. reptiles, birds. mammals. and hu · not form a thick wall and the ~ingle unit mcmbraiw sur-
mans. 15 Cryptosporidium pan,11111 in fects the intestinal tract rounding the sporozoitt!~ mplure, whcm these oocy,h arc
332

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 Cr:ypw.poridiosis 333

Paras ;oonorous vacuole

Paiasue pelhcie r1',crO\' lieus memo,ane

Mnas,on zone

Comb-hke suucturc

Figure 17.1 Line drawiog of Cr)'ptos//Ql1d1'umm a hos! ep,1hel1;;! c,i.ll

released rrom the host cell. These forms are invasi\'e ancl can cold rempcratures. may also increase mortality. Im munodc·
infect epithelial cells and ..o reinitiate the de\·elopment fidency may be impor1an1 in the de\·elopment or clinical
cycle, resulting in auto-infection. 11 dlsea~e. Severe Infections have been reported In Ar.:1b1.ru1
foals suffering 1rom 1hc combined immunodeficiency ,yn-
drome•8 and in humans with acquired immune deficienc)
Epidemiology
syndrome r \J OSJ. In foals the disease is more prevalent in
In livesiock. clin ical cryptosporidiosis occurs most fre- the first 1hree to four weeks ofliie but can occur in animals
quently in calve~. bu t it has al~o been recognized in pigs, up to one year of agc}J Per~istent infection in athymic mic~'
lamb,, kids and immunocompromiscd foals. 5· 2"· 11>. ~. 4 ~· 4'• and AIDS patk'nts demons1ratcd that T cells arc necCS<ar)'
Outbreaks of clinical disease arc most common in ani- for immunity to dcvelop.~q In ralvcs. a strong cclt-modinted
mals one 10 four weeks of age. Diarrhoeic cal\'ei. ma) ~heel as immunity develop~ after infection with an i11crcase in CDB-
many a~ IO million oocysts per gram of fa<1cc,. resulting In T cells. ,¼ ooc~ sts are produced in the sporularcd state,
severe comamination 01 the em~ronmcnt.'1 Cryptosporidia auto-infec1ion is particularly imponam 111 immunologicallr
may also be found in the raec~ of subclinicall) infected compromised hosts. re~uhing in per:;l$te111 infection.'
caule of all age,;. in human,. cryptosporidiosis may be acquired by con-
Oocvs1s arc resistant ro most environmcmal facrors. but tact with tnfccted individuals or 1rom animal smm:es. and it
the lime 01 survival outside rhe host is unknown. Chlorina- is a common cause of traveller~· diarrhoea. 311 The lo\\ ho,1
ti<)n of water has linle effect on \rjability or the orgnnisms. specificity, excre1ion of large numbers of oocysts and low
hence water-borne outbreaks of the disease are an impor- numbers of !locyst~ required 10 init iate infectio n in suscep-
tant public health problem.49 Under experimental condi- lible host~ impl) that animal species can be potential re<er-
tions infectivity is lost after storage for six momhs a1.J •c. 42 voirs for human infcctlon~. 10
Transmission is by ii1gestion of oocysts in faecal con- C1yp10sporidi11m par,•11111 can be divided into two dis·
taminated food or water, \\'hile aerosoi transmission mar 1inc1 populntion\ based on differem antigenic and molecu-
also o,·cur In both humans and animal~.zo 4~ Infection$ of lar marker,; and have been referred 10 as ·animal' and
the respiratory traet have been reported. which may indi- 'human' types. Human infection may be due to either type
cate aerosol transmission. ' 2 Calves that consumi: adequate \\~th anthroponotic ,human to humanJ transmission occur-
,·olo~1rum nnd have high serum gammnglobulin concentra- ring \\ith the "human· l~'Pe and zoonotic transmission oc-
tions are less lrkely to become infected.27 Recovery usually t-urring with the 'animal" type. It is nm clear if Lhc 'human'
occ12r~ within 30 days. 1There are repon-, where Cr)'prospo- ~'PC can infect :mimals.6
ridimn has bee11 incriminated a.,; the pnly cause of dla·
rrhoea, but Cl) tosporidia infecrions ofren occur together
Pathogenesis and clinical signs
\,ith other enteric pathogens. such a~ rota\'iru~. coronavl-
ms. Salmonella sero\'ars and emeroroxogcnic Escherichia C1)'p/()sporidi11111 spp. have a marked t!ft'Ccl on mt1mbrane-
coli. These multiple infections act S)'llergistically and in- bound dlgesti\'C enzvmes. resulting in poor digestion.
crease the mortality rate. Severe stres~. »uch as exposure to Malabsorption may occur a.s a result ofvillous atrophy.,:,

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334 ».1.-,.,.,, ""'"' Proiozoal diseases

Except when it occurs in very young animals. infection be demonstrated in the brush border of intestinal epnhelial
is inapparent or. at most. produces only mild disease.3 • 23 cells. ; 2 Care must be taken to distinguish organisms in tis·
The incubation period varies between rwo and seven sue sections from small droplets of mucus adhering to the
days.30 Diarrhoea occurs at between one and four weeks of surface of ep11helial cells. 12
age and lasrs for one to two weeks. Although the morbidity Cry1prosporidi11m spp. oocysts are very small but can be
rate may be high the mortality rare is generally very low.' 12 recognized in faeces smears flxed in methanol and stained
The onset of oven disease is usua lly sudden and cli11ical 111th moclified Ziehl-=-:eelsen4 or safran.in. The latter stain
signs are very similar in all affected livestock. species. De- may be the tn0$1 sensitive.8 OoC}"ltS appear a~ red ~pheres
pression and anorexia are followed by a profuse yellow wa- agninst a dark background and need to be dilferemiated from
tery diarrhoea with associated dehydration. Tenesmus yeasts. which do not stain re<l. High sensicivit}' has been re-
may be present. After a few days the cliarrhoea becomes in- ported using immunofluorescem staining techniques. 13
terminent and the !'aeces pas~ for up to JO days. 23 Loss of
body weight may be quite marked. Relapses after apparent
recovery may occur. 42 \"omiting may occur in pigs.46 Most
affected an imals recover spontaneously unless compli·
cated by infection with other cmeropachogens. Severe
chronic infections may occur in immunoc.:ompromised
hosrs.2

Pathology
Generally. the e;,ctem of the mucosa! injury in the gas-
trointestinal tract and the severliy of the disease are directly
correlatcd.2ll Emeric lesions increase in severity towards the
terminal portion of the ileum. and the caecum and colon
mar also be infected.'° The mucosa of affected pans of the
incestine, which often contains watery yello\1 material. is
intensely hyperaemic. The carcass may be emaciated and
dehydrated.
I
.... ..
Stuming. fusion and cross-bridging of adjacent imesrinal
Figure 17. 2 Crypiosporidia In epithelial cells o: ihe small i~testine
\illi are seen histologically:1 In affected pans of the imestine.
mucosa! epithelial cells are often low columnar. cuboidal or
e1·c11 squamous and. in calves, 1here may be focal necrosis of Techniques for concenrratlng oocysts in faecal speci-
epithelial cells in some areas. 15• 18 The very small ba$Ophilic mens, such as those used for the diagnosis of coccidiosis.
organisms. generally appearing as small dots. although ring· mar facilicate identifkation. Tht> final prepara1ion can be
or crescent-shnped forms may also be identified. are found e;,camined directly. or stained bra safranine-mcthylene blue
embedded in the microvilli or just inside the distal cell method. 17 28
membrane of epithelial cells (Figure 17.2). They are more Clinlcally, crypwsporfdiosis should be differen1iated
easily demonstrawd in tissue sections that arc stained 11~tl1 from other emeric conditions. such as colibacillosis. <almo-
Giemsa.5 • •10 Extra-intestinal forms in the gall-bladder. me- nellosis, rota11rus or coronavirus infections and coccidiosis
semeric !~mph nodes. 1rachea. lungs and uterus have been (see the re!e,-ant chapters). Under field condiri<>ns. however.
idemified in pigs and sheep. 16 diarrhoea is often associated \\ith a simultaneous infecrion
with rwo or more micro-organ isms.:!3 and this. 10gether
with che tendency for the infection to be subclinical. makes
Diagnosis and differential diagnosis
diagnosis diffic\llt.
Demonstration of numerous oocysts in che faeces of young
animals ma nifesting clinical ~igns of emeritis. or in prepa-
Control
rations of Intestinal lissue. are the mosr reliable means of
diagnosis. However. the finding of Crypro,poridi11111 does Chemotherapi' docs not ·a ppear to be effective.w bu,. in
not necessarily mean chat the organism is im·oh·ed in the most cases. lnfectlon is self-limiting. Adminis1ra1ion or Ouid
disease process, and it fs therefore imponant to associate therapy and other sympt0ma1ic rreatment is important in
the relevam clini.cal signs and histological lesions 10 the the pre,·ention of dehydra1ion and tlie correction of the
pathogen or other pathogens isolated before making n di· electrolyte imbalance caused by the diarrhoea. If managed
agnosis. correctly, most animals will recover unless the disease is
ln fonnalin-fixed tissue secrions. endogenous stages c:an complicated by infection ,,;th other emeropathogens.

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 Cryp1osporidio~is 335

Pre\'emion should be aimed at a reduction of the inges- should be kept in isola1ion. Although oocys1s are resistant to
rton of oocysts by raising young animals in a clean and dry many disinfectants and mos, pre,-ailing emironmemal con-
em~ronment. Faecal material should be removed from pens ditions. !.hey may be des1royed br e~-posurc 10 a 5 per ce111
on a regular basis. and animal~ suffering from diarrhoea ammonia solution or IO per cent formalin. 11

References
ABRAHA.'IJSlS, )l.~-. 1998~ Bovine T c\"ll re,P<)nS('-$ In Cryptfl)J)()ritllum ourbrt"J.k" of ncon.,ul culf d.iutr.bor:t.Austt<1Um1 \.'tu•ri,UJI)' 101,r,uiJ. S7.
p{Jn•um infection. /mernmfa,ull /ounUJ.1 ofPu,1ui:0Jow. 28. 1083-1083. -13·'-''35
, ,,oE~x. b.t:u l93i, Abom:iSDI nyp1osporidfosi~ In cattle.\ ·eu•rJntU")' -:o Jo:tR....,\TIUCK.C..LA MllJtU.. J.P.. :98.;. Cryptospondlos,1-.:. Thr
Pml:n/QJf)', 2,1, ?3!'>-?38.. Camp.•tulium ,m C.n,,t,nwng 1;Jt11cmio11 ;or 1(11tri11(ltJtmt, 6. IS-i 16?.
3 .,sotR.-<1~. •·• 1996,C')'ptosporidio51$Jn bovuic and hum•n he,hh. ~• U\1~,.• :\ n. t9B..; TJ.\;011t11t1)· :md rc\it",\. of thi: ((JccldJa.n ttenu11,
Jour,:n/ ofO,,io· .St·rett(I', 81. 303&-m l CrJptosporidmm, Pro1<>1un•.\pl<UmplO'• /01mr(ll ofPt0t<WJQ/u,:.·. 31,
9.;..911
.i AS(.U>, -.:.w, 1987. Crypc0t,pnndio>ismdomcs1ic nmm;1I, ;and humans
111 Proct:tr.1J, -17-19. 25 t.H'ISL, .o•• 19t$5. \ ~1rrmm1· l'rnttl.!Uf?iOJ!:,,'· ..\mt"S lown~ lo\,., State
Unlvtrllty Prt.....,,
,; ,~Gu:t. li:.W., ttlrmn, s.• Gll..\\', F...W.• \98u. lmc~,hrnl !~sJon, Jn specihc
J"1thog<n·hec lnmbs associated w11h a C:ryproswrul/un, from cnlvc< 26 usr,~.\\', l).s•. eu(,RUH.!\ tu ... ~usol.!n.,t\..,"S. c.A. i R:'l.tt."il, J.A •. l'.)8;
!',.

wllh dJnrrhcu V,•wn11t11j' i>lltl:ulo,o~ 19, 67-7'8, Clwn,oproph)l3"i< of crypto$porldlo.,, in thi~kcn<. usln&
hn.lfunglnon~. s.allnom),'rln, J&1,lodd. or m01umsln. lwtt!rlran /oJJmal of
$ \\\'r\1.)-,t:L-~iur.;.1. L "1., t\OIJIN50~. tt, .\.. l'CTRY, I,. r-111K>,,\I U, \· , I\·.,~-.. b,
Vt1url1111f'.I,· R,•.1.v,rd1, .Ja.l5-t-:i55.
"CA5~1ont!, n .. 1998. Diff'cnmilatlon bL•rv,·ccn hurnan and anlm~
2; 1or,a..., 1 \\ .\LI r,.,"· ~o .. Mnc:ur,u J. & Q:11",\;· \1. 1938. Romvirus and
isolate, of Crwros110ridi11m JJ<lMtm usloll mole~ulnr and blqlog,<".ll
matkeo. Par,;..,rotog,roi Resei>r<ir. 8~. 29;--301. CT)pti>sPotidiu,11 sht<lding in da1')· coif fueo,, und its relauoruhlp 10
ootosuum u-nmunt- 1ru1\'i:fet. fuunutl ofDlt:ry Scftnu. il, 1238· J2~H
MlJ'"K, ,.~.,, C.\l\&m.'Ct.t.. P.L tr,-i. Cl'J11ro1pcri~llum 11gni ~p.
n from
28 )1.uu;us. ,1.11. 1o nu,;,11 , n... 1~; Sri1ining: of coccidlnl OOC)-:~v,. 71rt
lilll1b>. ond Cr:,111osporldl11m h</:'i, •p. n f1om a "'111 ,,,,h ob,or,-.11on,
\ 'ttl'rmr,ry RrnJrtl, 12) 329.
on 1he OOC)">I 7.e,rs,'1rift fi,r Pormlrenk,md,, H. 2119-2~8.
29 ,100.,. HW ,. e11 ,:u·1r1'. w.1 .• t981. f<''7~ tran ..mir;:,.ionofc-all
8 s.u:111 n. +· &u,r,Fu. ~" 193'.). Sen,111,,·. rnp,d. simple methods 1or
ayp10,por:dio bet\\'ecn rnlvc,, and pip I ',,rrrlmrr;· l'nthology. IH.
dei1.ottln,; c:,,1nrn1,urfrllum fn r.n...t.ces. Ltmu1. 2.. t t,tlt,
248-:!.SS.
9 Ulf0;0\\'11?.,. ff fl,, \'.u.&..UIH, \\., 8UG\\,\ \ \, 1,,,.. 111-, I \, r., l~fl:fl,
30 \toom. D..A 1~. \hnlmizlng 1norbldity .M1d mortoU~ front
C1111rn>po1idfo,t~ In blatk So1ah African ehlldr,i~ !io11th,Vrirn11 c:1ypt<»porldlo,i. Ve1,•r/11tuy.\1edittrw. 3.. 81 I..,S!S.
\!.edln~I frmm(JI. 14, 2;2-2.73.
:u MORI\;, M. Li\lU\1~Rr s.,11.,\IJlf;ft.A 1..l\';'0. ?ntho.tog,ral:tnd
10 ftUJ,!11.\kl, ~- ,- J;MITI!. t(,\',. 1997, 0,J)IOSf)OriflllmJ /Nl(l'tmJ· OIJC)~t
microbioto;.:.1ro1l Ub$1."tvutlor11,1 mi\d<' un:~pontnncou.s caK~> of ~lcu1e--
t!l.Ct~tlon a.nd ,inbiUty in 1.":lCp~rlmL-nuill}' ln!1."Cl1.'d l;1mb-s. l·vui1.·1ttiologJ• neo:.u11ol c.llf dlQrrlroca. Q11wd1n11 /01111,nl of,~mptlf<ltrr'<! Med/cm,• .m.
Md /llf<er/tJJt, 119. 105-108.
218-240.
u c.,,u~ntu. J,. ·rt..woru. s.• >1u1·e1usa!'.. G. 1,,.ASC\Js, ,,;,"'·· 1981. f.lTt!(t of 32 ~i\CTRt. ,, .. 1989. ·\n1mill -omd humun cryp1ospoMdlosls.: OJ>pnnuni'i-1
dhinfoc1:1:mi(,n )UrYl\"'all oft.ryptO.ipnridlum onr)r::,.t!lo. flh.• \14't.tri1Jt1')' info,1ion,r P.Jthno~mcit} of the geou,. C,:.71ro,p,m1Jmm. CCKcidlu and
RNVJrd.111.414-115. lnie«,nol f.oet,dJpn,orphs. /'foffi!,Jmg.<o/rht'fiftir /memn1101wl
i~ ,oLTlFfll. 1. ,.w.. 1990. Facuit)· of\ e~trin~· ScicncC'. Uni\ ~rsuy of OKcufJ(JjfS Con/t·t,•ncr" lourt ;Frnnct; 1i-~O October l9~. lXR.·\ 1.;;
Prcconn, Ptr~nul communie2tion_ Ri.."t' de P'Unur('J'5U~
13 COH£S. )<.0. & SS0\\1>E.'<. ~ .. !!190. Cryptospondlal d,:u:h~ In foal< Fool 3J S~\tl 1-.A., flU~f'k, J.l)., J'\C,t, C.J.t., ltOUCliUI, '\I.A.. II YA.rtUlJ·\, f.rt 1 n
.\l,•,ff,·me lltlll Sllrgl!T)', 18. 296-3-0ti. Acute emerocoJlets in a human bl.'ing infected \\ith I.he J)rotO;{oan
Cl)·pro.~JJQr~dtum r~1.,t11>.·11u·,nfo,K>·. ;o, 592-598.
1-l ctffinr"'-r w .,... 1.9S5, CC)'P10->'PQndl~i, Jrnmml ,,jtlio.-!,1rwnc<m
\ ~u1r1,rary· .\f•tt.lfa,I J\ssoi"iminn. 1s; !334-1333.
1 34 o'oos{k:',Hut r.1 ... l':f85. C'.l'ypto,porldlum tnfection rn m:m, ,tnimah.
bl<d> "11d fi•h. Amtm/11111 v.n,rbmo· foumal. "2. 2>3-'.!58
1.1 f ,ra. If,~ u-.:<.iAtt. 1.. r-.. 1~e6. CryptosJ,JOrldium spp. and
Ct)p:o,p,,ridl<>.>I), .\/u:rol,lo/qglcal Ri!l·f.,,,,,. 50, ·ISIH6J. 35 P\:\"'1':11.JU.. l't.J.• n10,1,,s<~.,. 1u,.,. C"...\.R:\'rlt. F.,,., 197L Cryp1os:portdlal
inf«l!on in a calf I ,1,•rillor>· Pnrhoro;1.1·. 8. ~79-184
16 ft.CT,\., .• ~o\.l\(Hl~·Al.:.fDO, c_ Ct.A\"tt.., c.-QUtU?., J.. 1qt).5,. Oeu.'1·nc.m al
36 l'OHU'Sl, I, MOOS, U .\\'., tJU \'lU.[,. :O.,f,. 8f\181(:i,;, \\'.J,. 1~7ff".
Cryv10..,;,wridium otu;ysti; in t:xtf3•intt~rjn:\I ti<:'tu~~ uf ~hc;cp und J">lg.'-.
l ,r,on,rnry Pnnul/o/0/;;'', 5g, 201-200- Cryp1o,pondi~>1S a, a prol>ab!c toctoroi nc",)no1al dl:whoc~ tn col\'~
four,MI of rht Amrri,·mt V,.m.•rinnry .\1,•dt,al.A.-<SOc.~ia,fon, 172. 45:?-437
l:' FRIPP. P.f, ,._ 80ilt\\,\, )~ t,, t987 Cr;,·p;O$JK>rtflium a),..,-.-Ocl:ned \'1."ilh
J~ SL\\T'·•· n .. 1~5,5. Cry11:o.Jpotirllum m,:l,tpgrldu sp. th;1\' J. Juunml of
duarrhU\".i at c;a ..RtJn'ku,".:i ho>pwtl. 11h· ~mlwrtr A/ri.ro11 /011r,u1! "'
Compamtfw Pmlwlugy, h5. 262.~oti.
rp1tl1•mlo/ogj•,md /11ft<tfo11. l, -1;-i·
38 )~·,,~".... P., •.,,,u:-.:n. ,.,. e,,Mcr.uLc.,r:.,. \..L, 19:-8, CS')1\t05pori\i!('),i1ri m
t.8 HALL t,;,,\., JUl\'SOLn~ O.J .• fti\~Ol\S, i..ft., llt.ASU, A..P lo '\fOHCi\,, Mt. 1983,
immun<>d.:ficlcmt Arabian f°"J\. V~,·rl1111'}· Pmlwfo~·. 15. l:!-1 ';
Pnthaloi,,y of calve,; 'A11h diarthoe,l in ,ou1hern Briialn. R,.,,,,ud1 Irr
v~:rrirral) Sae11w, 1s. 2·1o-250. 39 su~. T.'- Tl!.tC.lffttfu•. ~.. 1988. Pn\tO'/(fal infoct:on,. In 1hc-ar.,111ltcd
irnmunodffich~nc~· ~}·ntt,omf. JQtlfJJ(J/ of Bccrtou .\lfrm.,·cr,/1y
19 11u~1. \'.J. """"11. 1.. 198,. Kl'}'PIOsporidlen ln(cldorwn b.im Kalb. T«Jtnfr(w•s.8. 7S..103
,ach,wl,. Vorkonnnen und exporlmontelle Obcr1rog11ng. B,irli11cr rmd
\/1mc/1t11e, 'rlrmr:11£cll, Wocl,e11,rorif1. 9,1. 289-2'!2. io ncc,,~_
s o,, 1u;. 1-qaa. ~~tal \"ru.•rmao· 1'a1hatog:.1• ·romnm: n,C.1->cckci
tnc.
:.!U lmll' :'\G, , •• ICOI "fl l\•,\:\1'111.R"if', \', "1£.PS[S, ..,,, lt-l8i, c~i,to&pt:)ndi~il.. ,\
,1 T\7.lf.11, 1.•.• 190: \ ,pmw.oan found in lhe p,,p1lc gl:md< oi1h~
tu.~ for nirhume 1ran....m.iufon.. 1Anet~1. 2,sss:n 27i-272.
c:t1rnmon mou. .c. PftX.trding10J1lt'1~«i<t).'of&J"tt~nmt'ntnl (JmloK>'""''
::, HOM,ftnt. F w .• 1981, Bo,1ne <l}'P!O.Sporidla,1'./011mal ofrli. ~,1111 \l•Ylicm,•. 5, !:?-ll
Vrita,1 \ ~h~rinary ,twx{nrlon, 52. :!al-2.~l.
.12 rnvo,u. "·· 1.983- ~ 1p10-.pori<lio)I:,. in anim.tl~ and h.lmt,uh.
2:2 JUtttu·r 1.,-.. • :,\!OIK,ft"-S~. t>,R,, 19$1 Cryp,ospqrldln M~iated \\ith 1\lim>IJ/0/opr,1/ l/,:•/,11~. i;,8,1-%

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336 '"'"''" n,,,: Protozoa! dl~eaws

>J n1roR1, s .. 1985. Th~ rclauve 1mponance of en,cric pathogens alf<!<:lin~ ~rimen,31 mfocmon of p1glctS wftll c,,-111a1JXJru(fom. Hrs.,.,.<•: In
ncon:ncsof domestic:: anunal., ,,Wr'f'lnt:t!s ;,, V1rcri,u1ry• 54·,u,r.c<' mul tr~1Nit:a')•.!ki,•nu. 31. 3S8--368
Comp~)rt1d1'VMl'd1<:.i1w, 29.103,.,200,
.ii unos. ~-'·'"<;u1mt:..\:T, l'\,L, l9tJS. nu~~p\!<'ll'SnfOyp1()$1)nndmm
.. J TI.Wt1R1, ~.. C\\Jrnru.. ,.. sur .ff\\.'OQU, o. & !>~ODGRAS~. n.R,. I ~ . \n {Apicomple.xa: Crypl{)!)poadiid~cl inlt:ttmlt anhn.ii,. frmma! OJ
ou1brt::ik o( calf diarrhoea atmbu«<I "'Cl)1H<>>PQn<li.\l ,nfe"ion. n.~ Pctrns/101010·, 71. 6?_;.629
Vrrtrlnn,y Rt•t,:tJrd, H)i' S7q-.atSQ,
.t8 Wt B!'TUI. "· \. 19-93, Molc-t:Ulill mtthOd"'- {or d<::lt,"(tlC,11 ;md d.1SSU1cu1.tf.ln
.t5 rLo•um . .... LAltst:-.. , .. ,..,nm. ,,. ~ 1.unL, n.• 1~2- Oh.u-rhoc-a u1 ,:oat IJib of Cf}µro,,xiru/1,1111. l 1nrrrJifr>/qg;• f¢a,•, R. 26.l-Z,>6,
.aurihuu..-d 10 Cryµrospl.Jrtdtum 1nf1..oedon. TIil• t'('ftti11ur:,· J«~·IJrd. t 1l.
.;9 /Al,,-~. f \'.'\:rt, G·t>.. f",\\'[R. fL c-- taU.IUtA"-1. fl 1 199;: '-r'}'JUO)pondfo);~:
»-39
pa1hog,metj~ and jmmunot~ Pt1m.ftcdl<J;:,• T,x/tly, 8, 2·i-Z-
HI T/.IPORi ,;... l>fC:C.\R1,.tV.1~, L\\\""O~. G.Jl.f.:.. &1\0\\L\.~O. \.r,..19..tk

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 18
Toxoplasmosis
Synonym: Toksoplasrno:re lAfrik.

JP DUBE\' ANO CG STEWART

Introduction globular to o\·al in ~cction, (Figure 18.28). Its anterior

lconoidall t!nd b poilllt!d and it:s po)terior end Is round. II
The protozoan parasite Toxo11lasma gout/ii is a common has a pellicle ,outer covC'ring,, a cono1d. rhoptries. micro-
parasite of many an imal speC'ie:. and human:. and lb distrib- nenws. a well-defined nudeu, in the central area of the cell
uted 1hroughout I.he world.Although most of the infoctlon< (Figure 18.3). and othN organelle-< found in rncc:idi;;n
of livestock caused by the parasite arc subclinical. toxoplas- rncrozoi tes. t ·" 2 "'
mosis is a major cause of abortion in sheep in several coun- The 1nchyzot1eenum, a ho,t cell In a \'arie!) of organ~ aml
lries.3· 30 4;;, :;;;. s7 ' 9 It is also kno1,1, to ta use reproducuv~· tissues b, active penetration of the cell membrane. ,\f1er en-
los.<;es in goats. t,. 'II!. 4~ In somhern r\frka there is a high tering the cell th<1 Lachy1.oite become~ ,urrounded hr a par.I·
pre,·aience of serologicall\ positive animals. bm reports of ~itophorou~ \·acuole (J:igure 18.3). Xumerous incra\11cuol.ir
toxoplasmosis in domestic animals aw rarc. 71 -_; In South tubules connect 11w parasitophorou~ vacuolar membran1110
1\frica namral infec1ions in ferrets kept at Lhe Onders1epoorc Lhe parasite pellicle {Figure t8.3). The tachyzoltc multiplies
\ 'eterina[\ Institute for experimental purpose:-" and losse:-. asexually within the host cell b) repeated endod)'ogeny. En·
in chinchillas in lhe Western Cape Province have been re- dor.Jyogeny rendo = ln~ide, dyu ; two, geny " progcnvl h a
poned.1t Because toxoplasmosis can affect virtually every specialized form of reproduction in whic:h cwo pro~eny form
warm-blooded animal ii b Impossible to summari1.e Infor- within the parcni parn~tw, thereb\ consuming it (Figure
mation for each animal species in thi~ chapter. but the r.Jis- 18.3). Tachywites cont inue to dMde by enuodyogen) u111ll
ease in animals up until 1987 has been reviewed .30 :\lore the host cell is lilted with para&itcs. 2 ~ which is rhe stage as-<o·
recent information is included in this chapter. c-iated wi1h arute toX<1pl3smo~is in animals and hL1mn11~ .
1\f1er a few division~. lachyzoltes induce the formauon of
a cyst wall in a host cell and transform inw brndyzoites con-
Aetiology and life cycle
iained within the wall (figures 18.2. 4 and 5). Tissue cy~ts
Toxoµlasma gonrlii is a coccidian parasite of domestic cats gro\, and remain intracellular (figures 18.2 and 4) as the
and other members of the Felidae. with other warm- brudyzoites encysted zoites) slowlydi,idt' b~•endodyogeny.
blooded animal species, compri~ing essentially all mam- rtssue cy<t~ 1 3f\' in site. Young tls~ue cyi.t~ may be ai. ~mall
mals anu birds. as imermediate hosts (Figure 18.1 ). II ls a5 5 µmin diameter and coma in only two 10 four bradyzoi1es
classified in I.he phylum Apicomplcxa l.e\~nc. l970, class (Figure 18.4). while older ones may contain hundreds of
Coccidea Leuckari. 18i9. and order lfoncrlida I eger. 1911. cresent·~haped ~lender organisms (Figures I 8.2 and 5). The
Opinions differ regarding the placement of f. gondii into a ti"sue cyst wall is elastic, thin r< 0.5 µml and argyrophllic:.
family and subfamil}". Various nmhorities. however. have Cysts in the brnm are round and up co 60 LU11 in diamcrer,
placed it in one or the other or the. famllit!!I Eimeriidae. wherea$ intramuscular cys1~ are elongated and may be
:-.lichin. 1903. Sarcocys.idae. Poche. 1913. or Toxopla<;ma- 100 µm long: 11\.: tis<uc cys1s in unstained squash prepara-
1idae, Biocca. 1956. The name Toxoplasma (toxon = arc. tions vary in size depending on the pressure applied to che
plasma= forml is derived from the cresct:nt shape of th<1 ta- cover~llp. ,\llhough lh('y may de\'elop in ,•hceral organ~. in-
ch}"l.Oite ,tage. There are three infectious singes of T gcmdii: cluding lungs. liver. and kidneys. they are mom pnwalent In
1he tach}7.0ites (in groups). che lm1dy-t:oitcs (in 1issue cuts). the brain. skeletal muscles and myocardium. Tissue cyst ior-
and the 'q)oro1.oi1es (In oocysts).30• 51 matlon can occur a~ early as one 10 rwo we('ks :ifter infenion
The tachy1.oi1e i$ often crescent shaped and i-; approxi- and cysts are probably innocuous for their host and l'an per-
macel> 2"611m In si1.e in tis.<tue.smears (Figure 18.2A) and sist for the li,nitth or its life.
33;

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 Oehnnrve
host
(cat)

U"sporu 1a:ej
oc;vsts
oassee n faeces

Cysts ingested
by ca,

Ingested
cysts in
mfeclive _
meat
(raw or
undercooked)
Tachvl'Oltes
transm,ttedm
ihrough ~
p,acema Crvtym
,n feec
::ate~
C!Sll!I
Con.amma:ed
food and ,ngesied by
water intermeniate
hn,s~
Sporulatee ooc, s:s

tnterme,:l'aie hOsts
Infected roe-n:s

Figure 18.l Life cycle of Tox('J{)lasma gondi,

n,e bradyloites are approximately 7 x l.5 µm. Bradrzoi- ingesting tachr1.oi1es or oocysb whereas near!) all cab shed
1cs differ siruciurally only slightly from tachyzoites. The 00<:)'Sts after ingesting tiS<>"\le cysts. Oorysrs are le.;;s infectiou:.
former have a nucleus situated towards the posterior end for the definitive host (cat) than to non-feline hostsF 15
(Figure 18.5B). where~ the nucleus in tachy-1.oi!cs is more After the ingestion of tissue c:ysts by c-c1t~. rhe cyst wall is
centrally located (Figure 18.3). The comems ofrhoptrielt in di:.solvcd by the proteolytic enzymes in the stomach and
bradyzoitcs in older tissue C)'Sts arc electron-dense (Figure small imestinc. The rele.ased bmdyzoites penetrate the epi-
18.58).:?·1 Brad)"lOites c:ontain several amylopectin granule} thelial cells of the small Intestine and initiate development
which stain red with periodic acid-Schiff (P.\Sl. lhesc par- of numeroU"- gencmrions of T. gondii.-u Fi\'C morphological!\·
ticles are absent in tach)zoites. distinct type,(.-\ to El of T. gondii de\·elop in intestinal epithe-
Cats shoo OOC')'Sts after ingesting any of the three infectious lial cells befort' gamctogonr begiru.. Type~ ,\ 10 E di\idc
.stages of T. gondii. Le. tachy-1.0i1es. bradyzo!tes, and asexually by endodyogcuy. endopol)'geny or schizogon)' di-
sporozoites. 33 • 34 • 37 Pre patent periods (time 10 the shedding of' vision into more than two organisms). The origin of gamoms
OOC}'$ts after initial infection) and frequency of OOC.')'St !<hed- has not been determined. but it is probubly the rnerozoires
ding vary according to the stage of T. go11dii ingesred. Pre- relt'ascd from schizoms of type.~ D and E which initiate ga-
parent periods are 3 to IO days after ingesting tissue cysts and mete formuuon. Gamonts occur throughour the small ime~-
generally 19 day:. or more after ingesting tachy-1.oite~ or line but most commonl)·in the ileum (Figure 18.6). Gamonts
oocysts,29· 3-1, 51! Less than 50 per cent Of cats shed oocysts after and schizoms are located In surface epitl1elial cells, U$uall}

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...
•~ • - .,(
• .,. •.
-"
-r~ ..
•

~
_,
r-
'
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,, , •
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Figure 18.2 -.c:ivzo1te$ and 1,ssue c1s:s of Toxoplesma gondi, 3,r a 20 ~m and apl)hes to all pllo,og;aphs ,n lhe comp:is,te /,gure
A Ex;:acellular single (arrowhead! and paired tachyto•tes larrowl ,n a macrophage. Impression s~ar of lung, G1em~ stair.
B Tat:hY?oites in a section of a l\mph ~ode Note numeto~s oval-shaped tachyzoites (arro1•,sl. Haema10xylln a11d eosin stain
C IWacellular tissue cyst In a <eura1 cell ,n section of cerebellum The ~ost cell nucleus ,s ,nd~n:ec (arrowhead). Note .r1n cvst wail (arrow) enclcsmg
numerous oradyio1res
D T,ssce CVS! in sec11011 of blarn Note dar~ly stained bradvz011es 1a,rowheadsl and unsta,~et! t•ssue cvst w.ill (arro,1J. Per,oo,c acld-Sci11ff haema1ox1:m
E Tissue cvs: ireed from mouse oie;n Note mm cyst wall (ar:ow) Unsia<ned

above the ho~t cell nucleus (Figure 18.6). The fcmal;: ·1: goll{lii may di~semir,ate 10 extrolmcstinal tissues. roxo·
gamete is subspherlcal and con1ai11s n single cen1n1II> plasnu, gondil persists in ln1estinal and extrainrestinnl tissues
loca1ed nucleus {Figure 18.6). !\la1ure male gamoms are of ems for at least several months. if nor for lhe life of the cat.
O\'Oid 10 ellipsoidal in shape and c<>main iO tl> 21 nude! Thus. a cat Is both 1he intcrn1rclia1e and 1he definitive host.
(Figure 18.6) . Each microgamew has two flagella. The lnrenncdiare hosrs bocom<, infected after the ingestion
microgametes swim to and penetrate a mature macro· of sporulatecl oocy~ts or tis~uc cyst~ containing braclyzoites.
gamete. After penerrarion. oocyst wall forma1ion begins In the lumen of the small intestine. sporozoite~ released
around the fertili7.Cd gamete.\ \~1en they are mature. the oo- rrom oocy;1s !Ind bradr1.oirc1, liberated from tissue cysts
cysts are discharged Imo the ime~tinal lumen following nip· penetrate the mucosa 10 reach the submucosa and regional
ture of the infected intestinal epithdial cells (Figure 18.7).33 lymph nodes where 1hc> first multiply (Figure 18.8) and are
Unsporulnted oocysH are subspherkal to spherical and 1hcn di~emlnated via 1he blood and lymph s1reamsis u ro
are 10 x 12 pm in diameter (Figure 18.7). Sporlllation infect a "ariecy of cell types in which they muhipl> by en-
occur11 outside rhc cat within one 10 fivt? da}s dep1mding dodyogen> 10 form tiss1ui cyst~. Sporozoites and brndy7.oites
upon aerarion and temperature. Sporulated oocys1s are MT. go11dii can he tran<mitred to the foems,ia 1he placenta.
subspherical to ellipsoidal and each sporulated oocysr con- This occurs in humans. sheep. goats and se\eral higher
tains two ellipsoidal sporocys1s, each with four ,porozoires. mammal~ when infection occurs duriJ1g pregnancy.2~ 10· '
-\s the enreroepilhelial cycle progres.~es. bradyzoltcs pen- I loweV"l.'r. In mice, infection can be tran.ml11ed lO the foews
etrate the lam Ula propria of the feline in1esli11e and mu lei ply from 1hc dam intcctcd lwfore pregnancy and fo, man) gen-
as iachyzoites.33 Within a few hour~ after infec1ion of cats, eration~ thereaf.er. 2

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-
.

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.,I
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•
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f igure 18.3 Transmiss:on e!ec:rM micrograph oi :acnvzo1t~s of f())Of],'aSmi/ gonai, ,r a par.;s,to~l'tQro1Js vac.;cle !!'vi o' 11~t cvropfasm in ire per·tweal
e ,Jate cf a mouse Note ,o~o,e !Co). eiecuon·lucent ·~op:,es !Rh). dense granu:es 10g,, .;mylocewn fAmJ. a centra ly ·~ated nucleus INJ, anc :he
,r.t-avaCllolar 1ubular nar:,or, ;n1·11 ,n ?v One :ach;'?O•!e is d1,~oing ,nto two by e<1iro~1'0geny No1e cono1ds c; ;ne :!a~gote1 or~~·~ms farrov,s1 are
~:;rser.t aoove me parent nuc e11s 1l'rof CA Speer College ot A9ncul1Ura! Sciences a,,o i\atura! Resources. !JnM,rs,r; o• i ennessee, Knox.ill!! aqc
J, s D F:,u. K Kueni and O .;us· ~g. USA.MRIIO Fort Ge:e•,c,., Mal'/land, USA are ac,nowledge•j w,th thar<S for prov,dmg this ligure and Frgi;,ei 'a~
:i and 91

Epidemiology solid 1mmunll)'to rcshcdding of oocysts afwr primary infoc-

Domestic cats play a central role in the di<-,emination oft. tion. \!though they may shed oocy,"t~ on reinfection. the
gondii in nature. 'i:01 only domestic cats Fe/is do111,•s1i1·11s) number of o<>cysts ~hed for 1he second time are fewer than
but other member,. or th~ l'elidae can i.hcd oocysrs.JO after primary i11fec1ion."u Oocysu, are not present 011 a c.it's
Oo~;.1s of ·1; gondif were reported from fo1Jce. of nu1urall~ fur th~rt1iore pc!llng a cat i~ 1101 the !,OUrce of T gondii
infected jaguar !Pamlrera oncoJ,8 ocelots Cl·e/i$ parr/11/is),1'-; infection for humans but handling faecal!\' contaminated
cheetah (,\ci11on.rx jubmus), bobcats (lynx m/11:,J. 62 and material. such as sand-boxes or garden soil. is.2" !>µorulated
• Canadian lyn~ (Lynt co11tule11sis}.' Outbreak. of acute oocy,ts are resistant to cm·ironmental factor~ ,md can
coxoplasmosis in human~ were auributcd to contamlnaric)n ,un·h·e frce1.ing :rnd drying. l(I
of a water reservoir by oocyst.s shed by fora! domestic Herbivor()us animals mainly bec;omc infocwd as a n:suh
cats' and oocysts inhaled in a riding ,table.· 3 ,\ cat may shed of faecal contamination of paMures. w111er and reed \\ith
millions oi ooc\'sts lifter ingesting u few (];;sue c,·srs or oocysts from ca1~ and 0thi.'r felids, and pasture comami·
· ."l, ~1 0 ocyscs arc"hc d ·m Iarr,c num bers ~,or 1wo
brady;,;01tt'S. nation mny be of more imporumcc than conserved feed.
10 three week" following infection. Cats usually develop n $porulauid oocy~I~ remain infcctlou~ for more than a year

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 Figure 18.4 i·a~srn,ss,cr ele.ttoo
mic,c;rapr c; a voun~ hss:..e CV!ii in
a mouse ora,r Note :,el!-csvelo~ed
cyst v.alr ra··~w1 er.cJcsing lour
bradvzo1tes No:e cono1d 1C1 and
postenorl·, lctatej ~ucieus {NI
in one brad·1zo,1e Th,s '!llure was
provided t:-i Pf01 C l\. Soeer and
Ors D Fr,tl K K~ehl ar.d D Aust,,.,g
v,deFigu.re '83

and. because of 1hei1· ~mall llize. are easily dispersed br wind Afrlc.i - 3i of 41 ll<lns (Pn11them irol, 1hree of four leo·
and rain. Transmission of T. gondii rn ewes b}' contact with pards (Pnmhem partlr1sl. :tr.id 10 of 23 cheetahs - has been
infec1ed placenrns and aborted roemses is not or practical repor1ed.
significance. Toxopla$111t1 go11dii is an important cause of abortion and
Toxoplnsmn gomi/F may be 1.ran$miued ro pigs noc <mly related problem~ in sheep in several countries including the
a, a result of 1he lngesrion of oocvsts. bul al~o oi uncooked USA. Am,1raha. ,ew 7ealand, 1he Ui-.. and Norway. 3 · ~ ;io J".
or undercooked swill. infer1ed rodents or other imcrmedi- ,~. 4'•· ;i;. ,~ M< Jn these countries toxopla,mosis is a priman
a1e hos1s. or 1hrough cannibalism. cause of loss in 10 10 20 percent of flocks which have abor-
There is possihly also an uctini sylvatic cycle in which tioll/Millhinh problt·ms. Similar!) toxoplasmosi~ can cau,e
1,ild felid~ ac1 as definitive host~. and 1hcir prey. such as ro- severe losses in goats. 17 :ia. "''
denrs and birds. ar!' imem1edia1e hos1s. In sou1hem .·\frica. The consumption of undercooked mem comaining tis-
in whil·h a large popula1ion ofwild felid~ and a wide range of sue cys1s appears lO I.>~ the moit common method ot infec-
potcmlal inicrmcdi.uc ho~ts exist. such a sylvalic cycle may rlcm in human~. Tissue cy~t.., have been lsola1ed from
play an imponam role in 1hc uansmission of T. gondii to muuon. pork and chicken meat. Caulc arc highly resistant
herbi\'Ores. to r. gmrc/ii infection and the consumption of beef is prob-
Very !lute infom1ation l~ available on lhe prevalence ably 11111 imponant in the 1ransmission of the infection. 10
of toxoplasmosis in animals in coun1rie-• in southern 1 luman mfoction can also occur follm,1ng rhe aC'cidemal
Afrlca.5 ff. b6. ; 2, 1·1 However. nOlwithstanding a high pre,-a· ingc~tion of ooc:y,t~ fr!Jm cat faece,.
lcnc;e of serol0gically posi!i\'e animals. th!' occurrence of In Cemrnl Amcrk'a and 01her countries with a mol~t
disease in domcs1ic animals i~ vcrv rare and only a fe" cases of 1ropic11I dimmc. 1he prevalence of scrologitally positive hu·
1oxopla$mosis have been reported in do~<>1 ;i • 77 although mans increa~es from about one )'ear of age to adolescence.
these might have heen due to nt•osporosb (see Chapter This is con~idered 10 be due 10 children in 1hese areas ha-
23: :-leosporosis). To.w,pltisma gondii has been reported b11uaUy p!aymi in ooc1·,;1-conramina1ed soil:w Toxoplllsmn
in a Cape hunting dog (/ ymon vicms) 58 and a cht.'e1ah 78 in g()lulii is an opponunisrk puthogen in humans \\ith
~ou1hem Africa. Re<:e111Jy. the finding or amibodi"'° 10 T go11dii immune ~y,tem d~~runc1ion. ln human lmmunodeficlenq
in 50 of 68 (74 per ceml of sera from \,ild felids from ~outh , irus infections Lo~opln~rnosis u;uallr presems as a

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3•12 ,.,L110~ t\\V)! Protoz(}n:I diseas~~

Figure 18.5 Trans,111ss,on e,aciron micrograph cf a larga : ss,e tvs, of Toxoptasme gcmf11 n a mouse blain
A Note thin cys1 wall larrow) aria many bradyzo1tes
B H1gner magnif,catton o' a G,aovzoi1e in loog,tuclinal s~tioo. Note cono,d !Coi. rn,croneires IMnl. elec.rcn-Clense rr.Ol)i•es IRhl with convolutaa recr.s.
and a terminally located nuctetis INu) (This figure was prcvided b\' Prof. CA Speer and Ors O Friu, K. Kuehl aoo D. Aust,ng: v1de Figure 18.3

meningoencephali1ls. It is also imponant for pregnant were serologically positive. the source of infection possibly
women as tach}7.0ltes c:an cross the placental barrier and being associated Wilh the handling or consumption of raw
cause defects in the deYeloping foetu~. or undercooked meat.
·" widcvariatlon in the occurrence ofT. gondii antibodies
J1as bec:n recorded in humans in Lropkal Afri<:a. 30 :\ limited
number of studil"!i concerning human 10xoplasmosis ha~
Pathogenesis
been conducted in soUlhern Africa. In Soulh Africa. in one \fter the ingestion of T. gondii b~ a non-feline ho~t al-
study. 37 per cent of 806 human ~era from Cauteng were though the following sequence of events can also occur m
positi,e.63 while. in anoLher. 29 per cent of the general cats). it.first mulrlplies in the submucosa of the small intes-
population in Zimbabwe were serologicnll~• posich·e.-2 How- tine and its associated lymph nodes~8 · • 5 (Figure 18.9). The
ever. -17 per cent of the slaughtennen in Zimbabwe (n =-90) host may die of enteritis and lymphadenilis before other

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 Agure 18.6 Enteroepnhelial stag~
of ro~opla,,"lla gondii in Ileum of a
cat six oavS a!ier reading !Issue
C'f'S!S No1e mature schizoms IS)
O'~fe gamor.;s IMgl. and fema•e
gamon:s !Fgl are rotated m
ep,til~ ,a1 cells abl)V8 ·Jie host c;ell
nucieus Zo,tes prol!abtv aeve:oc•r.g
scr.,iomsl ace 31so present tov,aro
,he !):us~ ~order TathY10ites Ii I are
orcsem ,r. tt:e lam,na oropria
Hae-natoiyt,n and eosm srain

I -

1@em
~--
1

Figure 18.7 Unsporu•ated ooeysts oi

ioxop/asrrta 9ondii ,n faeces of a ca;

organ~ are affected.30 Parasitaem!a occurs during the first develop subclinically and tissue cysts containing bradr-
week. tachyzoi1es being disseminated in lymphOC}'les. mac- zoites form . Tlssuecys1s do 001 usually evoke any hos1 rea.c-
rophages and granulOC)'tes. and as free forms via the lymph tion a~ ency,rmem protects the bradytoites from both the
and blood to the rissues and organs in which they actively h umeral and cellular immune s~'<tems. Inflammation may
invade. or are actively phagocy1osed by. host cells} 3 TI1ey follow cys1 ruprure and reacrh·ation of mrection may occur
multiply and, in doing so. destroy the host cell and cell- in immuno;uppressed animals or humans, such as those
10-cell rransmission may occur wi1hin infected organs~ 3 suffering from the acquired immunodeficient disease
(Figure 18.9). In this wai· small ne<'ro1ic foci are formed syndrome 1A1DS), 30
tha1 elicit an inflarnmato~ reaction in which monuc!ear In susceptible pregnant animals. cransplacental infel.'tion
cells predominate. In most animals a pro tective Immunity causes abortion as a result of either placentiris or foetal infec-
develops before tachyzoites. have had 1ime to cause serious tion. or both. Parasi1aemia and foer.al infection occur during
deleterious effects. The infection, howel'er. continues to t.he seeond week after ingeStion of OOC}'Sts b~· pregnant

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3-1 ·l ,t<.. tu.>\. '"\o: Prowzo.aJ tHscascs

Figure 18.8 Transmiss,oo e!etiron m,trograch of cel!s m tne ilea I lamma propna oi the small 1m.estmo Cl a mouse.:18 noo:s aflei leedmg ooc1s,s.
Note t/le tell on the lei, ,s a ne•,t:ophil showing specific granules tSg! and a parasitophorous •,acµole (Pv) con1aim119 ~....o 1ectiv1011es and a
:iell-de..,eloped 1ubulovesiC1Jlar mem!iranous network !Tm). or. :he r,ght side a sir1gle :iichyzo•\a 1s located w1tmr. an endothl!lial cell (En) that has !lecoma
displacad but ,s still connectt~ bv a narrO\Y cytop!asm,c tr1dge !arrow) with the cao,lla'Y enoottiehum fEJ\J. tlasal lamma ot capillary eodothellum •.Bl),
lumen (Lui of capillary· spet1hc granuferSgl of neutroch, 1ihis 11gure was prov,cted ov Prof C.A Speer and Ors D frn:r, <. <uahl ano D. Ausung: v1crs
Rgure 18.3)

~heep and goats. 16 The s,:,ver!t} of the lesion~ in llw foc1us sheep and goa1s during the earl) ~tage~ or pregnanq- res uh~
generally exceeds that in the dam. the effect of th<? focrnl in- in resorption. abonion or mummification of the foetus. In-
fection largely being determined l>y the age of the cuncep· fection b(:twcon 50and 120 days of geswtion result$ in abor-
tu~- Infection during the early stages of gestation may lead to tion or the birth of premature. weak lambs or kids. or in the
embryonic death and resorptlon, while infection of ovine birth of offspring suffering from subcllnical disease. Apart
foeruses at about 70 to 90 days of ges1a11011 is responsible for from aborting. ewes an<l does do not generally manifest anv
the death of a large proponion of the foetuses. and lluiy arc ollier e,idencc that they are infected. Clinical 1oxoplasmosis
either aborred or bt'Come mummified and an!' then cxpt'lled has. howe-\'er. been obser,ed in adult goa1,,?2 pig,~1 and
16 0
ai a later stage: Some survive the infection for ~everol chickens' • ,:1 but is rare. Ulindness and torticollis ht'lvc been
weeks but are born dead close to term, or else! 1hey are born described in cimaries.80 There is no well-documemcd case
ali\'c but weak and die soon thereafter. Infection of ewes of clinical toxoplasmo~b in cattle a11d horse~.2°· 10 Clinical
during the last 30 days of ges1~1ion generallr re,u!ts in $\lb· toxoplasmos!s QC-curs in dogs, cats and ;\ustrallan mar~upi·
clinical infection of new-born lambs. als. 30· io The clinical signs manifested by 100 cases of fatal
!he pathogenicit) or r. grmdii i~ closely related to the vim· toxoplasmo~h in cats from one ho,;pital in the USA 31 and in
lencc of the strain of the org:mi,m and the susceptibilit}· of the wild and ,mo animals~0 have bec,n fi!\'il'W<!d. Acute systemic
host species. New \Vorld monkeys. Auscralian marsupials and toxoplasmo~js in animals induces a wide variCt) of clinical
Palla~ catl> are ll1e most su.:.ccptible to coxopla:.mosb. whereas signs because or the many organ~ and 1i:;;.ucs that may be
Old World monke'>'S.. rats, c:mle, horses and buffaloes arc affected. !"he most common "igns observed are those result-
highly resistant 10 clinical 1oxoplasmosis.30· • 0 Other unknown ing from encephalitis. pneumonia and rt·tinitis II In spne of
fac1ors vaguely cla:.sified 3> ~tre,, may affect the outcome or the reprnduction of 1he parasite in the intestine of cat,. clini-
T. gondii infecllon in a ho,1. \•!ore severe infections are fountl cal e\'idence of emeric disease has not bcteh recorded.
rn pregnant or lacmting mice than in non·prcynant or non- ;\lost posmatally acquired infections in human;; are
lactating mice. Concomitant infoctions may make a host generally as)1np1oma1lc. but fel'c:r. Jymphodenopa1hy.
more suscep11ble or resistant 10 T. g<mdii infeclion. Oinical headache. myalgla and fatigue may occur."11• 31 Severe
toxoplasmosis in dogs is often associated with t'anine dis- illnes.s may develop in chose suffering from 1mnrnno-
temper. chrlichio~is and lymphosnrcoma30· 61 deficie1icy s~~idromes or who are undergoing immuno-
~uppressil'e 1hernpy. Congenital infeccion. on the other
hand. may ha,-e serious consequences, chorioretinitis
Clinical signs
neurological dbturbances and hydrocephalus bein(! most
In li\'C:.tock, clinical sign, are most commonly encountered commonly manifc~tcd. Clinical 10xopla~mosis In humans
after 1ransplacemal infection has occurred. lnfeclion or has been reported in !:>ouch Africa. ;o

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 Tuxoplasmr»ls 345

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Figure 18.9 .esions in :he sir.,11 ,mestir.e oi c;gs ieo To~aGiasrr.. go~d11 oocvsts
2@11-![l
--- . - •
• ~
L ·'

I\ ·~o,e transmural neoos,s l2rro::I and oaifoonmg of •:ill, tsrrow~eacsl

B H:gher r,,a3n1ficat1on o! a~ asea o' ,r.•,amma,,on a,c necros-s iai ro·,•, 1ad12cent :o rela!t\'?IV a~a'fetle!I ep11he11um
C High magnif1ca11on of a·,illus shO',·,ing oedema o/ lamina prcoria lar:owheadsi and exudafon imo :h~ lumen troll'! an ulte• (arrow!
D ~i;J!!e, magn,l.cat,on of lamina p1o~r,a showing :ac~1·,o,:es (arrowsI H;iemaioX'1ltn und i:gsm s:a,n

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346 <rcn..,, nvo: Protozoal disc:m,s

Pathology enZ)1nc-Unkcd immunoabsorbent assay (EUSAl, and im-

munoabsorbent agglu1ination assay test (lAAn.:m The TFT.
Systemic toxoplasmosis occurs mainly in young animals !I\AT and ELISA have been modified 10 enable detection of
and immunocomprornised hosts. The predominant lesions lg:\I antibodies which appear sooner after infection than
of syStemic coxoplasmosis in animals comprise Interstitial lgG and disappear faster than lgG after recovery. The finding
pneumonitis, focal hepatic necrosis, lymphadenitis. myo- of antibodies to T. go11dil in one serum sample from an ani-
carditis and non-suppmach·e meningoencepha.lltis. 19· 12· mal merely establishes that the host has been infected at
23, 30. 31· ~0• 54 , 61 , tiil Macroscopic lesions in the lung vary from ~omc time in the past; it is therefore preferable to collect two
irregular grey areas of necrosis to haemorrhagic pneumo- samples from the same animal. che second cwo ro four
nia. :-.-ecrosis is common and appears to be related to the weeks after the first. A four-fold increase in amibou} titre in
rapid replication of tachyzoites. The most striking and the second sample Indicates an acute infection.ln some ani-
parhognomonic lesions of roxoplasmosis in animals, how- mals a high antibody titre persists for months after infcc,ion.
ever, are in the placenta of sheep and goats.'1· 5 9 · ' "' r7• 30. 311· A rise in amlbody titre ma}' not be associated with clinical
39 5 6
• " • ' These consist of foci of necrosis varying in size from signs. The fact that titres persist after clinical recovery com-
microscopic to macroscopic in che COtJledons while inter- plicates tlle imerpretation of the re~uhs of serological tests.
cotyledonary areas are unaffected {Figure 18. lO). The char- The direct agglutination test using formalin-presen·ed
acteristic lesions manifest as white flecks or mulrlple white, whole tach)~toites and mercapt0e1hanol has proved \'er:,
chalky nodules up 10 2 mm in diameter and are found in ap- useful in measuring lgG antibodies in livestock, Including
proximateli• half of the confirmed cases. TI1ey may be sparse poultry?0,3i , 1,,14
or dense. may occur in any plane of the cotyledon (Figure Finding antibodies ln foetal fluids or presuc-kllng serum
18. 10) and may be confluent. Not all cotyledons are affected in ruminants is diagnostic of congenital infection because
to the same degree. It is often necessary to wash the cotyle· maternal a.mibodies do not cross the placenta.32 ~.
dons thoroughly in saline soluri0n in order to expose the Toxoplasma go11dii can be isolated from the infected
deeper lesions that may otherwise be overlooked. rissues. secretions. excretion~ and body fluids of animals
The earliest lesions are necrosis of meseuchymnl cells. collected either ante- or pose-mortally, by the Inoculation
and oedema in the foetal ,illi with infiltration of mono- of them Into laboratory animals and for 1issue cuhures.30
nuclearcellsassociared with hyperplasia and focal coagula- Using these me1hOd$ it is pos!iible not only H> isolate
tive necrosis of the u:ophoblastic epithelium . Older lesions T. gondU. but also microscopicallr 10 visualize the organ-
consist of foci of caseous necrosis invohing foetal and ma- ism or its DXA using the polymerase chain reaction (PCR1
50 60
ternal villi which may show mineraliMtlon. c\lost T. gondii rest. ' A rapid diagnosis may he made b, the micro-
presem in lesions are degenerated and difficult to find in scopic examination of impression smears of lesions . •.1,fter
haematoio.·ylin and eosin-stained sections. lmmunohls- d1yi11g for IO 10 30 minme!>, the ~mears are fixed in methyl
tochemical staining is useful ln locaring T. go11dfl' 0 (Figures alcohol and stained wirh a Romanowsky stain. Giemsa
18.lOC and D). being very satisfactory. Well-preserYed T. go11dii are cres-
Lesions in foetal tissues are microscopic and occur par- cent shaped (Figures 18.2.1\). In tissue sections, the
ticularly in the brain. liver and lungs.•· :io. 55 Lesion~ in the tachyzoites usually appear round ro oval (figure 18.28).
brain a.re found in 90 per cent of cases and consist of leu- Tissue cysrs are usually spherical and lack septa, and their
koencephalomalacia and gliosis. walls stain \\ith silver stains. The brad}"!.Oltes are strongly
Placental lesions have also been described in toxoplas- PAS positive {Figure 18.20). The lmmunohistochemical
mosis in cats36 and pigs.'2 ~umerous pale foci of necrosis staining or parasites with nuoreseem or other types of
were seen in the allantochorion of the cats while multiple labelled T. go11dii antiserum can assist in the diagnosis76
foci of necrosis of the allamochorlon were associated with (Figure 18. 1O). Electron microscopy can aid the diagnosis.
numerous tachyzoites In the pigs. 'f'oxoplasma gondii tachyzoaes are always located in
vacuoles; they have rew (usually four) rhoptries and often
ha,·e a honeycomb st rucrure (Figure 18.3).
Diagnosis
!11 cats. oocysts in faeces are concentrated by salt or sugar
The diagnosis of toxoplasmosis can be made by bfological. floatation techniq~ies.30 They can be dilferemiated from the
serological or histological methods, or by a combination of oocysts of lsospora spp. by che fact that r. go11diioocysts are
same of these. Clinical signs are not sufficiently specific 10 smaller (IO x 12 pm) and nre shed in an unsporulated stace
enable a definite diagnosis 10 be made because the disease (l'igure 18.7).
mimics several Other infectious diseases. Ewes are usually serologically positive at the Lime of
Numerous serological procedures are available for the abortion. and a negative test generally excludes wxoplas-
detection of humoral antibodies. including indirect mosls as a cause of the abortion.:\ positi,·e result. howel'er.
haemagglurinarion test (JHTJ, indirect fluorescent antibody is not necessarily diagnostic. as high titres may persist into
rest (IFT). direct agglutination test. latex agglmination tests. the follo,,ing breedingseason. 30

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 , ..
.· ..
•

.
•
~.,
• •
t.
.. ...,,,
•
•• < " .• ~t , -
l '
•
" ..... . ..
..' ..,.,..
....-~
..
..
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10mm ~I • ,-•• '\..;~

B
• •
..

• •

• D
Figure 18.10 lesions mth~o!acen:a o; goa!S and sn!tllp natura!iv niected with To~tasm1: goll(!11
A Cotyledon oi a goat w,:h ....tn,sh -yellow areas or d1scolo,won rarrowsJ. The m,e·coty:edonar, area 1s normal Ur.siaineo
8 M,cnmopic lesior in th'? p1a;er.ta of a goat No1e necrcs.s. mmera1i1ation. and dispersed T gondi, aot,gen {red colou,ea are2~J.
lmmunoms1ocnem1cal s1a1nin9 \'Jttn ant1· T gondii sernir
C Cl'l s;riace of a sheep cotyledon snowing yellowiSh·,•.il,te are;:1s 01necrosis in vrlli {arr~wst Unstained
0 H,gner magnlf1c:ation oi ,,g~:e 18 iOC sllow,ng im1!ct (erio::s11achyzo,tes andd1snersed i good11ant1gens (arrov.neadsJ

Dif!erential ctiagnosis bron disease viruses. ,\'eospora cn11inum, and Sarcocystis spp.
(~ee the relevam chapters on these diseases).
Foetal and neonatal losses in sheep resulting from T. gondif in-
fection should be differemlate<l from oll1er possible causes of
abortion. stillbirths or neonatal deaths, such as lhose caused
Control
by Chlamydophila abom1s (Clrlamydia psitmci) , Q 1mpylo· 111e comrol of toxoplasmosis in livestock ls aimed at the pre·
b&1crer jej1111i. Listeria mo11ocy1oge11cs, Salmonella serovars, vemion of infection in pregnant animals. Tbe perpetuation
Bruce/la spp., Coxiella burnetil, Rift Valley [ever and Wessels- of the life cycle of T. gondii in them depends on the availabil-

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348 ,wnn'< "'": Proto1.ocl diseases

iry of susceptible definitive hosts, the most important of munity to subsequent challenge. The level of monensin In
which is prohabl}'lhe domesriccat, and of infected intenne- the foed should 001 exceed 30 ppm a~ otlterwise refusal of
diate hosts. such as rodents, Cars and rodems should there- feed or toxici(1 may occur. Incorporation of monensin (200
fore be excluded from areas where livestock are fed, or ppm) imo feed blocks is a safe way of utilizing this drug. 10
where their feed is stored. If swill is fed to pi.gs, it should be The addrtioa of decoqulnate to the feed of pregnant ewes at
cooked, and cannibalism should also be prevented. the dnily rate of 2 mg/J<g body weight reduc~ the deleteri -
Treatment of ind ividual cases is u.suaily limited to those ous effects ofT. gondii. 11 A Jive modified T. gondii vaccine is
suffering from acute infections in high-risk groups. and lO marketed in Europe to reduce the p(evnlence of abonion. 13
immunologically compromised hosts. The treatment of Pregnant women who are serologically negative should
choice is potenlia1ed sulphonamide mixtures, such as sul- avoid gardening without wearing gloves. Cat sand trays
phadoxine, and trimethoprim.30 These two drugs act syner- should be deaned daily as oocysts w!ll nol have had time 10
glstically to inhibit sequential steps in the biosynthesls of sporula1e.
folic acid. >',-lonensin ac a dose of 5 mg/animal/day incorpo- Pregnant women should not eat raw or undercooked
rated imo the feed of pregnant ewes can reduce losses from meat and should wash their hands after handling raw mea1.
toxoplasmosis. 12 "xperimcmaily. monensin limits bm does Immunity follows infection so that a second abortion due to
notprel'ent infection; ewes 1hereforcdevelop protective im- t0xoplasmosis is unlikely.

References
,Ut..Ull~I. J,J.. STl".Plll.l.., C.. OUlll!V. J.P.• 199.S. 10Xop/.a1111n go,:d/1 Jn •• ouaiv 1,v.,191h. Toxopl<tst1lfl•indu«-d abortfor, In duirl g1><t1>. Joumal oj
VancOU\'er hl:tnd cour.if) CF1tlls t'fJJ1rulqr nmcomj1t1?n1fs . .,.crut~· and IIW -~mffl(tul 'lc•tc>rlrm,y l\lttdic,11 ,lt.Sl)(idl(Otl, li8:, 671--674.
nocy.,,
shedding. /oimtul of PflflLritOIOiJJ', 8-1, l3fl-4lll. t'; uul\f.\'. 1.P.. 1glh. Et)ii.botfc toxoplasmosi~ as~Jatc.d with nbonion in
:l flf\ r1t1.EY. J.t: .., .. 1959. C"..ongtmlta.l 1ra.n!imi~!tinn of1ox<,,pla.smn~i~ lhrot,tgh di!ll} go:ic5 In Morunnn,/ouma/ oj1hr:.,t.m,•rf(tm Vl'ti'rfnary \1':dkt1l
<u=ive genM!rloni nf mu,,. V11111r1, 183, 13411-1349. ,\$:lorlm/o,,. 178, l'>GI--G10,
3 JSE\'EJtl.El. u:....,. & ,,.·,\'ESQ~. w...., .. 19ts1. 0\'1nc ilbortion nn.d roxopJa~oS:is 18 uuut\\·. ,.r.. 19&4. E.1tperimtnral toxopJtt~rnosts m.she-<!p ftd Toxoptasmn
in Yorkshire.-. nu, Vlmm1t0')' R,"vttl, 1:i. 6--11 .. g<,mdll OtJC}StS. lnrt•rnnrfonrll (j(>(/1 and Sll4"J>p R,r.i~nrt·h. 2, 93-lO-i.
,I llf\ fJU.fi\, nc,A,, W\T!iO"\, \\',,\. k PAY''l:, l,M,. l97J, l"hc pnthologrof the 1.9 OtJEff'li., l,v,. 1986. Tox<lplar.m<>).is in C3t-~- Fe/mt• Prartin:. 16. l!!,,,-46
platemo ln ovinc abonlon due 10 toxoplasmos,s. T/10 Vertr/11110• Rtrord. 20 oustt. J.P.. 1986A.A revkwor'1o~op!,6mo,sb. lnci1111c. Ve1rri11Df)
88. 12-H:?8. P"m1<ftology. 22, 1ri--202..
:s MllVMutY, 1•.1:.A,.. \\'A'r:i-o~ \\.A," ~Prst:h J.n.. 1a-:1. The pathalom· or ihc
21 ouac.v, ,.,.•• 1986. A1evic\\' of to.xopla1mosi~ In pig;t. Vl!terinar,.,
fottu:> Jn ovine abonton due to toxople>tnosls. 77:t! Veurim,,,· R«ord,
Pmtt;/10/0!l',•- l9, 181-22.'l,
88, ti4-l 78.
!?2 a>uat-Y. J.r.. 1987. Toxos11a~n1os1s in 'l.JliHS.. Agri•Prttctite. 8. tJ.....'i2
6 UlG:\LJ.'.-1'., 11.n.• TOSTlS. a.c .. l)U ~·U.S.Sl'-. J.t,., j!IJ\SSON. P..\ .... '-1C-.CUU\. n.M .•
19ij6, Th~i,nhufni, of Tm:011/«.rr,m go11dii fr<m, forrcb In S011d1 \frica. 23 uuor·f J.r.. :989. lcs1m~ in goat.,: fed ·10:copJasmagoudllooc}'SI~
/oum/11 of1/1e:.or11it ,>.frict111 \ 'ei<rirtnry ,11-,/ir11/ .1w,.-lmio11, 37. 2·:3--2H. 1·~ctr/rwry· P11rasl1olo!l}', 32. 13:.1-! H,
.., .o.own.. W.ft. •a~\'O, AS, w1;ma:1<. b,11 .. IS.V.C·Rf.N'ro:,., J.t.. SUL...... 1:XC• .S.1'1,, 24 uu.8t.Y, J.r.,tft93, Toxcip/()Jmtr,, Nrt>~,W1(L $!1f(tJC)'>ll5, and c,nhcr- ti:llul!
\1.,\JUOS . .s.A.. 199:,. Outbreak of 1os:opll\$tnosis a.ssocf:\tcd \,·ith munjLip31 cy>l•formfng coccid!o ofhurru,ns anrl animal._ In, 1'1Ul"Jt. J.P.. :ed.,.
drinking Wl1t(!r. Um(ct. 3SO. rn--111 PnrasitfrPffmr..oa. Vnl. 6. NewYor',;; Academic Press~ pp. t-l5S.

a uuRKJt>GL, 1.1 .• 198<,. Toxopl~rr.o'!O!I /'he CmnfNJtdimn a/C~mi1111i11s 25 a~umv. ,.v., 1a94. Toxupln,.mo,.i~ /m,mul n/the ..\merl«·utt Vt•tfflllfl"'
Eclltl'(l(IQll/Or Vtl(Ft,Ulti0.111. 5. 4-56-162 .11"1/ral.-woci,11/1111. 205, 1693-1598.
s wxro~. n.. 1990, tMne roxopl~smo.t>. a rc,•ftl\\'. /ournnl oftit~ Ro)nl 26 oust\'. J,t-.,199,;. Duration ofi1nn,unt~ ;o >hedd.i1'1g of r,,.t0p!nsmt1
SIJri,•IJ·,;/.\fetlic/11,. 8-1. 500-:i! l g,,,11/11 oocyn, i;y cn1,.Jour,,a/ of Parasitology. Ill. , 10-I 13.
10 ILUX"fO'S, tJ .• nu,,,•m, O.A., r!U'..E..i, ,,.,,., ~1C'COl,<.M'\., c:. ,1,. RSu.,~():,,;, , .. 19,&8. 2r ouao·, r.P.. 1995. lnfe<:ti,•il)' orrd pathogonicir)' of ToxQµ/11:rrna gc>r:dii
l'unMr surdles In th•.,.., of mon,nsin In the tonrrol of e->'J'PrimP,11al rm~-st< fot<o1s.Jo1mwl ofPt,rlJ:$JtlJ/Qg;,·. 82. 9Si-960.
o<ine toxoplo,m0$,,./011m11/ ofCIJmpararir"1 P111hol<>gJ•, ~8- 225-2.'lG. 28 DtJMY. ,.r.. 199; Br:uJ.}'iOitfModuced munne ,o,.,oplwnoS'h: sm.ge
11 ouno:-.. n.. 8Rf.8Nl.R. t.. \\'1UGH1S, s.. ~M.t.b'Y, ~.w.. 'fH0°!\ISOS' t:..M, & com·ers1on. p:uhogenfiJS, and li$1U:C C}·,1 fom1atiun Ill m11:t- (L.id
,u1 u,m. 11;,. i996. l)l,coquin:nc nnd the eontrnl of t.•xperimt.-n1:,l o\inu brody:i:ones or d!O'crom srrofos or Toxoplt<Sml!go11dll. /Q11r,rtJJ of
IOXOplo.c:rrtO~i~. Thi Vq1~tlm1ry Rt4''fJrt/, 138, 431-136. Eukary0tir .\licrol>i1Jlogy. ,,.i. 59?-.602

12 uux10~. D., OOXALD, a....~: . ".nxu.\:::;r.>.s, , .. tf)t\i', Moncrtsio and du~ ,·ontrot 2'J uuu•~. 1.P., =•Oocysi shedding by cat, led bolot<'<i brod>7.oi1<, "1\d
of ~xperimemal O\'H\e to>;0pfasmosis: Asystemic' t.1'lloet. 1'ht Vtt:?rfmuy comp:iri;on of mfc,:r l\1ty of brndp:olteo of the VF.G srmln roxopla,mo
R~..ord. 120, 618-{;!9. gmullitu Cal~And mu:c~ JourrmlCJ/Par!4-JtQ/ogy, 8i, 2lS.219.
13 m,x t(J~. t>, N 1r,.;i.:-1':,, t-1'., 1995. :\ tommt""rclut ,•ac:dno rnr <>vin~ 1<> Dunn·, 1,P.• 81.ATITT. c.P.. 1988, 1'oxop/.(i,'ftno!.isofJ\11/11111/s and ,\Inn. Boe•
'toxoplasmo,i,. l'rtra$lrofogy. 10. Sl I-S16. Raron. Flor!drc CRC Pr<·$~. pp. 1-220.
IJ CUL\Dt.l!., M.A., SPC~tf:.lt, J..\.. 6,. .BU.trtlUA.X. P,L, L99,,
Sccopre,atenccof 31 nu1.r,·. 1.t1• • C.\ttrlt.~~R. J.1 .. 1993. Hi~tologiall)' confirmed clin!Qf
YelJ'$pora a111/11wn,nd To:mplmmn go1ulilln nnndomestlc lcllc/,; from 10>10pln,,mosiS in <'llt<-IGO rnsc< tl!J•2-l990), fm,,111,t of1h,.~m,•,irat1
S011rhtm Afrlc<L /otmur/ /J[Z/Jo 11ml Wi/1U/fe .1/l!<lir/11~. JO. 248--:?SI. V~r,rlnnry ,\1rtlfntl /U,(Q('uuitm. 20.'i, l.iSG-1566.
1~ rnmr.v. J.P.. 1977. Tnxnpln.cmo. 1/ummo,:dfa. &J111>I1fo. SiJ1to.q11is. iuuJ 32 ount\'. J.P.• U!O'so. J.P•. m:S':\10!\.S, G."& .~matSOS, w.n.. 1987.
oth~, w,.u~ qest-forrnlng coccldJn oJ tllllll and rutlmols Jw ""J£ft. J.P. Se.rodiuµnosls of J>Q$fnnr11Uy .:and prcn.n'2Uy indm."C'd toxopfasmu'.'i~ In
cd.J. Para!ihlr P.-orotoo, Vol. 3. 11:l~~York: ,\cudemk P«-ss. pi,. lOl-237 sh.t..-cp. \m~rlrcu:Joumal of'Vf!1t.•ri1:Q')' Ri's...v,rcl,, 48, 1239- 12.43.

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 Toxoplasmosis 3-19

33 n11»t.v, 1 11 1,1 Ml S!lit,, J,);,. 1g-;-:, c~~t i1td.uc,"d tc>:wpJ~mo.."''-' rn c;4, ... :\u,iro1han b1rd..,,Jownnf of l-"rtt·ri11nry l>u:gnmur J,;:re..$ll1!J-1tio11. J,
/ottmnl nfPmto:oolo~. 19. i;;..1;;. llii•l()~
3,1 UU!iln'. 1 P.a.. rnt..~UL 1,>,;., 1976, Feline cw:opJa ..mo~;,c; fmm .,,u1c;o~· .iS 1u.,11 t n. w.,.
k t..\U·k, r.c.,. 1~1;J The p..tholo~'Y of fO).'tJJ1ltu1111> infrc:11on
lnfoct,:d mkt• ~,nd 1hr dc\'rlopmttnt Qf fo.ro11/ru.ma c.1·..;... Ju:,nidl ,,f m th~ ;1:c.~.ant 1."\\t:.. U~11,,t1 m \ (f('fllin')· \dtm«• ..a, 1"di,,.ll2
JJ1v:o~oal1tgy, 23. :ni-.5..in ;;., 11.\Rruf\. w.1. .a. ,r ,h.. Jl,\l l. S.t 1c-;,:;:7. Tu,r,pl;M1in,i, n,i ;i c:.uw of O\int·
J3 DU8.t\. 1.P.. u.:,..OSAY, D~. "SPli.E.a, C.A. 1998. Stt\Jtturt• 01 Toxo:,Jns,,u, pu.nn.uo.J mot't.Q.fjty. \1t·u1Ztv,lmuf \(lt'ri1111r;,, /()tJr,mJ. 5. 119-12,;
om:dii uichy1.ulte--;, l>md~"'2brtc..; and •prm,1.oit~. anJ hinltt!,''Y .a.nd 5; ll\fll1C'f w.1 ,~on.F~ c;,<• 1968,.. Obse-n'.lUOfh on ;au nuthwak or ov1ne
Ck\clup:n~t Qf tl~,u«" ~,,~ CU11irol ,\flt1n/Jinto10, Rt't'fr:c-< 11. 26"7-2'9, cP:1\,:<rnmd 1•i~u-pt.\.,mufi1t,.. Amamliun \ i·tmn11,:.• /(rtlfmt/. .f4. 1OS--10'7.
0

36 UUtUl. J.P., \f\M I~ \S.f.. & UM,L-0~16. T 1' 1996. L~~on... c>lnl"'OnataJI~ .;,II 11arM1"• C..F.U., 1%ri J \W hunrlred '1.nd 1.•IJmt)•fC'lur itUlUJhll.... okt thL·
tndla:~ ioxoplul'in10,h, in ra;-"'. ~·t•1t•rtnt11'' Pmltblo~·· 33. 29Q...::?95. :l:atl<>nol /'1<li"!ll<al Cord<n,, 11M<>rl:1 /~11tnnl oft/I• .\fllllh ,IJr/,·w,
3; nu~n. 1.,., ,.,u..,.. "·'·,. 1 Rl:-."1.l. f,K •• 19;0. lhe f().,o,,lam:,t 8"""11 1.'cwrln11ry ,\faJ,,al Mmctatfon. 27, '2a3--u.?.
ooey·~,, 1rom t.'Ul (l,'t'.4.'' /uurnul OJ Experunf'mol M('fUcmc. 132, 6J6...&6'.? ~ IH'i\'J;, T. "rnuu \",,.,••• It)~ lt-rl"\'Oa.11.-~nt,1 "' 1·a.lf>/TIMmt, J{,(JU(lft nmlboJIR
38 PUM.\". I .P.. \llt.u:ft. s... Of.'\~!0:\"1~- (,,
nun I lJ /. ,. "A~n,n"'°'· \\
.Jt•• t!U:ib. In ><'rd of dninc<lk plj:> 11nd .wm• \\llJ ~.unnp.:cJC\ rrum /.imbJtm<.
TqXtJt11<uma gond(Hnduttd i\bomun ln d.tu)· g(ll'lt"", /nttnml nfthr /011,,wl or ,.,ttrtt}illJlhf.,.\', 85. :-s;:t-J7l.
:\m.t•r:cnn l·t'tl'rlUt1T)' .1d«ti"1/ t'.5«iallon. Uf.8•• ' '59-162 r.o !A«.oH,;. ''·"· !C.. ,1.-t~)'\. 1•.1t. 19;3. Prt..\ nl1.•11t'l> o1 To.'(tJJ}fm.,1m antibodii:~ in
s.., 11t1Wl·U l-,(' ~ ·'-'llll~~Cl\, w.11 .• 1ql\t,, CpU:outnlc,ga,:
39 DUISFY, , ..... MILIJ ti, Wllthcm .\fricu. Mmt1J \{ria111 ,\lt•1ht11I Jm,wal~ 53. 6!9-621
lm·ti.dgouinns-on a shct-p farm ,,;th roxo1,lm,,w, gqurlh·induct.'d hi n,ua, ir,;.,·.c- Ki "'\;Jl\V, r,r.. h " " ' ,,l,.~""·· 199.;:. Pmho/ogy fl} /)omt,lll(
aboruon, /(,mnud ofilh• Anurlcmt V,•wrinnt)' M1'tJicttl :\Jsorlur1mt. 18ft, ·\n/mal., ah 1,,.ldr: 1.,mctc,n: Ac-a-Jt.--m11.:" .rrcs.,
15$-158.
ig-;6. l'te\·;,aJ.entc or
62 '1\IIC.Um,·uo, .-\..'\. tllhl.l'''l\)a. O.W. h '1-~\UPJ\, Ct,O
~o ft•l.itL"<I tnf«t!on" /n;
tJl.tl:\fY. J.P.. " PtW:io.ll\'C"t k. ~~n.1'oxaplibtnf1.Si'- und ;m1ihutlu.•... to TtMfJpltw,m f_l(Jmfll fn wild and donw~tk nuhubh ur ;\,"\,·
i,.\MU~I .. ti.. P\lWM. M I. ,me~\~. , "·· 11.-d"j, Pnm.slilr l>IN.v'1$N ofU,'tld Mc\jto• .-\ri1.unn and C~olor:ido. Jon maI"' W,ldltfC' (){~.,,.~. t2. 220-232.
\famm1ds• •\m,-.;, Iowa Sam• \Jnlvcr..11i· p,.,..,, pp . .;ill-SI~
Cl;& \.1.\-,o,·, t-.R., J.\U.lll~- \1.11, "ff 1 llll!t•. f•.1 .. l'li.C, ~tn!n1,tJi;aJ '111'\e-)' o(
Al DU86\. J.1~•• HU\.'h ~,.o.. l'.A~l.'\RGO. i1.r...• ~m !II.:-.-. .• WII.Xf:\\., r.., K'\HU t0%0JllJ,mu.,i.; In ah-r. Tmn!'.v\lfll. !><111th .\frl(n11 .\lc.'tffrnl /1)11fll(I/. ·i!l..
o.c.H~"' 111owoz.. P., 1!!1'1-\. ~olog!c i..1nd p,H.J.,ilologJt ro~pon~~ of i;'U""-170!1.
donk"St{C thid:.en..: a!n1rorn.l inoculation. ,..;1h IO..copltu11w 15m1(1ii
r,.. -..:btm. 1.v, .. wutir~:,.. 1;,t•H & \\'1111.\.\b. ,1.r_ 1981. Sp:t..tic ~rcsh. in 1wo
OOC!r'S!S:• •.\11wtft:tm Joumul <1/ ti·ltrl1tu1'• R~1rth. S.i. 16J:i8-1672,
Uu,:rm;ih: pup, ca1J~\'1J b) JOX,,plttMn" p.aml(i. Jm1t1lh.l nf1ltr.'t(Jt1th
l'2 DU8l-'"i J.P•• .s<:tnAFlln, 1).11., OR.DA~. f.t,. ltl. ~ UNJ)S.\Y u..:,... 19:"JO. los.lons Lj'ric(tl1 \ ·N~rfour;· M·otlot/011 . .>2. 24~·10.
In (et.al pigs with 1rnr1iplncf'm31l~··indu.c.-nl rnX<>i>la~mt.bb. \'cu1ri1w"' n5 owr~.,.· '1,H- r I ARt.:SO!'ii1 ,,., h1~U>. ., J 1S98. Olngno:,;;i.s ni rnxoplo..,m.s
Pt11T10/ox,•. 2i. 4 11--l 18.
abomun m l"\'1\.'~ by 1x;1rmcrrui-<1 chJin roiictfon the le'Jt\rlnfll)' RtYOrrl,
,i3 nue:,-·r 1,,,. &SUAR.\ l<\.. ,.,, l980. i\i:ras.it~ma and u~~ue: inr('(tfon in 'liheep 10\2. H~--H8.
ltd TOA'ap(n$mll gmulH o•~lS Jrmmnl Of t>nrnsltoloSJ', t,6, l l l-1 l-i l;J., rA'.'\IU\ ,-.:.. "- \ \"\; Ks.\rL"\, t , l~2.. ·the ~ropn:vule:nct:- oru,xoplu~o,t,
..~ butU..'"l . , 1?.• ~r>,~.tt.J. ~··rntR.,~_,•. ~ µ.~r !\. 1·.u•• 1w,,
Jffn:s111-0~1. o, In ,r,hc,·p. goat~ and p1g!ii- fn Zlmh,thwc.Arl',,QL\ OJ J'ropirnl .\Ntficlu,·a111t
Sc.rologic and hisiologic d~gl'l{)~~ of io-...opla.."'n'l1c abcm1ons m >-ht!ep 1u /ltlf(IJ1mlo,JJ•, 13,;, 31.1-~I!\,
Oregon. 1,,1,rnul of (I,.• ,o\ua•rJoo,, \.'e.•wn,1mJ .\ fnli,a/ ,\5s<>cim,on. 19b. 6:- v~no, "'· 1ou-..:'io0,. <.L, tonr1111.0.1. .• \\·1tu;111. e.('.. & ,, ,..;1 :r,.;, 1.\t, 1t,ft.S.
:?91-Z:i-i. Lp1.t,i1>U1 hf to):.(1r,la"1mo•,fs ir, J,..;1n1;.1n,n!I, w:illuhiC'!J, :ind pot<1f<1D!J.:
~~ uunn I1•.. -.:PnN. <:.\.,,tu., .... )i.. KW<J.:. n,:.u. "11n,r. 1. \ 19~7 po)-.!ble 1r~.rum~,,on \'fo donu.~,;c ... i:j, , /or,rnul flf tlh1 Amc'rirm,
00~11.Hnduct"d murinc 10:xoptasmo,b,. ttfc rye!~. patho~t:nl,a) and \f('ttTit:ttl) ,\ll'<li(<11.-'u"'>('4·/mioli, 189.1166,,,,1109.
,tag-c com·ersJon ,n mtt"t"" fed ro.topT1u,,r" gmulii OOC)""'· }(mmm of o8 Jffl)J):\tl lfl·. r...l. U,\lltU:\· \\,f,. l')U'IJC\". J.P. 4 t.:f)OPt'H, b.\\',. l99J.
P,u11.."iu1h»gy. al. 870-88~. r,lthut~· "' ",.fH.'1irru:nu.tll,:,indur,-tt. arum ro;.,.,lpJn,,.mu!li~ Jn
~6 DUl\£\·, 1,P, ~ tOWlt. A•• l~lt TO'lit,plijm,O,I,. in .,_Jwc-p: :\ nu,:ttt~, nnd m11croJ)(tth .\ujmtlltm \ ·,,1,·/111111'· /mu,,nl, 70, oH,
IDlnm~tcd bibll1>gr,,phy, ,\liill/ltltlftlll• 1'1t/Jll(111f1111 Xr,, /Oof 111<• &J 10·.,,-. J.t. ~ bunt.l. , ,. . 1~& O,inr .1boulon 11nd m>c,n,11:tJ tJ1..•.,1h du1..· co
Common«wiilh l11sNWll·qfi'nrasllo!o~·. LontlQ,1, pp. 1-152, ioxapfa,mo,1.. in \fonl.,1n,, Jnrmu1! of llrt*.-\,m.•riNut \ 'clitWOI)' .\icvlfr:<Jl
47 DUftF\", J.V. 41 IJIH\A~. 1.r.. ,~o. D~~gtlO'-lS uf maMph,r:i•nt.iUy induced ,·l.><ud111/D11. 18-1. 661-131!,I
co,-opl.i,mo ..i~ in J'ig;.~. Amctrlt'i1,1Jo11m,II11/L-'ctt!rlna'}' R,.'$«lt,'h, Sl. ;o ,:.·,v:o..nU1..,. '-'-.., 1H.\TUtU{. c, x.. 19(13. Toxop!asm.,sism 1ht uduh. Souti1
!:?95-12:19. .·\fttam ,\!l"'lii:,11Jm,n1al, 37. 102.6-1029
r.,., 19t;1.,\11 ou1btcaloi
~· DU P~S.J-~. J.L-., 1UO,\.L"1~ R.O. s. GUlt"l.l·l.t, -;, -..,1n. ,.n., t~hl, Toxopta:,mosi-. m do&":i m ~nth Atrtc.,: ~-ven ns~
t<>•npl.\"1at<l, in rhinchil"1< In Sou1h Af:,Cll, /<>11mnl of1/11• ">u:h ,ljri.-(111 report.... /011rrtol of th1· MJmJ, A/nc.YW ,,~r,~rmnl".",' ,\l1vftcat A."Soc,mio,,, ~.
, ~ur:,w,:.• Mtrllml •.\$<0</alfor,. 38. ,~113. JJ9-J.;6.
.:9 ["",FL\....:n. t.\ ... WAU)J:l.A.~n. u .•.\....:DD£.~rs. o .. lOll:r,. T.. IIOk~'''-'· c ~ :2 ,.,,"~lN.U·1 .. 11 . t1L\r\.11Ut-1~, ~ 1r1 ,. LhTRAnnu ..... 10i4- An lfl\t'-\U)tatinn of
inuu.:,\... 1•• 1--999. Fol.'l,.d 101-s ln dafry ,to:iu: o.,i l'pfd(lndoln¢"C.U 4tUd}· kn ih~ rel~tmn.,.hrp tx·1w('(.>I\ infl"C1inn wuh rcaxo:>!a.wm gcuufli 4Hd ~ntatl
2.2 ht'tds, Small Rumtn(1m Jl-tt1·Mr,l1, :W,:J;-48 With t1lllffl~t .. 1/r.: C.'f'ntlfll .<tfri«m Jrmmul ,JJ 'dt,llnn,·. lUt ZOb-21 o.
so t:STF..IA.'\•Rl:1'0,\:1)(), I, 5- l~Xf.S. L'\. 1998. ()t,h.'(;ltOt\ of ToxtmlllH)llt gnudll ~J llU r\Clt. ~. \I,. IUl\!\~I ... tt.O, ,m..un. "-· llUll( \ , 1.1•. k,,-.i-:~.
If.tr: •• 1lr.'.f
in ti.SS\ie, or Sl1t:'ep ornur chnUtmgL·d with different dn>1..."tof 0<><;"'tt,. f.pi~i:mlc un1uplu,mu~f, ilt•.~CK'fatcd \\ilh ln!C'ctrd t;H!\, .\",•w t:u,glm:d
lmemnrio>tnl /011,,,111 for P11ras/1nloro·. 28. l4S!l-l466 Jtmr,u,,' 11/ \l1•tli'111r-. •100. m1S.-fi'~
51 f.l;,. 19;,1. roxopi~mo.si:s: Para~1tic lite c:ydt'!. pati,oJOSt'\· u.nd
.fftt'.'1.""\-0.. i -1 1mt,E.J1. r. \ ~ .• 19;d. l'oxopl,umoaic. a, ti pubUt: ht·JJtJ, h&.urd./m,r,;,,/ of
immum,to~·. In: H.\M:o.mso. D.~1. Sc LO\t., s,.1 •. ccdsl fl:'(f Cm'ridftl, :hi S.uuth .J.-fr,ctm \'t•tt'riltuf)' :\s.:«<lati.Dtt. .,-; 1~7-1.31
. . P~rl Pr~..,.
8chin1ore! Um\'ct-.it\"
:-s ruR,1Jt. t,.,-:, .19":'8. S<mwasp~ch ofthl" p.sthuscnc..-.1~ :ind cmui>~muiw
.5-2 S:~f\'Jll.. .\ .• UIJBLY. J.P., \o,\11Tlf, n.tl, .g ntt;X>;rL. p.,. 198.g. 0(K.')!~Hnduc:t"d ptttholo~- rJf 1ox<>riru..mO?;K}oumn/ tJ/rhc :,omh~\[11ca,, \ 1e.r1.•rott1')
Toxanli1Smago11d,1111rec-lions in c.ot..'- Jtmrmrf of t~t,rosuology. fS. /1.ulX'm:fo,: • .io. J....S.
""5()..755.
71.> tu.r.1., ~101.,,0.1 ,. n1,oc,, J.fl 1~87, lmnnmllhi,10chc.·1111t·n.l
A..,

.S..l G00D\\1S. '.\1 ..., •• DUUJ..iY. JP.&·Ji,\TKlS. J 1994 To~oplrumagondll d.lag1m,is nf m,npJa....mosi> in fc~~ d.11d feml mc111bran1."" o! ,h1,.'i.·p .
pe:tph"1'31 neurhis in ch1ckcn,.J,mm"I o/VNt.•rrmtr,· fJlfl>r'IOJftr ·\mc1rt,m, Jm1r1U1l of\ L'mro,ar,• Htth.'fltt'lr. 4ft. 3~a,..351
lm'f·.ffignrr'o,1. G. 382-385.
77 \'.\!\: HHll(.H \ J. 1,, \"AS ftt ,,mm,., I.D.f •• l~~' ftt\np!.i"lmth(~ in O dog.
54 lt.\.Rnn. W.J.lt ouu1 't, ,.v.• 1991. ratnl toxo1,1n.-..mo.isls in ~-nnw natl\!\" Jouma{ ofllh•.x,,ul, t\f'ricmt \'e-.r--.71nnry·.~/allo,,, :;o, 211..-11·1.

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350 Sl'Cno" n,o: Pro1ozon! diseases

;8 VA'\ Rt-XSttVR~, 1.».J . .,. ·\Wi$TO~l.. ,,.,\.,, 198-;. CQncom.h am feline -.he tnrecllon a~ n ceu~ of reprodlJ'tU\1." ?oss in .sheep fn ~OT\\'11)", Ar.Ur
infeclious peruoniu, a,,d 1oxop!:,.;mos1s ma chce1ah Uvillor1_1·.i: \ tum,:nru: Sau:dhlm•ica, Ii 412-4.?3
111bnll1$), Joumal of th,• South .\fricmJ Vt'trrinnryl1$$ac:.auon• .iS fto \\1U..1AM'- ""\1., JUL to~ ...."! Ali:\tltn. J.A. 'tA'~1UJ.l), Lt... BOUU)f' , 1.•
?05-207 BUndn~sand 1on1culll, ha, bcon de>CTibrd In canaric. ..~1•/1111
2001.
;-9 \\".\OE.lA.~O. tf., 19,fi,. TQXQpl.t>mu~Js in .. http. Th~ refali\"e lmponam::e of Diua,t~ i 3. 2.62-2U7.

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 19
Besnoitiosis
Synonyms: Elephant skin disease. olifanwelsiek1e (Afrik.l

RD BTGALKE AND L PROZESKY

lntroduction Bo,·ine besnoitiosis is a much more common disease.

than equine besnoi1iosis and has therefore been studied
Besnoitiosis. caused by the protozoan parasite Besnoicia. is a more comprC:henslvcly. Caprine besnoitiosls has recenrly
relatively common disease of caule and rarely of equids in received considerable research anention. I lowcver, it is re-
southern Africa. lnapparent, mild and severe forms of the markably similar to the bo\~ne disease. Bovine besnoitiosis
di$ease occur in can.le. The ~evere form is characterized in will thus form Lhe basis of the descriptions in ihis chapter
the acute stage by fever and anasarca. as well as orchitis in and be compared 10 the disease in goals, where appropriate.
bulls. This is rollowed by a ch ronic stage characterized by
scleroderma, alopecia and hyperkerarosis. In most cattle.
Aetiology and life cycle
however. the infection is clinically inapparent. The proto-
zoan parasite responsible for the bovine diseas1;? 8es11oiti(I Although the Hie cycle of B. besnoici In its presumed final
be-S11oiri was discovered b~· Besnoit and Robin .3 The infec- host needs further elucidation, lhe e\'idence obtained [rom
lion in equids is caused by Bemoitia De1111eui. other Bes11oilia spp. and ics morphological similarity ro
1\lthough bovine besnoitiosis has been known in south· Toxopla.sma and S(ln:ocysris spp. indicate its coccidian na-
em France since the middle of the nineteenth century. ii 1ure and justifyitsclassification in the phylum Apicomplexa,
now seems to be of more significance in Africa where its dis- family SarcOC)'Stidae and subfamilyToxoplasmatinae.2 ~
tribution is fairly widespread. It is or economic importance Based on observations made on other Besnoitia spp., 18 · 3;
in several countries. Including Israel. South Korea. Russia. and a single, as yet unconfirmed. obser,ation on B. 1,esnolti.35
Nigeria. and S0u1h Africa, where ii was lirst discO\'ered and it muse be assumed 1hat cartle serve as intermediate hosis
described i.n 1945 by Hofrneyr in the Rustenburg district in and contract besnoitiosis by the ingestion of mature isospo-
the :-.:onh-\Vest Pro\'ince.21 ran-type oocysts (c. 15 i< 13 µml shed in the faece5 of mem-
Although ca11le may die from 1he disease during the bers of lhe ca! family. which presumabl) serve as final hosis.
acute anasarca or chronic sclerodem)a stages. the mortality The sporozoices apparently enter the bovine circularion
race is low. Of more importance are the debility, me po~sible and multiply by endodyogeny in endothelial celb of blood
sterility of bulls (which is usually permanent if the animal vessels. particularly 1hose in the skin. fasciae, upper respira-
has shown frank clinical signs) and lhe downgrading of car- tory tract and restes.1 Merozoite-like endo-~oitcs {tachyzoi -
ca.~ses in abanoirs. due co trimming. or even the condemna- tes). c. 5.9 x 2.3 µm in size, reinvade adjacent or more
tion of carcasses resulting from the pres·ence of cyi.ts in the distan1 endothelial cells to produce further endozoites. This
fasciae and intermuscular connecd\'e dssues. developmental cycle is associated with the anasarca stage of
Besnoitiosis has also been recogni1.ed as being economi- the di$Case and is superseded by cyst formation.
cally important in goats in Kenya and Iran. where the dis- Cyst formation begins about one week after the initial
ease is characterized by scleroderma. alopecia and the cycle ofproliferacion. \\'hen actirnted. hypenrophic cysmzo-
prl!!>ence or large numbers of cysts i11 sites similar co those ite-comaining histiocytic cells become detectable in the
observed in cattle. 11• i:t. 3·1 same sites where the cndozoires were formed. The hisrfocylic
Clinical B. be1111e11i in£cctions in horses and donkeys. nnd cells are observed either wichin or in close association with
subclinicaJ infections ln a mule and zebra {£qm1s burC'helii) blood vessel \\'alls. The merozoite-like cystozoites [bradttOi·
ha,·e been recorded in South Africa. 9 •36• 39 equine bes11oitlO· tes) are c. 8.4 " 1.9 pm in sl7.e and multiply by cndodyogeny3;
sis was. howe\·er. first recorded in France by I lenry and :-.1as- in vacuoles of the markedly enlarged host cells with enlarged
son20 and ~ubsequemly in Sudan.2 ancl multiple nucllli (Figure 19. I). The characteristic thick-
351

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3 -.,
"- srcn,n.. n,n: Protozoru diseases

\\'alled cysts. with their conspicuous periodic-acid. Schiff- in Figures 19.3 and 19.4. Cy,1 fonnatfon b remarkably S)l1·
posith e hyalin e walls (Figures 19.t and 19.2), apparenLly C'hronou, in experimental ca~es and there is no evidence that
formed by the host cells. and eccemricallv placed ho~t-cell cys10;,oite, from disintegrating cysts give rist• to further en·
nuclei. reach a diameter of c. ~00 µm afler about ~ix weeks. do,.oice~ or cysts in du: same ammal.' Cyst fomrnrion is asso-
The ultra~tmctural morphology of tht' cy,.101.oite~ i~ depicted ciated \,itl1 the chronk ,derodermastagc of the tlbea~e.

Figure 19.1 Besnoit,11 t,esr.oi,, c·,s:. cysiozo,1.es in a t,$t a·e s,rrc:;nded

tr, a rim o1 cvtoplasm ccn;a,n1rg muu,p•e nucier c: 1ne Mst ,e,,
th,st10C'/tel follo,,ved ~\ a h;·alino :.all
~-
'

t
,·

.·

Figures 19.3 {topl and 19.4 (bottom) ~-:rastruc;ural morpno·ogy c:

Figure 19.2 Scle;oderma..•ma,ous oists in the oerm,s ..·y~to cites t'd,jJ'l01tesl

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 8(";noitlosis 353

Only the cys101oi1e phase of the life cycle of B. be1111eni goats and black wildebeest (Co1111ochacce.tg11011Jare suscep-
has been described. which is morphologically very similar to tible ro artificial infection with bovine strains of B. besnoiri.4
B. bes11oi;i. The cysts are slightly larger. reach lug a diameter s. ,e. zi;. 36• • 1 no naturally infected host~. olher th~n cattle.
or more than 600 µm. and the hyaline walls of the t-y~ts are have been found. Strains of 8tsnoi1ia isola1ed from blue
thinner than lhose of the bo\ine parasite.9 Gysroi:oiles are wildebeest and impala, although amigenically closely re-
ultrastruc:turally distinguishable from those of B. b11s11oirf.4 ' lated 10 cattle strains of B. /Jesnoiri, are relarlvelr nonpatho-
Besnoiria in b lue wildebeest (C<m11oclwe1es 1<1uri1111.~) genic and reveal viscerorropic rather lhan dermawtropic
and impala (tleµyceros mela111p11s) is morphologically and affinities in cattle and rabbits. to Consequently, it seems
immunogenically ~imilar to B. bemoitl, bur differs from it in unlikely that antelopes infected with lhese strains could
certain biological respects.25 The taxonomic relationship is serve a$ carriers for the typical cattle disease.
obviously close :ind hence the parasites of antelope origin Whereas goars were successfully infected by inoculation
are regarded as being 8. besnoiti. 10 with cystozoites of the caprine parasite, auempts to simi-
larly infect cattle and a variety of laboratory animals (rab-
bits, mice, guinea pigs, hamsters, rats or sheep) faiied. t3 . 2~
Epidemiology
On the strength of its apparent host-specificity for goats and
Besnoiria has a fairly wide host range and several Remoiria som11 uJ1rasuucrur.tl differences from B. besnoili of cat!.le.
spp. have been described. Frank disease caused by Besnofiia the parasite of goats was recently named B. caprae.'!ll, ,o
has only been recognized in caule. goats. horses and barren- Experimental and circumstantial field evidence indi-
ground (Rangifer 1ara11d11s groe11/a11dic11s) and woodland taces that bo\'ine besnoitiosis can be transmitted mechani-
(R<mgifer 1artmd11s caribou) caribou. 15· u bur clinically cally from chronically infected cattle 10 susceptible ones by
inapparent infections have been detected in blue \\1lde- blood-~ucking insects such as 1abanids.4• 43 Although this
beest, impala, zebra, donkeys. a mu le and a warthog method of transmission probably plays some part in the epi-
(Phacodzoer11s aerhiopfcus).23 demiology 01 the disease. it does not seem to play a major
In Soulh Africa, bovine besnoitiosls Is of economic im- role as only clinically inapparent cases \,~th small numbers
portance mainly in the 8ush\·eld and Lowveid of the Lim- of cysts are produced in cattle in this way. 4 The severe form
popo. :>:orch \Vest, :,.,tpumalanga and KwaZulu,Natal of the disease can consistenrly be produced e,q,erimemally
provinces. A few cases have been recorded in lhe ~orthem in caule immunosuppressed with corticosteroids prior to
Cape PrO\ince and western Free State. It has also been re- artificial infec1ion with cyscozoites or endozoites.17
corded in S\\'aZiland, Zimbabwe. Botswana, Namibia, An- It seems logical 1hat a carni\'orous host muse exist for B.
gola. 7.aire. all the East African countries, and Cameroon and /Jemoiri. The wild cat (Fe/is lybica). incriminated as a host in
Nigeria In West Africa.6 Outside Africa It has bc<:n reported Russia,35 has an almost colmtT)•\\ide distribution in South
in Israel. south-western Russia, South Korea. Portugal, Africa. Domestic cats, which could be fed raw, infected meat
France and Venezuela.6 by humans, do not. however. occur on all infected farms.
The disease is more prC\'lllent in less temperate. and Moreover, ir is difficult to understand how a small "ild cat.
even subtropical climates. In South i\frica, the majority of such as F. lybica, can become infected on such farms where
new cases occur during the warmer. moister monihs of the caHle are the only known potential source of infection for
year. This is apparently also the case in lsraei.43 them. It therefore ~eems probable that a hitherto unknown
All breeds of cattle and both sexes seem 10 be susceptible, cycle or transmission in\'olving a carnivorous host occur. in
although infection in cah·es under six months of age is rare. A nature, although it has not as yet been proven ei-peri.men-
survey on a farm where the disease was pre\'alent revealed tally by feeding muscles and connective tissues containing
that lhe highes1 incidence was in animals 18 momhs co 6 numerous cysts 10 a variety of carnivorous animals.';
years old. Of 5 018 cattle sun·eyed in an e ndemic ru:ea by ex- Aaempts to transmit B. cnprae co domestic cats have also
amination of the scleral conjunctiva for cysts. 427 (8.5 per produced negative results.t7
cent) were infected, of which ; 4 ( l 7.3 per cent of infected ani- Capr!ne besnoitiosis is widespread in northern Kenya
mals and 1.5 per cem of the tor.al number examined) showed compared co bo~ine besnoitiosis. which appears to be lim-
clinically detectable signs of chronic infection.4 Thus. com- ited in its distribution in that country. 28• 33 !n lran, ourbreaks
paratively few animals develop clinical signs oflnfecrion. The of the disease in goats have been limited to the south.3~
passive rransfer of antibodies from chronically infected cows Equine besnoiliosis ls a rare. ,·el')' sporadic disease and
to thtir ofispring" 3 may be an important factor In the above- its epidemiology is obscure.
mentioned ei:,idemiological features of the disease.
There is ex-perimemal and circumsta ntial field e\'idence
Pathogenesis
that chronically affected canle barbouring large numbers oi
cystS serve as sources of infec1ion of bovine besnoic!osis.4 13 The endozoites associated with the acute anasarca stage of
Although a variety of labonnory animals. such as rabbits. bovine besnoitiosis proliferate in the cytoplasm of endo-
gerbils (Merio11es tristrami shmvi1) and hamsters. and sheep. thelial cells. causing vasculitis. and sometimes thrombosis,

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354 "'"no, '"'" PrmozoaJ diseases

,.

Figure 19.!i Chnica· case oi 1ne St!aroderma stage cf oesm: 1,os,s Note pl101ophob,a and sa•,va;ion

-tdl!"
•

Figure 19.6 Marked sc!erodeflT'a cl hmdquartecs ano l!!9s Figure 19.7 Stlerooerma of dorsu:r, of nose

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 Besnoltiosls 355

tha1 lev~l for a week or more. Pyrexia is accompanied by

progressive inappetence. culminating in complete anor·
exia in severe cases with cqnsequem loss of weight. Af·
fected animals are generally listless and prefer 10 lie do\\'n;
are photophobic (Figure 19.5) and hence seek shade; and
have an increased respiratory rate. They walk wirh a slow
gait if forced 10 move.
Hyperaemia of the muZ1.le. periorbital skin and scrotum
is noticeable in light-skinned animals a few da}'S after the
rise In tcmp('raturc. Anast1rca nlso usually appears at this
time and may vary from nothing more tha11 a slight swelling
of the face. or thickening of skin folds or the neck, back or
chest regions. 10 subcutaneous oedema over the entire body
in more severe cases. The testes are swollen and senslti\'e to
palpation. F.arly anasarca often goes unnoticed and 11.S pres-
ence is only revealed when oedema fluid accumulates along
the vemral pans of the body such as the brisket, sternum.
prepuce and leg$. The anasarca ~tage of1en goes unnoticed
in the field. In cases in which il does occur. there may be a
clis1inc1 break berween 1.his stage and the ne~,. wich disap·
pcarance ofrhe above-mentioned ~!gns by the third week of
the reaction. or the two stages may be sequential. Animal~
sometimes succumb 10 the disease in theanasurcastage.
Figure 19.8 Scletoderma close·Ull o:severe dern>.atttts CIIJStcsa The most characteristic clinical sign of the scleroderma
stage in severeli•nffocted canle is progressive thickening. hard-
ening and promlr1em folding and puckering of the skin. which
particularly in smaller veins and capillaries in the skin. sub· may be ,,idespread or more localized (Figu res L9.5 to L9.8J.
culis, the mucosa of the nasal cavity. laiynx. and u:achea, The skin of the legs is often hard and the limbs may be mark-
and rhe restes. 1These lesions. together with a possible toxic ccUy thickened (Figure I 9.6). Jn such cases. movemem is slow
effect of the parasite. cause an increased vascular perme- at}d restricted. upparemly due to pain. These feaLures usuallr
ability. resulting in oedema and some1imes small haemor- appear three 10 four weeks after the initial rise in temperature.
rhages and foci of necrosis in the afore-mentioned sites. Progressive loss of hair. accompanied b)' dermatitis with
The circulatory disturbances are probably also respon- hyperkcrarosis and dermatitis crus1osa. also occurs (figure
sible for the necrosis of 1.he epidermis, deeper layers of the 19.8). Parts of the epidermis may be shed in patches, leaving
skin and of mucous membranes of the upper respiratory tract greyish, hairless. seborrhoeic areas. Sit-fasts are commonly
in the scleroderma stage of the disease. Very large numbers of seen over bony prominences as well as on the ventral por-
rapidly growing. thick-walled cysts, especiallr in the dermal tion or the scrotum. Often a sero-sanguineous exudate
papillae. fibrosis and granulomatous reacrions around C)'SlS, oozes from the skin. This dries to fom, large scabs which
oedema of the skit) as well as hyperkeratosis and acamhosis sometimes become secondarily infened. Deep, raw fissure ·.
probably account for 1he typical sclerodenna. 1 which usually become maggot-infested, sometime$ de\·elop
bcrween foldsoflheskin in sit.es such as lhe breech and over
bony prominences. The superficial !~·mph nodes are in\'ari·
Clinical signs and pathology
ably swolle1l. A mucopurulem nasal discharge, which forms
Two sequential s1.uges of 1.he clin.ical disease are recogn.ized crusts often cloggi1tg 1ho nostrils and causing stridor. may
in cattle. namely an acute anasarca stage, which is asso- occur in severe cases. These signs are usually accompanied
ciated with proliferation of endozoiles, and a chronic sclero- by an undulant low-level fever {which persists for several
derma stage (Figures 19.5 to 19.8), which is associated with weeks). poor appetite and emaciation. :vlore pronounced
cyst formaLion. fever and anorexia occur in more severe cases, some of
AR incubalion period or four days was recorded in cattle which may be fatal.
aJter oral infccrion with sporulated oocysLs offcline origin,l• Bulls invariably become asperma1ogenic, dut> to the
whereas Bigalke.i reported a mean inci.1bation period oi 13 presence or a usually persistent and severe orchitis. fol·
days after experimentally induced mechanical transmission lowed by uni- or bilateral tes1icuiararrophy and induration
of the infection by biting flies. in the chronic disease. Acute. subacute and chronic orchi·
Fever is the first clinical sign and early-morning rectal tis accompanied by pronounced spem1 changes are abo
temperatures may reach~ to41 °Cor higher and remain at observed in goat~. 31

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356 ,~cm,~ nio: Protozolll dl~cascs

,.•
• -·r~.J;,:...,..,..,.,
..

Figure 19.9 Numer®S cysts

mlhe mU(osa of the Mbi~ates

Cysts become visible 10 the naked eye in the scleral con-

junctiva and nasal mucosa (Figures 19.9 and 19. LO) about
six 10 se\'en weeks after 1he initial rise in temperature.
Most animals smvive. but convalescence is slow and scle-
roderma and alopecla may be pem1anem. Gradual improve-
ment in the appearance oft he animals seems to ~ associated
\\ith the degeneration of cysts and a subsequent decrease in
their numbers.
Surveys conducted by scrutinizing the sclentl conjun<:-
tfra4 and by serological tes1s 19· 22• 26 indicate that most ani-
mals which become infocted in the field co111mc1 a clinically
in apparent form of the disease.
Only the scleroderma stage has been recognized in
horses. The disease is very S-imilar to that observed in cactle.
but recovery from the skin lesions is usually more rapid in
horses, and 0U1growing hooves. apparently due 10 laminitis,
are sometimes obserl'ed.9
Only the chronic scleroderma stage has been obs<"rved in
narural cases in goats, bul ast1bcU1aneous oedema has been
recorded ii) artificially infecled ani mals.29
The pachologyofbcl\ine besnoitiosis bas been described
in a few publications. 1 2 1. 25 · 36· 33
Ar necropsy. apart from L11edinically discernible lesions al-
ready described for the anasarca stage, ,,~despread petechiac
c111d ecchymoses in the subcutls. oedema of the lymph
nodes and testicular parenchyma. and a catarrhPJ to mucopu-
rule111 rhinitis and conjunctMtis are often noticeable. ~ lild w
prominent oedema of the lungs occurs less common!~··
In che chronic stage, cysts are ,~sible macroscopically as
small, whitish granules the size of a sugar granule particu- figure 19.10 ·~um~,ous vlhite cys:s c!ear:, i;s,b!e It! th!! mucosa o;
larly in lhe muscles. imermuscufar fasciae. 1endon5 and 1urt11rates

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 Besnoiriosis 357

bution of cyst> and the pathology of lhe disease in equlds are

similar 10 those in cattle.

Diagnosis
Endozoites of B. l)e.snoiti that are free in the blood or in mono-
cytes may be demonstmted in peripheral blood smears of
caule for several days at the height of the febrile stage. but
they arc scarce and difficult to find (Figure 19.11}. Blood
smears made from the ear or tail of chronic cases often
contain cystozoitc~ originating Crom punctured cyi,ts. The
coccldian-rype endozoires and cys1ozoitcs are merozoi1e-like
in appearance. and are lhereforn not diagm>stic for besnoi·
tiosls.
The clinical diagnosiS is easily confirmed in sclerodcrma
cases by demonsrration in skin sections of large numbers of
the typical cy~ts in the dermis (Figures l 9. I and 19.2).
• Cysts of B. bcsnoiri are ,~sible 10 the naked eye in the
scleml conjunctivae of cattle from about si.x co seven weeks
Figure 19.11 1'1!r1p1Jeral blood smear coma,ning emlozoites after infection. Cysts are plentiful at this site in rypical sclero-
derma cases. They may even be detected in small numbers in
the sclernl conjunctivae in catcle which show no other i!vi·
tendon sheaths of the legs. synovial sheaths. perios1eum of dence ofinfection/ 8 Their Identity may be confirmed by ex-
the mern1arsal and metacarpal bones, testicular paren- amin ing smears prepared from single cysts isolated from a
ch~nia, mucosa of the nasal turbinates, phary11:1: and upper tissue specimen with the aid of a dissection microscope. or by
respiratory tract, bulbar conjuncti\·a, endocardium. cardiac microscopical examination of tissue sections.
vaJ,·es and intima of the more peripherally located larger At abattoirs. or at necropsy of can le with typical sclero-
veins of the legs. head and neck. 1· ~;. ;\11 derma. large numbers of cysis are visible at the predilection
:Vlicroscopically. in the 11cu1e stage. varying numbers oi en- sirns.
do1.oites are found predominantly in endothelial cells of capil- The indirect immuno11uoresccncc test and the enzyme-
laries and small veins. and less frequently in what appears to be linked lmmunosorbenc assay are useful aids in studies on
mesenchymal cells in the imima of veins and small arteries. the epidemiology of the cattle dbease 1s. n . 26• .a. • 3 and on
The vessels in the dem1is and subcutis. nasal cavity. larynx and the efficacy of ,·accinacion. 16
trachea and res1icuJar parenchyma are most severely affected. l\lost of what applies rn bo,ine besnoitiosis ls also true
Parasitized blood vessels show necrosis of endothelial cells and for the caprine disease. However, no serological test has
fibrinoid degeneration and necrosl$ or their walls with result- been developed for 1he latter.
am oedema. haemorrhage. thrombosis. and necrosis in sur·
rounding tissues. :'-lacrophages and a few lymphocyte~.
Differential diagnosis
plasma cells and eosinophils occur pimvascularly and in the
walls of aifecred vessels. 1 Particularly if no subcutaneous oedema is present, the acute
On microscopic examination of the skin of chronically febrile stage of thE.' bo,1ne besnoi1iosls can be confused with
affected animals. nw11crous cysts are discernible panicu- other acute febrile diseases, such as heartwater and redwa-
larly in the dem1is and subcutis but also In the intermuscu- ter. which also occur in the C)Jldemic areas. Acute photosen-
lar fasclae (figure 19.2). Cysts in blood vessel~ are mainly sith~ty should be differcmiated from 1he anasarca s1age. ln
located in the inti ma, but may be present in all pans of the cases of bovine be:.noitiosis where there is oedema of the
wall. Hyperplasia of the endothelial cells and o<:casionally legs and \'cmral pans of the body. cardiac failure and acute
sclerosis of the intima may be e,~dent in the immediate lumpy skin disease should be considered.
,1cinicy of the cysts. 1 The skin lesions present in the scierodcrma stage of bo·
:Vlost of the cysts are enclosed by a thin collagenous cap· ,~ne besnoitlosis are sometimes ,·ery similar clinically 10
sule. When the organisms die, they elicil a granulomatous rhose seen In dermat0phllosis (Senkobo disease). sarcopric
reactioo.25· 311 mange. lousiness. subacUle to chronic swea1ing sickness.
The clinical signs a11d pathology of the disease in goats lumpy skin disease. photo~ensitMty. and poisoning by mer-
are remarkably similar to those in canle. •i. 11• i ~. 34 cury. chlorinated naph chalenes and hairy vetch (Vicia spp.J.
Repons on the lesions of besnolliosis in horses arc lim· Severe dennatophytosis, onchocercosi&. and mange can
ited and deal only with chronic besnoitiosis.2 • J 9 The distri· be difficult to distinguish clinically from equine besnoitiosis.

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358 1rL,10:1 me>: l'rorozoal diseases

Lousiness nnd mange should also be considered when wildebeest strain of B. besnoiti grown In cell culture. The
dealing \\ritb skin conditions in goa1s. vacci11e is Issued in the frozen form and is recommended for
use in weaner~ and older animals.:\ :,inglc dose of ,,accinc
pro1ec1s canle from 1he clinical form of the disease for up 10
Control
four years but doe~ 1101 entirely prevent suhdi nical infection
);o ~ui1able drug b available for 1he trea1men1 of natural Ca$!!$ from taking place. The indirect immunonuorescent test has.
of besnoi1iosis. alihough oxywcracycline has been found 10 howe\'er. revealed that level~ of amibod}' to a homologous
have a dela)'i ng elTect on the course of artificial infection in anugen start 10 d ecline at 180 days afte r vaccinaiion. 16
rabbits, if the drug is administered concurrent!} with the Annual revaccination for 1wo or three yea rs is therefore
organism:0 Long-acting OX}'leuac.-yclbte at 200 mgtkg gave advocated.
complete protection to e>.-perimentally infected gerbils. ~2 The S}'Stematic annual elimination or all chronically in·
Te1racycllnes. wound dressings. control of secondary rected caule. detectable by examination of the scleral con·
infections and Infestations. and P.rovision of shade and junctiva. in a closed herd on a farm in an endemic area over
nutritious food and water ad /ibimm. are advocaced as a period or several years. appeared to resull in a decrease of
supportive treatment. ne\\' infecrions in the herd.4 Until the epidemiology of the
A live vaccine against bovine besnoiriosis has been devel· disease is berter understood. it will not be possible ro deter·
oped by the Ondersrepoon Ve1erinarr lnstilmc.; Each dos~ mine wich certailll)' whecher prevema1.ive measures based
of vaccine issued contains c. I x 107 organisms or a blue on 1hese acllons will comrol the disease.

References
a.,sso~·. r.A .. MCCULLY. R1M. & DIC..\1.££.. k.O.. l!}fO. Obstn'3UOn$ 0:1 Ute U CHU:\IA••\Jt, & tObh\XL.\~. F.. 19':"'9, Bestloltto~i<. In Wtld .and domesttc
p3Lhogent:sis ol bo\in~ and nmt"Jopc., .>Sr4ins of Ewsnoitta IX's11nlzf ~oats in Iran Comrll \t'IL'rf1111n.11n. 69. ng...168.
(~larorN. 1912) !Mccdo:\ In c>rde •nd r.1bbl"· (),uJcrs1,·poortJmm111/ of
li CHO()UtTTI V,t fHIOUC!II IOX, I .• MIUJ fC,, J.:,C.. cmrns. n.c. "C'.OU61!\"£U.\,
h:lf'ritUlf)' Rcn,arc/J, 3":.105-1.:!.6.
1.r. .• t•l6S, Besnohi<~ls in b:irrcn,j:rouad canbou in nonhtrn Canada.
~ au:."-:.-t.TI s.c~, .• 19.33, GloMtlwm Ulf&euo1\S in ch.: Sudan /oumat of ('.4muJw11 \ ttt.•rim,ry Jot111t(J/, 8. 28?-28f
Comp,m1fi1,· Jlnrllofoe:,• nmt Them~111/c,, 46. 1- lS
lti ot. W;\AL D.T l938, Vc,tNinruy Rc.snf<'h tnstmm.•, Ondersu~poon.
3 HE$.'110n, c... • fe()UI~. ,·•• l!fl!.. "ia~o'>poridll1~ cut'1n~ chl"7' u.n~ \.':l<.'ht.• Unpubll•he<I dom.
Rt~'lh! \,.t'lt'fUmlN. rort{l)Un:, 3i1 6.J!Hi63.
lj n1t<c;1~r., l )lf\·o(lK':\, A,0., MAlU..fllli..t\, LR.. R1\Ul n. \_, rJP"iXO, 1 •• nr
..,, mc.;.,u.,:, n,n.. J.9&l, .X<.o\,. cont1.'pt! ()0 the tpl<km1o1uglt3l !..•.ttuc"°' q,f a.- PO'TC.11 l"l!ft, f :t,. 1968 &.rsu'1llfn iK~nt>i ri: Sludlc~ on the
Wl\.\t. D.1'.
bo\.inc: bc.isnoit\n~k u~ determined hy lnbbrtaim')' .ind field dcfinmn, hmn ;uu1 cx11t'rimc.·m.1.I m!i:cuon.."' In t·nul,• Purasl10/ogy
irnosc!saclon<.. Omlt•1'$1~JXIOn J(J11mfll of\le1<ri11ary R<1A't>rd1 35. 3-1311. R,~,'wutt'IL 75, 1 l•l-1 t7.
.5 n,GAt.Kt:. tu>.• 1972. \ C'h:rtnary Re~~areh lnstiturc. Ondrr.su:poon. 111 1mun\ r.r.• 1q77. fo:m1,lnsnt<1, Hnmmoudla. Ewsuofttn. Snr<OQ'Sris¥nd
Unpublhhed dactt. other tli$ue cysl•forming rocddfa of ,nan und onim-.tl~. In: t,,.RHf.tt, 11•..
6 WGAl)Ce. 11.0.. 198~ Be.11oioos~ ~nd glob1dlo~ls. l11: ncsnc ,,. • Mcrsn·IUi, 100.), l'ora.wie Prcmr.:on, Vol. Ill Xew York: Academic l're:ss.
1., ltdsl. Oi.<eO.«'$ ofr.tmlr 111 rl,e Troprrs. The Hagur. Boscon, I one<>n:
19 (AH o,lA,, ,1,..& f"IMXrt1 6., 1983 ~i:roJogicnJ .siudle-.s oo l>ovrne
~larlinu.< '11jh!l!T Publf<lwrs
bcsnmtiOSI> in lsr.icl. l'ro1m·11/ ,\11fmal l{~a/tlt am/ Prod11r1iq11, IS. 32-38.
i Nm\tJ..t:. n.u•• U.\:,SQS-. P..A .. \1(;<:UL1Y. K,M .. 80$MA"i, P,fl, ,-.:sctoEM,\~. J,11 ..
20 nr.~u,. "·-. ,IA:,...;os-, r.•• 1nu. Cuc-d by 1'Q:..,, J,\\ •• 1!~. Studlc1.on bo\1nt•
197,t. s,udicc; in r.otdc on thcdC1,c!opmcn1 ofa livc,11ccinc agnin.s1 bc~nniclo~,, ,,1ch special ttfrrcncr co 1he•c1lolo1,,y. 011ders1r1wm
bm.1nc heliincncu>~>.Jouma/ of1lu.· Stn1tl, A/r1C(l11 \.'tu:r11u1ry·Asi«iatlo11,
Jrmmtil IJJ \:tli•rowry Hb.!onh. l8. 255...350.
4E. 207-209.
;z1 HQr ,in"A', t.l.11., 19 1s. lflobidfo<h. m C'olttl<". Jor,mnf nftltt Soutlt ,\frium
8 PIGAL t,;:t. 11,n, ,.,_ :,.Au1>1' T,\\ The crmgn01tf:c \.lluc of c-y~1.$ i.n th~
1g&2:
C'nt•rlnury• .\frdltnf -'i$$()fl11tion, 16, 102-109,
M:kllll roniuncclw in bo,inc b<'>11Dhlo,1, /ntmtr1l1J/tltr&Jm/1 Ajnttm
\·,~1,•rfoaq .\,lft/!tal JUjDC111tio11. 33. 21-27~ 22. 1A.~JTKJU;J:, ):. UC VOS, .\,I, k IIIG,\I ~-i RJ>,. 1984. SL-rodl:ignr,~lsnfbovine
g a1G.,U,'1:., n.o. & sr.1n.rm, .\.P .. :~6:. \let~rinat) Rtst.':atch tn..-;wutt.
btl~noitiosri. h>· ELISA a1)d fn1munofluore<-cent:t' 1c~h Omlt•r:.rtputJrl
Jnumflf o/Vt11•ri11a1y R,•st.!dt<''1 51. 21ca...:?43.
Ond!-r:,topoort. UnpubU<hed dnr3.
;o 8J{;,\U.."'£. H...O., VA,\ ~1u:~. 1.w.. au:s()!I., P.A. A ,1(':CUU.\'. k. ,, •. 19'6; a, i1;.u1• ,u & OA<..."-0~, r.A .• 1~n. Bffnout~fs in a w:irthog tPhncoch<>ft'us
Srudl{•s. on tht• re}:ntoiishlp hi:nwl·n '81'j./Willn or hluc \\ildcbt-..·~t and 11<•thtopi<usCU\'l,·r 1822 ,. /Ottf1ta/ ofthe So/Ith ,1/rfr,m \ cn•rlnary
ttnf>:i.l.n. J.lld &'StJOi(/tl bdtJtJlti o( C.lUft. O,ult(.i.(t!l)QOr/ /OtJrtt(I/ "' ,\.<>villltlon. 4-l, 287
va,~ri,uuy R.ekarcJ1. :w. 7-28. .24 uv~:-.'1·, , n 198;,. t/~1,,mfir;.• l'totc,:t>0fo~·. •.\me\. Iowa Sum.• Unh·'-'riit}'
n aw.\..'""Oi\MOI, o .• 196~". A preliminary repon on the finding.oi lk1.noitlo PrC'SS.

b,.,,,u,;u in goat ->kin, .i.lTl!'t"t\.."<J wilh dirnpt~ in Ken~.i. B:,llt'tf11 ofE11f:(J(J((c 2; ,,rc.m.1,." "·· M:,...,os. ,,.,•• '"~ 'J0.1',Mh, 1w." nu;.,1..u:, 1t.o.• 1966.
D1u,.-t>f:\frlr1t, 15, 263-~71 OD>e1\'0UIOM on 8rJ,iul1tn t:,')h tn th\• eardiO'r'afvC'Ul,u :.yitcm of ~om~
1:: llWA:'\(0....'1fjJ, o .. CARLI:.:,•..\.n lit \\A~nlRA1 rN.. 1989. \n cpidl•rnlc; of wUd-n.n;~IQp(.'1 und dom(.'-<tlc cauh.•. Ondtuh•f>ontt Jtwtt'tttl t>/Vt>t~nnof)
rk.s.nomo~,s in goa.1~ fn kcnya Th~ V~tcmuor)' R«ortl. !2.5, 461 Rr.J.ivm·h.13.145-?;5,
lJ ftW,\Mi.\\l('U. o. >.:AMG-Aa. 't .,_. \\',\\llUlA. ,.r. .• 1996. Be-inohlo!'i:ti In 1:C):il'I: .!.ti ,t.tr:.t,., :,.t .• 19n.. StroJoiuc.11,:unM1 1'1( Besnoll:n hf-sm,ltl (.\1:1mt11I. 191.?J.
and sh~; ex~rim<nial and 11,ld obs•l'\·ntion<. 7.unl>nbt<Y 1·,1erh1nry lnfection in tc.rtu,•I by imn1unnfluort1'(cnc.c-. A•mmlblm1 f11t
Jaurr:aJ. 21. l-i. \'11,r11111m1Nl/.:J11 II, 19, 391-J•r..

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 Besnoirios1s 359

~1 -xo·.\."-<t.a.. C.1, 41: li..\$1GiUE, ::i,. 1 19',;. Capunl! l,t-6!'10111(>:tC.: :-.tudl.es an lf> r.w .xgl)Q. s,udicj on b<>,1nc- besnottic,s.1\ i,i,h .i,ptc;i3J rcre1cn«: u>
11t)Ui.
L•xpenmcnuil inutmiccrau.-. hosts and U'le role of ,he domC$11C cai m rht ao1iot~·- Omtrts,(fpot'}tt Journal o; \ ·~:cn,mrr lli·wnrr:h, 2&5-3.56 za.
tran~mt~u,n \'rrtrlmuy Pom..<ftotog;-. 52. 201...210,
li ""'"'u ,, .19;a \"tri:Iolchcnd,· t>amcllunRdijr EntwlrUungibtol1>g1e
:a "f\(.t. o.,., ~DAMTIII, C"'., 'l.t, r. ,. • \JU'.\"(;,\, L ... J99'J:. An tpld1Jmic
""'"('""· di:r Ci.u1un(.:t•n Sarron·,dt. r-r~ukt•lw. h;,1-,JQm. Cy.:tolu,,,1J<Jrn1
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 20
Sarcocystosis

M B MARKUS, J J VAN DER LUGT AND J P DUBEY

Introduction Dogs, coyotes, foxes and possibly wolves. jackals. and

raccoons are the definitive hosts of S. cruzi, whereas bison
Stucocysris is ubiquitous worldwide. The proco;,;oon com- (Bison bison) and cattle are the intermediate hosls.:?9 The
monly and characteristically causes a chronic. subclinical definitive host becomes infected by ingesting muscular or
infection in the cardiac and/or skeletal muscles ofli\·es1ock, neural tissue containing mature sarcocysts. Bradyzoites
bu1. on rare occasions. Sarcccystis dc,es give rise ro dissemi- (synon~mous \\~ch cystozo!res or cys102oic merozoices811 )
nated acute disease. encephalitis (which can occur in the iiberated from the·sarcocysr by digestion in the stomach and
absence of disseminated infection). abortion or premarure gut, penetrare the mucosa of the small intestine and trans-
birth and. possibly, eosinophilic myosiris. In southern Af- form into male (micro-) and female (macro-} gamoms (ga-
rica. cases of acute (pre-muscular) sarcocysrosis have so far metocytes}. Within SL'( hours of ingestion of infected tissue.
been seen in goats and canle 112 (Figures 20.J to 20.3}. gamoms are found wiU1in a parasitophorous vacuole (PV) In
goble1 ceUs near the tips of,~lli. '.l•lacrogamonrs are ovoid co
round, 10 to 20 µm in diameter, and contain a single
Aetiology and Jife cycle
nucleus. :.Vlicrogamoms are ovoid to elongated and contain
The coccidiaJ narure of Sarcoeys1/s firs1 became apparent one or more nuclei. The microgamom's nucleus di\ides
approximately30 years ago as a result ofwork by Fayerio the and, as the microgamonr marurQ~. the daughter nuclei mi-
USA and byHeydorn and Rommel in Gennany. The classifi- grale cowards rhe periphery of the gamont. :V!ature microga-
cation, history and nomenclamre of Sarcocysris have been moms of S. cruzimeasureabout 7 x 5 µmand comain310 ll
discussed clsewhere.3 1. 40• 79· 115• 135· 13;The organism is cat- slender gametes. TI1e microgametes are about 4 x 0.5 µmin
egorized as follows: phylum Apicomple.xa (Sporozoa); class size and have a compact nucleus and two flagella. '.\licroga-
Sporozoasida; subclass Coccidiasina; order Eucoccidiorida; metes liberated from the microgamont active!~· move to the
suborde. Eimeriorina; family Sarcocystidae: subfamily Sar- periphery of the ma.crogamont. After fenilization. a wall de·
cocystinae; genus S11rcocys1is. This generic name is derived velops around the zygote and the oocyst is formed. The en·
from the cyst stage (Figure 20.4) iu muscle ('sarco' = muscle, tire process of game1ogony and fertilization can be
from the Greek word for flesh). Sarcocysrisspp. have the or· completed within 24 hours and gamoms and ooC)>sts may be
ganelles 1har are characteristic of the phylum Apicomplexa found at the same lime. The location ofgametogony and the
such as apical rings (also called conoidal or preconoidal rype of cell parasitized, vary with the species of Sarcocystis
rings), polar rings, conoid. pellicle, subpellkular microtu- and stage of gametogenesis. Sarcocystis differs from otl1er
buies, micropores and micronemes. coccidia in that no asexual mulriplfcarion takes place in rhe
Typically. multiple species of Sarcccystis are found in do- intestine prior to sexual reproduction. ,
mestic animals {Table 20.1). A two-host Ufe cycle is involved In contrast to od1er coccid.ia. Sarcocysri,S oocrs1~ sporu-
(Figure 20.5). The intermediate hosrs - water buffalo late in the lamina propria of the gut. The inner mass of the
(Buba/us buhalis), cattle, goats. pigs, sheep. or equine hosts OOC)'St {sporont) dMdes imo rwosporocysts. Four sporozoi-
- harbour the asexual stages of the parasite, whereas a tes are fom,ed in each sporocyst. Sponilated oocysts arc
sexual cycle takes place in the definitive hosts. such as cats, generally colourless and thin -walled (Figure 20.6). Occa-
dogs or humans (Table 20. L). 111e period Of time required sionally. unspornlated or partially sporulated oocysts are
for completion of the life cycle varies considerably. depend- shed in the faeces. The prepatent and patent periods ,:ary
ing upon the species of Sarcocystis:10 In the following de- (the latter is usually longl, but oocysts of most Stm:ocys1/s
scription, the life cycle of S. cru:I will serve as an example.29 species first appear in faeces between 7 and 14 day~ afler
360

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 Snrcocystosis 36 I

Ingestion of sarcocysts. Curiously. both sporulatlng and

fully SJ)Orulated oocysts of SarcOC')'stit have been seen in
skeletal muscles of baboons tn South Africa. which are fre-
quently both intermediate and definitive hosts in terms of
the me cycle. 8·1
The imermediate host becomes infected by ingesting
sporo~sts. Sporozoites excyst from sporocysis in the small
intestine. The rate of the sporozoire from the tlme of inges·
lion of the sporocyst umil inirial development in arteries in
t.he mesemeric lymph nodes. is not known. First-generation
schizogony (merogony) begins in endothelial cells of arter-
ies as early as seven days post-inoculation~9 {DP!} and may
be completed as early as 15 OP!. Second-generation
schizoms (1neroms) have been seen in vascular endothe-
Figure 20.1a Acute san:0<:ysws\s in a goa,. natural llliett,oi ScJ7.zont rn
lium from 19 to o.16 DP!, predominantly in capillaries but also
a vascular endothelial cell ,n the brain
in small arteries ,irtually throughout the body. These
schizoms seem to be most numerous in the glomeruli of the
kidney (Figure 20.7). Cell division in schizonts is by en-
dopolygeny.10 which is a specialized form of schizogony.
The nucletLS becomes lobulated and divides into a number
()f daughter nuclei (up to 37). Merozoites (tacbyzoites) fomi
at rhe periphery. The s hape and siie of schizoms vary con-
siderably. Both first- and second -generation schizoms de·
vclop within the host cell's cytoplasm and are not
surcounded by a PV. Merozoites of Sarcocysti.s have the same
organelles as do Toxoplasmt1 gondii merozoires (tach}-~Oi·
1es). except that there are no rhoptries40 (Figure 20.8).
Merozoiles are fou nd in peripheral blood from 24 to 46 DP!,
coincident \,irh rhe maturation of second -generation
schil.onrs.r. Merozoites in blood are extracellular or locared
within unidentified mononuclear cells. lnrracellular mero-
zoitl?li contain one or two nuclei and some divide into two.
Figure 20.1b Natural lnfect1cn glr,merulus in the Kidnev oi agoa, apparent!} by endodyogeny. The number of generations of
coma•ning schizonts in vascular arv.:otheiial cells. (nssve oy cc.nesy oi schizogony and che type of host cell in which it occurs, var-
Or J A Neser. Onderstepoo<t Veterir.a,y Institute, South Afr:ca1 has. depending on the species of Snrcocysris concerned. Xe,·.
eriheless. trends are apparent. All species of Stircocystis of
large domestic animals form first- and second-generation
schizonts in ,·ascula r endothelium. whereas only a single
precystlc generation of sci1izogony has been found in Sarco-
cy:s1is species of small mammals (mice and deer mice). and
this is usually in heparocytes:10 Merozoites that ace liberated
follo\,ing the final cycle of schizogony initiate developmem
of sarcocys1s in muscle. Cysts of some species of Sarcoc}'sris
a lso occur in neural tissue. whereas those of other species
a re possibly noi formed Lhcre. 3· • 0 • 67 The incracellular
merozite surrounded by a PV becomes circular to ovoid.
forming a metrocyte which undergoes repeated divisions
(see below). Evcntuall~ . the sarcocyst is filled with brad}rzoi-
tes. which are lhe infective stage for the p'redaror. Sarc:ocysts
generally become infecrious at about 75 DP! but in this there
is considerable variation among species of Sarcocysris. Sar-
cocysts may dlsappear from a host with passin1i of time.:w
Figure 20,2 Acute bovine salcocystosis natural infec:ion Vas:~la•
sthflon:s in enrfoihelial cells and some apparently lying free in :l'e tum1m Neither schizonts nor immature sarcocysts comaining only
OJ lite vessel (Tissue bycourtesvof Or J Kitching, Reg,onal Ve1e11nar,· 1nerrocy1es give rise to intestinal infection in the deflnirlve
laborator,. Allerton. Pie1errnam2burg. South Africa) hosr. There is no lactogenous transmission.

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362 SK110;,111,0: Protozoa.I di~eases

,,,,,. -• - .-

Figure 20.3 Natural infec11on

eklctron micrlllJraph ol a Satcot)'St/S
sthilont in avasC1Jlar endotnelial
cell in the kidney of a goa1. ~ 12 000
Note the absence of rhol)lnes and ot
a paras11ophorous vacuole. nss.ue
by courtesy of Dr JA Neser,
Onc!emepoort Veterrna;y IJ1S11tute.
South Afr1cal

Sarcocysts vary in size and shape. depending on the spe- red in the periodic acid-Schiff reaction. Even matur.e sarcO·
cies of the parasite. Some always remain microscopic, cysts may rnntaln some peripherally arranged metrOC}'tes.
whereas others become maeroscopic. Sarcocysts usually The uluasm1cture of the mamre sarcocyst wall is of taxo-
rake a long rime to de,·elop to a macroscopic size in the mus- nomic value. bur the line stnu:ture thereof may vary l\ith the
culature of horses 100 and other livestock. and so, for in- age of the C)'!it. Sarcocyst development begins when a mero-
stance. il rends to take more than a year for S. giga111ea :mite emcr-s a muscle or nerve cell, a.~ e.~lained above. The
(Figure 20.4) and S. medU$iformis to become macroscopi- rnerozoite resides in a PV and is ,;urrounded by a parasito-
call) detectable in sheep.53 • 114 :O.licroscopic sarcocyscs vary phorous vacuolar membrane (PM ! which appears to develop
from very long and narrow to short and wide. Macroscopic imo a primary sarcacyst wall (P\Vl. The P\V ls the P:>.l plus an
sarcocysts appear filamentous, like rice grains. or globular underl)ing eleccron·dcnse granular layer sit11aced immedi-
(Figure 20.4). Sarcocysts are always located within a P\' in ately beneath the PM. Sarcocyst walls have been categorir.ed
the C}'lOplasm of the host cell. More tl1an one sarcocyst may into 3, cypes trypes I to 3i) to aid~peciesdeterrninalion:13 ·10
be found in the same host cell. The sarcoq·st consists of a Species of Sarcocys1is of domestic animals are generally
cyst wall that surrounds the parasite·s merrocyte or more -specific for their imennedlate hosts than for their de-
brady?.oite stages (Figure 20.9). The stmcture and thickness finiLiYe h<lsrs.40• 4'· 144• 14~ Certain "~Id carnivores can serve
of 1he cyst wall differs among species of Sarcocys1is and as efficiem definitive hosts for domescic dog-rransmined
within each species as the sarcocyst matures. A connective species of S0rcocys1is such as S. cmzi, S. rime/la and S. capra-
tissue wall (secondary sarcocyst wall) surrounds sarcocysts cai1is. However, none of the species infective for dogs are in-
of S. gigamea. S. hardangeri. and S. ra11gi/eri. Hismlogically. fective for cats. and ,,ice /'ersa.
che sarcocyst wall is smooth. striated or hirsute; or it may Carnivore~ can harbour Snrcocysrisspp. in cheir muscles.
possess complex, branched prorrusions. Internally. groups but the host specificity of such species has yet to be deter-
of bradyioites are Crequentl) seen to be divided into com- mined.15· 4o. fl, In southern .\frica Sarcocystis occurs com-
parunems by septa that originate from the sarcocyst wall. monly in lions93 and several other caniivores.16· 87• 91 1·13 h
Septa afe presem in all but a few species of Sarco,:ysris. The has al:;o been ~een in the domestic d()g in South Africa. 113
structure of the parasites \\ichin thesarcocysts changes with
tlle maturation of the sarcocyst. Immature sarcocysts con-
Epidemiology
tain metrocytes (morher cells). Each metrocyte produces
two daughter merrocytes by endodyogeny. After what ap- Sarcocys1is occurs common!) in the muscles of cattle, sheep,
pears tO be several such dhi~ions. some of the metrocytes. pigs, horses and goats in southern Africa, as well as in \\ild
through the process of endodyogeny. produce banana- ,·enebrates in the region. i. u;.is. 76· " cr..9,. !12 9; 9,, 98. 11&. u1
shaped parasites that are now termed bradp.oites. They MJ For instance, die pre,'lllence of S. cmziapproaches 100 per
contain promii1ent amylopectin granules that stain bright cent in the hean.s of adult cattle.66 It Is imerC!;ting to note that

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 sarcocystosis 363

that sometimes occurs in sarcOC}'SlS in non-human

prlmates. 9-
The long patem period of several months in definitive
hosts contributes 10 the high prevalence of Sarcocystis infec-
tion in livestock as it provides ample oppomtnil)' for con-
tamination of pastures "~th sporocysts, which are res-iscant to
environmemal conditions. 101• lO?. tr l.ong-cenn production
of sporocysrs b>•definitive hosts is enhanced by the fact tha1
the lauer are susceptible to repeated reinfcc1ion. ll is not only
roaming cats and dogs but al~o wild carnivores which help 10
malmain the life cycle if they have access 10 dead animals.
SporocystS of Sarcocysri.s have been seen in the faeces of the
black-backed jackal (Canis mesomelas},62 which may at"1 as
an allemntive definitive host in sou1hemAfrica for the p,uho·
genie. dog-tran~milted speoies of Sar<'oq.~ris of livestock
(Table 20. I). :-.:evertheless. the main definiiive hosts in
southern ,\frica are cats, dogs and humans. Dispersal of spo·
rocysts by coprophitic anhropods. such as Oies.115 is probably
of con~iderable epidemiological importance.

Pathogenesis
The pa1hogenici1y of the different species of Sarcocysris in
llves1ock \•aries (Table 20.1), acute clinical sarcocyst0sis
being associated with the schizogonic phase of the life t"ycle.
The pathogenesis of the acute di~ease has yet 10 be eluci-
dated. but ascites and oedema in tissues are probably
caused by h}1>opr0Leina,emia and desrmction of vascular
Fig.ure 20.4 Two examples cf macroscopic Sa1cacys11s g1ganree eysis
endo1hcllum during schizogony. \\'hereas the fever wh ich
1n ovine Oi!$Ol!hag, The age$ o' :ne cysts in the respecb\>e oesoph~, are
different. the larger Q\,'Sts are older occurs is most likely related co 1he release of pyrogens from
burs1ing schi1.on1s. which either act directly on the
h)rpo1halamus or stimulate the release of prostaglandins.··
because of its high prevalence in the skeletal muscles or io. 10Therc is evidence 1ha1 prostanoidsare imporram in the
cattle in South Africa. Sarcocystl$ was for a short while pathogenesis of acme sarcoq~tosis. 22 a nd it Is speculated
mistakenly considered co be concerned in the aetiology of 1ha1 destruc.ivt> immune responses $UCh as local delayed
borulism.t>l hypersensitivity reactions could be involved. TI1e
Although no active anempt has been made co ascertain mcchaniMnS responsible for the anaemia which is a
which species of Sarcocysris occur in livestock in southern prominent feawre of acute $arco~"itosis arc 1101 known. 10
Africa, the following have been identified and life-cycle nor has the pathogenesis of Sarcocys1is,!nduced abortion or
studies have been carried out locally o.n some of them: S. (lri· the manner in which the parasite alfect.s foe1al health been
e1icanis of sheep: 90 S. bertrami of horses; 18 S. capraca11is of clarified.
goats: 9• S. cruzi of cattle;llb S. hinura of caule: 91 S. gigamea A complei- and incompletely defined imerae1ion be-
and S. re11ella of sheep:86 and 5. miescheri<ma of pigs.91 Sar· tween nutrition and the endocrine and immune S}'$tems is
cocys1is homi11is of cattle and /or S. s11i/1omi11is of pigs. both responsibie for the growth penurbation that is often a con-
of \\'hich have humans as the definitive host (Table 20.1). ~equence in animals which have recov<.'.red from acute
must occur in southern Africa because sporncys1s are often sarcocys1osis. Connective tissue breakdown and musc:le
observed in human faeces:' A kind of cys1 which close!~· re· wasring migh1 be correlated \\ith 1he abnormal nitrogen
sembled S. suihominis has been seen in pork from an abal· me1aboli~m that appears 10 result from rhe malabsorption
tolr ,in the former Transvaal Province of South Afrtca.91 and internal recyding of nitrogen. 18 The host's immune
Other species of Sarcocys1is will unquestionably be found in response ro the: lnfectlon leads 10 changes in the concemra·
domestic livestock in southern Africa if looked for. Human tion of 1he cytokine signal. There is thought 10 be a tumour
muscularsarcocystosis is known in South /\frica96 but. ultra· necrosis factor alpha response by macrophages. which is
srructurall), the wall of a sarcocyst in the person concerned localized within tissues and no1 rcllec1ed by a rniscd level
was found to be unlike those of domestic animals. being In lhe plasma. I formonal changes occur, panicularly in
more of the general type with elaborate cyst waU protrusions re1,,ard 10 secretion or growth-associated honnones. such as

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364 ,~cnn" nm: Proromal diseases

somatostatin, thyrotropin-releasing hormone. in,ulin-like are visible in the ce.111rc of some lesions' 1 56 and there ap-
growth factor-I ,somacomedin CJ and gro,,1.h hormone pears to be immunohistochemical suppon for their occur-
(sornatotropin}.48 which evenrually lead 10 inhibition of rence there.62 Sarcocystis-specific rype-1 h)'-persensitivi~
growth. Imerleukin-1 and other C}1oklnes may also be in- may be im·olved in the pathogenesis. 02 However. when ex,
vol1·ed in gro111.h retardation. amined b~ light microscopy. many cases of eosinophilit
Sarcocyslis has traditionally been regarded as the cause myositis are found not to be ob,·iouslr related to Sarcocys-
of eosinophilic myosltis. This is part Ir because sarcoC) st~ ris infeclion.8 ·1 Conscquenlly. I.he sarcoq•stic aetiology of

h
G.
---- -
I
;'
~11
~
I
~~
J

1-------------- -- --- I
'

& '
(/ b

/
;-- C

,
Figure 20.S life cycle of Sarcocr,ws
a = sporocystS f = sa·cocv,1 rn muscle
b = Sj)OtOZOile g~ oradyzo,c mero1011es
c am! d = parasites undergoing schi10ij0rly in endo1helia cells of h = maci09ame1ocy1e
blood vessels I = m,crcgameiocvte
e = tachyzoic merorertes !free or inside mononuclear cells. I = OOC~St
in the bloodstteaml

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 ;,arcoq•stosis 365

eosinophilic myosi1is remain~ unproven. panicularly since sporor:ysb at any one time. 1~· 2-l 29- 38-4o..u. :..1 109 120. 1zs il is

the condition has not been observed in any animals \,hich when numerous sporocysts of a pathogenic species or Sar-
have suffered from experimemally induced disease. These cocysris rrable 20.ll are ingested simultaneously in feed or
ha\'e. howe,•er. usually been roung individuals 1hat were water comaminated by definitive hosts tha1 clinical disease
infected on only one occasion. It remains a possfb!lity that develops. Under narural conditions, clinical sarcocystosis
the development of eosinophilic myositis is dependem has been documemcuonly in caule! 0 shcep12• 50• 108· ui and
upon repeated ingestion of sporocysts of Snrcocrs1i.. goais.81 Concurrent infections might predispose animals 10
Some ~pccies of Sarcocysris can cause rrans.placemal clinical sarcocy~1osis:u1
infection: and congenital sarcocysrosis has been induced in >-amrally occurring acute sarcocyswsis ha$ so far beeo
caule, ~heep. goats and pigs.4° Animals infected \\ith large recognized only in goa1~ and canle1·1• in southern Africa
doses of sporocysts during pregnancy may abort, have s1iU- (figures 20.1 10 20.3).
binh. or give birth 10 weak animals. Sarcoc:yscis schi,:oms In the acu1e illness, 1he body temperature may rise to
ha\'e been demo nstrnted in foetal memb~anes and foetal tis- over 40 'C be1ween two and three "'eeks after infecrion.
sue,;. The exact mechanism of abortion is not kno,m be· although an intrease in remperature ha~ been noted in
cause the presence of schizoncs in the foetus is rare. experimental!} induced disease in horse.s IO to J 2 days
certainly in expcrimemally induced infeciions.~0 Under after they were given large number5 of sporocysts. 99 · 123
natural conditions, neonatal death has been recorded once Apan from fe,·er, clinical signs are not notiC'l'd until abou1
in a goat in Australia81 and in one lamb.32 There. are. how- 24 days after Infection. when animals may exhibit anaemia.
C\'er, several reports of congenital infections in cattle11 and d iarrhoea. anorexia. weakness. mild dehydration and loss
these reporrs up to 1988 have been summari7.ed. 10 In a of weight. 11 · H Other signs ma~· include excessive sali\'ation
planned survey. Sarcocys1is was idemified in only one of 498 (a prominent fearure in affec1ed cattle). muscle tremors
aborted foemses ln Callfornia. 1 indicating that Sa.rcocystis is and loss oi hair. which in ~ome caule is particularly notice-
1101 a common cause of abortion in canle. Earlier repor1s able at the tip of the iail. 14• al). ~9 Pregnanl animals may
may have been of misdiagnosed .\'eospora. infections (see abort or gi'"e birlh prematurely. If the discas<' progresses.
Chap1e r 23: Neosporosis). many atrected ammals become recumbent, showing signs
of central ner\'ous system involvemem. and die. The dura·
tlon of clinfcal signs varies from a few days to several
Clinical signs
weeks. Growth ma)' be retarded in individuals chat suTYiVe
Virtually all Sarcocysris infec1ion~ in lives1ock are subclln- an acute infec11on 20 • i1. ,o."5 · ,11. 71 and wool growth maybe
ical. because animals do not usually ing~t large numbers of adversely affected .

..
Figure 20.6 Faecal flotatioo irom a d:,,g naturally infecied with Sarcoc,s;is. A. low magnificaiion. :, s'1ow the size cf a Sarroc,,s11s spo1ocvs1 !SJ
compared \·,,tn a To.~ascarisegg IT1. x ~30 B H1gn magnif:ca,,or, 1how1ng a Sarrocysris sporoc1st {S1 a"d \'NO Neospo,'8 or tiamm01Tdia-hka oocySi'S
(HI, x' 250 1From rafe1ence number 26. w,th perm1ss1onl

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366 StenO!-.'lWO: Protozoal disease.~

Table 20.1 Sarcocys1is species in livestock

INTER· SARCOCYSflS OEFINmVE PATHOGENICITY CYST TYPE LIGHT ULTM-STRUCTURAL PERTINENT

MEDIATE SPECIES HOSTS INUVESiOCK IN LONG· MICROSCOPIC APPEARANCE OF LITERATURE.
HOST STANDING Al'PEARANC1' PRIMARY CYSi WALL INCLUDING
INFECTION OrCYSTWAU PROiRUSIGNS REfE!lcNC£1S.
IN WHICH FlNE
SiRUCiURE OF
CYSTWAUIS
ILLUSTRATED

Cattle S,C,Ul/ IOog. coyote. fox. Pathogenic Microscopic Thin Filamemous, rlanened 16. 27. 118
jackal raccoon. ever cyst wall
wclf
S. h11SUl8 Ca; M•ld!y Macroscop,c Thtl V1llcs-lilil! constricted 28,37
pathogenic at basi;. laterally
exparded rn ;he middle
and tapered distally
S,hominis Human. other Mildly Not known Thie~ Palisade-like. nor 37. 103
primates pathogenic constncted at base

Sheep S. ariericanis Dog Pa1hoge,1ic. but Microscopic Tom Filamen!Ous. long and 6S. 118
less so than flar..ened 01e1 cyst wall
S rer.ella
S. renella Dog, coyote. fox Pathogenic Microscopic Tn·ck Palisade-like 16, 36. i 18
S. giganree Cat Mtldly Macroscopic Th•ck sgcondary Cauhflowe.r-hke 16. ilO. 111, i 18
pathoge:i,c cvs1 wall formerl
S. meduSJform,s Ca: Unkno.-m Macroscco;c Th:o Snake-l1~e 12, i 1!, 118
S. mmoensis Dog Un~nown Macroscopic Th·ck Pahsa::e-llke 123
Goat S capracams Dog, covate. fox Pathogenic M1croscop1c Thick Pahsaae-11,e 3S. 4>;, 6S. 118
S. hucicems Dilg Possibly Microscopic ltru1 Thm Filamentous. long and 70
pathogemc cys1s can be uo ffaneneo over cvst
10 2,5 mm long wall rrne cvs, .van is
similar ,n appearance
to !hat of S ar,e:,camsl
S moute,· Cat Non,patiiogemc Macrosccp1c Thie~ seconda!)' Ca~llftower·lll(I! 58,68
c~'St wall formed

Pig S. miescheriane Dog, fox.1ackal, Pathogenic Microscop,c Thick Palisade-l;ice 60

raccoon
S. suihominis .. -luman. 01111), Pa,hogen,c Microscopic Thiel< Palisa:fe-lilte 71, so l()i!
pnmates
S pcrcifelis+ Cat Pathogenic Macroscopic Not ~nown Notxnown 26
Horse S.benrami•• Both 1h1ck- and Villus-like conical with
S equlcanis,,.. Dog Mildly Macroscopic thiri-walled cysts corrugated ou,line 18. 42. 46. 95, 117
S fays,~ patl1oge11c have been seen
~.va:er S. btJlfaloms Cat Un1<nul'l11 Macroscopic Thick P.;'lsa~e-llke 72
buffalo 7,
S dubeyi Un~own Unmowr. Macroscopic Th c~ Palisade-li(e

S. fus,formis Cat Non-pathogenic Macroscopic Thin Cauliflower-like 103. 105

S. levir.ei Dog Unl<nown Macroscop:c Thin Filamer.wus 25,73

In the aosence of 8\'lllenre mat 1wo cat-transmmen species or Sarcocysris oa:ur 1n goats. S cap11fe/is'-.; 1s prov1s10~!1)' cons,dered to be a svnonvm
, al S moulei
L1 a.1d Uanec noted possible_ b1ological and sm1crura1 differences between tile organism they studied and S su1nominis in Europe
+ fne validity of thts spec;es is uncartain and confmnat,cn oi the pi~al life cyt!ei.. s; ,s needed A mwocys1 m awanhog (f'h-acochoeros
aethiopicus) from the former Transvaal Province m Somn Ainca had cauliltower-like cvst wall protmsioos It differed ultrastruc:urally irom a
rn1crocvst in wanhog from Namibia KwaZulu-Natal Province in South Afri(a, and Z1111babwe This microcyst was no, unlike S miescheflet>a ~'
However. theie is rwt as ve, any evidence that e1the: soeciss of the warthog 1s transmissible to the domestic pig Tv. o species of Sarcoc)'sus have
s1nte been descnbe!l from war.hog in South Afnca; 3
+.- It ts not ye1 clear whether ine m1clc-walled cysts ,n horses betong 10 dlffeient scec•es o~ not

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 Sarcocys1osis 367

Clinical myoparhy. which may be accompat1ied br oedematous and congested. There may tie small erosions
chronic illness, has been observed in horses \\1th muscular or shallow ulcers in the mucosa of the buccal cavi~,
Snrcocystis lnfocrion, bu1 there Is evidence 1ha1 there ma}' oesophagus and obomasum.
nm be an associarion berween the presence of sarcc,cys1s The predominam microscopic lesion in acute sarco-
and clinical myositis.5 5• 1• 1 cystosis is inflammarory rather than degeneratlvc} 0 · 17
Intestinal Snrcocystis Infections in definicive hosts are Medium to large numbers of lymphocytes and macro-
usually asympromatic:10• 114 However, a dog in South Africa phages, together \\~th a few plasmn cells and eosinophils,
that was clinically W (cause unknown) was shedding vast are found perivascularly and in the interstitial Lissucs of the
numbers or ooC}'S!S and sporocysLS. derecrnble upon direct hean. skeletal muscles, lungs. liver. kidneys and, 10 a lesser
examination of faecal specimens. to; This massive intestinal extem, 01her organs. The predomlnamlymononuclear cell
infection was probably 1he result of repeated meals of Snrco- infihrate is frequentl)' accompanied by haemorrhage.
cystis-infec1ed meat. oedema, multifocal degeneration and necrosis {especially
in the cardiac and skeletal musculature and kidneys), vas-
culi1ls and. less commonly, thrombosis. The haemorrhages
Pathology
may be generalized, and are often not associated with
A haemolytic, normocytic. nom1ochromic anaemia, \\'ilh inflammation. :Vluscular lesions in acu1esarcocys1osis vary
few or no re1iculocy1es in evidence, is charac1erlstlc of acute in severity from muscle 10 muscle and within individual
sarcocystosis.u-lU, • 0 The packed cell volume begins 10 fall muscles. In the brain and spinal cord, focal haemorrhages
berween three and four weeks after infecrion and reaches a sometimes surround areas of necrosis, and scaucred foci
level of below 20 per cent in animals wiih moderate 10 se\'ere of gliosis and of mononuclear cell infiltration in the
disease. Inf11Ction with as few as I 000 sporocysrs can causc mer,inges are.e"ident. Enlargemem ofJ),mph nodes results
reduced haema1ocrits between four and five weeks after from l~111phoid hyper.plasia with attendant distension of
Infection in lambs. 10!1 Clinical pathological findings have medullaf')· cords. oedema. and his1iocytosis of the sinuses
not been uniform in the experimental si1uation as far as in the medulla. Schizoms may or may not be present in
parameters or blood coagulation. such as partial thrombo- organs and. regardless of location. are not always asso-
pla~tin time, pro1hrombin time and thrombin time. are ciated with tissue changes. particularly not in infections
concemed. 1!1 with non-pathogenic species of Sarcocysris of livcslock
The serum bilirubin level is elevated during 1he acuce in- (Table 20.1).
fection, and ln 1em1lnal disease the blood urea nitrogen Some species or SnrcoCJ1sris fonn macroscopically visible
level also rises. S<>rum levels of enzymes such as nspartate sarcocys1s in the muscles of chronically infoc1cd Jives1ock
a minocransferasc. creati ne kinase and lactic deh)•drogenase (Table 20.1). ~lacroscopic cysts of S. gigm1cea are some·
tend 10 be elevated during acute and early chronic sarcocys· times seen in sheep, being particularly conspicuoµs in oe-
co~is. In pigs, an increase in creatine kinase ac1M1y is de- sophageal. laryngeal and tongue muscles. They are oval or
tected from 28 days after infec1io11.2., Senim creatine .kinase round (Figure 20.4). Meat with macroscopic sarcocrsts is
le\'els in horses increase 4- to -10-fold some 63 days after oral generally condemned in abanoir,. as unfit for human con-
adm!nls1ra1ion of sporocysts and may remain high umll sumption, despite the fact that a zoonotic risk has not been
approximately 147 days after infection. 12a idenrlficd.h· 52 -~a
:S:utrition is affected in acute sarcocystosis. There Is a In chronically infocred livestock animals, sarcocysts
marked reduc1ion in retained nitrogen, percentage nitrogen occur within myocytes of the myocardium and skeletal
retained per unir consumed. hydroxyproline excretion and muscles. and sometimes in the brain. 3• ~o. 114 A tissue
creatine excretion, and an Increase in the ratio of3-me1hyl- response is not normally apparem in muscu lar Snrco·
histidine to crea1illC excreted b)' parnsitizcd cattle.46 cystis i11fec1lons.;u, s,i Harely. a degenerating sarcocyst,
The pathology of acute sarcocystosis has been well de- or sarcocysts. may be encountered in muscle. The result-
scribed, \,ich much of the early work having been done <>n ing reaction may be macroscopically \~Sible as one or
cattle infected with S. cruzi. ln all livestock species, gross le- m<ire dull white foci about l mm in diameter. The dead
sions a1 necropsy include serous atrophy and necrosis ofiat, parasites are surrou11ded by mononuclear cel ls, neutro·
pallor of the mucous membrmies, an icteric appearance of phils, eosinophils. giant cells or a combination of these
rissucs and organs. h}1dropericardium, hydrothorax, ascites. cells. However. the cellular response is usually mainly
and widely distributed pe1echiac and ecchymoses (Figures mononuclear and neutrophilic, a different reaction to that
20.10 and 20.11). especially on serosal surfaces of the \~s- ofeo$inophilic myositis. How sarcocysts arc removed from
cera, and 1hroughou1 the ~keleral and cardiac musculature, host tiSS11tl is not yet clear, but there is phagocytic in\'oh-e-
which may also show mu lei pie pale foci of variable size or me111.~u. 120
haemorrhagic streaks. Skeletal musclt:s are often pale pink Eosinophilic myosit is is a specific and app.iremly sub-
with dark red mo11li11g. Lymph nodes are enlarged and clinical inflammatOf')' condition which is sometimes
oedematous and often contain petechiae. The lungs are encoumered in livestock {particularly canle and sheep)

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368 " t n,1, r ,11~ l'ro101.oal disease-s

figure 20.7 Second.gene•at,on

s-41•,oms in ~ct ion o1llovine ~,csey.
za days aner 1ngest1on oi S CILJZt S1Jorocvsts
;,, An immature (aitow~adl sth·1,nt wnh
nuclear lobes and 11•,ti sthizoo:s trat a:e
almost mature (arrows). x 830
B toor.-,dual merozoites (a;rowheadsJand an
immature sehizont {arrow!, K ' 000

slaughrered in abattoirs. The macroscopically \'isible Diagnosis

l"'sions consist of greenish or yellowish areas. patches or
granulomas (measuring up to 5-15 >< 1-3 mm) in card iac \dendfication of the spe<.ies of Sarcocy$//.~ that are present ln
and skeleral muscles. There is no predilection for particu- muscles of chronically infec1ed ti,·estock animals (there is
lar muscles. The presence of lhe granulomas often resi1h, frequend~· more than one species in the same indi\idual
in condemnation of the affected carcasses. Early lesions can often be ha$ed mainly 011 the t1ltras1ructural appearance
consist of numerous eosinophils (which are responsible for of the cyst wall (Table 20. 1). ldemifica1ion of Sarc<>cys1is 10
the green colour) lying between muscl e fasciculi and fibre~. 1he specific level is, however, usually an academic exercise
E\'enrnal degeneration, necrosis and di$appearance of rather than a ma11er of practical imr>ortance; the reason
muscle fibres occur, their place being taken by eosinophils being that chronic Satcncysris imections in Jh·estock are in-
and connective tissue. TI1e granulomM ha\'e a central area variably subcllnical. The pre\'alence of sarCO~'$ts in the
-coma[ning necrotic cellular debris. which is surrounded by musculature of domestic herbi\'Ore.~ is high, the occurrence
eosinophils, epithelioid cells, fibrocrtes. giant cells and of mamre sarcoc)'sts in an animal wilb acute sarcocyscos1~
lymphocytes. normally being an incidental finding. indicatil'e of a (pre\'i-
The granulomas of eosinophilic myos[tis should not be ous) chronic infection. It is difficuh to diagnose acute 5arco-
confused with the sometimes superficially similar cysts of cystosis beciu1s<! 1he clinical signs of disease are 11011-
Sarcocystis, S\lch as S. gigamea of sheep (Table 20.1), which specilic and there are. at present. no definiti,e te~ts ro
may also be macroscopically ,isible (Figure 20.4). confirm the diagnosis in 1he lhing animal. Parasitological

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 :.arc.ocysm,is 369

Figure 20.8 Experimental bov,ne

s.a,cocys,os,s.. e·eciron m,crog'l!ph
ot a s~n,;on1 10 a ...ascu!ar
er.dntr.elral ,e:I Ill ;he ora,n., 6 030
Note that ,hon1nes are absent and
:ha: nc c.a<as11ophorous vacuole ,s
\'1S 1bl8

... ,·
' .. ' ' ..'
'.. ' ' ' •
' -. ' •
1

. -. -. . . ,.
'
...
... . .... ' '
Figure 20.9 iyp,car
' '
• .. h,s,opa1hcloglcal appearance of a
- G

'
•
' nat,re sarcocyst in striated muscle.
Cvsts octur cornmonlv m the

. ..-_:: -·~ -
'
muscu:,m,,e 01 ll~estock
....
(By courtesy of Or JA Neser

--
. ......,..... -. -' Onde1s,ePOort Veterinary J115;,w1e•
Soutn Aincal

confirmation of clini,:ally suspected disea~c is based on the nccmsb. rl!.spcrti\'cly. Schi1.om, may occur in va~cular
finding oi schi:t.oms (Figu tes 20. l 10 20.3 and 20.7, in endo- cndoth(:lial c.clls in the: subcpithelial connective tissue of
thelial cells of blood ,•essels. on microscopy ufbiopsy mate- placental CO[)~edons. llltrnstructurally. I.he mero,mites ol
rial (such as muscle or !)mph node), or. more commonly. on ~ITO()'Sris arc similart<> orht!r apicomplexnn'l.oltc:s (see Dif-
his1opa1hology al'ter a necropsy. However. schizom~ or free ferential diagnos is}. exccpc that the) lack rhoptries. lmmu-
mcrozoi1cs arc noL always detected In infecced animal,. ei- noh,srnchemistrv is helpful 111 idemifymi:schizom,S in Lissue
ther because the>· are too few in number 10 be found or be- section~.
cause schi1.0111s ha\ e ruptured by the time that the infection The parasilacmia duringacut<! sarcocystosis is not u~eful
becomes clinically apparen1. It is particularly in foetusc;s fm diagnosis because ch!' de1ec1ion of parasites in blood or
that the parasites arc not consis1en1ly pre,cm. and th!! buffy-coa, ;mear,, b 100 time-consuming.
chances of diagnosing Sarco0:sri.s-indue1?d aborcion arc im- Because of the size of rhe sarcocysts. a diagnosis of
pro\'ed !fa rnriecyoffoecal tissues :ire examined microscopi- chronic infection wich species or Snrcocysris such m;
tally. These tissues ~hould indude ,amples rrom the brain S. gigm1te<1 (Figure 20.4) or S. 111ed11siformis can readlly be
and placema. where sarcot1-smsis gini~ rl~e to gllosis and made macro,copically in long-standing infection$ in older

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310 S1:C110 1' l\\'O; Protoioal diS~ilSl'S

Figure 20.10 b!per1men:a1 bcvme

sarcocys1os1s: petech,al and
ecchymo;,c haemorrhages m the
k,cney

Figure 20.11 Expei1mental oovine

sarcocvsiosis'. pe1ech1at
haemorrhages in the brain

animals. Cys ts of some species of Sarcocyscis. such as animals from the same farm are compared with [he titres of
S. cntzi, never become large enough to be clearly \isible to Individuals \\hich are ill. R<'search h, bch,g carried out in an
the naked eye (Table 20.1 ). au empt 10 refine techniques for the tliagnosi<: of acute sar-
J\llhough various serological tests !indirect fluorescent cocystosi~:tn, ;,. 63.15. 106. 112. 119 l:!S. 130. llJ. 136, 138. 139
antibody test and tmzyme-linked immunosorbent assay) i\ search for the small sporocysts of Sarcocysris in the
have been described , the)' are of limited value. Fe\\' labora- faeces of cami\·ores (partlcularly dogs, by cemrili.lgal 00l3·
tories keep antigen as it is not commercially a\'8ilable. Re- uon. using a super-saturated sodium chloride solution. can
sult.\ are difficult 10 interpret beC'au~e subclinical Sarcocysrls help in the aetiological evaluation of suspected acute sarco-
infections are very common In li\·estock. Furthermore, there cys1osis. \\'hat should be taken into consideration is tha[ the
is cross-reac1ion in serological tests beiween pathogenic prepmem period in carnivores is one 10 two weeks. and that
and non-pathogenic Sctrcocysris~pp., although Sarcocysris i~ the duration of 1he pa1em period can be several months. Al·
largely antigen icall) distinct from other coccidia.31 · -io. 83· 8 ·1 though it is possible 10 stain sporocysts in faeces (they are
Antibody titres can nevertheless assist Lhe making of a diag- acid fas1. as are C>ocys1s of Cryp1osporitli11m and other coc-
nosis. especially if those in rhe senim of clinical!~ hea lthy cidia89). thls is not necessary because they are clearly ,isible
 Snrcocyi.tosis 371

on ligh1 microscopy a1 x-100 (o r even lower) magnifica1ion in However. free1.ing of mear for i<>ng periods does n01 neces-
fresh preparations, provided that the illumina1ion used is not sarily kill Srm:ocystis 13 and is. 1herefore, less elfective as a
100 brigh1. The internal s1n1cmre of a sporocys1 (contain ing means of interrup1ing transmission.
fow sporowites and a granularsporocystic residuum) can be To effective!~· control Snrcocysris infec1ion on farms. car-
seen when an oil immersion objective is ltsed. casses oi dead lives1ock should be buried or burned. Cami-
,,ores tha1 ea1 uncooked mea1 of domestic animals ~hould
not be allowed on to pl'emises where their faeces can con-
Differential diagnosis taminate the drinking water. feed or bedding ofiivesrock.
At present. no \'accine is avanable 10 pro1ect livestock
Known parasites of livestock which closely resemble Sarco- agains1 acute sarcocysro~is.
c;ysris morphologically are Toxop/asma g911dii (see Chapter Although prophylactic administration of some drugs
I 8: Toxoplasmosis) and che newly described Neospora ca11i- (amprolium, salinomycin and monensin) cun reduce the
rwm, which has been reported as being a pathogen in canle, clinical effects of acute sarcocys1osis. none of these re-
sheep. goats. dogs,, and horses (see Chapter 23: Neosporo- gimes Is pracc.ical because exposure toSnrcoc;ystis is unpre-
sis). In his1ological seccions. Sarcocw1is can be dilferenci- dic1able. 3·1· 78
a1ed from these proco1.oa by the presence of schlzonrs in For chemmherapy, emir halofuginonc has been used
vascular endothelial cells (the)' often spuriously appear co with success in li.vestock but the drug is toxic and can1ful
be free in 1he blood vessel lumen). whereas Nenspora and dosing ls neccssal') .1 lalofuginone (0.66 mg/kg body weigh1)
Toxop/t1,m1t1 can develop in a ,vidl' range of cell 1)'})OS. The given once or more often has pre\'·emed deaih in goa1s and
presence of schizonts in numerous endothelial cells of sheep ½1th acute sarcocys1osis.66 Some of the po.temiated
blood vessels is suggestive of a Snrcocy,ri; infection. su lphonamides are effective in elimina1ing both immature
On electron microscopy of St1rcocys1is schizonts. mero- and mature cysts of S, nwris in mice. but their effects on ~ar-
;mites are seen 10 be free in the hos, cell cytoplasm (Figures cooysts in liveswck have not yet been swdied, 121
20.3 and 20.8) (i.e. no pamsi1ophorous vacuole is evident).
Groups of Toxoplasma organisms, on 1he olhcr hand. are
Sarcocystis species with unusuallife cycle
surrounded by a parasitophorous vacuolar membrane. In
or biology
Neospom infections, a parasltopnorous vacuole may or may
not be present. Merozoites of St1rco~1is do not have the or- Sarcocxstis species are rarely fa1al 10 \,~ldlife.$3 Howe\·er,
ganelles known as rhoptries which are observed in :veospol'a S. fa/caw/a, which has an avian-opossum cycle. is known
and Toxoplasma 1achy1.oic merozoiles. being more numer- to be biologically different from other species of S<1rcocys-
ous in the former (dMding tach)'7.0ic merozoites contain 1is. it appears to have a wide range of intem1ediate a,·ian
fewer rhoptries chan non-dividing merozoites). hos1s but appnrencly emir 1he opossum (Didelphis spp.)
Immature schizoms wlrh multiple nuclei not sur- has so far been idenlified as the definitive host. 122 There
rounded by discrete cytoplasmic ari>.as are seen in sarcocys- arc se\·eral repons of fa tal acu1e ~arcocystosis in passerine
1osis. Mulrinucleared scages do not occur in .Veospora and birds, a1 least some of which are likely to ha,·e heen caused
Toxoplasm(I infections because di\.'ision is by endodyogeny, by S. fa/ca ruin. An unusual feature of pathogenesis is chat
each parasite giving rise ro cwo daughter organisms. schizogony may lasr for l•P to ~ix months. Pneumonia is 1he
Tissue cysis of both 7':eospom and Toxoplnsmn lack the predominan1 lesion. with ini!'llVM<:ular schizonts heing
1.')'S\ wall protru~ions which are characteris,ic of sarcocysts. present.
Cys1s of some species o[Sarcocysris (e.g. S. cr11zl) are divided Another group of St1rcoc)'Stis-llke organisms with un-
into comparunenrs by sepra, unlike tissue cysts of .Veospom known life cycles have been reported to kill na1unill) In-
and Toxopla,11111. There are meirocrces (ovoid, rapidly mul· fected capercailliae, minks, chinchillas. sea lions. harbour
1ipl)1ng organisms) in young sarcocysts (and somerimes a seals. raccoons. monkeys, black beats, skunks. sheep.
few in older ones) but these do not occur in cysts of caule. horses. cats and dogs. HO The disease is often con-
,\"eospom or 1"oxoplasma. fined to the central nervous sys1cm or liver and schi7.onts
Sarcocysrl., pnrasi1es can be distinguished immunobis- are not intravaset1lar. SarcocystS of these parasites are not
rochemicnlly from Toxop/nsma. Seospom and other api· known. The l!fe C)'cle of one of these organisms in the horse
complexan parasites. (S. 11e11rona) has been partly resolved (see Chapter 24:
, Equine protozoa! myeloencephalitis). Sarcocys1ls ca11is is
anotherprotozoon belonging 101his group. 3 'The parasites
Control
affec1 mainly hepacocytes, and dogs. cars. bears. chinchil-
The life cycle of S(I rcocysris is depende111 upon 1hc predator- las, a horse and a sea lion are known to have died of acme
prey relationship thar exists be1ween its definitive and inter- severe hepatitis. 24 Reports of organisms causing myeloen-
mediate hosts. C,msequently, if meat eaten by the definitive cephalitis In sheep:;· •1. 124 and caule34 Ul probably belong
hosts h~ been cooked, the transmission cycle is broken. to this group,
372 ~.cr10~ n,'O: Pro1oioal diseases

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COSR:UJ, P.A•• 1991. Ncospom,Ukc prot0zoan infl'Ction os o maJo:r cause or acu« sa(cocystio.is in growing p~. ?ara;iro!ogy Restar<l1, -s. l 1.5-118.
abonion in CaJifomf3 d.tiry c:aHle,Joumal ofliteA.mnt«m t'Ncrfnll.l)' 23 Q,.\UCi>SCHIES.~ .• SCflSIIDER, T RO°M'•.ta. ~f. t< 81CKI<AlmT. Ji;., , t988. The
M«IIMl i\s..Vit'intitm, l98.2•11-241$.
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2 a~m~. ft.~ .• O()~l:o;G, ~•. STOt.rn. M.. Q\l,t\NO'T, $... h BOC..l~'HAAl)l\ r., lS98, weigh, gain pt srre$s--sensitive aud sttes.s·m~nsilivc PlSS· V~uri11ary
Three new San;oc.._w,u'lpede.s~ Sarcocy~·tif.giraffa~. S. klmerluruiS, n.ndS. l'tlr<111l10/ogy, 27, 2:l t-22~
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3 a1c;;AI.,KE. R..D.&TUfn:>:, R.<?,, 1.960. Theocc.urnmce of:L<')'St ofSll-n:ocy111s
~ l.'ltS$A!\.UJ,.'1!.. A,"!>, .a.. li,I'\,"", 'S,P.. 1918. S1udit!'° on $(,rcoc,•.ui,rtn ~tala~ s!:s... I
1.ankC$1Cr 1882 In the cer~bdlurn of .in ox, /m1m<1/ ofthr S,,1111, ,\frlr1111
Sarc«ystls /ev/11~, n. op. rro:n cho wa,cr buffalo R1</Jnl11sbu1Jnlls.
Vewrinal')' 1\J,,ruca/ Assoc,ar,011, 31. 2il-27<1,
atrsthrlfrft1r l'arositc11~,mdr, S.S. 12i-138-
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!16 ou••Y. 1,1••• 1976. A "'"icw oiStircotJ':lli.f uf dom'1$tic tmlmJb and of nthor
glardla.4. So111h Afrlain .Wt<llail/011mal, 70,436.
cocddfo a( cat> onrl dogs. /011r11al of1/1t.~m~rlcar1 \ltr,1r/11ao· ,\/,-d/«11
s cA1.oow. c.1., cwLOw, r.n. t sr.Hoa;. 1.., :woo. Cllukal. palho!o~cal nnd ~iarion. 169, 1061-1076.
epldomiologicnl findinis in thrc~ outbreaks of a ..i'nc protozoan
2; uun1EY.1.,•.. 19a~. Quanlit;uJv~ parasittmia in ou, I!'",. fee! $tr'('()C)~ti1 rru:i
myelocnctphalftl>. ni.·
l'Cll!fl/U1ryR,wml, 14f>, 7-10.
~poro~"Sl$ from CO)'Ol8'$• .:\,ntrletm Jo11m.<1J o/V<u>ri1tnry Rt'11!flftil, 43,
6 CUAMBE.RS, r,Gv 1987. Carcass ilnd offal c:ondemnntion.s tu meat 1085-1086.
in,-pectlon in Zim\>ilbwe. :timbnbtw Varc·rI,wry/0<1r,M/, 1.8. 11-18,
28 DUUh"\', 1.P.• 1982. Dev•lopmen1 of th• OX""'1t q'cle of Smco<ystil lurs111a,
r COl.llrR\". P,. 1987, The pathogenffis of af'1.ue b-O~ne ~.ucocy,s1osis, 1. PYoi:1,et//11$' of tile Hclmmt/iolugitt1/ Sod,·l)I of\l'a,/Jihgt1111, ,9.195-3().1..
Cllolcal sign• 3nd anaemia. Iris/I \1ewt11my fo,rma/, 41. ?73-230,
29 DUBEY. J.P.• 19112. DC\·~lopment 01 o,~oyote 9·de orSnrr:oc,ws rflt:i.
8 cow•,·· 9., 198g. The pathogcnesh; or a,-ucc bovine <nrcoeJ1$t0>is. II. Jouma/ ofProm:011logy, 2.q. 591-60 I.
Bone marrow ~nd peripheral blood whico cell respQn!e. /rlsll \lwril>IIJ)'
/OllfllDI, 42, 33-39. 3.0 tHl6EY. 1.1~.. t983. lmmunny lO sarcQt)'$to.$!$: ~1odihcadon of int£~um::~
coi:cidiO<ls. •nd dlsnppear;,n,-c of.<a~'Sh in do,ry goais. \1':urlrmo·
g co1J.ERV, r., 1989. The patlwg<t1,',\i• ofocll<e bovint ,:arw()•>tosi,. Ill. /11 Pttff/$(tolo,.'V. 13,2:1-.11.
1•irroocudles on I.ht lnternctlon•bem~n tif')'1hrocyie.~nd mon<>ncdtar
phagOC)1C$. lris/1 Vereri11no• Joumnl. ~2. 69-7-1. 31 i>vet.-r. J.P.. 1993. Toxopfasm"· Vcmpom, Sorcuc;i•,5t,s., and other tJSSue
~St·fonmng coccidia of humans and animals. tn: xarutA.1,1•.. ted ,,
lO Cl>US.AV, P.• 1989. Th•pathog.,,,esisor :.<:Ute bo,inc saiCOC)'StOSl>. IV. Pam,11/c Prortr..oa. 2nd edn. \'ol. 6. :-;.,, York: A~ode,mc Press.
Theeffilea ofoonicosteroid the rap)' on thecoursc or the Ollaemia lrislt pp. 1-158.
Vt!ttirina,yJoumal, .;2, 85-92.
3.2 ou1n:v. J.P.• rr.Ann&. w.,. f.. ,MO)t.\:>. 11. r.. 1989. Fatnl pennaml
11 cou.o,v, P. $' we.Ave~~. e., 1981. An ou1b1eak of Sarcocy$t/$ In calves in
sorcc,syStQSTS in A lnm9./91murl ,,fPnmsl:ology. ;$. 9.8C>-~.
Ireland. Irish Ve1ert,1ar,•Journai 35.159-162.
33 ousa-. r.r.4>-ooE:1iNG, 1..., :001. Tnxopla1nlosl(nnd reJ3:ed fnhicdons.. /rr.
1: co1.i.r.-.,~, c.u.., ATXt:-."SOx, e. $. CttAALF.STO~. W-.\,G., 1979, Smdf~ oa
MMUEt a .. l"\'flUR. M. ~K()CA-.. • .\.:.t., (Nb}. P(lf(~ltirt)i~csn/lVlld
S<lrtO<Jf$t/s spccios 111 The mac,oc~tic specius o! ,he,p. Vew Ze,1/wul
.\/ammals..Am<:>. Iowa: Iowa Stole IJnlveislci• Press. pp 4i&-:H9.
\le,erinao•/oumal, 27, 204-206.
34 U1.18ll', J.I•.• l'EHM,'I', A." ~11;,.~lil)Y. \i,I 1987. Encuph~ltf~ couo;td by a
13 cow~,. G.11 ... CIIARLrsrDS, w ••.G., 1980. Stud!~ on Sam><)'Jll$SpL-cles Sam><J·$:is-ll!u,organlsm In a s,ccr Jo:muilof1fi,Amcri<an l'~wrlnal),
VII. The effec1 o!tempemu~ on che vl~l>llit)' of matC<>C)-;sCJ (Sntt<)Q~l/s
.\fed/cal .'JJO.:lmiq11, 191, 231-23?.
giga11tta) of sheep. Ne11· Zeo/and v,wfnary /oumal. 28, l8!H91.
35 DUSh-Y, J.P." ~•tEII. C.A•• 1991. SatCOC)~tis Mllltn. sp. l,\plcompl~!(a;
1.; C:OJt.,•t;R,A..lL, Ml'TCHY.U.. O., '\\k:..\0)1 It.I. $,TAVt.(Jft. P.A., tg(i3, Dallnc:nr
Snrccx:ystidae), the etiologic ogent of gcnontllz,•d coec!dlo,~ In do~~-
dlsea.c. An infection of cu11lt pre,umed 10 b• c,11,s,'<f b)' en unldcniifl<.-d
/011mo/ of"/'ams/10/ogy, ;;, s22-S2:,
protozOQn. Canadltm Vtirrinary Jounml, 4. 2.52-264.
36 ouetlY, ,.,•• $Pr..t•. c.., .. c.u.u.,;, G. ,..i1u~-r. J.A., 1982. Oevtlopment oflh•
IS CU>O>m<G.~. C.A., KOCAN. A.A.. 8.-\RKER, k.W. & DU8J\\", J.P•• 2000, Musculru
,h..,p-canfd cycle of S(,rcor:y,;1is 1L11"1/a. C:Onadln11 /oumal of/.cQ/r,g;•.
sarcocyoto.,I~ ,o CO)Olc< from OlJah<>ma. tnrm,nl ofW//dllft Di#!f1tel. 36,
q(t. 2.;&1-2,1;.
il/1-763.
3; l>U&fY. t.P., ~t£lt. C-.A. "4HA9U:!l.'f'O,. \\I,~ a.. 1989. Ultru..,iructural
16 t>At.'11'. or.pt. ~ MARJ{v~. M.1t.. 19So.Sepnrndo1, o( Snre.oc;-sti( species
d1ft'ere,ntiatl0:1 bel\ve-ensru'Coc}"$CS o! Sarcoc;•stlJ hirj'flttl and S~·W
!Proto:tOO! Sporozoo. CoccidiaJ by means of cyst \\';ill ulJ.rastn1tlutt-
/romlri/$. Vt:Mr:llry P11rmlrol.ogy. 3-:. 153-IS,.
Proc.,d111g$ofll1~Elrcrron ,\f!=p;'Sodel)'ofS,,t11ilm1. lfrita. 10, 95-96.
38 DU-9'Cl', J.P.. Sl•UR. c..,. & 1:PUSG, a,.. 1'}82.. '58.rCOC)f:§.lOsiS In nC\\'bom
1; typ~ of
DAL-Y. T.J.M, &-MAJU.'.U.S. \tb., 1990. D~criplfon of n di$tinet~t
r.sl\!CS Ced St,rcocystitcru.:i.sporocy.srs from coyntcS. J\mtrrtcan Joum11l of
SMco()~tis (Protozoa: Coccidiai U\ wnd ungulates. and its slg,tifieante-.
1·,,..,1nal)· R~s,arth ,13. 21,1,-216-l
So11rlt ,!fr/can /ouma/ of5ciurrc,. 86• .Ji.
39 OV8£\". t,P•• SP!ER. C...\., Jal.lJ.N<;~ G..P. <& ll.U>.'T, f..\.. 198,;;, sarror;•uis
18 i98,~. S<lr<QCJ~lJ.S of d<>m1:$1.k und
DAL.\'. T,J,!-1 .. Mi'Qi.;US.. M,6. ti .81~, H,C,
envracanw. Dt.,·elopmcnt In go:us. dogs nnd <O>'OU?S. Jm~martor.al GO/It
"ild equlno hosts. Prm:wdin~ ofthe l!.ll.'t:<ro11 MicrosttJpy s«;,,ryof
nrrd Sheep Rmartl,. 2, 252-265.
Somiwr11 A/ric11. 13. il-72
,;o ouer.v. J,P.. sJ•r;Ett.. (:..j\. ~ l'A,'.f'.M. 11 .. 1989". SnttQCJ'StO$i:t ,,fAnfmnls <md
19 OAUGSCIIIF~, ., .. i\l Tl7ELD~ ~. (.- RO;\nlEt,. ~t., 1989. Hemos1a11c aher.. rlon.$ tn
."WoJL Boca Raton. ~1orida: CRC Press.
p1gs-fedsublelhal dost~ of Sorcrx:;'$ris mi'~cltcr:'n1111. \lcttrutm)·
Parasir.ow10•, 34, 1-13. 41 DUBE\\ ,.r,,-N."1i£R. C,.A., \IU~01\'r', u.t "'UP"l(;Q\11.f, T..r .. 1989. 0\1ne

2o OAU(;S(':Hll:S. A.. Ht~1"%. , .. 116.~~ISG, M. & kQMM£l. ~,.. 2\)(H). Grtmih

spo,awan encephalom)·elius linked co SnrCCX'.)~risinfe;:Bon. V.>1rri11nry
performance. meat quality ond acdvlt!es of glycol)<tk l!n?.ym0> In 1he Para;rrol<Jg.1°, 34, 1o!l-163.
blood and muscle 14$ue ofcalves infected 1,i1h Smcoc;J~I/t mal. 42 OUB£\', J.P•• s;nu:rrEL R,lt,. S'TRO)rtlERG. P.C. II TOUSS..'i.XT. M,J... \Iii-;'
\f,terinary• Pcv~froloizy. 88. i-16, Satc9Q>tisf(l:,,rhp. 11. from 1hc ho""· Journal ofP,m1,<11<1l<,g1, 63.
21 tMUGSCHIES, .\,, MO:.tMEL. >!.. fft'SSl~(;t M. a. KAl,LW!t'T, £ .. 1~90, Slt(CO()'StU 44341;
mtcschnfana: 1\uswtrku"1len derchronlsche11 lnfl'ktlon nu! 43 OU8t'f. J.P.• UDTU}AN, 11.:-..s•. C\."'-~0~.1,. .&; U'.'iOMY- o~.
Condemr.orion Qfbecf l,ecausc-oi ~1,1/s /J/i;ura inf<'<!1lon. /r,r,m,1/
'*·
Flclschb<!schaffcnholts·mcrl:m•I• und die Gc,,iclu~on,"1cklung bclm
Sch\\'t'ln. f'l,/srhwinscl111ft, iO. 200-205, ofrh, {m,r/con l'i-lfri1wryM<!dirol ASJ«lml/J11. 196, 1095-1096.
 Sarcocysrosis 373

..;..s DIJm·. l,P"•• WOMlkOOl:.. 5..(•• SP£P, C.\ &cstl.Aa...tA,, 'lo.P•• 1981. ~rCOC)"SlO~ Chemopropil)'la.,c und Thetopie dcr aku1en S.ul;,osporidiose, B.<tfm,r
in ~oor,, Clinicol Signs nnd potholo~ and hcniatologfr finding,. und M/Jt:chM.r lltttir:1/icl:e Il'otl1,•11sthr:fl. 9~. 229-?34.
Joumnl t,ftl!c Am1•rienu V1:tcrltztU)· .\(tdi<al AJ.'iOefdlfo,r, I :"fl. 683-699. 67 HMYIXII<.'<. ,.o. "i..,IW!II. z.. 1986. 7.llr S<hi,ogonie ,·on S<U'rtX')c<tis
4~ £Mflf.R. -., , nuR\>J;AK1, :.1~, 1981. \\'irtscbuhhcheVcrluMcdurch c.wirams. &rtm~ imd .\fit11d11mrr rJ~mr.:lliclie J•, '<Kir,•11$Cllrifr. 99•
.,..,,~o,porid!os,, ($ar«>CJ'$d$ o,,1car1isund S. spec,) b<-i dc,r Mast ,1;1n UIS-159
SChaillln1mcm Prtt//rls<lr< Tltraru, 6Z, ,122-IZ•I & 126-42,. 68 11noon~1. \.Q. 4 .:1a,1»1. J),. 1996. bol:ntun und experimentrllc
10 >.,,u, w. •
utJkl·-V, ,.r.. 1982. Oe:\.-elopnu·m of Ytrco<:)·.#u /<t,ni in lht Obcrtrngung \'On S<1rco<,1~lls moult'l NC\·cu·l.cmalrc. 1~12.. &•rllm:r u11d
equine. Jnum«I qfl'nra,/tolog:,. 68. 85&-aoo. Milnrholfer 1,rrlir:tlid,e 1,,Vochenschrift. 109. 440-145.
Ji F.\UR. "· "uu1u,v, 1.r .• 1986. Bo,ine sarcoeysto'iia.. Q1mpc11dlum 0,1 69 UhYU()j\S • .\.0." \\£HUJOR,~. H~, 1987. Finc~U'Uct\llC ot Sctrtot)'$ti$
<.0111mu111g lid11m1io11j()r tl:r PrncrW11g V,tm11arir111. 8, PIJ0-1'1~2. 11r/etitt>r1iS l!cydom, 198$ In 1,~ in1ernn•dlatc ;tnd fin~I hqsL< (•heep and
J8 t.\\'tll,. n. • l:.l...<.A~~t!M. t ,H., 1.991. Bo\'int snrC'Ocy:StOSIS'. HO\\' p,1,u.,]{et dor,I. 7.i,nuall)/ntt far /J/1Aurrorog1,..\tll<roo/o/og/~ ,,,.,, Nygltn,, A21,,l.
negau,elya!Te<:1 growth. Pnraslrolog:.• T01lny, 7, ?S0-255. 353-362
?O HEVlX>lt.S .\.O. 4 U',;TfRHou-.:1 R, , •• 1983, Z.Ur Enn,id:Jung \ 'On Sarcoc.,,-ns
,& F!S<:11£11.5 A O::lY.NISG, K.. 1998, Char.,c,c,rt,.. ,fon of 00\11\C Snr~n,
spttJ~ t,y llntll)~i~ of their 18S ribosomal ONAscquNlCd./nur1u1I of hudronis n. ,-p. &rllncr umt >.stmcht11erTfM1r:.rUrite Wocl,e,u-r.hrl/1. 96,
PnNWIOIOIQ', 8,1, .!I0-5,1. 2iS-28?..

50 IITIOFMU>. <.O,, l,\.'<OVITl, t.e.. l'I\Z\CO!'. "-!I.. D1JO~~·, 1.• I, MUIU>U\' D,A., ;1 .\.o., \,f....:1Gr.J1. J.11 .. 1988. El."'IOllisf'iner .\kuu:n Sart.·tx:ystls
11C\1>0S\."I.

159~ Sllreo<;ystosb "1d1 in,•01,-.,mcn1 of the een1ral nutvous S}'Stein in

su//lomlnis,lnfoltlnll 4uf di• M:mlolnung ,'lln ,\l,;,11JerltelrL &-rl/1,rr
lambs.. Touma! Q/Vc.terinnry Diagnosrie ln:•('StiJ:arum. s. 29J-29il. 11nd \ffit1d1M1·1 Tl<'rllr:tlldi, \l'or/ien,chnft. 101.JOl'-310.
72 IIUOSG. L--t. T., OU8[\', J.P.. ~·•m{k'll.A, t. & Ut'.CL\, ,I. .. 1997, Sl.tt,'OcyMls
5l FO<iG1:r-.--. ,~M•• 1980, Sarcocysri.s infection and grnnulomoiou-s :n)·os.td~ m
C'11tlc In Zimbabwe. 7,imbob11vWrtri11n,y /oumal, I I. lH3. bullnloni.f n. sp. lPmtt)J.():1: Snrcn(~-stldae} from the w'tlter bu1T:'11o
lBubalutbubolL,1 m \'it1nnn.1.Jo11rnal ofP,11asi1ofogy, BJ. 471-l'N.
52 rn~u. c,.t., 1974. l'rc)'·pi<'<laton~rnlsslon in 1h~ c111'1001loldgr oro,;,u,
surc~tidlosis. At1SJm/uu, Vt1trlnor;·Jo11mnl. S0,38-39. 73 uvosc. l .T.T.. OIJ8l!V, ,.~. &-UCGlA, \ ,. 1991, Redcsaipuon of Snrcocyst:$
leirinrf DisS(Jnaikc ana K.,n. u1;a !Pro10Ma; S.rcocysudae) of 1ho w;i1er
53 R>RD1 G_1;.. tg&G. CompJcuon of the eyclo of uansmlsslon of
buffalo <B11boh<i lml>nlisl. Jourtwl of Pnra.rlrolo[O', 83. l J.JS-1162
,..,co,potidlosl• bel\'<'l:ll cau and <hrop ,~ar,,d ,pcdfie puth<>g~n r,.....
AullrO/iDII Vctrrilllll)'}Ollf/t(IL 63, 14,,z.. 7J nt.tosc;, l-T T. ~ur.<-1.A. A.. 1999. Sa,coq'stU dub.r)'i n. sp. (ProtozOL.
Soreoc~,tidne) in 1ht \\'3terl)uff;uo 811/x1/1,i bubalis>, Journnl of
.s.: roRE\1'. 1.·• •J,.. l986. Ewh.ration of dec:oqofn;'!;e, ra.sa!ocid~ ;tnd mon\:n....in
Para..<ttolrm . 85, toz-101
J.ga.iru1 Cq,><lrimentall)' induced san:ocystosb. in cah·~l)../antl!rlrtm
73 10.\CUJM,.\, Tt.N'fT;JI, , \1, IEFFJUJi$..A.C, • JOH~w:,;, 1\.\t., 1996.. A
/oi,n111/ of\ ,trrlnnr;· RnMl(/r. .;;, 161ol-1676
RAPD,PCT\ denwd m•rker can dlffcrenlinre bctw,cn p111hog~nlc nnd
55 FA.\..'-Sl'."\, I.LA,. PtGM\'$8. A..D.AX VA~ MOt.. J,;J,,C. '° 00),S. I ..A../1.., 198';'
non-p~thngenic :;.tr(0()1r,sspee,e,; oh~eep. Molewlnrond C.,/111/1,r
sa,ro.,~r.:,·und chronlsche Myopa:hlcn bt1 Prerd<n. R•rllrw, m:d Prot,,.., 10, 165-1-2.
\1fMr!1t•n,•rTlerlln1/ftllt t~rocJun~hrift. JOO. 229-232.
76 u.. 1:983, 5Drt(X.1$tls inrcction In \..itd .. outlu:m
lot\hJ.'Jt, I.A, &- MAkk\J.S, ~,
56 CMJADH<\N....., .. YATM, w.o,G. f< AU.CS, J,R., 19!7, ,~odution of
Afrkon bird<. Somit AfrlcM /mmwl nfScience. ;9, ,171H 71
t'<)s;nophilfc niyosi1iswithan unusual 511«ics ofSarcrJ<}>t/sm a beer
cow. Cnnadlon Joumttl (1/l'n~rii:nry Rcsr.arc/1, SI, 373-3'i8. ;; KAJ.:iER, t,A," \I.UO;u~. M,K., 19tt.;. l">cpt1.nm1:nt of Zoology, Unwi:niit),· o!
1M \\11waiersi~n<1. Joham,e~buri;. Unpublish<'<I dota..
57 r. \>ft.A1Ull1', 1.e•• .;:1otrtl!1'. fl'." FA\TR, Jt., 1.98-1. Hurnottal a,n d (ellulJr hnmum:
responm In cnttle ~nd shocp Inoculated \\ith Sar<0<.J~/Js.,\mmm11 7! ut~. R..G. /. t \fiJ11. R.. 1980. Ampmlium for proph)"li!.'<.ls Of t,vin&
Journal ofl'cttrl11ary R~s.«rch, ·15. 159~-1596. Sttrc(J<Y$II•. /01111u,I ofJ>,trnsirolol!J', t;ll. 100-106.

r.a Citt.U'f-Att f.A.+ Hf'YU01'~. ,..Q. 4. )t£lfUIOet,. H .. 1989,.111(' fine 1-INt;CUIC"Of 79 LEn~E. "'°·• 1986. Tht 1<llConumi of sar<09'ftis (Pro10,oa, Apicompll"-lJ
q,t> u( Sarcccysrls mt1ttll'I from goats Pnmsito/QJ.:,' ll1':st"t1rclt i.5. SP1-"d:e-.S. /011mal ofl'artultoltJfO'· 12. 372-382.
WHIS. an u, Y. &-U.\.'<, :,,, 1911&. ~tudlC$on m'1n-p,g cyclic infecnon of !x1mx;•,,;,
i\f GILE). R.C... lfl.\\10:0. n~. A.. ~ [ L , W,L.. \\'llt'f.\ta;R. ~ MIK~CH. n•• BflV,\,"r Juih~mlnufound in Yunnr,n PrO\in«'. Ch~.Mln 1.oologti;r, .S(11/(o, 32,
u.,,. •-t AnR. n., 1980~ S;srcoqistosls in ,-.1ute in l\cntu,k:1. Joum1Jl oftl,~ 329-33,1.
Am<!rlrtm l'11<•rl11«r;• Mcdlral Atiodnhan, I ~6. 643-5,Jfl, 81 l-f.\C);:[£, ,., ;,., nuon· t "·, 19:96.Congentr..at sarco~'5(0~is in .1 S4:HU\cn
60 obsn.. 1.. XAt:J.. M." tROLR.. ,1., 1978. tlcht• und clcktroncn• goat.J/Jrmrnl of Ptira.,iwl(Jg>•, 82, 350-351
mikro,k~pf-,he llnll'~ud1ungort rnr F..1m,it:klun~ ,·<>n Musk<'l")'<l~n 82 ><ARl:l/S. M,a .. 1911. Depanmf11l of Zoology. Unl\er,ity of rre101ia.
vori Snrtl.>t:)1tls sutr,mi, in Hn\JS$C'hwt:.incn no:ch cxpenmc.n1cllcr Pretoria. Unpubllsl1e1J dalo,
JnM,lion. 7.mtra/b/1111 /t1r Bakuriolog/t. M//dQ//lo/ugh• 11>11/ Hygi~nt,
83 ,aw:u,, ,,.e.. 1973, Ser<>IO(.')' or ""oplasmo<ls. i~poros!s 011d
·\2~ I, 368-383.
<ar<oSJ>Ond,,:,,ls \'tu• E1111lm1d Joumnl of 1/,d(an~. 289. 980-981.
t;1 GOW8~0\'. \'.I~ HVUAl.l"O\'St:lL 0.\'.4r,J.:IS1JA~O\'A. i_.1 .. 1q.74 C3.ffll\'Or~~'t~
8... \L\R).US ,$.ff .. J918. SaraJC"J~rU and satcocystoii~ in tlomL-Stte l.\ntmab.
~u.?Ce o( ~\,..,ni.., 6-UfCOc)'St infts1ation.. Vi!ur/t:arlln. no. 11 for I 9-.:.
1:md mi\n. ,-ld't fllltf!'S fo './tu·rimlfJ' Sdtrrc~ and Cornpamh'w .\fttlfrfm•. :?2..
a.;..as. In Russian!. 1~9-tql,
62. GM.SSTft0l1 O.E.~ RIOU'.\".l\.S:H \'AO, 9 .. C[RSH\\''IS. Lf. & OA.l(".C~ O.J.• 1990.
8, >L\m:U._" ~- 1980. FIJ,., os oruurol twtsport ho,15 of S..rr"')>tl.r and
lminunofluorf!'$t<.-nl locaJiz3rfon ot Sarccu.,ystis rr11tl a.ntlgt'fls. ($?:C. •md
01horcoc:ddla. /oumtt/ of /Jnrrufrotog;r. 66. 361-362..
f~f- In 1e-!!olon, t.1! eosinophillc myositls il1 cnUIC". /lJUrna/ oj\ ·,1i•rltu1r;·
D11tgntm1t /mJ1mfgtt1/011. 2, 1~7-149 86 MJ\R~vs. M,A .. 198i~ Satcocysws,~ In dO.me:'$tlc Ml.mals.Jour,ml oftht
Souti: ..\fritilrl Vturfnllf)',t\S.$(>l'lntion. 32. 350.
63 11r<:>;O\OTII, A.I\. Xi f)l.~'ft:.1'. ,\.M,. 1999. Oerclopinent anct \'llHdtHiOI'\ of
~P.l'Cit-<·•P•"'ilie ncsu,d Pens ror diagnosi, or acu10 sam>L')'ouosls In 87 ~.\JU.:.U~ !lt.u.. 1982. S-t,rrocysos m African bmb and mammrits: bpecificit~·
shtep. 1,11emmlottal}oumal/orPomsilolOCJ•. 29. 13-~I-1:!19, for the int<'l'lllcdl~1e ho,1. Mol«U//lr<111d Blocl:Mnical />arr,s/rol~. 5
tSuppJ,J, 3,;5,
64 Hf..D1Gt.B. 1:..., 1915. JJothotogJcat fnc:--lt£ig<1tio,,simoln1,i;t,•l.ri?. Prttori~;
GoYcrnmem Primer. 88 MARXU~," u.. 1967 T~rni, for cocadl~n mero>.oltes, .4nnnl~ o/Tropl,:nl
,\/udlrmimuJ Pt11'(1Jifl)/<©'• 81,463.
6.5 Jln'DOR!lo. A.O. fi HARALAMfflOI!,,, s. 1g82. Zur i;nt\,ic.klu~ \o"On Sartocysris
<aprt1t<111U Flicher. &•rim" un(l ,\ll/11cltt11" n,r11r:rlldw 69 .)U.k.;us.. M .th-N su~tt, 1.s.. 198;.Stalningofcoc.:ddl.il oocy~L.\. The
l\iK:herisr/1rlfl, 95. :?65-2n. Vete,int1r:,• Rcn,rd, l ?1, 3:?9.
66 Hf\'DO"' A,0,, IIAAALAM&IOtS, ~ .. MATUSOOiA. F.R., 19$1.Zur 90 M,\A).1J,, ,: t\, I. ISU!itl1 l,K., 1987, ~partmt1U of?.ootO,b.')\ Unh·t.~Jt~·oft,h e
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91 ,: un.::u~. '-tk. ~ t>u , .• t .1.~1 .• 11,S.-:.:.. Dc!pa.runcm of Zoulosr?• Unr,~...rsit}' or
the W1r,,\,h~n,.r.tnd, fotwnn<"-burp., UnJmbl~slu..-U datol ttS om '-IS\',. i-; .. t'-198. Tiw prl':,,,C*nl ,:.ice c>t sp«H.~·.s\'\lt'mntir.s In .SnT\"on-s1i1.
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of :ittrtl>t')'$li'!i of tht• nttmnuun J.l!br.l &11111~ ~httl lu,rrmtumn~. Sm1tlt
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98 uhuto (~;.p.~>..PP• i\." fin;1I
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1AA1 Anti~'1111~ c-h,,mctc:rl,:ulon or monodonal ,1. 198;. Zur KJinli:. l'. nzun-;d.ti,·ltJt uni.I .·\mll<orperhlldung hl+t
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107 ,:O<iltl, o.J. &- ,1.,.1tt..:U1t# ,1 '8., 1o181. Oran1;e-Gr0\1: Vctcrtnncy Ho~pha.l.
129 SMnu. ts:.." m rcnun. t.,· .. 1Wlb. t:q)ttim<>rn...il nllCmcysc Sarrv<)·s1;s
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Pal'tbilo/1>~. 22 :;03,,.513
U2 \'A.~ DlR.UJG'l 1-l,, \t\R>.-us..n.n,. ,;ra...'"Hl\t,;.. J.P.~-0..\l.Y, r.J..';I.. S9£H
l3fS Tl'TLK, .\, \1 ~ l('JH~"SO~. A,\S .. l~l. (:0mprtn..~Jl OF m 1r11,n trtUhfat1on Xc.'C'to:ic t.•ntt.•phJlitb. a_~ :t nmn1!c:n:ulon nf o.cuu! ~rcocys1osis in CA ti Ito
product5 ot S<lrcc,ry.wis a,wmr1.'l1 und SarcQc;)'-Stl.$ ll'llttlln.1111,mmlioual Journal afth,• Sl)ftt/t ,1Jr/rw1 \ ,1.ritla,y ,lsso<ln1/011 65. 119-1 Z I.
Jmmlfrlftir Pl<ft/,<iro/oyy, 22. 1r,3-11;.i.
143 \\T:~MD[)I. JI.It om ,t:-.:t,, K.• w.u..TrR.c;.,. ancxtu1urr. 1.. l995, The
u; n-,ntt. .,. ,, u rm1X"-OX, A.M., 199;. Phylogeny ()I lh(' tl~su~ t1'Sl· f1.,rmtng hlnrk-ba,ked iurJ;al (C:>rnl\<ira: C:anldn•) In '<on11bi:s: tn1crrnedla1r h<»t
co<...:ldf,\ .Mwmrw i11 Pm11.,fra/1•io·, 39, 69-139. ol 1wo &,r'-"h.")'.'ifk-;oipttle\ Wrol07Am: ~areocy.:i1idac). l'ltraJ.ltt'. l,
138 n,'TfJt, ,. \1., \'lt,l~UYfift. (' .• nttJ~:,ttL p, ,II ffO\tMU.. ,1., ,991. Ot!lB'CUO!'l of 391-3~-l
,pt•rit-..,,:o.J1t-clfh.· ;md -.·tO"S.>•rt-:i.rth·~ cpl:optho. in .5<u(«J"lls cyit()~ltt•, b~-
\·,x .. u1w•• n., un,s. x.w .. u11;,. r. £.21.L\.X<.i. v.v., 2001
144 \'A~<,. t.t).. iuo.
mnn<X'lonal ,rn1ibndh:>, P11wn1n1~, 1/J?sulrrh. 77. 212-:?lt;,
ldenillk3tiun or Snn"O(J~tls lto111f11u-hke Protozoa: S:ucocys1rua,1 ,,.,.,,
lJ~ rt~TI R: ••, ...1.• ll~J!>1hN\1.\:\. t:..L.• fORSSOS . .\.M •. 1991. Sl-,,.irJU(U'I of In watt< buffalo 1Hlllhd1u /111/1,11/.ll based on 18S rlt'l,\ g,,ie s..oqucncc,,
,.UUJgun, from 5'JTt'04.,,$IU ')petfes Osi.og (hrom-,tofnt."U,itt~. Jtmm,,I o.f Jour,u,/ ofPar,u1t0/0l!J·, 87, 93•1-9J7,
Ptumitalo.~·. Ti i2i-'i36.
1-1$5 \';\,""'G. I~~-, /IJU, '1 .:< ..
\ \0, \',G,. C:Ht,, ).,\._., \'.\'.\"C,, Ct<: 6' :tKA~G. \,P,, !N'>OI,
140 n,.~11. •U•• UUOLV. J.P. w~.as. D,)1, . WL.\..~Clti\JCJ),i.w mcrn. I flUT'I., 0.4' \n•l)<I; of th• 18$ rR:-~ gene, of Stlrt0c,,<1i, sp«ll"' <ugi:1:,1~ ch~11he
1n,,~." ,,•. tg~g. FntoJ \isccraJ -.ind. neurnJ 1.1rco~·s1o~rs in Uog':",,, Jaur,u,I morphologic.all) "mllarorg:ini,n» lro,n ca11lc and Mll<'r bu!Tnlo ,hould
t1f Cflfllpl1(0/lt" PnrholOf!J'· IZl. I ;9-18-1. be cons.!dt'r,"tl du• ~me ~J>e(lc.:,ii. A{ot,•culurt1r.d Bi«-lttmrt·at
U.l TR,\\J.•D,\RGAT7., f,l.... ~r.llUPf, f.W,l,. CiM:\~TR())f, f>,t-..., f~r.,t..\'\1,f.T • Pnr,m:oh,g:,, I IS, ill:l-:!86.
 21
Balantidiosis

B 'f SPENCER

Introduction subject of incidental idemifkation In histological $!"Ct ions of

the colon ofpigs.'· 111
Baltmtidium coli is a ubiquitous protozoan of the class Cili-
aca which inhabits the lumen of the large intestine of many
Pathogenesis, clinical signs and pathology
pigs. but may also be found it, other species of animals, in-
cluding guinea pigs, dogs. rats. non-human primates; a,id Under normnl circumstancer., B. coli li\·cs in the int<.'snnal
birds. as well as in humans. It isacommensaJ but under cer- com<.'nrs and on the mucosa of 1he forge intcMinc. and docs
tain circumstances, may apparently act as an opportunistic not appear t<> have any deleterious efl'ee1 upon 1hl! host. On
pathogen. occa.,,lon, it may fnvndc the ime,tinal mucos;, when 1his
becomes damaged by. for example. a primal) pathogen
(Figure 21.1 ). In sornc animals 8. coli will penctra1<.' tO the
Aetiology and epidemiology
submucO"il or even :i~ deep as the muscular layer. 1 The
The ,·egctarive form. or trophozoite of 8. mli is a relatively organbm produce:; hyalonurid~e which facilitate, ii>
large. motile, oval to ellipsoidal organism val}1ng in size penetration in10 tis,mes. 11 The resp<mse 10 the tbsue
from 30 to 150 pm in length and 25 to 120 pm in width. [ts lm-asion i,; lnfl:immmion of the pan concerned. \\1th the
surface is en ti.rely covered "ith cilia arranged in slightly ob- de,·elopmcm of supi:rficial and deep ulceration. Cllnical
lique rows, and internally \\~thin the endoplasm are two maniftbtation, in :.uch affocted animals are 1hose as.ociatcd
nuclei, a large kidney-shaped macronucleus and a much with mild ro ,everc. 11\'en fatal. emeritis. 1 The faeces maI be
smaller, spherical micronucleus which lirs in the notch of bloods1aincd. In piglets. anaemia. emeri1is and decreased
the macronucleus.9 The nuclei are readily identifiable and growth rate ha\·t· been reported. 5
their presence assists in confirming the identity of the para-
site.1The parasite also contains one contractile ,·acuole and
several food vacuoles. Reproduction is generall) by 1rans-
- ,::;
' .,
t ..

,·erse binary fission. but conjugation ma) nlso t3ke place."

Cysts are produced. These are ovoid m spherical and
measure -15 10 65 ~im in diameter. ihe peroxhome, of the
uophozoites of B. coli found in subclinical cases are $maller
than 1hose isolated from pigs \\ith acute balantidiosis.8 The , .
organism can be recognized within the cyst by the macro- • p \ ..
nucleus.q \
Both cysts and rrophozoices are passed to the external
environment in the faeces. Generally, transmiss ion is by in-
gestion of ~-ysts which may remain viable for days or \\'eeks
.. '
in moist pig faeces, but crophozoices may also play a role in
this respect.9 Trophozoites. howe.,·er, ure much less resis-
tant than the cysts.9 The faeces of sows are the major source
of infection for piglets. 5
..
• ,
...•~.
,
No reports of clinical disease caused by B. coli have been figure 21 .1 Balant1dios1s ma p,g· !Jai;in/ldium roh paras,tes n lhc
recorded in South Africa.1° but the organism has been the mucosa o' ·ne colon
376
 BalantidloM~ 3"

Infection of human~ Is usually acquired by the contam· genesis of 1he di~easc process must be borne in mind: the
ination of the hands or food with pig faeces.9 In some !act that the parasite i~present docs not nccessarilr indicate
humans harbouring the parasite. 1here is s<:'1 er.• diarrhoea that it b the primary cnu~e of the condition unless additional
or dysentery. Ulceration of the large intes1ine may develop. e1•idencc can be provided.!'
which can be complicaced in some cases hy hnemorrhage."
perforation of the gu1 wall and peritonitis? The ulce~ arc
Control
similar to those of amoebic dy~enter, .i
Tctracvclinl:'• have been used both for the rrc:nmcnt of clin-
ically affected animals and for the prophylac1ic treatment oi
Diagnosis
pig$ so a~ to pt'el'ent transmi~slon 10 humans. The addition
The diagnosis oi frank disease caw,ed by 8. coli iniection is of 100 g o:-.·vn:1racycline per tonne of pig feed appea rs 10 be
ba~ed on the clin kal :.igns. lesion~ and presence or large eft'tciem in clearing the faeces of B. coli cysts .5 Me1ronlda-
number~ of organisms in affecu.'d area~. Before making a z11lc at 30 mg/ kg twice a day for live day~ adm inisteccd per os
definitive diagnosis of balnmidiosis, however. the pntho- can be used for treatment or dogs. cats8 and humans.

References
,\LMl:Tlt\\ 1.. , Ht<i3- Propagarfrm t1f b;1l,1mhJlum 1r1 the porclne tntc.•,rlne i ,\,t\1t,\ H ...\ .• l:l•t:.c)J1\H\ A,11 ,i. PflV.. \.,\...J~l,. lO\OJ11:Ut1tt1,h,
\i.wrinn')' 11.-dfc/t,_.., 16. ~. halnnlidfa~~.. .and :ame-bfa"ihamong_ ,oo.animilh, und mnn. iUSlnt
'..! o\~atRS<>s, W.,\.1>., 1971. /l(UJ,c,fc,gy, 6rh t.tdn St l.t1Ull\-<: n,~C \ ~tt_h,h~ VJ.1l••Tlm1ry 'l•'11i,ttf Jm11t1(.JI. 29, J29- I:{5.
Cotup.111) II ,~.,, ,~c ,_,., a .. 1997. \Jlln<-'l<U(turnl ,md <11c>rnt'llllrnl id1•t1tllk,11inn or
J WO;\'., r!=t:99-. 8f1ftmtUltu.m c(Jti h0nwpag". lm:f/cnl.,·cuipt!nn.cdu,; ~:a~· p<·-ro'<hf)nll', ,n Bnlrmrfdlum ,·uli. Cillphora.. f.'CJJ,n bioloJ:,fro, ·15
l999. ll'i-120.

,t corrT.rn. J.,,.w.• 199.:t. F,1,u1t} orVt.:1trlnt1ry ~l\'n,i:. ~,m,.x•r.,1~· of q ""m -"D\. r...J.L .. 1~~. Jlelmi111lt:r..-,nh,n{N>th mu/ PrrUtr.A.lii rJ/
Pretoria, Unpuhti,hoJ dato. /k/1/WJ//t'(llls/ ,lllhllOb c;1)t Cdlt, l.OlldOI\; 1lall1,•r,·, 11nUl\l ~nu C,lS.Wll
I tel.
5 ti·'~"'· "·"·· en.'°'·... tt.n .• ,tf ,"t.1:u,.-G. "'-' .. PCI\'\"\', n.11.1... $<'B<"H.t. , 4·
~l'R.\\\ 11., ,g.a,, l)L,,.'fll•·~(J/S:, .. ,:r~S1h cdn ,\mt"\, luwu lo\\,, '\taJ<"' 10 ... rr,nu.11.1 .1~ Fai:uh)·o(\'Nl.:on,U') St:h.•ncc.·. tlnl\l•f'il) ofPrctnria.
Unh·t~it;· Pre)..,., Unp11h!l<hrd d•••
6 1'1'.\ffE.UIO~ .\f.fl\C- UM\. i\t "V[ u,u.'.\: \ 1 Jl)91 1:Jh.tl ~.-;,,.-.e nf ~nt~1inaJ 11 ,a '1J1t.11r. . c,11 . ..... 1 IN ,l;n, M.(, .1957. Thl"'prodw.-unn ofh}'Jh1f011hfaie by
bauittldia.,l, Rm•/J;ta tlfl Soc/,da/c 8,t«lft•u,11/c ,\lrditind Trlip1tYli. ,3 &,Umtftliumcoll. J::r,>t1rtm,•11wl P,11w11vlo~ 6. 31-36.
1''3-176.
 22
Leishmaniosis

J J VA :S DER LtJ GT AND CG STE\.\TART

Introduction malian hose and these are engulfed by phagocytic cells be-
fore transforming into amastigotes wl1ich multi pl}' b) binal')
Leishmaniosis is caused by obligatoi:1 intracellular parashic fission. Healil~· parasitl7.ed host cells eventuall} rupmre,
p rotozoa or the genus l.eishmr111ia that affect various mam- and 1he liberaced amascigotcs invac\e other macrophages in
malian hosts, but di,ease occurs most commonly in humans which they may be spread locally or syinemically.17 In tis•
and dogs. Cutaneous, mucocutaneous and visceral fom1s of sues, amastigo1es of alJ I..eis/11nt111ia spp. are morphologi-
!he disease are recognized: the visceral t'orm being chronic cally similar (Figures 22.2 and 22.3): they are round 10 oval
and of1en fa1aJ in humans and dogs. bodies. 1.5-3 x 2.5-6.5 µm in liiameter. containing a single
l.eisllma11ia spp. are widely distributed in Cen1ral and nucleus and a rod-shaped kinetoplast. 17
South America. l:as1 and \\"est Africa. Asia. che :Vfediterra- Leishmaniosis has rarely been reponed in livesmck.
nean basin and some neighbouring European countries. bu1 There ha\·e been reports ofinfection in calves from Africa in
they are rare in southern Africa. The cutaneous form of 1he 1he eatly Ji1erature. but no details are given. 13 The cutaneous
disease has recentl~' been diagnosed in rwo sheep from dif- form of the disease occurs frequently .in horses, donkeys and
ferent localities in South Africa. 1s, ~i mules in endemic areas of human and canine leishmaniosis
in Venezuela and Braiil?· ' In 1989 and 1990 cutaneous
leishmaniosis was diagnosed in sheep in South /\irica - in
Aetiology, life cycle and epidemiology
one instance in two adult ewes in a flock near Barberton in
~·!embers of rhe genus Lei,Jmumia, which comprises a con- i\·lpumalanga, and in the other in one sheep near Greycown
siderable number ofspecies, belong 10 the order Kinetoplas- in KwaZulu-Natal 15• 22
tida and family Trypanosomatidae. 11 Identification of Canine leishmaniosis is endemic in the Mediterranean
species within the genu~ is complex 3Lld is based on clinical countries, and isqccasionally seen in North Europe and in the
and pathological features of che disease, geographical distri- USA in dogs thai have previously been in e11demic areas.:?O
bution. morphology and behaviour of the parasite, and 1he In Souch Africa, visceral leishmaniosis has been diagnosed
resulls of biochemical and immunological 1es1s. 0 C\,tjce in dogs: iu l~64in a dog from Durban whose life histOl")
Diseases caused by I.eishma11ia spp. are trat1smi1ted by is tmcenain; 18 and in 1987 in an animal from !he Free Stace
blood-sucking sandnies of the genera Lwzomyia in the :'Sel\l which had never left the cotmtry.2'3 In Namibia, 34 cases of
World and Phlebotomus in the Old World. The disease in cutaneous leishmaniosis in humans h;we been diagnosed
dogs is caused by L. inftmwm. since 1970.q \vhile only a single case of the cutaneous form
There is usually a primary reservoir hos1 for a given Leish• was reported From the Nonhem Cape Pro1~nce in 1979. 1
ma11iasp. in a panicular area \vhere the parasi1e is maintained The epidemiology ofleishmaniosis in southern Africa ls not
b)' a cycle between sandOies and the mammalian hosts (Fig- known. In '\'amibia, npnn ftom humans, l..eishmanin parasites
ure 22. l). Secondary and incidental mammalian hosts also have also been isolated from the skin ofhyraxes (Proawia cap-
become infected.? Both domestic and wild dogs (Lycao11 pic- e11.#.v} and from naturally infeeted Phleborom11s rossi sandflles
rus), as well as rodems and other \,~Id animals. are reservoir (l~hich are possible vectors of !he human disease). The isolates
hosts. but occasionally humans may act as the main reservoir. made from humans aiid sandflies am idcmical. bm are dis-
The parasite occurs in t\\'O forms: a promastigo1e stage tinctly differem frcun those recovered from the hyraxes. The
which is found ln the sandfly: and an amastigote stage which two Kamibian strains of parasite are also dissimilar to
occurs in the mammalian host. Duri ng feeding. the infected [_ rropica and /_ mnjor which ~-ause cutaneous leishmaniosis
sandfly injects promas,igotes into the dermis of !he mam- in the Old World and are afforded separate sta tus.9
378
 l.ci:.hmaniosls 379

,~ ,
. (i''

Figure 22.1 Ve cvcle ol a

leisilman,asp
a: mamma!!an hcs:s
b= Pillebocomussp
c = promast,go,e
d = macr011hage containing
amasugotas
e ~ amaS'Jgote

Pathogenesis, clinical signs and pathology in the course of a generalized dissemination of the parasite.
Chronic renal failure due to an immune-mediated glomeru·
Leishmaniosis in humans comprises several syndromes de- lonephrids is the main cause of dead1 in affected dogs.20
pending primarily on the species of Leishma11i11 concerned. Clinicallf. canine leislunaniosis is a chronic wasting dis-
The clinical spectrum ranges from aS}mptomatic infections to ease accompanied by anaemia. imcrmittent pyrexia and
those \dth a high monality. Lesions may be confined to the generalized lymphadenopathy. Skin lesions. of which four
skin (eutaneous leishmaniosis) and result, inter a/fa. in nodule fonns have been described, are common: symmetrical
and ulcer fom1ation. known in some pans of the world as the alopecia with dry desquamation; a generali;,;cd nodular
·oriental sore·: to themucocurnneous tissues (mucocuraneous disease: ulcerative dermatosis: and a fonn characterized by
leishmaniosis, sometimes referred to as espundiaJ. in which the presence of multiple pustules. Keratoconjuncrivitis.
the organisms are found in endothelial and mononuclear cells epista'<is and diarrhoea may also be present. 8• 12
of the skin and mucous membranes of the nose, mouth. phar· At necrop:;y, in addition to the lesions visible clinically
ynx and elsewhere and may result in severe disfigurement of in dogs. there is gcner'dlized lymphadenopathy, and
the invaded tissuc:s; or they may involve several tissues espe- hepato· and splenomegaly. Microscopically, large num-
ciall~· the liver. spleen, bone marro\,, and lymph nodes Ct.he bers of macrophages lilied with the amastigote form of
highly fatal ,isceral leishman.iosis or' Kala azar').2 L l.eis/mul/lia are presem in the spleen, bone marrow ancl
ln dogs, the parasites multiply within macrophages and lymphoid tissues. In non-l}•mphoid tissues, a granuloma·
other cells of the mononuclear-phagocyte-system and cause tous inflammatory infiltrate composed ofparasitized mac-
chronic inflammatory processes. 5· 1, Ca nine leishmaniosls rophages. plasma cells and lymphocytes is evidcnt. 12
is a systemic disease, and, in contrast to the simation in l.eishmaniosis was diagnosed in cwo adult sheep born
human cutaneous leishmaniosis. the skin of dogs is affected and reared on a farm in Mpumalanga. The clinical signs in
Figure 22.2 Le,shman,on
11 a sheep: numerous
macrophages in the skin
containing amast,go,es a:
a leisr,mania sp.

Figure 22.3a Group of le1Silmama organisms 1n a macroimage Figure 22,3b H1gner magnilicauon Ix M 000) of a s,ngte Leis~man,a
11 '2 7SO) !By courtesy of Ors J Soley and S Ya1es. Oeparttneot of oisanis11 1By courtesy of Ors J Soley and S Vates. Deparunen; of
Ana1omy, faculty oi Vetenna~ Sc1enta. Universuv of Preto:,a. Ana1om1 Facu:1Yof Vetc,tinary &.1en~. tlni\e1s,w of Preiona,
On:le!Siepeortl Onders1ciwor.1

both cases were similar. The pinna of one ear of each animal Another Ca$e was diagnosed in an adulc sheep from
wa~ sel'erely swollen and the overl)'ing skin diffttsely thick- Kwa2ulu-:"\ntal. It $howed alopecia, hypcrpigmemation.
ened and covered \\ith a crusr.y exudate. Small crusts also htperkeratosis and crust formation of the skin of the
oc~·urred in the skin around the external nares and eyes. and dor;um of lhe nose and both ear~. and around the e}·es. It
in that of the upper lip, muzzle and lateral aspectS of the also displayed a focally disseminated a lopccia and the
face. Both animals reco\·ered spontaneous!~. presence or nodules in the skin or the ventra l aspects of
 LeN1maniosi~ 381

the limbs. abdomen and base of the tail. Its rec:ove~ was indirec1 fluorescent antibody test and the enzyme-linked
spontaneous. immunosorbc-nt assays the most suitable. Serologr is of
less value for the cu1aneous and mucocrnaneous forms of
ihe<liscase.· 11 JG, ,~.211
Diagnosis
The diagnosis of leishmaniosis is based on identification of
Control
thc agenr and serological tests. Demonstration of the
amastigotes in Giemsa-stai.ned smears of splenic. bone mar- Pcnia,·alent antimonials. cspcciall} mcglumine anti·
row and !)'mph node a;.pirates. and of skin scrapin_g,,. give;, a monate. are 1he mos1 effec1i-.c drugs for the treatment or
positi\'e diagnosis, while an indirect immunoperoxidase canine leishmaniosis. With therapy, 1hc major!cy of cases
staining technique of paraffin-embedded section is also in dogs show rema rkable improvement. provided renal
available. If the infection is oflow grade, detection is possible glomerular filtration is not seriously decreased. Treatmem
only byartemp1ing in 11itro or in 11i110 isolaJion. tsolated l.eish- Is, howc\'er, expen$ive and relapses arc the rule, \,ith the
ma11ia organ isms are characteli7.ed by isoenzyme. D.:-:A and development of complications such as iridocydilis or end-
antigen methods at present. Serology is the preferred method stage renal failure ht:ing C'Ommon. E~Yec1ive vaccine<: are
for t.hcdiagnosis of,~sc.eral and canine leishmaniosis \\it.h tl1e not a\'ailable.i0

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•.-<>s. 198.1 77,d,.,/J/:11w11i1U<"J Wo,l,ltfoahh Ofll"nl,ation. c'.tntl'&. 13 1.t1-,.U'I (..J•• 1~6. lnjt'(tuJ11~ ·1 rwilNII fJJ>~v,,a"/ l)nn11..·nrc .\11/mnls. Q~'(..
Technical Repon Scrie> 101 England: f.9ngnmn :.c,rmlric « Te!'hnic,,l.
~ •:.o~ 1990. Crmtrol of ,1,., 1..,lsJ,manlosiJ.. World Hcahh OJ\tan1ta11nn. 1-1 ''"'""\.' ''Tl 1·.• ~,tco:,,., ,u ..-POU,.,,.1993, Comp,'ln~onbrtwee:um
Gcnl'\si. Tcchnfca! Rcpo,i $en"' 79"3. ~nz)·me•Unked 111unv119wtbt.·m tb").lY u~mg a dca•rge.in•solubtr
3 ,\QlJJL\R C.~1 •• R.\.S(;tL J_f. I. OE.\M L~1-, 198t;. (;U11UlC(lU:\ lt.<khffll.lflii.L'oi:,. L.•IJimrd>1it1 ,,,pmwm A.nt11ten and J1rcc11mmuno1luo~cncc ioc the
b frequent in c-qulne.s from un ,ade,11ic ore~ In Rio <I,, l>-'1eiro. Br.uil. dl.a.gnostS of c.,ntni., lci!i>hman1o)i$. i ·c·t,,..rwu.,~· l'nrasitulog),·, J9.
,\lem.Orlm doJ,rsutuu, Osn..yddo Crtt::. 81 471-172. 13--?I
.,: DQi\r\:Offf,•(iAf\RIOO. P,, MtU:Sl)C/' LC,, TUJ'-IU!S, ti.,\~, MUIUU O. ~.C: 15 \l,\R.")IUI 1. c·•. •Mo. Mcd1t~.,1 Ut11\'t!r:SI~ ofSourhi,rn, \frir:a. :Wkdun~, .
i.\ H:Rl'tWSOO, ('..(;. ~- UHl),\,\l rA.. 1.. HJ81. E.rlzt)Olic t."<1uine C U I ~ Sour]; \frlca. IJnpuhh.ticd oh«:rvutlori,
lui..<mmnniasb h1 \'c1ur1.utla. T111,i.•mc1trm1, o/tfw Rc>ft,I Soci'ery of Tro11fCl1l
16 \L\'flll~. ,\, OIi t}U,L\J.I ~- 1•.. 19'95. PCR ISnlf fn t•;no C'Ulli\•UUOft ror deltctlort
.\1i!dfctnttmul 1fyg,·c111c. 75. 411
of u.,;slum.mu, 1-pjl, in drugnosuc fflmpll~ ftom hum.J.1t., ru,d <log,..
s cn,,1;. it.P, 1·0:-:c.., n. & 11&" k.~.. 1985. Rlology or l..c,,hmmnaand /numnl of<:l/11i,nl .\1,rrt,f,/otogy. n. 11-15--l !4~
les..;hmanfa~l!o. /11: OM ~r.. ~r. 4, an.w, ~~-. rc,dsJ. t.u,sJunrm/tlJ;~,r·
17 MOns1~ 0.11 &. AlllJl.11:•n:,,.uJHlK. n. t!;J8:- Morphotu~:. ultrru,.truc1w~
Arn>1er<b:m1 EJs~viN SClencc: PubJu;her.s.
.tnd life C)·tlc'. /,r: t•Ln n,, w." MU.let..~);£ '-O>tJL \;. 11 .. (t:d$), T11t
u oc.,.,,tA~. \\"'. 61 HA~SO~\ w•• ,9&.2. tei~hman1a!il\. In: c;Ru,. c.. :L:d.u /,;>11/Jmnnln<t Ill Rltllqgy mu/ ,\Jtd/dttr ():Janda: .cadc·mk Pres-
C/1,:,a,/ Mlrrobwl<>f:!•u11d 111.t«:Nou.s OJSt«s.h ofthe v,,g 011d Ott,
l'hil.idelphln. W.B. S.1und•r<. 18 Nl'""'· 1.r. .. , ~ Stnc~ \•t11mn.irfan. P0tg;et¢:'T\h. "iou1h ·\!nca.
Unpublr-h<d .i..10.
P-lRPEJt, L, P--'\8.\~'At.. k.:,it., 00,11.,<,,0, ,; .. lt."iO~. J.\ ~ tO,Q.£\U..\.. 1). 1~83.
ldcnnficmion of l.t:uhmr,m<1donm·mn-ama)tigo1t-s in cunme u~u,t!S b\' 19 1rn.1.a, L 19!tl:l D!;1~n,>,h, of ,mtlnt."
ftOUJI..\. ~. '-.\..\011,. \..,. n

immunopemxidase Sl31rung. Rts«1r(h lit t~trrittmy ~druu.. -n lt!hhmun\J...,T> f'1 J pol~nwr-~h~dia.Jn re,-;,cttnn tl·chniqul'. 1'1u· Vetf!rir:i1ry·
!9-\-196. /r,'C'Qrd 144. J6:!-2t;4
IS HRMflt, t., fl.\flf\"1r\.l,.. H.., fO~Of\'tlA D RA)IO:,,••••, .. 4' oo,11sc;o, ',.f., 1 ~ . :to ~t.APl'f."-:l)(J,... rt.I- 11JS&. Um1nc lt.'t"l'ho·um~-,;, nk' ~ •''""'""' <Junrr,~I},
SI.In k,lon, In atnlnc lelshmania.,ls, /ormml o/ S1111rl/Anlm1t, Promc..•. ID. I-In.
:?9, l8!--3811.
!!1 <;QtlL<tob,·, Lt.L.. 19&9 Jlelm,11111,. ,V1i1uJIJ()d, mid Prvrfl:JHt f>/
9 <,KO\"i., ~ .... , 1989, Lolishmnnlll1tl~ tn South WcM Afri~n''=.unlhla to date DQt11c.~ticar(d .,t11im<1h ,~[tlh f~llrton t.>f,\tnt1mK~
S0111/1 ,lfrr,011 M1:1trC1,/ to,mwl. ;;, :!!l0-:!92. t..tttrlnt11J Hdm1mhr,ftJ~·p1ul Emamf11hGJ • lohdon: 8o1'11f~tc.
10 1..us,o~. K..& .!-MAW. 1.1 .• 191';. E\'Vlmfon.das.slficution nnd ~1.•ographicaJ nnduli ,... ("a_...~d.
1'hr
di:,.lrihutfnn. /u: Jll"T1 IL~. W. &, £1U1Ct.".·S:0:01UO:. lt., {00$i u \"i\'\ urn I vcr. t.J • r..,tn \1J~. J.1 "01: \\'A.\L n.t .. lif!;C!. Cmant.*<lu,
1~iJ/mUluitl$/, i11 Bil)fos._,y,wd .Hrtll(i11,. Orlandn, t\<adc'fltk Pro~. tel~hmanin.,1, In o sht:cp, /m,r,wl <>I,,:,, ':-4111/J .·ifri«m t'ttrrfttnl)
U LI:\'l~t S".D,. C<)R.USS.. f,O. cos.. f..E.G. ltr .u., 19-UU. A ne-,vly r1.1,i)1..~f ·b.«1nttti<>1: 63. ;~ ..7;
classifiri111on or 1hi, prrno1.oa. forimul ofPttutmml~l{J'. ;t;, 37-58, 2$ \'UTT·' · ... \lUliUJ~~J~t., II ,H • SUU'l f, f, '1 U\~lt\'.\tUQ, 5,..,, l~':IQ. fatuity
12 I UXC.:t"T UH~,.,. ti t.o,. M .W•• tsR$. 1.elV'lmantn-"h Jn dt,g-,.,. ll!Ia."rina0· o(\'c1erinory Sc!cnc\!'. lJnh1ni-il} or Prc-11,rin rretori.J \Jnpuh!tshed
.-tnnt"11. 2.S, 358-36i. uh,'lt.'f\"Ut.CJn".
 23
N eosporosis

JP DUBEY

Introduction arc similar to 7: gondii tachyzoi1es except that the rhoptries

r.
in :V. ca11i11wn are electron-den~e whereas in go1l(fii the}
Neospora ca11in11m is a reeenllyrecognized prolozoan pata- are electron-luccnl42 • -i
sile of livestock and companion an imals. It is not a new Tissue cysts are often round 10 oval in shape (Figure
parasite,1 2 but until 1988 was misd iagnosed as Toxo/)lt1s111a 23.4). up to 107 µm long, and have bt-en observed only in
gondii because of structural and biological similarities.34 neural tissues (brain. spinal cord. nerves :ind retina)34 11ilh
The disease was first recognized in dogs by Bjerkas, Mohn rhe single exception of a solitary tissue cyst in rhe ocular
and Predsthus 15 In '\orway in l984 but the parasite was musdt>S of a foal. 72 The tissue cyst wall is smooth and up to
not named. Dube)' and his co-workel'S''14 subsequemly 4 pm thick. presumably depending upon how long the in-
described the parasite and proposed a new genus. ,\'eo- fection has existed. In most tissue cysts, the cysr wall is 1 to
spora, \\'ith .l\l. ca11i1111m as the type specie~. //1 vitro isolation 2 µm thick. Septa are absent and there is no secondary cyst
of the parasite in cell culture inoculaled with tissues of w11ll (figure 23.5). Bradytoi1es are slender (6--8 µm x 1-1.8
paralysed dogs led co newer knowledge of ics biology.lll pm) and camain the same organelles as are found in
tachyzoites except that there are fe11cr rhoptries and more
periodic-acid Schiff (PAS)-positive (amylopec1inl granules
Aetiology and life cycle
in rhe bradyzoites. Tissue c.-ysts may degeneracc and cause a
Neos/)oraca11imm1 is a coccidian parasite, <.losely related to host reaction (Figure 23.6).
Toxop/(1sma gondi i (Figure 23.1). Dogs are both an i111erme- Dog, fed tissue cysts ma) shed unsporulated oocys1s.-6
dia1e host and a definitive host.34· 76 In addition 10 dogs. Oocysts can sporulare omside the host within 24 hours.
cattle.31 sheep,37 goats, 1°· 33 horses,'lff dcer"19· 101 and water Sporulated oocysts contain two sporocysts each with four
buffaloes (811ba/11s b11bt1/is),31 • s-,. 59 are also inwrmecliate sporozoi1es. Neospora cm1i1111m oocyst$ are 10 to 11 pm in
hosts. Antibodies to S. cr111i11111n have been found in foxes diameter (Figure 23.7) and are morphologically hidistin-
(\111/pes vulpe.s),23 coyotes (Canis la1r<111s).69 camels (Cam- guishable from Ha111111011dia lzeydorni round in canine
e/us dromedarius) 55 and white-tailed deer (Odocoileus llir- faeces, and Toxoplasma gondfi and Hammondia ilammondi
gi.11/a1111sJ,39 suggesting that these hosts may be natural in cat faeces. At present, nothing is known regarding !.he
intermed iate hosts for N. ca11i1111m. Cats. mice, pig~. rats. frequency of shedding ofoocysts lhe survival of the oocysts
foxes. gerbils, and monkeys may be induced to be expe- in the em·ironmenl, and whether canids othertha11 domes-
rimental intermediate hosts.3 1. 66 Only an asexual cycle tic dogs are also definitive hosts for A'. ca11im11n.
occurs in the intermediate hosts which consists of taohy- Susceptible hosts can become i11fec1ed by ingesting food
zoites and ti.s sue cysLS. Roth 1achY'1.oites and tissue cysts are and water contaminated 1,ith .\". ca11inu111 ooc.-ysts from dog
microscopic and lmracellulnr. Tachy-,mites are ovmd. lunate faeces. E."perimentally. animals may become infected lacto-
or globular an d measure 3-7 µm x l-5 µm. depending on genically.26· 98
u1e stage of division (Figure 23.2). They dh-ide into two
zoitcs by cnd odyogeny (Figure 23.3). In in.fecrnd animals.
Neosporosis in cattle
rnchyzoites are found in many cells including neurons,
macropru,ge~. fibroblasb, l'aseular endothelial ce~. myo- Nllospor1>Si~ affects both dairy and beef caule.4 • r.a ~6
cytcs, hepatocyms and dermal cells. Tachy-1.oires are usually BO\ine .V. c<111i1111m infection has been reported from
located within the host cell cytoplasm within a parasito· Europe. Ausua.lia. ,\sia, Africa. New Zealand. and the
phorous vacuole. Ultra.structurally. N. cani11um tach~·1.oites America~.ii. 3 1. 7~· 5 1. 82 96 Jt is a major cause of abonion in
382
 Unspotijlate.d
• OOC\'StS -,
~-as$ed in faeces J

Tachyzoites iransm,11a! tnrOIJ~h piacenta t Ootysts 1n

food. water.
or soil
'

~
+ ~
Contaminated
food and water

g ..

lnfetted f01l1US Intermediate Hosts

Ingested trf
m:e:!ia~
,
~
Sporu1a1ed oocvsts

Figure 23.1 L1fo cyc10 of Neospqra c,minum

dair) cattle in the USA, 1.z. 8 • 96 l'\ew Zealand,9 ' and the :'\eth· abort than seronegative cows.·8 · 8S.1C>3
erland:.. 103 Based on serological surveys. up to 100 per cent Neos1x>1'(1 rn11i1111111-infectcd cal\es may be botn under-
of caHle in some herds have been exposed to S. cani- weight. unable ro rise and with neurological signs. Hind
num.~~- 8 - 101
t'-eoporosis has been reported in cattle limbs and or forelimbs may be Hexed or h)11erex1ended and
from South Africa. t.3 and Zimbabwe.1>1
62
neurological examination may reveal ataxia. decreased pa-
.Veospom cc111in11m is efficiently tran~mitted venically in tcUarrcllexes.and lossofcon~dous proprioccprion.11 · I:!. 1"Ex-
catde. eqm ror several generations. but horizontal transmis· ophthalmia oran a,,ymmeLrkal appearance of the eyes may
sion is necessary co introduce new infection~ in the herd.3 · 1• aha be o~erved.
Xeo.(pom cnni1111111 can be cransmfned transplaccruall~ rhc prese.nce of spt:cific amlllodies in scrum from a cow
m narurally infected cattle. sheep. goats. deer and horses. that has aboned is onl)' indicatiw or exposure to ,\. cani-
and experimemall~· in several otht!r ~pede~ of ani mals:' 1 1111111. Several !>erologic:al te!>r~ con be u~t!d to dt>tect .V. cani-
Transplacemal infection can occur repeatedly in the same 1111111 antibodies. including enzyme-linked immunosorbcm
animal and through its progeny for several generations. TI1e assay !ELISA!. the indirect flt1orescen1 antibody cest (11'.\ n.
mechanisms or primary and repeat congenital transmi~sion nnd the direct ai:glutination te~t. 0 ii;. 1" 26• ~ 0 • n. fib. II-!. ffi;. <>J.
of Infection arc unknown. 101 · l(>l Reagents for some of lhe~r. te~ts are availabll' com-
\\bether the repeat congenital infonions which occur 111 mercially. An lgG ,ll'idiry ELl5.-\ \dth the potemial ro dis-
dog& and cattle are due to relap<e of the primary infection or criminate bemren recem and chronic' N. ca11i11u111
reinfection is unclear. infection, in canle ha~ been de,cribeu.20· JI It has recently
:\bortion is the ontr clinical sign observed in a.duh been found by the use ofa newly de\·eloped F.I.ISA 1hnt there
co"·s. 31 • ~~. 103 Cows of any age may abort from three month~ are ~erological dift"erenct.'l\ bet\,een .\ .•·1111ir111m-as!'>ociated
of gestation 10 term. Most :\'eosporn-lnduced abortions epidemic (abortion 5torn1) and endemic (~cattcrcd) abor-
occui" at five t0 six months of gcswiion. 1• io., Foer.uses mar rions.'ll \Vichin the group of amm:ils thm cesred seropositive
die i1111tero and be resorbed. mummified or aborted. or ma) hy !FAT and immunoblot. dam~ from herd~ \l\rjth 1\. cani-
be sr.illbom. born alive but diseased, or born clinically nor- 1111111-associatcd endemic ahonlon~ had slgnificaml)' higher
mal but chronicall)' infected. Within herds. abortions may ELISA fnd1c~ than did Lhosc from herds with N. cn11i1111m-
be clustered. sporadic or epidemic.'· eu. 96· 11' 3 · tao .\"eosvom associa1ed epidemic abortions.
ca11i1111m-induced abunions occur year rourtd. Cows wirh A definitive cul-off cit re for scrodlagno,tk purpose, has
N. c<mitwm antibodies (seropositivc) arc more likely 10 noc been established for caul<> hccause of Lhe uncertainty nf
384 s.n:.1tA't-!'\: 1\\'r,: Prow,;:.oal dijseas-C'S

•
I
...

•
Figure 23.2 Tathyl.OHSS o' Ne!Jtp:Jra
e3n,num
A lm01ession smear Nore organcsms "·
di.,a,ng tarrowheadsl ,mo two are ......
tligge• than single tar.hv:o,:es
1arro1•,sJ. Giemsa s,aio
~
.. " •
8 Tachvzones in paras<t®horo,Js
• ,\.
; '".
vacuoles 111 cell cultute G,emsa
sta,n
C S'6C!ions of the sk,r. ct a cog. A
Nore suppurative ,rl•amma,,o.,
associated w,~~ two groups o!
:achyzoites (arrows) a~d in,fr;;dua!
tatll\',Oites {a11owheads1. .~ •
..
Tathyzo11es in Sl'.!C'Jons are m\Jth
smaller than those in smears
Hasmato~ylin an!! eos,nsta n 1
2opm
..-
•
s~
serological diagnosis in chronically infected animal~ and the cal C\'aluation. Although .v. rrmi1111111 infoction can cause
limi1ed availability of sera from r1on- infectcd cattle. In ~ero- le~ion~ in $evcral organs. foeial brain is the most consb-
logical assays, Litre and absorbance values are dependant tently affected tissue.1 u, 11 rhl· most charactcriStic lesion is
on antigen composition. secondary antibodies 11nd other focal encephalitis d1arac1erbwd h> nccrosi~ (Figure 23.8)
reagents. Further. cut-off values can be arbitrarily selected and non-~uppuratlve Inflammation (Figure 23.9) ll Hepat-
10 provide sensitivity and specificiry reque~wd for the itis i:. found more commonl1 in foetuses aborted <luring
particular application. The age of the animal may also affect epidemic events than in those in which abortions are spo-
selection or a given cut-off value. For example, an IFAT litre radic.10c Because most aborted foetUSl)S are lik•dy to be rap-
of 1:640111 or 1:200·10 has been considered indicative of N. idly :1u1olyscd, e1·cn semi-liquid brain tissue should be lixed
can in um infection in adult cattle, whereas much lower "'11- in l Oper cent buffered neutral formalin for histological and
ues I:80) have been selected as cut-off values for samples immunohhtochemical (Ii IC) examinntion 1 :: (Figure
from bovine foetuses.a The situation may be different t'or 23. l 0). Tlwrt-arc no significant gross !~ions ofncosporosis.
adult cattle sera probably because they have been exposed Although a presumptlw diagnosis may be made by
to a very great diversity of antigen,., •\I though.\', cm1i1111111 is examination of hac:matOKylin and eo$il\ (lffi) stained ,cc-
closely related Ill T. gondii. Stircccyi:1is ~pp. and other apl· rion,. UH. i~ nccei.sary because there arc often only a few
complexans. cross-reacti,~ty has not been a major 1s~ue i11 .\. cm1i11u111 present in autoli sed ussues and these are often
animab experimemally infected with J\'. tw1i1111m and re- not 1i$ible in H ~ :.wined ,l!c1ion5} 2· ,,11 Only rareli are
l.aced apicomplcxans. 13 Recent evidence ind irate~ that even .V. 1:r111i1111111 organ i~ms suflicieml\' munerou~ to be found in
low IFAr titres (1:25 are specific for Neospom ,nrec::uon. 10<• each hhtological MIC'llon.12 1 l11e sensirivit} of e,·en the
E.xamination of the fol!tu:, is neeessary for a clefinith·e di- mo~t t'fficient method (lltC.J to dl:'tect \' cm1f1111111 in tbsue~
agnosis of neosporosis. !dealt~. the entire focms should he !~ low.
submitted but if this is not po~ible then samples from the Finding,\. ca11i1111111 an1ibody m foetal St'Illlll or prerolos-
bro.in. bean and liver should bt:! examined for histopatho- tral calf <tlrum indicate$ ini1:1c1ion. hu1 a nl!gmhc rc:,ult i,l a
logical changes. and body fluids or blood serum for serologi · fol.llUS Is lc,ss useful as antibodi production in the foetu< Is
 );cosporosis 38S

,;... ,..., .. ... . . ..~.:. .. . . ..,.. -~ .,

~t
. ,, ,
.... ' .,,, " ....,
.... ,,:
• '
- • _.
~

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• 4
,
,r
,.,.,.,,~
..· \
... ... ,,
.
~

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•
.
.,, ' ••

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'

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. , ... 'l\
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Figure 23.3 T,ansm,ss1on eieciron m,crog:apn oi d,v101ng racnyzoires ,n two paras,tophorous vacuoles {Pv\. The Pv nave r.>,m, tucu!a• nel\vorl<s
/vrows point 10 cono1dat end o: dc111ghter ;ach'(zo1tes forming ins,ce ,he r.lJlther 1achyro,1es Note electron-den~ rhoptr,?.s till. mlcronemes IMnl.
m1crc,~bules 1M1l or subpellicullar M:u!es. conoid (Cl. ~os; cell ~~claus (Hen!. CerdiOl)ulmona!\ ~iter; 1:ndotheli31 cell culture 1Counesv of
Or CA. SDelli. Moncana S1a1e Un,vers•rv. Bozeman. Montana. liSAl

depend em on the stngt oi gesrmion. level of exposure. and Thu,, enm a lo\\ II A'I titre of 1:25 should be considered spe·
the time between infection and ahonion.9 · 1" 2 In one smdy. cifk in focm,6. In congenimlli i11fe<1ed calws. precoloM!".il
!FAT titres (1:80) were found in 50 per cent of neosporosis• scrum from live cal\'es. and brain and spinal cord from
confirmed foetuses and in only one or 64 foeruses that were dead cul\e~ are the specimens ol choice for establishing a
aboned due to other cau~e~. while in another low IF:\T ri- diagno~is.
tres 1:25 W!'re found in G'.\ per cent (31 of 481 oi foetuses Isolation of \'vosr>ort1 in crll culture is rarely possible
1mmunohiscochemically positive for K caninum and because most orga11i~ms in bo\'inc foernses die when the
la none of the 39 foctu;cs aborted dur 10 other caw.es. 1<> host cclb bl'<"ome autolysed ...!- Se1cral polrmera,;e chain
 An unusual case of visceral neosporosis has been
reported in a 10-year-old Appalooba mare which had a hb-
tory of weight lo~ and anaemia. Lesions associated \\ith
N. canimun tachyzo ites were confined to the intestines an d
mesenteric l~1nph nodes. Thb b the first reported case of
Aleospom-associated enterirls In any animal species,ii3
:-Seosporos1s has been diagnosed in ;rn aged Pinto mare
suffering from a pituitary ad,moma whic:h had a history uf
hind limb paralysis, b izarre beh:wiour and mastica1ory
dii.ficulty.:io Lesions were present in the cemral nervous
~)'stem. peripheral nervci> and myocardium, and consisted
of mul1ifocal non-suppurati\'e encephaltHn)•elilis. polr·
rediculoneuritis and mulrifocal rnyocnrdil is. Seosporn
tachvzoitcs
. . were demonstrated b,,- !HC
and tissue C\-Sts
examination in the• brain. spinal cord and peripheral ner\'es.
Neosporosis has heen observed In a 20-vt>ar-old horse suf.
fering from a pi1uirnl)' rumour.~:.
,Yeospom orb'llnisms were idcmitied rccenli~· In the brain
a nd spinal cord ofa Ca li(omian hors<' which had manifested
tleurological signs.73• 74 The pru:asiw was isolated in cell cul-
turt-s inoculated 1vith neural tissue· of tlw horse, and was
named.\'. lwghest.74 Neospora Jwghesi has also been isolated
from the spin(!] cord of a 13-~ear•old horse from Alabama.-
.,.
.-\ntibodies 10 N ca11i1111m were found in 23.3 per cem of
Figure 23.4 Tissue cysts lai<n·:,s, of Neospc:a camnum 296 horses in the USA slaughtered for food for export "° and,
inanothersrudy, in I t,5percem of536horses in the US,\.u
A Sect,on of a calf brain. Note small ussue cyst w11n 1hm cvs1 wall
Hae!lllJIOxylin and eos,n sra in
S fntraneuronal tissue cvsi ma s3e1,cn ol :he sp,naf cord of acalf Neosporosis in goats
Hae'!latoxvhn ana etis,n s;a,n
C lntraneuronal ussce cvsi ma !etuon oi spinal core or a call Abortion associated with .\'. cr111i11111n ha.. been reported in
s,
No1e sit,er-positwe c,st wa i ver stain
herds of pygmy goats in California 10 and Pennsylvania/'
D T ssue c•1s1 In a brain squash o' ar expenmenta~iy in:ected mecse
and in dair, goats in Heredia. Costa Rica.'r. in the Califor-
~iote thick cyst Y.all fopposrng arro,vheads). Unstained
nian episode, a roe111~ from each ofr\\'o different flocks wa~
eli:amined histologically. Tissue cyst, wel'c prt~ent in en-
cephalitic lesions in both cases.
reaction (PCRJ methods ha\'c been reported with which One neosporosis-infccce<l foetu~ from a small nock of
C't111i1111111 D:S::\, ;s :.z. :;7 but have not yet been
to d<itcct 1\'.
p~-gmy goats ln Pennsyh'lUlia 1113$ examined. l he nock had
c,'llluatcd cri tically for the diagnosis of N. ca11in11111-
experienced abortions. stillbirth. and birth of weak kids. The
i11duccd abon:lon in caule. foenis tha1 was examined was full -term. and the lesion~
present were predominantly in the brain, e~peclallr the
Neosporosis in horses midbrain in wh ich on encephalitis charau erized b: numer-
ous microgliaJ nodules with minimal nccro~h, and severe
Clinical neosporosis has been reported in seven horses. all perivascular Infiltration of mononuclear cells wa, noted.
from th~ USA,24 30• 48• :..'I :.s. n. : 4 lhe first of which wa~ in Numerous tissue L·ysts were observed. some of \\'hich \\'ere
1985 in a rwo-months-prcmaturc aborted foetus. Only lung undergoing dt•genera1ion. Tach)~toitcs were nm noticed.
tissue was available from this case and numerous ;\'. cr111i- In Heredia. Costa Rica, the farmer had a lso experienced
1111m tach)0loites were ob~erved in the ie:;lonb that were an undiagnosed abonion problem in his !lock of goms. The
present. ,ij Recently \'. c,minum was identified by PCR in an foetus that was C:l'.amined was of3,5 months' gestational age
aborted
, foetus in Francc.90 and a Saanen-Toggenberg cross. Lesions were found mainl)
:-!eonatal neosporosis was diagnosed in a one-month · in the brain and were characterized grossly by hydroceph-
old Quarter horse fiUy in Wisconsin. 72 Shl' had <:ire.led her alus .ind a hypoplastic cerebellum. and microscopical!~ b)
dam since bi rth and frequently tripped over obsmcles. If menlngoencephali1is which manifesred as a ventrlculltls.
separated from her dam she could 1101 find her way back.. gliosis. necrosis. mineralization and va~culitis. An unusual
Encephalitis associated \\ith tissue cysts was re,·ealed 1Jn fearnre of this case was lhe presence of large numbers of tis-
histopathological examination. sue l'YSts .
. and the rnri1v oi tach,-zoirc~.
.. .
In a.dd[tlon , mvositi~
 -.:eospcn·o~h 387

. ' --
••

. ,.

.. r 1
/.r
I

\
~ f

,_.,. ., ... )\

I
!1~m
_;t ..... ' .
Figure 23.5 Transm1ss,an electron !:l:c·cgraoh of a Neospcrac,m,r.vm 1,ssue cyst in mOlise !lrc:r, Note ai,ct srrr.oth c,SI wall tC::I. absence o: sepra
m,c1or.e11e-s (,In) perpendicular to :~e oradyio,te p!asma1emma• .;my 0Gect1n granures !AQI tnat s:ain rerl with PAS electron-ceiise rhoplfles IRI, and a
Sl;bterm.nal nJcleus {Nl (Cllunes-1 o' Dr CA. Speer, Montana S:a:e Un,,e,s,ty Bowr.an. Moniana USA!

a:-:,odatcd with the pre5encc or tachy-..:o ites was noted. The were in rhe spinal cord which shO\,·ed a myel itis, 1milateral
foctus·s dam had an N. m11i1111m IF•\T ritre of I :800. Fi\'e t>f reduction of grey maner in the vemrttl horn, ,u1d focal ca\'·
;; mhcr does In this herd also had JF,\T antibody titres to itation and were as~ociated \\ith intact and degenerating
i\". cn11i111on; cross-reacthit)' with T. go11dfi was obtained. .\'. rnni1111m tis~ue cysts: iachyzoires were not srcn.
E.xperimcmally. goats ha\·c bi;en pro\'ed 10 be susceptible 10 Based on the result; obtained in one ~1udy, Neosµora in·
S . ct111{11um infection.• 0 fcctions arc nor con~idcrcd as a cause of abonlon 111 sheep in
England.8'3
Neosporosis in sheep
Neosporosis in deer
There is onl}' one report or congenital neosporosis in a ,heep
lamb ·17 n,e affected lamb was born \\'eak. partially a1a.xic. l\:eosporosb ha, been diagnosed in a 1wo-mon1h-old female
and died at one week or nge. The most prominent finding~ black-.ailed deer !Odocoile11., liemio11w c·ol11111bim111s) 1har
388 \lr., u\: .,...,.• : Ptotb1.oal dh,L·U<il'\

",-.,, found dead in California. lJS:\. aoz Lesions and stillborn brown-anclered deer (Cen•11seMii si{lf11e11sis: from
.\'. c-r111/111m1 were present his1opa1hologica!lr In sections or the Pari~ Lou, l'rnnce.'" \"l!Ospom n111i1111111 tb,ue ,y~1>
11w lung~. liver. and kld:ie} ,. Tachy.r.oiws were 010,1 numer- a,soclatt'd ,\ith a non- <uppurath c l'ncl'phaliti~ wen.·
ous in the lungs and were associated wilh interstitial pneu- presenc. :'\eospornsis was 1hough1 to h<• the main disease
monia. while in 1he kidnt'}, an inter,1i1ial 11cphriti~ hl\Olvcd m 1he dcdine the J)opulminn of lhis endan or
.i,wclatcd with their pre.\eni.:c ln tubular cplLhelium and lu· gered ,pi:dc, in t iw /.OO. ,\ntibodies 10 N. ,v111u111111 were
mens was noted. fai.:hyzoile~ wi1hin hcparnc-y1es and found in Hl2 of '100 (40 ..'i per cen!J whitt--1:1ilcd deer
1-.upffl'r t.:~·lb wure found in 1he Ii, t'r. (Odot:()i/.·11$ 1•irgirifr11111~· in lllinois, US.\. 111
'-:<:o\porosi" w,1, aJ,o beeu rm·ountc:red in a full-1erm

Neosporosis in dogs

• ! log~ of an) ngl! may bl' aff£'cted. Fatal neo~poro~b ha~ bt'en
reported in a. to 15-ycnr-old dogs.3-;. 41 llw disea,e ha, been
reported in dog., in South Afnca.60
Subcllnkall) Infected bitche!'. can transmit the p:m,~i1,•
to their foetuses, and ~ucccs,Jvc litters from the ,ame bitch
OH\) he born infertcd.7 · • \\11ether there ,~ brcl~d pre-
dispo~itlon and differential sex susccp1ibili1v to neosporo-
sis in dog~ is 1101 known. Mo,;i de~cribcd en$e, ha\·e been
in Labrador RcLiie,·ers. Boxers. Gre)•hound,. C,olden
Re1rien•r, and Bai.set llounds.42
Only limiwtl infom1a1ion is a\11ilablc- conrcming th!' pre-
rnlcncc or \'. rn11i1111111 mfection 111 dogs. Using an 11'.\T lil re of
I:5(). 0.25 10 ~9 pc.>r cent of dogs in i.everal ct>uniric~ ha\ e been
found 10 h.s\·t \. m11i1111111 antlbodk-s.1 · 1" ai 1·• w,. "·'
0 e The moM ,cvc•rc caws ol ncospomsis occur m young. con-
genilally infoc1cd pup:.. The di.ease in young dow, moM fre-

, .~6·
~
• qul!mly manif<~•~ ns hind limb parcsi, (Figure 23. 1 I ) that
de\·clnp, progres5h'<'ly imo paralysis"· :t9. 34 • 11• (;() " " 9 '''' but
..
~

~ • .•· .. .....- ' • a

-• ':. t•
"
20pm
' Ii.. ,
" ... • •I" • • • r
'
';
Figure 23.6 De~e!l8lat,ng :•ssce M: lar,owl in :he 1:!ra,n o' a gca:
, . •
••
'
... '
•
• • ,.
~ 0
(

'
r,. • a a rnlls and moro~~c1ear ce!!s surrounding the t1s~~e ti•SI.
r1.iu,"atoxylm and cos.11 Siain
•
•,._•. " ' ,,. ' •
~ ~
....'
. . ... . ... - .
#
,
}
•• •
..
,......
..,

..
, -
...,• .. .1 - .
:_ .. ' \ ~ ......~, :-
.,;,~ '
}-
'' • 4, • ' \ • ~· •

~
'

••• I "'... -~ lJ!'\' •• ;-•~' _:_ •

• .. •.. '\ .4-
~ <t--...·\ ~-
... . ' \, ~,.. ~
••
l "
I
~

. ...r.. . .
..
I
•-

...
.... .c

~
, - .. \ -:, '
-,J:.""-
......
•'
\
\ 1 • '

, ,. • ft,• "• ., ""' ...... , \ .....

' . - ... . ..t•,

., • • .,.- .... ..<' •• , '
• . . - •. ,,. ~ ..I • ~ . .. "';-- •
...
'

, •. . .
... ...•. . ~ • • '
f f :,, • .. • r,J' .-~ -., '-
• .. \C,,,J 1
A "4, i
..
'
... . . \ ..
• •
n • ,• • .t; •
..
, • • ' • 'I~
B •
.. • .... •
••
100 f.l"l, " •
Figure 23.7 ·JOMts of .Ve::spora ~nmurn
• • • • • ., .
;. = 11-1:p.:; ·~ateti figu(e 23.8 ··,:.al ll0~1ne nra,n ,r•ec:e~ ·;:•:n r,eospora ca11,1..,-;,
8 = Sp0r111ated Nc,e a,.,.,,, •ecrot,c tows,....:., Ql!a· cells a•.:i rnonontltlear cells a:
No;;; ;,,,,1) sp:irocvst~ lar•o-:.-,1 a :! spcnJzciles !a1 ro'l..'ea~sl l.nsta;ned
0
:he pe•,pne", .iaernatc,ylin arid eo;,r s:air
 \cusporosi, 389

1he neurological ,ign, are d.:penden1 011 lhl! ,lie pam:.itizec.l that high 1imi, (1:121100) have been ,hown 10 per,;b1 for at
. ,.
in 1hc l"cntral ncrVt>u, <ysiem.' • The hind limb,; at\." gcncr· leaM four 1·ears in 1wo clog~." ,ind clinical neospc,ro~i, n•ri ·
ally mar(· sc•1 cn:!y .ill'cc1tid lhan 1hc fore limb~ and often ficd l11<wlogicallv and by isolation of l11c parusilc) was d;a~-
exhibi1 rigic.l hyperl'X1en,ion. The cause of 1he hind limb 110, ed in 1wo dog:, with II YI titre, of 1:50:33 Thim.- h nu e
hl'p1m::\ttmsion i, rnH kn0\\11, hm is most likd> cau~cd by a informa1ion rm Jg~l production in pu,1natally infocted dug.,.
combimnion ol upper molOr neuron paralysis and myositis. In 1wo congc1111ally infected nninmb. lgM was not detect·
\\hkh re.uh, in rapid!> progr<"ssive fihrous comrac1u~ ofllw able a1 :i !:HJ ~erum dilution.:•··
mu•dcs 1ha1 ma}' c-ausl' fixation <>f )Oim~. Other dysf1111c-tion,; rhc deicction of V. cr111i1111111 in hiopw ti~~ucs or ntc>·
which occur include dltlkulry in swallowing, paralysis oi the logical prcpura1io11s etlll c-ontirm a diagnosis. :Veospnm
jaw, muscle nacddl1y, mu,clc atmphy and even heart failure. ,w1i1111111 ha, been detcl'ted in hiop\le~ of mu.<cle~. lung
Pog., \\'Ith hind li mb paralysi~ may be alert and survi\-e for aspiraw, and tkrnlu l pu,1ulur exudate lmrnunohis-
months. cochemical s1ai11ing. however. br('quiret.l co differentiate 1he
The diseail' may be lo(alized or genen?lized and 1irmall~ par;isite from T. gmulii b,il·au,c 1;ichyzuite< oi rhe two para-
ail organ,.and Liss111:, ma> be in"ol1·cd, including lhe~kln. ; •5 , ircs arc indistinguishahfe from each other by llgh1 mk·ro,-
Derma1i1is may be sewre and large numbers of.\'. ctmimm,
tachFoite, mav be as,oclatt:d \\ith it.
For the ame-monem diagnosis in young dog5. 1he clini-
. ...
cal sig:n of an ascend in!! pnralvsi,. panicularlyif several litter
mat<!$ are affel'ted, :,hould arou,e ~u,picion or neosporosis.
•
•
''
I laemawlogic:.il valut•s are usually not a ltered but there may
be increased IC'vels oi 1hc serum enzvmes a,sociated \\ith •
necrosis of myoc-ytei- and, occasionally, hcpatocyte:..
The finding of N. cr111i1111111 amibodit?s in scn.1m can nid
r
1he diagnosis. The Jr \T has been used mos1 often to diag-
nose <.,mine ncosporo,is and a high titre< 1:800) is o~en ln-
dic.itive of an ;1cu1 C' infection. It ~hould be 1101ed howc,er.
.>•

' .. ... #,ff

t
•' • •
I
y~&
~ti
A ..

• •

Ql
• i '

I
B
Figure 23.9 Sec?·on of 1he sp,n~I C():d e:r a calf congennalf, m;.i~~ ::i:.h figure 23.10 'levs:,,,, c11~1m1m ta:t., ,O:!es and a "Ss"e CV$; s:a,r..::
,mmunoh s:,x1';;rr tall/~, th Nc,n,num an:·boa,
N ~.anm~m N.-,10 glial p1ohfe1t,,O!!, pet '<'ascula; ,nflamma11cr, .~: :!''=e
:,ssu;; cys:s ,arrows! lmmunoh,stoc~e111ica' s,ain w,th N can·,oum A Numt!!OUS tac/wro,tes wrowheadt ,, tr:e nr, ci a tfog
am,booy 8 A us.we tvs, (a:1ow1 in a noumr, ,n a co,ige1\;'aily n1etied caif
390 >t•:110~ mo: Proto1.oal di~ea~~

figure 23.11 Neospore caninum infected dog.wnh hind limb paralysis The fust 1so!a1e of N e2ninumv,,,as cblaineo from lhe neural tlss~es
of this deg in 1988. lmmuno!iisto:llemit,11 staining, howe,er ,s required to d1ffe1entia;e the paiasite from T. gondiibetause 1achyzo1tes of the ;wo
parasites are indistinguish~b,e from 1::acli other by light m,ciosco;iv. Cerebrospm~I 'luid should baexamined for tachvzo:;es in cases manifesiing hind
limb µaralysis

copy. Cerebrospinal fluid should be examined for 1ach)'ZOI· disease when treatmem is initiated. Trearmem has been
tes in cas~ manifesting hind limb paralysis. successful in some dogs suffering from early neosporosis-
Post-mortem diagnosis is based on demonstrating the induced limb. weakness.6• ;z, 75 Sulfonamides and/or py-
parasites in lesions of affecced dogs. Gross lesions of rime,hamine, and clindamydn have been used with partial
neoSporosis that have been reported are necrosis in the cen- success forthe treatment of canine neosporosis. Acombi.I'la-
l ral nervous system and li\'er, granulomas (up to 10 mm in tion of trimethoprim and su lfadiazine ('fribrissen®. Coo-
diameter) in visceral tissues. yellowish 1,·hite streaks in pers Pitman-Moore) at the standard combined dosage rate
muscles. particularly those of the diaphragm, cerebellar at- of 15 mg/ kg twice daily. and pytimcthaminc (Daraprim©,
rophy and ulcerati,·e dermati!fs.'12 Jmmunohistochemical Wellcome} at 1 mg/kg daily, all for four weeks has reversed
staining using specific antibodies will confim1 the diagnosis. N. cani1111m-associated paralysis in some dogs. Various
The parasites most likely to be confused \\ith :,·. Cl!Ilinum preparations of clindamycin alone or In combination with
are T. go11clif and Sarcoi.")'Stis ca11is. The latter dh1des by en- trimethoprim plus su lfonamide have been used co treat
dopolygeny imo many organisms. whereas T. gontlii and ncosporosis in dogs.6 · :is. :s. ' 4 There is no treatment cur-
.\'. <:an[1111111 divide into two organisms by endodyogeny. rently available that will prevent a bitch from transmitting
The respc>nse ro treatment b dependent on the stage of .V. caninwn to her offupring.

•
References
:\XUff\M.)~. M.1... nusr.n,UtD, s•.<,., DA.MM:. o.r.• l)UBf'\', ,.,... JCOFOtAN, R,.L,. JU;.\(). D.!I.. DUUF\'. J,J••• f.OXR.\O. p_\. ii B.,\Rn. B-C,, l99S. Evaluation of
l':OXH.\D, P.A., &991. .'\"t°1)1porn•like- pro101..<>.tn inf~iona.~ :1 maJor ~use ot nborum:.,. tn e1utlc1 :utrihutothlc .io n"°'p,no~i.i In ~l·f~cN! dalt>· hl.'rcb: in
abortion lo Cnllfomla dairy a.nl,•"/011mol ofthe ,\111,rlw11 Ft;,r1,1my C:allfomla, Jrwmnl 11/1//, Am,•r,t'.(m l',1tri11ary .W,d/«JI,i,.;«111rlo11. 20,.
\./rrJ/cn/1\$socinl/(JII. 1~8, 2.:1-24.;, 1206-1210.
2 .,xorn ...oM, M,l.., t•..\U1LA., c..w.. UIUkMO~u. M.C'.. , P1rx\ffl, ,,,,,. llL\NC.IL.\Rt), 3 A.,OttlSO~'• )S.J.., HC\?-!(.)Wt,,, f.J.I .. RO\\'L 1,0.. ~\,"ERLO\\', ),!,\\'.. 11.i\(~lfA\1, A.t;:.,
P.r... tHtFITMC\'CR, P.E., L\\'TO,, \.\.\',, ~,cM.USt'CR..)1. 0,4.JJT 8 .. ~UL u .. 8\A~ll.t, lf/CO~RM,, P.., •• 199i- E\'ldi!ll<'t" nf1.crllcal 1~nlfrni.;;.. ton r,(
 :wosporosis 39 I

Srmp<1m ,p. intcc:,,on In dairy CJllk Jarrmnl ofrhr Amrrtmn l',1erl11m;,· ~l'f>,poro .. h .md bm,nc ~,bortlon in !'rntland n11• i1.·1~i11n,y Rt~rrl.
,\{rdta1/ ·l<.~m·1111ion~ :?JO, 1lfl'9-I Ii:!. J.11 649-651
&'1\UUt, ,.~. G.'\S>t"•R, "·"·· H l I>. I,, IQ a.lltl. ,,.r•.. M ,111 I '"· ll. "" flll·t-\. ~.i ISU\Ul\. U. '1 \U \. -..,w . t'A.,1t'Ulf-1 t'.I•.. "kOt'.HIUt. a.-'! l~\L-'- t.,\ .• 199-•
.,., .. t997 Pr,:\•a lcnct· uf anubodic~ to \'tl'tlJ)t,m rm1111u1111n dif!'crcnt £.\u.n1h1,ulon af r.:d ru\c" 1\ '11/p.:., rulJ]t.\. from Bul~um lo: .intthody ti0
o.inld popul;1t1or1,. Juun,a/ o/ l'rtru.•lrol~gJ: 83, IU5&-IOS$. Sto:•JJ(l1n ,mrform, :md Tot1J/t.'amu1gt1ru/if, n,r i ~111rJ1M1)· R«<m.l. l.& t •
5 Oi~uitnmon Cit'
tr.AJUtLJt. I'!,~, l1 \\'Nf... JCJf1,~ C.L .ti flll·£t.. ,q,, I~{,,
.\03-309.
~Js/>mn rnainum 1.,·i1htn 1h,.-ccn:r..il n\~n-Qw.. ~'1-lcm i1n,J HI her [i.,,..ut.•> :t,.J t"IIIAtJ1.f. \.~~\. U">,"-A\. U,'-., ltO\\'t. '- 1>\l.."SJ1\.\. < C., \\'tllt.\.\ts. \t..\.
of ,l:f. d<Ji,, wirh clintcnl neo,;,oru•l•.Jc>11m11/ <>/S11111/I .~m,,m/ PrnNlrr. ,v,s,:rtt.. I ..\ . TOl.\1t'f·~l'.'\N\lt:.A~, \1,.\, USl " IJ ,. SI \\70S, ht-, nntMt \~
.r,. ~Si4. ,, o. taur...,,,ttvR~. ,u 1fl!m. fln.•vdlr.nc1.· nf ,int1h<>d!1.... 10 .\'t•mpum ,p. in
hon,.,.•, trnm o\Jah.1m.1 .me.I th3n:cu:rl.c.sutm or .in l<-~late rctO\t.'ft."1 trom
6 s.uun.-. J.,." ntl-l'"', A.I,, 19,t;. Cllnh:aJ .1,Ju.•,~1, of27 (U.'i-1,.•, of O\"IJ\f)Ut(l\i"'
., nntutall~ sn!t."CINI tun,se. h11f•rnmta111t! Jo:1r,u,Ifm Ptr11uu,1follJ ?9
in dog~ n,(' \ ...,...,llu,ry H.«ord, 13", ~),-,HJ.
rn~-,s.i1.
; R"1'Jla., 1. <. ~ mtr,., ,\.I .. 1993, :S:JIUrall) ll~etUTlng \lcrtl!'.;li 1r.u1"nh"on
!!S ruu . II.A , lJ;\"l)~\ . JJ '·· tlllH! \. J.£1, , lJU(..BlJR:\. S.L 1993. D~tn:tion u(
of .\"-.!'Os-porn t1mlmm11n du~ lm,-rnarumnl k,untttl.{or PnrMrolo~·. :.!ft;
:174'.
,V,-osJMril ,,mfnum in ,1~~,,c
\toetlun.'- \h,ln\,;.t m,,rtnc 1non()('lon:ll
.1mlbtld~ , Jounirrl nf \ ·,,uriunn /Jlf'8'1oulr lmT.,rtRfltlon.5. 57'9-5$4
8 ,i, C'tl\;ltA1J, I'.-\,, J!,t91 .\~b'J.:t>ra•Uk"
8i\tt1L H,f;. •.\~Ul<lt~t)N, M.1., UIIIUl'\, J,J•.
26 cou. ft.\., U~b...',\\·, n.~.. hL\GhUII,.,, 8.L & uuuu. J,I'~, 199.;, \'cn!C':,1
pm1n1.oul tnft.."(IIOno:: .L'\SC')t.iated \\ith bovine- ~•hoilinn'- \'1!1.•1i,111ry crun"'rnt~mn t,I .\·1.·0.1,·p()rtt ,·,wimm, In mfi:C'. Jcmrnnl fl/ Pam,:{Wff.1(,J.·. R1
Pntfrorogv. 28. l!G-116. 730-73?.
9 llABR. a.c.. .\ "Pub,..:.,~. M,L. ~\ULQ\\ , K,\\' ,1,, WSMO, ,, .\, 1~:;. Oi.,JW"!0'-1~ 27" t~(>:i"H."O, ,~ \., ll.\llN, II,[,. "'-\'l,ltl.OW. );.\\'. .\\ULHS(>:-.:, '1 .. 0.\1"1, U,. »(1!\0L It,.,
ofhm>fnc fot.,I NtVIJ/JQf(t infccc.lnn with on hidirr.;r fluon,m,nt nnrilm~\ DUa(\', J.P. '-10~'-t,)N,' lf.\SUMSS, \ .• 19')3, ,,, t·ttro I.SO\;\HOO n.nd
te<L Tlir \'~tur/11my//rcord. 13,. 61 H\13. mn.rncwriz.nnon ot' l:I ,.t/0$/'JQ!ft sp. (ft)m .,bont.'CI bO\'tO(• (n,ctU:(C$.
10 flArui. 8.t:•• ANIU,lt~';\. \f.l.,, \\CXllb.. I w .• ntHU.\', ),t•.Aot~u~St,\f), ,; ..\,.1'.)(Jl. Pfm1:,.'i1ofvgi•. 106, 239 ..2,19.
,\'trmpnm·Ukc pr"(ttt>z.o.d inftntlon, 3.,<;f1ti.atccf whh «1h.1nlun in gonh. 2'8 COSM.AD, r ..,. S\ 1JilUW, l... .\'Silt~()~. M,, fttlWL. I, l'H);\;IJl}Ri\..,,. ll. Tim-Jt.
Joumnl of\ 111,•ri,u,I)• Dtr,gmnrlr 111,vu1gmlo11. {, aGS-a~; G,, lU'cfmff\ EJI, R., f'U.'11-lt, 1,., tHUft\10\:0. \.f,, \ROA~:,.,;\,, l lUKt.''r. J.P.,
U fl,UUt, it,C., C."O:,.J,C;\U, l;A, llkJ,Jt\)llH\, It,, '•.vl·.RLO\\, 1'.• l.'-il.f,lbO', ,1,L. llUJl ·\ ,\ll I,,•. 4,. H,\llf(, "· 1993, l}eU."(tl\JIJ of ~t"fUin ,Ultlhod~· Ct'>J'IOO~('-' in

At'\~'ou,~, f., C:UA\l\'ltt, J\.t•. OVDl\. J,l•....U!.U,\.~S. ,\,A,. 199.t, t .rngrnhi1I 1 ctnifo wl1h 11atur.1. ar cxru-rimcmal \'t'OJ(,lom mfoctions. /uur,ml fJJ
Nro,/IOl'tl !nCcNIW1 In rw,·1•,bornfrom ctm~ rltn, hnd pnl\Jou,l) \'411.•rumlj' DJ.ttiuo.w·r lm•cstl,~tul()II, 5, .5;2-3}8.
nbom!d WO;pDrn-fnfcctcd fcw,u, four C8$C> 119~·1 ~92f./011r,m/ o; :!.11 t:\IIU)Q~. , .. Lt,. u.~ .. nC.l\\')1.\S, f),I), U:0.1)).\\", n.~•• ,\\IW'll, ft:•• ntJ:'I.G.\'\: ,
tint ,.,,wrlnm \'ewrim1r,• ,\Jed(wl..b~txfrufnn, 202, 113-11;. 11>., DI I..\UU'.'lil\. A., 1 .. 0M'U~<;~; f. !,\flfR, '\f, C.TI0Pf% n,. all~Ci. r,.\t,,e.

L.t. lhltH. 11,C., (0:o,,ilUO, ,,•., •• mmn·. J.P. 4 A'-nt M!,,O~. \J 1~. 19iu, A'tih(NJrl~·liJ;('
1mm., , .••.• 1!39:l \eo.~"'m 1.·mwmm infoction m t·.ngH,h ~ruinsr,
~nccphalom,•elltls In~ cnlf: pa1.hologv. ultnmrurmrr, ~n~ :\paniel Jitwrmatn~ Ui;a~m,>~uc li\-a.luatian ;md Or&am,m 1$4,l~nlon.
inununortv,irth.i1y. /tmmnl ojt'1'tffln110• nr,,g,,m•ric lm·,'fttl(mir,n.. 3. Joumnl o)' \',•1e11nm:~· t,m•mnl ,\f,,fln,:,, 6. 3.?S-3'J2.
39-16. Jc) U1\t'l, tl \1 ,, lf,HolR, ~ t
UII' 1'-~·:,. ~..,....UUO\\', "" 19!,II). N,w,H>m
~nctphillanay'fhti,. aml po1yt.t.dkulont"wi\.-,. In ,ITT .1ged m:uu \\ilh
13 ll\JI•. U,C., IU,ll'l, 1,11,. ,n.ttU>\\. •-"'·• HIJ>IOUlt'"'· ...... \ll!>.IM, .<.\
Cwhfng'::; dhct»e. Equlnt' \f<'Jirlllfll.'\' Jnunml.18, 2-iU-243
oU\u. M ;\, • r.o~AAD1 1•,A. l91.'f_4 F-,pcrlmc.-nml u:puxJuctJon r,f bo\1nc
(eutl .\'empuro lnfrction aisd dC""Jth \,ith 11 bo\·inu Nrl>$p(lm lso!mc. :.s1 nuft1.,. r,1>.. 1999. RC('("m .ith·nncc, \n \"fo11J(lra,,nd n,'Q,poro~!)...
lo:,mal c,/ \'nrtlt:ttl)' f>fttg,i~nc Im ,•s:fg,1lirm.1;, 207-:?ll. \.', 1N.rrn,r')' Pizrmfmt,,~••fW. 34ff-1,r," ~

1.; r.v ,1...-..:0\\U.\.. U.P•• nmi.n 1,f\.

IL.\~1.1.11. f•"'· ,.,, ..
\t.\1ftt<o,. 4i n" 3:1 ut.mn· 1.1•.. Utblrt. ti .• n ·u-1•1 H. ,u."' n,w_,,.~. J.F~ 199H fl~·dtoccph.sl~
~L~Cia1~ with S,'Osporo Cd11J1111m 1nfoctitln in Jit3bort,-d bl.win\!" rt'tLJ!t>.
~cn,,·.u~. T".r ., 1996. Sero?oglcal dfllt;,nO>t..,ot bo\ine neo>µ~ro~!,, b~
,\'cwpora canimm, monodonnJ anllbody..bMC'd rompetith·c lnhiblllon /ou11111/ <>J Comparm/w PIJ/110/oJO'. 118. 169-1 ~
l lJ~.\.Jmm1<1/ of Climcol ,\llcr{)/.>1()/()gy. 34. 1·123-l•l:!tl. 33 ovon· l ,P..\Cf,.\SD, U,M. kll,\.MIR. \ .... 199: .\'t'O!it}Oftl(lllllflllfll
r,\p1f.:ompt1:,~: ln .i ,anhorn JtUiH. Joumrtl (,[f'ma.,,;itofogy. 111. J32 .. J.3-; .
1;; iJJt N,.\.,. 1 ,1011=-', s.f-. .- 1•tc.1:St"lftJ>, 1., l,:8..i
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l'nra.sitc,lam·, 75, ~.J-5i, "-''I.\ um1,, ~, .• 1irJN, l)~·,cripciiln dt- un ,....i.,a ,h.• uhorto hft'-'tno o1~o< 1,11.fo
Uo 1-'<~"0!I.'.. w.. 1)£ 1.\.llll~f.\. ,\. \l>~SU, J.. f()Ol'l n. u. ,., UIJnl \", 1.1•.• 199.l~ ,1 lnft.•tr.:K'tn por \'r0SJN11t1<oi;>.en \1~,,co. ~~1,·nu11rit1 .\,J(,xiro. 2ff. i;3-35;

.V,'Of/)(Jf(t ,.,,,,,,,,,,,, in 1,,111 adult d~ Wilh prUhl'l......... i\'C Ct.!rt•h1.•ll.tt ,iN,n .... rm ,1 .. Ull I). J , ... UIUII"\, 1.1•.• ,,ni lhO:\ M.t. I 114\1 \U ., \AH,1 JI . .\I J ....
Ptt>gr,~'< i11 Vitturint,1)' ,\'t'llt0logy, 6, 124-l 2.7. \.1 JC.t.k, n.u .. 1~~~2~ N..•tJ.(ptJ111-Ji'-c prutc,1.oan m(ccuon o, a co.u....t: uf
 Neosporusis 393

nborriun in dairy (DUI~. Ummal of\ 't•i1.tituu·y CUag11os.1i< lmvsrf:,:mion .. Sl'rolrig1C' diagno~s of Scto$/~ra c:,mmum mfccuon f'amsimlo&,'V
.:. 223-2Z6. Ri't,'f!rc/1, 6-l. 50-!\3,
&l OtlL\:{.M)Rf, Q.~ MllflMV, s .. \'lU>H\f!(o. C, ,w:ri.oW, 1J.L~ OUHl\, l.r,. !JJ '.'1,\\\"\ll.\, 'U .• r.,n~ (: u.. 11.0~IMJ, 11..MOIUI,\. T . '.'111\UO\, ..... \".\)t.\\'t.
199~.•\bonion cuu~d by TI\"O~porm:i.. m -.·,mlt-. ·\t4s11aliw, \ -.,.,._.:-,,H~fj· ,. & U\tUIVN.'\ T•• 1998. Sc;orolO)!l('UJ ~Ufl:l')' of antibody lQ l\'l'UIJJ(Jnl
Jorm11Jl -?, 11 ;-1 11:1. ,,mtmm: in J.l.P,Jnt.•i.r dog.... Journal 0,1 t L'trn,wry• .\{rt(wal ~Nttt;<· M.
8'i3-85l
&:: urn ff, .t.•.JHt-~~,. '1 GUtHnu, I .t,.. 4o DUtir\, 1,P,, 1995. -\ '""'1'' oC th.r
irtddcnc:t or .\'(05.JIOm t:tminum in!tcnon In :1bont.•J itnd r.t1Ubon1 g r 'l:lUJU .:,.. , •.• PflUt~. 't" WllkM, n.. t.,P.;\t.\~!\. t; Hl"''1Nfi, .... ,f.l rtt,KAUl ... ,
bo\inc fc:iU:i;c.s In fngJamJ .and \\'4Jt:.... lh,· \ t't.1tmlll)· R«twl, ~J(J. ...,. 1g9:- \;•()~a,rmuawz,·cn1Nlch1 \bont• in-.·mcm Jttnderbesumd
,;()2...;()b, m Nordrh~n,\\'4:'ifalcm. n.•ur,rlri- ·1,~r,1r:riich,· l\'r,1dwm,tluift. Ul1,
I'll orn R...\.. \\'UM)'.\\ U,\\',, sr.11or.b..., f ..... JIM ftt.\, "·· HLP,H( ... " · t, n,u, .?06..l!-
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. . fgnific:ant cuu,.,.c of O\-Ult' .1bon1.on tn !!ng.l:md aruJ \\':,h~ Th,• t·,,.,nmmy {ill.Mf u.t:.t t.nUJIOrl:lt. fl., Ml rn1,:o,., c ., 111 , ... , i.,,. A <;:O!\ik.\111~. f ,f,, lftt1r1
RMQrd. 140. 175-177 SerotQJ,:1t"I diUta•ntf..~ in Al'O,,p<1m ,·mwzum•.l.s.,oc1awJ epidemic ;Jnd
~ l',\~•t'-'!, \.L. S\'tR.LO\\, l... \\·, - t.O\Jl.UJ, 1•. \., LOO\U.;.. L'...L. kl'll\\'L, J.D .• &.·mh.•Jnft' ubumun, Jt,rtrflnl of t"<1rt1.,;1oiCtJ.r,', O'.'>. ~(•<:-' .: ,
,.1...
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,A.
~h tmi.,nu. J;P, ,, uunL · 1,r.• 1~9 \:-.-o,.poro~1 .....Hkc:-.ll,uniun 111 :i herd nf
ag.gtutinudon tl.'11t c~1r .\'t>W/1t>ra ctmiuutm Ut\ ch:,pmtlm. opttrni:1..1tion, d;l\ry cotdt.~. rm,mal ,Jj \ 'vtrrmn')' VIC1.g11r,mr lm•,,.~:gotion~ I ·io.>-zr~,
.u,d comp.tril\>Cln to the indlrtct flucnc~«nr·,:mribod~ ,~t Jnd
~; TliUP-.:10~. A.S. TilO\U,\.QX,, .,, • UUNl\ J,J• \lj~I. Scwi,Huu aht11don ln
eruyme•Unk1..'<i lmnmnow:b:'tnl ~..a~·. <.'llmrnl mul /.?itt8uus,,c-
~ew Ze,UOJ1d f;lUI~. \·('ft•7.enlmttl H·r,·1vu1r,• Jr,mnal. 39, 129--133.
lahumrcn')' Jmrmuwlog,·, S•.,,i-fi.3
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..1udy o( Strupom rammuu 1n dru..1·y herds. /our,u,l n{11t,•,\J)Jt1riam
ncon;ua1 t· ,11\-f..''t lntt•m11/lu,t11I )f>Urnal /Cf l'rmli>itu/t"Jgr·, 28, l ·Hti-1..:72
l'erm,wn· .lfrtllr11/ ,1,,,,..,,,,11111. 213. 159$-1598.
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86 P.\R:t.. J•• HIEl"Af.A. s.~ I rtlUR,\10'.\0. \.I.C.. 19~:i . An
t:nl}'nl-C•Unkt'd
IIO(Htl , , .lo:., 1-901.J. .\1.•mpo111 UIIIIIIIWt tnfl.~tlon it1 the d~; l)J>iC.J.1 -and
irnmunosorb~Jtt ~\· (ELISA for ~rol~r-.ill d~Ooii!'>,of ,·f/l'J~1mro sp
.1typlcAl ,o;'t<\•\, 1'/nnmJ fJw;h•wu•t>tlwulii: I ,fdscJu,ji. 65• .32&-.:<3~.
rnft.'o("(ion in cuttle. Jmmral of\ ·c1.rri11t1t')' Diagnob'tic- tm1~tigA)tion, ; .
352-3;9. 11)() \ t SI UMISr. \l,C \'t~IUflJSI. L, lt.\U(•\I.UVl, n .. \I \f.'lll 1r. \ . \1., r.LllA1Df,, I ,

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o,\'--.O, \'

rnm,wm infcciion rn dairy cattle :md ~cu11<.."d c:dfhc,nd monal1t)'. ,11. 1~9. ,~a_,,,ou,
«muwm 1nft!'l!uom Jn bo..-inc IOcttnC!i> .11ntl tl.11ty
Oma.dinn Jtmmt.1/ r,J\ -,~Jt.•rinury Rffl'IJuh. GO. llJ...139.
L"nw~ With ~1hottlon~ in .\t~cnuno.. lm,·watlount Jv,wwl/ar J•urnJ!wlog:.
Jq, liO>-lt08.
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l'turl11t11)' /lty,,rd. l·W. 328-131
89 P,nrutct ..... , .. .\1.u,Y, \I.ft., Jl>M~, tt,ll, &-(.H\HI.J.t.tn,._, \\ .\,C.,. 1~,.
ProtQl<>W t·rH:cphJlnmrcHtls ofdOJ6,t in\'ol\'irlg .\'ros1J,l)tn mmmun ;:md 10~ Wl)()OS.. l.W.. \..\:1:>l -so~. ,1.1•• '.'1,,,1,,... P...K. t
.w r ALO\\',~,,•.• 1994. s;·,tctnic
Jru:oplnsmn ,t!,ond/1 tn \'C\'·: /..cahmd . .\'tu• /<>al.nnd \ ,11rrmnr)' Jut,r,wJ. nl"O~pnro~!, !n 01 CJ.llfrt:'01,,1 btnrk-talll'd dc:-c.•t ,:o,t11wt/1•u.~ lt<,mtnrw.r
.;a. 2..11-ZJ; colut>1bfm:m1, Jo11rn11/ nf \'ct,•rf11111')· Ui.t.1Jlnt.A.,tfc /111't1s!;gt.,titJn, ti • .508--510 .

90 1'1trl,u~, s .. l'lfH. fl.II., JUl\L\..xt>. c.... llJUWLL. r .. t lJUOUt.RT. C & w. 1998. \4'o$JN>m ;1bonion 111 c;ull<!. O.'l>l'K"Cb oi dlag_no.~-. 1:md
103 w1..1,m,.
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France i.U-r un Q\'Orhm ~quin• .i.nal}'"~ Cl r>4:11pt•t:1h:~. Ptt.1tl1111t.• m, wmm.,, w OVllf\', ,.r- ,.. J• ~..:ix~. M.c 1997 'icftllog.ic.ijl dfngn~t~ of
Vt/Mi'IJl:r~ Hquint, ~ I •.ll-31. bo,1nc iet~l neo,poros.f, Jommrl uf f'<m1Si1fJlag:,•. 83. 5·15-517.
91 hF1cm t.. ,u.,, t99n. Pl't.'\·.11'.mc(" nr :.\(~JMfll ,lHlihnlliO"i In Xe\,· l<=~l.1t1tt 105 \'.\(!.~. \J,J., ~U\\\'"0-\\ t->t\tll ', {'I. f. JI '>I Ol•'ioll f \, P . l~,J•.Vf0.,{/10((1
dulry cattle .:ind dog~ Ntu, Ztlllnnd V~lt..·r/J,nry /01111111/, .ab. 38.. ,a,bnrtlon :-umn inn midwt.~<ern d:m)·, /ourunl_of \l,•rr,rfr,nry IJ!d_gmmir
92 ltC>~t,,o. ., ruuu1u, P-., ou~f.\. 1.r.. 1998. UirN·, aaglu1m~mm1 Jl."lt lnr lm;-.11i:,,1in11, .,, ;;(Jo-508,
 24
Equine protozoal
myeloencephalitis

JP DUBEY

introduction In the fat!ces. :;poroc:ys1s contain four sporozoi1es and a

residual body.
Equine proto~oal myeloencephalitis (EPM ) h, a n.-urological Horses become lnfec1ed by ingesting rood and wa1er con-
disease of horses in the Americas caused by Snrcocystis 11e11- 1aminated with sporocysts. The only stages of S. 1w11rona
ro11n. The clinical syndroml? was rccognited in the U~A by round in the hon.c are schizonts and merozohes which are
Rooney86 in the 1960s and protozoa were lir~t demom,trated confined to 1he hrain and spinal cord27 (Figure 24.2). The
in lesions in 1974.z. 1~- ~0 lnitinlly, the protozoa \,l?rc consid- lime period and 1hc route of disseminarion of the parasiw iu
ered to be To.,;oµlnsma gomlii. 1~- n 93 A protmman \\as first the horse are unknown. ht an experimental aberrant host, che
cultured i11 Pit to from the spinal cord ofan attected horse in Immunodeficient gamma lmerferon knockout (KO mouse,
1991 and was named Si1rcoc_vsris m>11m1111 by Dubey et t11.i1 ~l)Oromitcs cxcyst in the small inwstine and can be found in
TI1e del'elopmcm ofa serological tesr in 1993 .,6 based on an- periµheral blood withi n one day of feeding sµorocysts. 22 23
tigen prepared from in t•itro cultured S. 11t·11ro11a merozoire~ St1rcoqsri.< 11euro1w lir:,t muhipli~ in \isceral ti,-.,;ue., and
led to the di~covel')' that S. 11eumm1 infc,tion is c<\nunon in c,·t·nrually resides in the cemral nen.ous system rc~S).i:i
horses in the Americas.·' 0 · 1 •1 ' " · 11"' The histof) of EP:'11 and StlfCOCJ·sris 11e11ra11a ,chizom:.are found in neurons, \ari-
it~ biology haYe recently been revitwed.J' ou~ glial cell, and residenllnacrophagc:1 in the brain, 2 c• » q.,
and divide by endopolygeny.~• Jn endopolygeny. the para-
site nucleus becomt>s muhilobcd and cventuall~ each
Aeliology and life cycle
nuclear lobe is incorporated Imo two budding mero1.oites
SarcO<)tSt!s 11euro11a Isa coccidinn belonging to the famil~ Sar- (Figures 24.3 and 24.4}. St:hi.r.0111s in tissue section< me ap-
cocystidae. Opossums of 1he genus /Jide/phis D. 1•irgi11ia11a proximate!)· 5-35 x 5-20 µm in size and co ntain up to -10
and D. al/Jii•<'Wris) arc the definitive hos1s nncl the horse is an meroi.oite,. often arranged in a ro,ette sonwtimc~ around a
aberrant intermedia1e host (Figure 24.1 ), ?I• .,s. 1~- ·'"· • ~ sz SL-a residual body Ii (Figure 24 4). :'1Iero1.oltes are approximawlr
otters and ~everal ocher mammals are al~o aberrant intem,e- 5 x 1 µm In size. lJhrasrruclllralty. $. 11e11rrma schizoms are
diaui hosis.:17 located in the host cell l"Yll)plasm wlrhout a parasit1)ph<>ro,1s
The na tura l imermed!ate hosts of S. llf'llfOll(I are un- ,·acuolt< (Figure 24.-1). :-.1ero1.oi1es contai n conoid. micro-
kno\,11. The sa rcocyststage of S. 11e11ro11t1 and an expC'rimen- nem~. a rrndcus. a prominent lipid body posterior 10 the
tal lntet medi<'lte host were recently ldentified, 11 l.aboratory- nudcus. but no rhoptrics3 7 91 (Figure 24.4).
raiscdcats fodsporocys1s from a nat urall} infected opossum
developed sa rcocysts in their skeletal muscles. The sarC'o-
Epidemiology
cysts were microscopiC' (abom 700 µm long) \dth a I 10 2 µm
l!iick cyst v..11ILThe bradyzoites were slenderamJ tlnv (abou t Equine proto1,0al myeloenccphalltis has been rcport~·d onl)
5 µm long). Labonuory-raised opossu ms fed infer1ed car In horses born and raised in che Americas, Including
muscles shed sporocys1s. The sporocys1s were IO , 8 µm in Canada. the US:\. Panama and Brazil. 1 ;.K. 16 ~t.. -'11 " " ~... ' 8
size. st. 3;, :..- bl ..,~ -::- - ., . Rs h• -\!though fatal EP:-.1 has been di -
.\ sexual cycle occurs in the lam ina propria 01 thC' intes- agnosed in other pans of the world, including Somh ,\f.
tines of opossums. Aiter game1ogony, oocysts sporulate rica.B> it occu~ only in horse, imponed from the Americas.
in the lamina propria of 1he small intes1ine and are excreted The dl,tribution of FP:-.•1follows the geographic range of the
394
 Equine protozoa! my~Ioenccphallli> 395

O;ioSS>Jm" M,crogamoni _,.

ingests sa'COt:VS:

I
' •·'
Matrcgamont
Sarcocvst[?J
•·•I. ·e, .,
Intermediate hos: f'J

Sporulated
00C'/Sl

'
SporocysL

rfarsa
hges,s
socrrllC) s:

Figure 24.1 life cYtJe of Sarcocysr,s naurona

opossum (D. 11irgi11ia11a In ~orth •\merica and It,, rolall\'e 24.5 and 24.6). mink (M11s1ela 11iso11 .'2 skunks 1Meph/1is
D. 11/11/1•.,nrris in Cemral and oulh A.merica. 36 Serological ml!pl1itis).-'0 a domestic cat. 33 Pacific harbour seals \P/loca
sur\·cys indicate that about 30 to 50 percent of horses in 1ht> i•iwli11a richardsi).1;:; Southern sea otcers (l:11hydm lwris
USA, Brazil. and Argentina have been exposed to S. 11e11- rwreis). ' 1 a· and a rhesus monkei· (,\lacaM 11111lmw1.64 The
rona?· '· 14• 45• 99 Although antibodies ro S. 111wro11u have significance or the~e hosts in the life cycle of S. 11e11ro11a is
been found in horses. donkey,,, and m ules, 118 bona fide clin- unkno\\11. Sarcoc,vstf.i mmrona infection in a se:i 011er was
ical disease has not been reported in an imals other than confirmed byli-ola1ion of the live paraslte.; 1 bpemncntally,
hor~l.'s. wi1h rhe exception of a Gram's zebra (Equus lmrch- an EP)l-like di.ease can be induced ln immunodeficient
elli bohmiJ in a zoo.40 • 811 The prevalence of S. neurona anti- mice.~ J~ J- -; Up 10 the present S. 11e11ro1ui has been
body increases with age and is not related to breed. s<?x. and recovered from the CNS of only horses and sea ouers: the
cawgory (race/draught) of the horse.llll Ailhough horses of other reports are based on immunohisiochemical c~-amina-
any breed or age may suffer from EPM. clinical disease oc- Lion with the excepcJon of the parasite in tbe monkey which
curs most frnqmmtly in three- to five-year-old lhorough- was not confirmed by this mcans.:!9
breds a nd Standardbreds. There arc no documented Ca$eS Opossums are also deliniti~e hosts for al ltJast two oiher
of congenital S. 11e11rona infection species of S(lrcorys1is, S. fafca111/a and S. sµeeri. 21. lS. 37 ;~
S"l'COCJ'$riS mmrona-like organisms have been found in Sporoci,-sts of all three species have not been morphologi·
the c:,.:s of raccoon., (Procyon lo1or):!9. 31 • 41 · ~6 • ,., (Figures calty distingui~hed butt hey are geneticali) relmed. 13· lo.r 9;
 .-.'A... .,
'•

,... ,c
' ' •
~""\

/
~ ~
... ~ .... ,,.,.
...
~ ~
... _>
I

,o I''!;
.. .o
Figure 242 Sateocvs11s nevrona ,n ~ect,ons of !he sp,ra! ccrc oi a norse Hae•1ia1:~· ,r a'-d ecs n stain
:,. ?er :a:reular ,nftlira:,on ana nei;os,s Two mfee,~~ r.eu·ons ar'O\\'S} Dul paras,;.; a•e :a•a', v,s,!Jle at 1n.s ma;n,f,ca: 011
ii Neu•on ccniaonmg sel'e•a sch.zon:s larrowsl
C Four sth11oms farrowsl n·,a• ~JS .rages of dc,eltomen
D lnir-acellularand extt1Kell~lar ·rd:v,dual me'.oloites (ar:o\-. n:,adsl

The pre,alencc of S. neuroun in the fMcc, of opos,11m~ i~ over a period <,>f ,\·l.'cks: hor;c< arc kncmn tu have cot!ap,cd
unkno,\11. In one sunc) ::.arcoi)•snsspecicssporocrsrs were while racin,:
found in 51 per cem of 47 road-killeu opossum,.. Bawd on
bions;.ay-s of23 infecce(l opossums. using immunodeficient
Pathology
mke and budgerigars (,\/(•/opsruac-11s wul11fwvs,. 21 had
S. ft1ka1tt/(I, eight had S. 111>11ro11<1 .tnd eight had S. ,,peer,." l.i:sions in hor~e1- urc confined to the CN'i but O(',ur more
In another survey. S, 111wro1w was idcntilkd in theimcMines commonly ht the spinal cord th,m In th!' brain. In chc hmin.
oi 19 of72 opossums from ~fissi&-ippi. U~:\." 5 tho.: bram ,1cm h- al'focuiu more lrl'ttucmty than thi: cere-
brum. <.,ros;, le,ion, consi,1 oft'ocal haemonnage, and area~
of di,colorauon (from red 10 dark hro\\llJ I to 30 mm in <!:O.-
Clinical signs
lcnr (figure 24.7). ,\ny region of the spin.ti cord may be al·
Because S. rroumnn can infect an)' re~ion of rhc brain and foctcd. and le~ioM ll1(1y be confined to one segment or tht'
spinal cord, clinical ~ign~ are related to tlw region af- $pinal cord CJr be dl~p<.'rsed. - '· , ,z ~,, i ... 1~
fecled:37· - 2. 7'1 For e:.."11mple. EP'.'.l lt:!sion~ in 1he :,,p[nal cord \hcroscop1callv. an 1mcephalo111ycli1is characterized bv
ma} result in demarcated areas of spontam'ous sweating or 11acmorrhage5 in. and nccrosb of. the ncuropll as well a~ an
ln lo,~ of reflexes and cutaneous sensalion. There mny be· infiltration of mononuclear l'l'lis in the parenchyma and
foot dragging, difli.cultv in backing. or am'.'(ia of the hind perivasculurly. .:; obsrrved. Chronil' lesions consist ol glial
limbs or of all the limbs. Horse, \\~th lesions in the broin nodule:. and ~c.tttcred areas of mononudcar <"1?11 infiltra·
stem may be dcpws,cd. and ha\'C vestibular nerve dysfunc- lions St1TCC1CJ>s1is Ht't1To11a pora,iw, au.• L1~ually a~soclatcd
tlon.-:? There may be mu~culnr atrophy. panicularty of thl' with thl' h-,i<>th and ma\· be scam or numerous. blll somL·
glute-c1l musclei.. Clinical sign,; ma} bl! sudden or progress may he lo,a1ed :iway from the lc~ion,. In !llOi>t cases ot laP~I
 l'qulm· jlni101.oal m}do~ncephalitls 39T

figure 2q,3 -·a :sm,ss;on e!e:tro~

m,crcg·cor c' a scn,zont ol Sara.ic.-s1,s
monccyte snt'.'111"';
t:61,1u, t) 11iCJ ti.l.rar!e
mero.o,1es t":1ing trom re1.,awa 00,.~1
{R~l Same o' :ne me•ozo,tes are- s;il,
. a112Clle~ ar:o,·:r.eaasJ 10 the residual
'
~

.. :- !.. boor. =·ct {Col: :iar.se gran~le, Dgl.

hos; ~ell m.c:e~s {f'.tnl;
• -. '.i, '
' m1:ochor.Cnan IMil: microoeme lf·/inJ.
m:crop,Jre 11.\p~ n-.cleus of merazo,te 1N,

only a fe\\ organisms are found in thl' ll'sions. Schizont< and bands of molecular weight~ l,l.5. l:l and 7 KD are considered
ml'rozoi1e, arc more genl'r.illy detected in tho q1oplasm of diagno,tk for S. 111mro11a rnfoctlon.r Becausl' 01her species
cells !neurons, glial cell~ and manophagesl and rarely in or $(1rc·ocyst/,· in till' hur~c (e.g. S. fayen· m.i) cro,s-react ,eru-
,·ascular c1\do1hcllal rolls. In ,N·Lions stained with haema- logkaUy In the indirect Jluorescent amibod,: test 'lF,\T' or
toxylin and eosin !I IEI organisms are no1 di~tinctin:. lmmu- other tc~1~. thc,c an: 1101 rdiablc for the diagnosis of EP:'>I.
nohbwchemisrry Cl I I( re\'eaI, many more organhm~ than The polymt-rnsc chain reat·tion P<.R ma) also be used
In ,ections srnint>tl only wi1h HE !Figure 24,8). for the tletl'ction ol S, 11P11ro11a 0:\.\ in CSF ~ Althougil the
1e~1 is ~pecifk. i1 has low sensi11vil"y. I-low ever. PCR may be
posit in: in acme ca~c, b('forc antibodie~ are de1ectable in
Diagnosis and differenlial diagnosis
the (:.SF. !he combinatic,n ol immunoblot and !'CR in-
The ante-mortem diagnosis <>t !:P\1 is bused 011 the clinical creases ,;ensim Hy. Several 1issu1? culture cell lines support
sign~ and tht' dcccc1io11 of.'>. 11eum11<1-~peclfic antibodies in 1he gro\,1h of S. 1w1tro1111 for i11 11ilfo i~olation of the parnsi1c
the cerebrospinal nuid (CSl·J.' '· J 7 s. ":I, "3. ij:i The dc1cc11on from equine tissue~.,.
of antibodie~ 111 scrum only indicate~ exposure and 1101 nee• At presem there rs 110 commercially available S. 11e11-
e~arily dinlcal dbet1w. bu1 their detection in CSF indicates ro1U1·spedfic ~erum for the idemltication of S. 111mrom1. Be·
a local production of sp~'Cifk ,tntlbodie, due 10 im·asion of cause polyclonal Sf/rcoc:y.~li$ sera produced against ocher
rhe C:-.!S bv Lhl! organbm. or their non -specific pre,<mnce re- species ol 5nrcocr~1L~ {e.g. S. 1·mzi of caule cross-reacr \'1th
Milting from the comaminattnn of CSF by blood or leakage .'i. ll/!llro11a. these polvclonal :;era ha\·e been used 10 identify
of amlbodies from blood a"odawd with increased perme- S. 11e11rc111a-Hkt>or~anism,.;• Reccml}, high titres. m!llrrma-
ability 01 the blood-brain barrier 'raused b} e.g. trauma. or :,pecitic scca ho\'e been prepared in rabbits that spccificaJJr
encephalitis or myeli1is due tu oth(!r causesJ.11 ;.: A know- stain S. 11v1m111n in equine li~~ue,. 2'l
ledge of the prevalence rate of subclinical S. 111111ro11a infec- Other prot01.oa 1ha1 ma) be confu~ed microsropkally
Lion in 1he locnl equine pop11la11on as,is1s in the e\'illuation 11 i1h S. m•11rm1t1 Ml' other Snrcol")·.rri$ $pp ..r,o .\'~>osJ)()rtt

of,.erolc,gical result;.. ,pp. 1' ,;: • 7• 7H and loxoJl/a~ma gondii."'; .Snrcocys1is ,pp..c1
The diagno,1k tes1 usrd commercially 10 d1nee1 5. m'II· $Chizom~ arc g,mcrull~· found 111 \'.l~cular endothelium. ;i
ro,w-~pccific antibodies ls an immunoblot using culture- Nt:ospor(I ,pp. di\~dc hr endodrogeny and there is no
deri\'ed 5. 1w11ro11n mero£ohe amigen.37 58 ,\mibodies 10 schiwm ,1age, wherca~ Sarn>cy.<1is ~pp. ,chir.onts divide by
 1q.pm

Figure 24.4 In 11irrode\'!!:opmentai s:c1ges of Sarcm:,•s,is neuro11e ,n bovme wrb,na:e cel!s G,,;msa stain
A immature schlzontS 1·,,;h progressi~el1• d,fferenua~llli nJt'e, ficm 1110 i; The .1ucieus 1n sand in
b ccn:ains a single n~,;:!aolus ,·,re•eas 1n c iO e me nucle1.s nas become looeo ana ra$ several nucleoh (a:10-.·.s)
8 Immature sch1zont wnn mul:ilooed iarrrowheat:sl nuc1ei;s
C Three schironrs with menno,tes a:round a residual bo!IV tarro·.~sl
0 Mature sc11,zon1 w11h :;eriphe•aily sitllllled me102ci;e; ,;;;rowheadsl Some merolo,tes ate a: the cen1,e of ,lie stM~nt
E Free merozo1tes (arrowheads

'•,,' '
•• '

~"'. , ..

•
Figure 24.5 Oegenerat1ng r.cs, cell
1.,tn e:ght 1nc1vidua1 nierczo::es and five
de,el()pi119 schizonts (a 10 el ,.i:h single
le·ge n~'Cleus. Seteoc;sr,, neurol'.a
mero1(11,es can form r.ew schilcnis
w,inout 1eav1ng 1he hos; eel, \f:cm
Dt:~v et al.,:..Q)
 Fq11in1• pro101ot1I rnyeh:,enceµhalilb 399

Figure 24.8 Sec:,c~ of a spinal core o• a n~ise iea\·11•; infectea w,:r.

Se1clJC)·s:is •1e..1on;1 N,11'.etous me·ozc::es ra,:cl',SI and sch,zonts
1anowheads. a·e prasent in neurons tar:o\'.S :.11 b!ac~ a:eas are
S ne111a,a lrr:mur~h-s:~hem1ca1 stain\',,:~ S Muronaam,body

J•

Figure 24.6 Transm1ss1on e.leWon microi;raph of a Sa1cacvsr,s r.i;.ircira

merozc,te .r, '.he brain of li?e racciw, m,,gure 24 5.1..ong,tu.!mal sect10!'
oi a 'T!ero10,1e Note the conoid le). m!cronemes (ml cllld nucleus lnl but no
r~.o,lines !From Dubey er at")

Figure 24.7 f",oss,secuon of 1ne so,~21

corr: of a 0 c•se :ea: It.id suliell!d from
cqu,11e pr .;,o,o,;! "l,eloencephal,1is ·~ote
area of d:~,cratior-!arrow}
400 ,1.,110~ mv: Pro101.oal disca:.es

endopolyge n}'· A.~ yet. no clinically confirmed case of to:.:o- p<Junds ha,·e no t been specifically tested agains1 .:>. 11e11-
plasmosb in the horse has been diagnosed. 41 Sarcocysris ro11a-mduced infec1ions in horses.37• 7.!. 73· 101 Current
11eur011(1 schizonts in the horse are confined to the c:-:s. A recommendations are 20 mg/kg of sull'adiazine and I mg' kg
case of hepatic sarcOC\·stosis has recently been recognized of pvrimetha.mine otall\· in liquid formulatior1s dailv for
in a horse: 17 this prmozoan did not react with anti-S. 11eu- 90 days. Prolonged use of these drugs may induce toxicity oi
ro11n amJb<)d ie~. haematopo1e1ic tissue:.. and ;upplementotion \,ith ,O mg
Equine protozoa! nwl"loencephalitis should be con,;id- daily oi folinic acid may be needed in horses suffering 1.rom
cTcd in the dif!crential diognosis of any di~ease of the horse such toxicil}. Didw.uril, an anricoccidial agent used in pouJ-
associated with Impairment of function of the C~S. such tl') has also been used empirically to treat EPM ..;~ The do~-
as ncosporosis (sec Chapter 23: Neosporosis). t,. ,;2. r:. ;a age rtue recominC'ndcd is 2.5 10 5,5 mg/kg of diclazuril for 28
neurological involv!'mem as a resu lt of ct1uid herpes,~rus- l day~. Prophylactic treatment ,,1th diclazuril pre\'ents clin-
infection, rabies. equine encephalitides caused by alpha- ical FPM Infection in KO mlcc. 28 Sulfadiazine. pyrime-
virus infect ions. and ceivital su.mosis.n. 73 1hamine, and dklazuril ca11 pre\'Cnt muhiplication. or even
•
kill . .5, 11e11ronn mernzoitc~ in a cell culture system 67 61s but
the!} have no1 becm scientifically e"'aJuated in ,,fllo in animals
Control
known to be infe1;1e(I with S. lll!ttrom1.
Horses suffering from J;P.\·I have been treated empirically There is no proven rnccine 10 control EP:-.·I but there 1s
with sulfadia?ine and pyrimethamine, although the~e com- work in progres::. to identify protecth·e antigens.J7 ""

References
IIARKC)IC Uf. UMtf+\fU-lO, C. ~. ll.\Tll<OS Dl '-,~., ~,:-.-10;, DU!- \J, .,_ 1~'3. ll 1'1Hll R, 1 J, \\,\f'.tr..\Y, -~-J.. c,1,,.1•.t .. Gfll'.l:\J It LC. l'O~Tl.fl', A- \0\\UL.
Equine pro1010.1I myclo~nC<phallus in <ou1hom Br01!1 {he t ,•termaf)• c.,,. ~ cw,11:. 1.a.. 1999. \n.• ~mu:p:ls 1tt11rmia.;ind ~.fi,!t:nwl<t
Rtv:tml, 119, 283-26·1 :;,vnc,n-ym(.lu..,! \ hor"'t! ,nfl.ou.a:un ch4111.mgt.'. Joumnt o[Parrull,>log_\. 8!':.,
:: ffEt('H, t ... noun. o.t •• 1,,.r To.,r1p/mmn,Hlu.- l!-11tcphnh,m)dtth in the, 301-30:\
hor,,c, I etl'/1fl<ltJ' P,,tlm/oir:,, 11. U7-~. 11 nnu H, 1 ,f.. \\.\f ""''• Jl.f_ r.t;\, ,,. t. r.mu I!!, ¥.S, 1 ,\, 1,..,us1 K,

J nL\.tl, ti,<, .. r.M~~utnM, n.r ,., ,r.,,,r-su,. , ..

t~f. St!roprC"\'Uhm«- nr s.,, n<\Mf... 1.u. itc,111r1s111. t:,c :tc>01. lmmun0<,.'Qnlil'l~m11 ~J.Jll"":
anttbndl~, in S/1,r°"';{'" mmmmt ln hor~"°\ r4:."iding Inn C'tlt1m; rJf ~rro..]':,ti.t nt•utt~n/1 in mu m~l .tnd ~XJ1mhasanc-tron.t«I ho:-~
<o11th(.'U,.l<"01 Vi•nn,.yl\ Jrri,,. /ot1n1t114Jj'lhr A1111.•rlr:an \ ·1.·urim1ry.• \.fNl/c.111 c-halle11g..\ff \\ith s.. 11,•11n:mn ~pc>rocy-sis. \ ~·tnl»d.t')' P1,ro.~ilol~1~•. 95.
.\.(.$.ot'lttlftm, 210, 517-S Its. 3-11-3:il

"' DL\Tnr.. LI. t.ftAS,TMO'.\t. n.E-.~ HAS!o.T"N, tu•• \\'AIKnt. LL. R..\ll!UTI'. ,. » is ll.u-,_ tJ,M. "''"-"- 1u~., cou 1'-S, , . .s.: ~\·1 JU.<J\\'1 K.., 1v9(), Nt!osp<m1
,·1 ,,wt"·,. ,~;. M:upr~,-J.ll!nn• or wttibt.)dic<. to Sllrtocyllls llt'Utrmn b, Mlc~phuJum~ l)lhb~ ~nd pol)·raaicutonf'unn,\ )nan ng<!d m-arc ,,.11h
ham·!'< rttMtUng in OrltRrm. /ounuzl of th~ Am~rlc.·an \.i'r..•r:',ia!}' \JtdiMl r.u..hin~, di~:a~. li1111im, \ l'f~'"nt"'' Jum,:af. 28. 240--243.
.~~inlltm, 210, S2~~2:". th "·"n 1.tt.. ,11\t:K.,t, x ,.• lU\\l u .. c::..\ , nrru k.. q., 'tAk(n
S 00\\")t\:-, l),[l., CU."il.\11:\(,... J.I. nw,~..s.w.. fJf.1..AHu,,-r.,_ ,\, OUPt)', l,t•.. ,. ~ c,~r:::,.i 11. 1.c... 199~ ~,n-<,c:\'.,ni/hkarul,, from p.JS:Serinc ~nd
!o.tJTUC. M. \1., MO\VI..A~U. 1•.11, 6' C'ON~r.Jt,. o.t.. t992. C:httr3Ch:'ri7..ation nt p~inacme tnrd.,: '} nt>r\)-my "ilh S(,f\'ll\!~'Sli$n('11rom1, .)gtnt of
Sottot~'Sr~ 11t,11101111 (tom o :horOu$.hbted l\.ith l•qulnc proroto-Jl t•qulnc psotoioa.t nwcJOl·nccl,>halitts.. /otmtal ,J[Pllffl.firofo,o~ 81.
m~·c.·locncciphlllitt~- Corn~IJ t ·,·1L•rint1rl.11.n. 82, 4 t-52 930,-935.
fi no,, ~u; .. G.\U.l<-AX, o t. & 01,1 fl,;., -r.1.
Proio1.oaJ ~nC't'p1udom}'c1iti,
ig90. I; l>.\\'S5, C'.,Pt,. G.'\Hrt. f",C J1 ,~c 01 I,"-.. OIA'\:U[A, II.I,~ OUBI'.\', (,'P,, l!i~
m ho~~82 (-.t"'-'., (l'l72:-l9&5 /"1lf1UJI rJ/tlwAllb:rfrrm \ ·~rriJ:tll)' Hcpnuc 'Sa.mx.1·~1ojt~ 1n ahorst./(Jllft:fll ofPurrl$tto/ogJ•. $5, .96,.:;...9(~
,\11.'lllcnl .o\'s$Qli11u,m. 196. 632-4;34
18 u,,, .. . ,,\ .0,,11. n.,1 &ouan.r.r•.J99t.Wc0()1tisn,111m1:ncuhur~m
; unt,\t1t, n.u '"MM11Ew, u~ .. N~8. MutttrOC".al ncurolu,gacdi~.1!1<" in ,1 t•,rmlfom .t horwwllh C'qume pro:010.U myt'iih!I>. fq1m1t, \·a·rtrinury·
horw. Jounral qj F.q11m,~ \ t·lrrinary ~nrun1, ft, 302.~0.; Joumal. i:t Jn-..,n.
a r 1,, 1•\110~. t.. ,, tUii Pu.11,,1.(tJI unct!phal.1Jm}clJ1is In horse,-.
1'1ffV\\:,. , 19 11\\1., ,\\ .. ')f"t--Ur, ,- _,
"'uuan·, f.r-... 1s91. /n ,'irrocuhn'Dtlon-0(
Juuntol uf tit,• Anwri,nn ~ ·,•1t•rl11n,.,,Mfflft11I Assor:mton. t 71 • .;92--4'12 ~">Orr.·,~-.,~Hi1 m·urr,,,a Imm the sp"in.al cord of o1 horse with t!'qulnc
U {111.Al>I.I, ~I.>\ .. us'n:,..\\. b.:r.., ft(')WY '·· Dl'XSl"ltA. C.C., \\11.tt,\.\~S. )L\., prumzoal mJl•Hh!>. fo11nmla.fPur<1...:afJlr1~·. j;, 78!1-79?...
,1•t,..r-..l!n. , .., •• TOrvtO·>.l~XUCA., ...,.,\,. Ll :i.:.t. :,:,o., '-f.\\'Tl'),, J,f.. ; km \\t.\, '1,uar.", I.I. h WITI0\,\1 0 CI., 1~80,
:t0 UO~H, I I., lllv~I~,, 11.1, U,\Wl>U, C..\.,
\t.u. & 81.AG.uun~. B..L 1~9lf. Pl'e\'afonc~ of 11n11.hodlc~ lo ,.t'OJf>Orlt. sp. tn l'ro1010"1 mitlocnccphallih In hnr>e, In C.:alltomla /Ottrn<tl of,;,..
hor-..•, rrom Al~b.1m• nm/ cl1•r•«er11,4lJ011 ofru: ko!arc rcco,<>wd from .-\mrnam hftt•m1my M,•dh:al .~s<K'U:WQII, 185, 801--802..
a nu1urally in re.red hor<e. /111,•r11arro,111/ Jo11m11/ of/ltrr1Uf1c,/ng;•. 29.
21 mmn.. r> 1-0()(). l'rc\lnlenrt· of '5nrc(Kj\'Crs '>l)~c,ts ,pcn>cp,1~ in wild
15li-1S-13
cm:stht opo!i<.~Um'li f/Jld<tfplu, :•iQ.¥JnimU1), .'oumnl OJ f>ttmsltt1lqg:.•. 6t>.
to c.uso:. t .(... 1'()\\'NST \:n, 11.r,.(1 I,, :i,,~.:1 XI.Jr, o,;.1., 1981. E~Uine protOi':0,11 iO.l- -lO
m)rdtk..•nc.:phulnl..,. ;t '"-'Pnn or n,o e;,s~ from \\'~Crn Can.'l.da.
Ct111ndfo11 \1et."ri1111rr/01mral. 22. 1.;0,.1.a.1. ~ ttUPt.\· 1 r. :woJ MiJ,;.r.-.dou ,md,h~\,clopmcntof.S.--m:oc,,·,'ffis11i•f1m,t11ln
U~\ue~ of mt('C' kd ~poroe) ,!1t lrom ,1 :unurally ln!col'd opo,.~µm
H conF.,. x.n. Ci(,\\,~,, k..f .• 191r;·.. 1merprenngfmnn.rnoblot resdngot
t".4,•t,mna,,-· /Jauu,wloro,, 95 J·f1-...J51
t~m:hrm-]>in3J lhnd for ~Uillt.' p10:01.0af myCIO<.tllCl'phaliti.S, n1f
Com11t11dium "" Cormn11111g F.dttrflnOPI for rlrt Pmrri.tmg \'e.r,•tmarinn 23 uu11t\·, 1.tt.• ;:i>01 P(trti)h\"mh, J.nd t.~~· :i,i\Jc migr.nion ol .S(utot:;in>
l!I, 117~1181 n,•,uq,111 tn tntcrfcron g..,mm.1 knocko:n mice fed ipQm~.,_~ Joumttl QJ
l~ CUMi.);, t',).,, ~tlJ~" .,_ \t,. H.\\lll l(J\. n.1• ~ II \M')f'\'fUt()()i,;., u.r•• 1974
Pmnsiu,ft,-,o·. m prt.•>,
Tnxup1a ..mp-,i., in two h•n~~ Jou,n.11' nflh~ .1mcrican Vt·rl'ri,,,u;,• ll Ll\JUI\ 1.t1 ,.._ II'-' MIi, c..r,, l'3~. TOJ:OJ>;fllllJOd.:JofAuluwts ttntl \/o•t
.,1,•rltral /UJot/m,on 161. -;;-ao. IJ.oca Hilton. Flond;:r, C:JtC Pre,--. t-.?.?'1
 F.quin~ proto1,oal rnyelocnct!phalitl, -10 I

:& DUftEY, I .... RL\CS. s.,.. "'~""""· l,C. .. """ '111,\L ,.,,. u,0,.w........ J:\ nua~· , J.I.. 'f'f-fl.ll. t \. .., U'.\D~r\\ . u.~ .. 1993, liot:ition ur a thlrd ,.pc.-c1es oi
,tsf.:\. s )!.:•• i;w<n:. o.r:.n•• mms1, "· • JVi.~1-1,mMt\\l;:\, A. 2000, Prmalcnt,• Satf0<,,1,, in lmmunodelicknt mlet, fed fou:> lrom opo"um•
<•r c.«n.Yx./"ltU ncm.roun ,-pottK") "" 1n opossuins Dltltlphil nrg1111n,m1 Didrlf)his l'lrg,1111mr, •nd !1><!1ffrrenun1lon from /iiir,·,l<)'stCs{,1/mwla
rrorn rural ~li<>!.slppl. \•l!tcri,1111)• 1•1,m,lw/lJl:Y, 93, ~!IJ...293. ind :,nrra<y,tls 11<•11,01111. /uumul of P1111nl1ol<Jgy, 8~. 11511-1 1r,1
2-6 OUUtY. 1.1•.. DA\")~. G.\\',. 1-:0t:S'J~rut ,\, ~ Mll\ u. }... l9(-t.. Equlnl" -44 OUbt\·, 1 r•., ,$11 hll, C'.,\., HM11R:, A.X., t0J:•l•• 1M, M,,., ttMO\\'~, C. A Rl.111PkU'1rT,
t.'11.cl'ph.nlom),~lltl,j. due tn a protoxnan r:irJ1'U'-" rescmbllng 1oxt>pUumn c.,.. 1991. Otv!·lupm•ni of• Sortocy1lli·ll!Ji •plcomplc,•n protozoan In
~,o,ulli /oumnl cf:lh• Amniru11 \l,1111rin11ry ..\J,:d,i--al AuocJtuim:. lla:,,, tho brain oh n,ccoon il''tJ<>·orr /u1or1 Jqurrutl u/llft'lfrlrnln1i1<1log1eo/
2.1,t-a~. Socleryof\\'mhtngu,,r. 58. ?50-25i.
a,; llU8£'r. I..P.. D,\\lS. S,\\',. SPU.A. C.A.. l\()\'\7'V,~. n.r>.. Of L\JfU~T;\, ,\., 4-."i ouaxv. ,.,., \!S11JflL'\'.I. \I.(.. ,,,.,,a,,'E\'.S, ltP£C0AA.kO, .. , •• lt)l.l,9
..unn.. \1.\L
G.1t.\~'Sl1t0M, U.£... TOPPER. )1.f•• fl \)Ille. A,S .. CU!\!'.\IISG~. J.1-..:. Prtl\'3lencc of .lntibuJ1et, :o Snr,:ocyttu. ntutoru1. To~o,,lasmn l?(JtUflf !nd
L9'!1 Snrrn...).•t£t ,icmonnn. sp. lJJrorn,n:i~ .\p:c:ompl~t 1hi.- t.'tiOlogi< VN,,.pora,,wi,wm in hm~e~ f:0111 Arg<'olin:t.. \'Nrrim,r.· Parnsiro/()lfJ.
..i~.:nt of cquill(! prototo31 1:wdu,mc:t:ph::ll.itl<.. Jounwl of Prrr!Ulto/c,gy. 8ti.59...r.a
77.212-218 -&G fJV':tCJ\.,.\, C.L OC.IJS8lt-~ \f,1-~ PODu.L. W.. Mt 11 I:,, 1~11.." J1f.£D. ~)I 199..Z,,
~8 OUDCY, J.r•• flUTZ, u., U!'-i>SW, p,'l:., SHI..:. 5,1)- KWOK, o.c..n ... llCO\lf'~~>S', St1iurc-O.C!thit}' ~~ocrnu•d \\'ilh cqume. protoZ:oW. mrclocncephuliris.
...., , ~01 .Oic.:l;w,rt1 prt'\'t'ntivr chcrnpy of ~mma imcrfon>n 5.:noclout Prog,eu in I <1emt11ry ,\,:1110/ng,•. a :10· s,1.
mi,c ft.~d Sarc:«j•#is mJUf(uut spol'OC)'StS. Vtttf,nnf) PattUltOlog\'. 9 l,
·•7 r,,Yll.ll1 A..,.. ouv.~· p•.. 1~. Uevelopment.uf 5'HrQc'J~t1J fn.n•n nl 1h~
257 ?63.
1.-qu.in~.Ju."rm.1/ of Pflttlfil/QIOJ...."i, 68, ~;6-,.860,
29 outtn'. f.P 4< HAMTR•.\...,.. :?O(K).
Jmmun<th1sl()chemlc:al continnauou of
18 un. R... ,1.wmw.1.G .. •uau. J.o. ot.u.. ,.c-.., •utu:~1A,. pt rox.1.c
f ..
~'\lt(((}(')1ris 11euro111,,1 mf1.-ctions m r:tccotm,. mmk. cat, ~unk and pom·.
tll('"'.(;lSS.JU.. m::iu. ~ . .t •• kU(J ..-Y:.
w.w.. S\VEL-.:u·. R.,W ..... nr-nu.. P•• .isgo.
Joumnl qfP<1ra,110!0',,zy, 86. 1150-11~?
Epidemiology ol ,·qulnc pco101.oal myc.-locnccphahll~ in '\onh Ame.rfca
:~o OU tit\ f. •• .. H:A\Hft.. ;\.:0-',, :,.1FJ'~1M, M. l· IIVPP"Ctlt t. Ci. . t$96, A based on h,siotngically confirmed en~. Vtrtarinm)' lfllt/mnl ~!cdlrmr .;.
~rc<X')~tiJ ua,ror:a:,Jike: organi,m .1('-tH:httcd \\ith eni:epbalh1\¼n :t S4..S7
stnped <kunk fMep'1{1i, mvplr/11.,1. /~mrtal ()f P/na,<ltulog;,· 82, I 7:!-17-1.
49 M:SGFR. C.lt, C...AA!'-t'>Ulfl\t, U.t'.. , t;, \tADll-ul, A,i\,. \\'II I IAM~, ,·.:i.1. \td..klW,
ll l)U!U.Y. J.1• 1-1.\)UR. A.~ .. UA.,~, ()\'., (!.,\,. nw-s•rn, '-1,J, '-' Nt1rr1n·f.ttT. c.;.r... S..\.. ff \MPUt. <.,. L.\M,l\1flt1'. J.L .., UUHtl\. '"'" l~i' li.'itpcrim,•ntw
19!/0, F,ulll noct<>wl11g oncophaliti., in II m..:oon 115.~l.u•d wilh n Induction of oqufno prmo,t>.1J m,~lo•nccphalhi< in hor>N usini:
liam><")•.<lu·li~ pm1<>1.oon. Journal of V,1r,/n,u;.-Diag11011« Su~,., ,p. ,poto<;si• from 1"1: opoS<um 1V11M11ltis 1•ugln/m:a
lm'f'JtiCation. ?.. 34S-34~. Vc.'trrlnary• 1'ar11st:ol<>~·. 68, 1U9..213.
?=t ouuo . J.t'. &. nmsrno~t. o.w•• i993-, '.\.1t-nint:t,o,i,,ncepha1iti... in mlnJ: $0 H -.:cu., C.)...• tHt\S\tllO\~ U 1 .. U...\;GE.MEIER, 1,L. t:..\JAl>IU.11:•.\,. t:.On!Ri\.~,
A:$AOCltth:-d \\ith:,. Saft'OC)·Sllf 1u:urtJJUJ,1ike orµDism./011.rnol of c.. Ttt.4,,to, ti\," 11,. itA-Mi'lW. '\.... » num \', 1.r.. 199-1, Ph~ IPS,J:nll;ic
\·i:termnn· Dla1,:nos11r luhtnlgar!o,,. S. ·i6i-&4 t. rcli'l.tlnrt'Shlp of ~,r~•stit. nc.urona tu oth£or mi:mb<'!f'S o1 the fa.mil)·
33 uuun, l,J•. IUGGEXS. R.1.. '1ARN. 8.C.:., SPAS(:U.R. w.1.. , KOW~. B, Ir ~,coq-stidtt• lla"'d on the w~UMCC of 1he small rlh<)S<rm•I subunit
JOf(c,L..::0-1 N, t ....~. :994, 5arrot)"ftlJ.•SIS.SOCi3t£"d m<"ningoc-nccrph:a.lo:n) clitb- ~ene. /01m:al ofPnrasiro/r/;r.•. 79, 966-975.
fn o cat. /OJ."nal o/V;11urinary D1atno..mc hw11,.i.1lgnnon. Q, l i8-l2.0. St f..L"lCt:f(, CJ:.~ GRAs~·tRO.)l~ u.r., IA,\;Gti~iuru, ,.1_ 4i: ~TA~lP1,H, ~-, 199;.
3'' uuu1.,. r.;o. KtR1J'O. c;..c ~ VfL\'.\<srnoM. n.L. 19,s, .strologic prcvQJt.'nC\! of Epizoo1it' of cqulnt.• protOZ()31 myelnttn~phnlltt.s on n farm. faurm,t of
~rro..j•stis r.n,rnttQ, To.w,plasma gondll and .\',YJSJJor<i (',tminum in 1l:uAmtrf(a11 \ ~tt.rlm1ry· \frdicnlliss«iaJl(m. 210, '+2M~-.
n1nroughbred hot"Jes f-rorn 8r;uil. ftm11ml ()/ thi' .4mcn·tnn t ·e1Nmt11J· - lmM. n.L lA."\:GbJtill!Jt. 1.L. ~, \MP(lt. 5. t>OS'..UIUl,
S:!: n_XGl!Jr. C.11:.,. GR.\~;)o1
.',t,·diwl A$$JX:iolirm. l15. '97()...!fi'2.. J.M.. PA1T1'll:li0~ J:,... t;..,J.llllJUR. \.\ .. \t.\lt~IUJ.. ).\ XIU!\11' 1 4

35 uu-ur\', J.1•.. .a. u:--ti5'A\', D.&.

1998. l,olntion tn lmmunodcficitnt mkc t,I ooan 1.1•. 19~ ldem,fic..Uon of op~ssums IDtdrlp/1/s 1•1f81nltmu as
S<,r«IC)·Mis nrurt1na ttom OJk>!>!>urn (Di(ltl1'11i~ i'irgirttlma; fatCt.'\ and h.s 1ht! putati\·e tfoftntt-i\c hoi.1 of Silrt«)'J.'tU mturn,ur. Jqumnl of
dJfretemiatm:, from Strrcvt::ttth/alc,1111lu l11trr-11mfanr1l Joutttalfttr 1'11111sJ1otogy. SI . 911,-~19
P1m,,1w/Qro~ 2a. 1823-1828.• al fRfrt.. D.L & DU8f\. 1,1,.• :tor.u. P3thfllogy,>f SU('('()tJ'i,((,1 m~tlfOllll In
as UUhf\, ,.,,,. U-!\"D'M\', u.~ •• ti 1rnMc, t,r.•• MS.\I, s •• N~'\:A, 11.1,.T., >,'\\'OJ:. imc-rferon·s.i~mma knc)ckc>u1 mfcc. l't1rri11m)' Pmhology. Jn pn.•,<.
O.C,IL "u ., '!,,)i., &t RO!o!'!\'TIL\I li.M •• 21)(H, Flr&I lsol,a.1lon ur S<JlliOC)'Stis S-1 c;.QA~"STRo,,. o.r .• 1995 Rc:<cm .,d\'0.flttS in tho IBborn,o[),' d1!\SJ10}h oi
,r,itr11nn Iron, the SouthAmortc.in. Di1Mpliis 11lbfr,11rrlJ. Crom Bra,il ('qU:fl~ paroh,!tlC: d&s.f".s... t•oJ. \ l'lt!rfrt(tryClinit'S qf W1rlf1 ,,m,'TiM: Fquirt,'
0

l,'etc•rwar:,· Paraslt.olt>gJ'. 95. 283-293. Practi~. I J ,13;-4.;z.

3; DUii! \', J.I••• LISD>,\Y. OS. "-<\1U.r. \V.J.A. llU.(). S" OJU ""'"°"· (),[, • nuu-..v. J.P.. CUMUl. P.F. .i,. \\U.U .\.\l'j, ~ -,:..
55 CR.-\:-.'STI«},1, 0,(... ,\.L\'\IU.!I'... 0.
)PEl.11. r:.,., 2001. A rcvfo·w of StJ1t'OC)1$W 1uuronaand t'qUine proiozo.:tl
1992 E.qume prom1.Qil1 mye1ith in P-..mamo.ni.m ho™'j ind 111Ql1u:c,n of
myelocnc..,phali.tis ·EP~H. Ve11.1rlnlll)' /'tU'a$lUJJo1;l. 9.5. 89-131. Sar«<Jlftfs 11e11ro,iu, J11urllfll ofPlltQ//lrt>/OffY, 78. '1()9..,t Z,
3b ouon-, 11 1
• >!ATTSOS:, o.c.. SPI Ht, C , •• IIM:,1,,.Jt. A..J., MlJUtOQSU' D,,
;G <•Ml<STR<»1. n., .• """" ,.e.. o~'1>,>-"" M,·1•. R.. ,ox, 1.c.. roos ..:11,.
mN~QUL~t. $..J,. H.A\l)k,A. & G(RROS, T.r:•• 1999. Charactctrfl~tlon of 0
..-c:o,u:R, P.f .. 1993. f.qu1ne,proL07.001 myelO('OCCphalltis;
1',11.. C.11.E~. R.C
Sartot)•s.'U ,tl!uron<t isolmc lS~GJ from 3. n.owran~· 1ni1.1c.ted hor$e from tultigl.'n ~nal}~1ot ol cultured Sa,coq-,rls nffmono nti:to1.01~. /ourm,I of
Ote~on.Jouma/ of E11k11ryolfr M/cro/1/0/q/lJ'· 16. 500-506. \'lltenmuy Dfr;gm,sm: J,zr1:;ttg<Uiorr. ~. 88-~.
l9 DUffn,,,P., ,\ l.\lTiOX,D,l!.,$Pl:fM,t..,\,,UAMH<, .\.~. Ll~~M 0.\..,
5-; CRA~~rtt()M, D.1:1 .. WILf'.. K.C , 1UTrt.r: P.A. 1 W)W\:.-1~. '\'. \t,, j'()()'.\.,\(lt,\, U.,
ROS£..,~ tc. \I ;.,'\\UK. o.c.n. ll,\lr.UC., H.J .. MULKOO~E:\", D~\S., TOR~Qvs,;:r.
nnt!Tn-):UJU'fl\", ,:.a, tl't\MQ:'\'"TI!\, ft.It, ~Wt1'CZU;. T .\\'., ICOSG, C.S...
)., &CiLRROS. T..c., ::001. Char.1c,eris1ic:-s of .a rt.."Ct'nl tsoli:uc of 5<:fC~'lttt
lmrnunohJ~tochcnuc:at
RU.ABE~. c;,&. 1'0:'\""" f-. 1 • .,. DAVlXi'E, J,H,. 19.9i..
lttmtma ~'.\"'7' trom ~ bo~ and lo,.--suf ptsthogc:11dry o: is.olate!, S:\6
dtagnos-i\<.if ptotor.o.an p-1rd.'-hc\ in lts:ion" ol cqume prt>to,..o.;d
Md s:,:, by pa<;.>1tos in cell culture, 1ttori1u,ry•PM~itolog;·. 95. 1.35-t 66. 1t)ycloenccph.i.litl.).. Joumal r.,/Vi!t(rit1af)' Dir.gnust" lnt'(,.(1iJ;a1u111. l.
40 uv1n-, .r•. • ~uu.m;. ~.. 1986, Equine p11>1Uwol mydotnctphall1,< In• ;5-7;,
pof,y.Jo11maf oftht'Anh'rtrnn Vnl~rinnrt .\lt-dfrr1I .'1.S.S'()('intto,r. 188.
58 (i.M.'UTftO)t, O,t \t,\C't1•ti!R..,lo.1J.S, l,M,. C.\Ji\DflAM. A ...\.. DlUU:\'. J.1 1. ,
UI l-131Z.
tMAM,(>:r-,1:i. R..k3>1',\>,:PUt. 5.., 19,i, OiU'crcntfo.tion ol' S<lrcQC)~tJs n.~urona
.;1 nune\·. ,.,... S.\\.':Ul'. W,J ..\. U~OM\'. u.~.• ~41(.H, A- \\-STA.,...,A, J.f•. '\rtFn. rrom eight related coccrdl• l>)· rondorn amplified pol)'morph,c O.-.:A
('..\.. , MO!iE.'.\,H,U. 8,M .. SJO~U, W1, tr."\\'O>,. O.C.H .. '-10"'\'., ...,k i tu:to. Ii. \1., ')S-.a)·· ,\fo/tt11k1ra,11/ tc/111/0., Pro//~. 8, 3S3-35&.
>ooo, C:omplt!tlon of life cy<·le nf S/1tfll<)>II.< 11t11ro,1a. Jnr,mal Oj
S9 (j;R,\.,~'TltOO,I, t).f '-1•.CKt~ ~ . \\1Jlr.f-\tkOtU1l, C., &\$i..LJt, C..ll,. C.Akll X1
Para;110/ogy. 86. 1276-l~O.
\\ . • H.,\f{J;J:,.'.')-, TOD:\, ' tr"A\'IU! W.J. .,,r. Oi.l:J.u.uril and ,:quln~
n... t9119~S(-lf'l't>(".','J:torilof.4n1malln11d
~ 0U"FY. J.P.. :iPUR., ('_\. II f.\\U, protoz.aal ~clocnc cphahti~ PriJN°,!dlnp Of th.: ,..\ss.ot'i111fcm ul,·lmcrtcr.n
.Wm. il<>c• Raton. Flotlda: enc Pre,$. 1-215 G111,in.-Pr,wlonm.43. 13-1-1
6{) KAM!I', A.s. ca RROS.. r.c. t.: Quarr. 1.r., 1991. Pyognanulorn:uou,- 79 'S.\B-SH, A t IV,"RR. R,C: , nu I "'()$Kl, M.. (;13,Ll~'l!H, E. l\A.Ml'.. J. &- (()'\;ft \I),
c.ncrphnlJus a.ssoc1att-d wtcb an unid~mf!i{-d N,,.,'f1C)'.HU nt•urona•W:f' r•,.\. ts...q;. In :•Jrro culu\.-nH<>n .,nd cxiw:omental inocuJonon 01
organtsm in II horse.Jo1wwl o/1°dumwry Diaf(nMti< hwr.i!t~11w1, 9, 5'.lftOC>"Stufalcmu/a and 5arcot:),·.rti.s nt.urontr mero1oiu.~" tnio
331-333. l>Ud~cn~~ f,\ft1IQps.JtUJt'lU undulorusJ.Joun:al o/Paro.~rolo;o•. 83,
6J Jf.\~llll ,\,~ .. \IUSUC. (i,, (;,\IJJ(oj\S, U,T,, D.IWI~. '\".\\"., CiR.\~),ftU~. Jl.t, 4r
1189-1192.
outt1 f. J.r 1993. lmmunohi.i1<>diamlcal "-tmir to dt.·mm,~trntc 60 ~IAll:iH." ·- IWSVffl;. ,, .. mu. Ef•• \JCflH.AXt:Y, ,\l,tt,,. lfflJP);JJ;Wlr.i. J,V,.
Sarr.'OC}·s1,s11r11rona in cq111ne pro1or.ual m~olooncepll"11as. Jo11mal of srnw.,Jtr. ,. " TlLL. a.... 2000. D<:tection of :,orcor.')"Jru nturonn In the tm1in
Vt•rerilWl)1 DiJ1K11<'$fir. Tm,esr,gat.iott, 5, 4l8-a22. ofa Gran,· <ttb:n \F.,/1111, /Ji,rc/lrlli bohmiJ,f()11mo/ of7.<>o ,m,J Wi/1//1/,·
,;;a 11.\Mm, 1\,:-i •• Ton,.,v,u,'.'o'f, ~., .. (Wk(<O~, r.c•• l~PPTJc, '.\t,r . .q OU"'8f>'I. J,'P., i998.
,Wcdlc111,, 31. 82-M.
\'twpcm a,m,mm·~c:i;1t'!d equine pro101v:1l m~·c-loenct·pha.tiu!<-. th ,tASJll, M,Jl. w1•1., lU ,\IP,\,,. a £1Uff..E!.\', , .,,.. 199~. SarCtJf.)'illl
\'1'1em1ory J•orasttofog:.•, 79, 26!1-274, nctt1fi1111i•3!)5(}C&3ll'CS ~uni;l in hm~-s m Bn:u:tl. Ve1..-rutal)· Ptlrrm:o/1,g:.•,
63 n~., 1997 f qutnt- ~1roti>1:ua! n1;·...tm·ru:t.1>halilb.
,.,..,T1tARU1, ~-" UNO!->:\\',
•H.311•31',
fqulnt /'rawct. )g, fH3. $-: ''"'11tw.1.G,1,GR11:•..:aJ1. Lt.. i~tt Pr<il(M(>ol dl')4.~ Equmt protcW:o-,1

i~ Ktu,,vr ....4 ,\'\'.OJ:Jli:i;.oN, D.(".. , .M t:CLURt H ,1 . kOUhn J ,,.. igg..:

myeloenceph.Wri) h~1m,wr, (.'flrrtcs o/.\'orth .-,m~ri'«i: Fq,1inr l'rii<'tic.:.
?.~3~59.
Enc:cphfttr,m~·eJhi, du-t~ tQ a 5arcoCJijrls 11-·1~ronn°Uke protc,ioan 1n .1
rhc.s us mflnkcr (,\!m:n:ca nmltut<l.} infcc.'tl"O wuh ~mmn 8J ,tooJH:. o tt.. GJlt.A!\~nm\l. l>.f·. 1. '4.1 i-u, >..l.f•• 1995, Oiagnasl) of ~ulnc
fmmunockfi,tcnt')' ,'irus. 1'11(,rfcan /c,unwl oj tropical .\(ttl{tutt.· n11d pro1010:tl my•I04..,t•plt~li1i, :ind ee"ical ,1enok n11,·lnp;,lh). lit,•
/(;11i,nr. 51. 332-338. Compmdium un (,Qrtllnulng Educrttf,;m forrht frnr:,smg \ et,.,.imirf(lf,.
17, ~ ls-128
fiS L,\I'Ql~TL f,M .. OUTGXA~. r., .. MMIS11'1 ,., .• c.UUASD, r.M fl'\RRt 8,C.,
SM'tl\.,,, D.K:,, lr:1~,C. r-,.~., fl\f\MA~. CA.. HU:'lo'TI~(.DC;ll'I:~ fi.A. ~ L(m};"N~n ... r,, 8,1 ,1ac.no LA. 11>11:\:-S{)!'I,, r.1 .. )1(!.»f"R, ,., •• u l!\'l". tr:.I • r:RU\W, L"· ,. ~:,.um,
1~1 .. 1998. Mtl\lO){Ot'n\"=cph3Jili.s du~ to a .')(lra1()'lti.$11ttur<m"•hkr J,R., 199;, ~1~n3$rmvn1 ofhv3d,ha,lng In three ho!"><'' by ,re,11mcn1 for
pro10roan in Pacific.harbor seal\ V'Ju1a11•iwlma rtthartl.sl).Jounwl of pmt(Tlo,11 rtl}L'locru:~h:tlhl!r-. 1'111J V,-Jerinary /lf;'CY)rtl. l,: 1. 2G:-26i
PDrasuoloi.~·. &1. 1181•! 1119. 85 R0'-1:.S. ~ _1992. Puu1tht" tctulnc. prot~>al myetocnn:phallo.> in au
b6 U\~(;. ht.. (,R\S:,11t()M, u.t.. 7,UAO, x.~,... n~~o:,..n' ....... 19~ F,1dt-nc:c- ,mpnn•d ArJbl•n ~ll).founu,/ t>f1htSm11/1 ,\frlt1J11 ~,·1.-rit11ll\
1hnt .,.urfnce proteins Sn14 and Sn16 of Snrmcy,m.s ,w,«ona merozoHes As.ioelllflUII, 63, 78-79.
.are in-.·olved in 1.nfccnon :tnd hnmun1ty. lnf«-rioa anti Immunity. 66. 86 KOO,L\ •••••• P'RlCl-rrt. \1.£.. OD.A~H'r hM. 4.:Ck.OWL r ,\\ ltr.O. f oc.;sJ
1834-1~. m~~Hri.;,efl(tph:t.llti~ fO hor,l"<,.. Cc11u:ll \'e1t,rJ1111rian. SO. 4'9.:-30J.
67 t1,~\\'. D..$." ouun. J,r.. 199g. Ducermmauctn of ll,t 3tUn1y of 87 =,1:1.•.,.. nRO\\'S. S.R.. t0R:<Q111>1·..., .. s,-mut. $.P. c:,it,En. "·"· I.
pyrlmcih.11.rnlnc. 1rimcthuprim an<.! ,uUonamido and comh1natlon::i. of ftl \Tiff. Lr. t\)99 f.nttf!h.1lom)'~"li1i.s: :tss0<·t.,1r!d with 3. Sar(Ot)WU
p)'Tfmc1hamlne nnd sulfonamides ~g;dnst Snrcr,cy,tu ncumr.a In cell n111,rv,zfl•llke(lrg.uthn-t in :1 ~t~t 011cr./cmma/ of,1,,, ,;mcrf(tln \4•:,,titut,;•
cuhurt.~, Vi~rermal)· Pam.ruoloiIJ·. 82. 20:t--22t At.-dira/. bw.:ia1u,11. 21.;, 1839'-IS.12.
6,C, t rsu...".1>-" &DUflUY, JJ•.. 1999, Arti,,ty of dh:l,tzuril 8tt:dn.1iil t.:.<1n"Oc)>Sris 88 S..'\\.'lU..f', W.J., tU ti), :-~!.t,.. (at,,,,;'STJlO!tf. D.E., HTS'C:UCUt I, )CW,. lt:(U", C!.\'i ,
th'Umrza and &1ra,q~1i.tf(llcutuloin bovin~ turbimnc «•II cuJtuh!5,.. WI rro,1. T,t. • $TA.,1,sR. s., 1997, S<lroprcvnlencc of QO!ibodl,s to
Joumol 11/Pnrrtif1uf<>/O', 86, IM· 166. S11rrt>n'StIJ n~u,011,> in hurM:'!oo rest<lln~ in <)hru. Joumtll oJ th~ ,\m1.•rfcun
~ U~DMY. n,,,.,, llUl:JM ,.,••• HOH'1'0:0,:. ):,'M ""Jj(>\\"\u,. o.o., l.!$9. l ",•r,,rim11)' Mtdit:111 AJ.U)l·lmion, 210. 519-524.
!)CV<'lopmenl of 'i<11""')'>t///flltnrr,ll1 In ,~u cullurc, d,•mon,tm:e, 1ha1 89 S..\\'IW W,J,A., 'o10RU ,. 1° S- Jef.O>. S.M.. (iltAXflf:.O~t. or.- );r.llf"I:, ~M-
It l'- dUJ'crvnt from Sarcocy1,ri1 mtunma. Pmcuuotogy l IIJ..?27-?33 m:'lo't'II\.UU·. c.,,. J. wtn-u\C.. T.r... 2000. Evahuui11n of n;;. r..r,o~
70 U~l>\....\', D.~.. ~XBt'1t<-. U., DU&lfl..tm, ~.)I•• , . \J~\b• .S...t>., k()St:U:, U.M,. a,..._-«,ciat«I hith cllnital improvement and .$lln'l\'al of ho~ \\1th cquin....-
\fll.Lt.R, Jt.L ,., utA.c.,o:u&\, 1t:.1.... 199t,, <"...cntr.11 ntr.ou, ~v.:,.1cm neosporo,i,; pro10>:03J myclocm.·cph:ditls,/01111u,lo/ tl:t..:lnJl!rirtm Vt.•11.•rl,:w~
In a foal, Jo11mal ufVi!M'in11ry D/ognos,/c lm~Slfgofl'on. 8, 507-510. Mrdir.nl:\..<.((11:iillion. 217 118J 1185.

71 UN"OS..\\', t).~ .. 1'UOM.A5. :'\,I,~ OU60', l ,•. , :!Ot>O. !Jicitoglca.l chMllC'lt,dl4lion 90 S..\\'UJ.E. \\.,J , .. ,ruo. ·fM .. ,~oRtti'f. r.s ... oAA?-"'STkO>,f. o I! .. ,~cm,. c;:,\\.,
of Satl«)'Sti.~ ntuf'tJtlJJ from aS0u1htrn se:J. otccr r &1Ji}'dt11 lu11U 11~r&J. HJ,UIWH. C. \\. ft. \\'tlTUM, T.L. 2000. Annl~"S!5:.0I n,;. f.s:c:10,-s-lu: th.ct
lm~mmlon.al Jourlt.tll for Patnsifology. 30. 617-t.~ di,,·tlupm,.,n, of equin\l proto1.,oal my1:Jvcn«:phlllitls in ho&~. Jo:,nud
of 11u,,Amcrft11n V~tt·rfr,ury• Mt·tltt:ul Ass«ior/011. Zl i. I Jt4-1180.
12 \t.M:;U\ N.J. ,997, Equine protozo:i.1 myclocnrophahtb. Vt."Wlfttan
C:linie.-. of/','ott/J Atnerlm: Et1uin<' Pr<u:tia. 13. 79,,,96. 51 StlAMArr, w.K."' Wt\l l,\O., M,I\., sggJJ. l)iug.no&ts and 1Mm,g1.·mc:nt tlf
equine- prourr,1;11 !lly(·1U\'.nccphnlh\s ~nd concurrc•nt tnfccdnu-. d1~\·~'I'-'
TJ ,1,,c.1vu N.1. o.,,,.s. $,\\', 1; ouurv. J,P,, 19»2. f.qumc: prnu:,zcm1
nl}"<tlocn(;t'ph-a.Uds:. C.ompe11aiw>1 011 <:<mttnulng l::dutttuo,, /o:rtht
in,,, n hQrr,es•.Equilll! /tractit,(,', 20. 23-2-'.
/>t(ltn.st'ng Vth'flfttJ.rian. 14, 1359-136';" s:z: ::i.r-u1u1tH, ~.A &. R\MIAr.T., '°' . 1996. \Vhnt Is rour n~urulogcr dtap,oq~?
J':qulOl' pru11,1.o:il m~·C'l0{'nc1.1ph.t1iti~. Journal oftire ..\nw,iran i·,·1rri,mry
-;-1 \t.\OICi ,:,;, u:
& JtUi<Us&. R.J .. 1987. Fqulnc pro:o-Eoa.l mreloencephnlitrs.
,\teditt,I \.-f,'tOC'ratio,r. 209, 903-905
\~:criruuy Clinlb· ofNnnlt Amtrit1t: Equ1n,'Pmc1in-. J. 3!:ti -40;S.
93 M,~.,.~~ c.F." >tA,'Hf\\ 1.G., 19Su. E\1c,hmc.c for Sarc«,~•11.~4 ... :he
f5 \L-\R!'>H, .\.[., &'\RR. B,C.. tAQI.ITI., J.. ~OflUH.\U!i>t'.\, ft,, \UfUC,\.\., f.E. A
etlo!og,c •gent of ,'lulnc pro1olt0'1! mrclocnceph•lltls Jollmfll uf
ce>:,;K.\U, p .., .• 1n97. Exprrimt,ru..;11 infl.!('lic:>n of nudt.· rn10-! u-. :i n,odtl for
PmmVJ()/s>~·. Zi. 288-i9l.
SOrox:ystis neuronn·a.~oci;ued cncepholltls. /'ams1rr.,loXt· Rl"R'n,rrh. 83.
70o-71 l. ~4 W>-l Jt., , &- UUHI, ,.,., :2001. Ulrrasuucrure of ,chi,mw, attd mtro1.olt~

;Ii ", •• .,, u .. '"""· u.c. . .,,o,r,,i,:, ,.. 1•.-.ir,, t ~ c'""'"· ,,.,.. 191i~. a( St1,c.,,;.11is ndtronn. \lqu•rlmil)' Pm,~si~olog,·. 95. :?o3-:!.7i,
'><quence an;dysl~ Md polymcr.isc chaln rcac!lun ompllft,alfon or ~mall yr; ~PlUl,C..\ nunn ,.r."'° ''"''rS<>-,,. tu .. , 1999. C:<:it'np,3r:itin,d~,~1orm\•n1
, '.JU bu nit rib1JWmill J>XA from 'iatcOC)')liS 1wuro,U1,;l11u,rlc11n Joumctl ()f mid nu.'roinlH" pruducdnn Qf two (\Qlnt~of Sarn:H:o·srls nt•1,rmu, and
\lermmrry R<.,.,,rrl1, Si. 975-981 S1,rcoc,~11, f,1!,;,1111/<1 lncultuml cell~. Joi,r1111/ ~fP11ms/11>/oi::, 86. ZS-32.
i7 \IA.MM-I, .\.f .• t\AH.k, o.c.., \t,\OlG\.\, 1.. 1..umn,.. J., ~OtOJH.\U ..,.,. "· k 96 sm1·••~r.1 ,. o.A. i. ouui,·, 1.•.. 1091, A /itlrtor)<III ~P .111.:., pro101.o•n nr.d
COS'AAll, P ..A.. 199(:; ..,•cosporm;i~ il~ ., C"llUW nf t'((Ulnc proto1,o:al concurrtm c-amnc diqcmtter ,1no infection 3$SOdatcd wi1h
m)'tloenccphaUth,, Journal nftlw ,\mertcm: Vettlrionry• ,\ r,'fli<1,t i:tl<Cphalith in a ru.ceoon tPt<>t)tm rotor). Juurnal o/ H'ifd/Jf..• lJIM"tlkS 2':'.
Au«rmfo11. 209, 1907-1913. r&-li92.
78 \IAK-•1-t. \.~ a,ap, H,c... PA(1';Hl\'1, A.tr- -..msAAn, P.A,.19.9.8, 0¢k'rip1lon of 97 r .\,-..H \t1111IR, S. ,~ \'C)\VLU., C-\,, Ulll.f"Jt. 1.J .. (,tU.ISt.R~LC... \li\CI..\\', K.J, i
• n('w Nt0<p<-rospocl<'> Cl'ro10tio3: ,\p1complo,..1. S3tc0<.·y<tld:,c). OAML l,b .. 199~. Muhiplc DX,\ mark~r-<liffo~nlia1eS/lmx;,,r/s n,·urono
Journal of Para.1/111&,g,·, 84. 983-991. a.nd ~rro,;•,ri.1 /ulct1111ln. Jo11rr1nl of Parasitolog;. 85. ?21-2.."11.
 Equin~ prot07.oal myeloencephalltis 403

9t:I TifUUN I D. Glt'\.~Sl'Rf}\t, D.f.. CE.l.SERG. H.B., \tOR'TON O,G 1Af-~C.H. M.A.&> IOOTRA\'Ur, o.~.. f:OFJ-\l,\:',;, I k •• \\OORl'.• ,.~. ~:,, CLSOS-, S.L. 1g:78, Pmtoz.o::ll
Gu.a.,. R.t.. 1992. Concumnl proto1.ooJ enc:cphalitis and ca.nine mi~loencephalms in ;!bllng ho1,;es, To:mwl of Equme .\kdlc111umd
dis-temper \1ri.Jstnf«:tkm rn a raccoon tPrrJ<:J·,m !otorl, The \'t~u,u:nry Surgery.:? . .Jli-42.8.
Rf'lOft/. 130. 162--164.
101 \\T.U;<:u, e,s.• ,99l. Tr("..1mem aiequinetorprotoxool mrctgem:cphallu$...
99 TIUOTSOS, F;,. \ICC\H, t-,•.J .• GAA.NSTROM. O,F-, 0A~T1. D.A,, ,,intt. >t,O. {,. Compi11ulium ()1) Ulmmuing Edurmf(Jt:for tht• Practi1in,H \ (utriumion.
1n.,va·oARGi\t7., u ... 19~. Sero11rL"VO.lcnce or -.amibodies to Sorcm,·s1i." 13. 1599-1602
m:urona m horses residing m nonhetn Colorado, Journal OJ F.qumr
\~•i,r/rmryi,<wr<r, 1,. 12Z-126.
 BABESIOSES

Babesiosis is caused by infecrion with species of tick•bome. borne diseases such as heanwater, gallsickness (anaplas-
intra·erythrocytic and generally host-specific prot0zoan mosis) and some of the theilerioses (e.g. East C<>ast fe\'er.
parasites of the genus lJabesin which belong 10 the phylum Corridor disease. and Zimbabwe ,heileriosis) are or consld·
Apicomplexa. class Sporozoasida. ord.er Eucoccidioridn. erable economic importance. The true importance of
suborder Piroplasmorina and family Babesiidae. It occurs in equine babe,io~b in the suhcontinem in Africa has not been
a wide variety of \'ertebrate hosts. including humans, and adequately assessed. but it remains an important disease. In
has a very wide distribution around the world. In soutliem addition, it is. a hindrance to the free movement of horses, as
Africa the resulting disease is often given one or more collo- many countries prohibit their importation from southern
quial names in the regions where it occurs, such as redwater Africa. Outbrea~ of porcine tmbesic,sis are rare and are not
or roohvater (Afrik.) in cattle. and hiliary fe,·er or galkoors of economic importance. Bt1lwsic1 motasi and Babesia oiiis
(Afrik.) in horses and dogs. The general disease manifesta· infections in sheep and goats have not been recorded in
tions are similar in all vertebrate hos1 specie~ arid, as the southern Africa. Biliary fever in dogs caused by Babe$ia cm1is
popular names imply, tend lo be characterized in va~ing remains ave~· imponnm, ir not the most important. infec-
degrees by imravascular haemolysis. haemoglobinaemia or
1io11s disease dogs and is widespread in sou them Africa.
and the development ofhaemoglobinuria (\vhich i$ respon- On the other hand. the prevalence of Babesia JIiii, infection
sible for the urine being coloured red). and icterus. Bnbesia in cats is low and encountered mainly in tlie coastal parts of
equi is thought also to have ti stage of development in lym- KwaZulu-Natal and the Eastern and Western Cape prov-
phocytes of the equine host. inces of South Africa. Babesiosis is a zoonosis, but cases of
The genus Bnbesia is named after Dr Victor Babes who. in disease in humans caused by lJabesi<1 spp. are rare and usu-
1887, established the aetiology ofa ca11le disease in Roma- ally occur in immuno·compromiscd (e.g. splenectomi:,;ed)
nia associated \\ilh haemoglobinaemia. The disease in patients. However. on 'liantuckel Island and the island off
caltle is particularly prevalem in troprcal and subtropical the coast of Massachusetts and i\:ew York in the USA, babe·
countries \vhere the vector ticks are widespread. siosis caused by the rodent parasite Babesia microri has
In southern Africa. bovine babesiosis and other tick- been reportrd In immunocompctcnt humans.

405
 25
Bovine babesiosis
Synonyms: Red\\·acer. Asiatic (European) redwater (Babesia bovi$ infection).
African redwater (Babesia bigemina infcccionJ. rooiwarer (,\frik.)

A J DE VOS, D T D B WAA t. A N D LA JACKSON

Introduction masked by a devastating East Coast fe\'er epidemic (see

Chapter 29: East Coast fever}. The campaign 10 control this
BO\ine babesiosis. or redwater as it is commonly !,a1own, is a epidemic by intensive regulatory comr-01 of thevector Rhipi·
tick-borne disease caused by the lnrra-erythrocytic proto- ceµha/us nµµendicu/atus lasted half a century and had a big
zoan parasite Babl!Sia. Four species are known to infect effect on the single-host 8oophil11s i:pp .. ensuring that red-
cattle in southern Afric~. namely Babesia bovis.39· 48 Babesia water remained or secondary importance for many decades.
bigemina.311• ~ Babesia occulumsftS and an as yec wmamed 111 the latter pan of the twen tieth cenrury. relaxation or
species.4 9 Of these species, only B. boi•is and 8. bigemi11a are breakdo\\11 of 1i,k comrol measures and the ad\'enrof acari-
economically importan L. The acute disease is characterized cide resistance in Boophilus spp. caused red\l'ater to in-
by hnemolysis, circulatory disorders in the case of B. boui!,~ crease in pre,·alencc and significanc!-' co the point where it
a nd death may follow in some instances. European breeds became recognized as one of th e most important livestock
of caule are particularly susceptible although the disease diseases in southern Africa. 8· ·18• 86• 112 During 1972 the dis-
can also be economically important in Sanga and Zebu ease caused the Joss of JO ooo caule in the Transkei region or
breeds. A clinically inapparem fonn or the disease is com- the Eastern Cape alone39 and contributed to the death of a
mon in young animals, and recovered animals become la- million cat1le during the war in Zimbabwe.I 11 A sun·e} of
rem carriers for \'ariable periods. Recovery is followed by a (armers in the heam,·mer-endemic regions of South Africa
lasting immunity to the infecting parasite. Cros.s-immunicy in 1993 indicated that losses of0.3 per cent of cattle were a:,.
becween the two organisms is limited. cribed 10 babesiosis;u (sec Chapter I: Vectors: Ticks).
Bt1b1>sia bovis was first reported in southern Africa in Although the di$l'aSes caused by B. /Jovisand B. bigN11i11a
19-l l 11 0 although clinical e,1dence of its presence in South are clinically very similar. it is imponam to separate tl1em
Africa was recordt>d as early as 1905Y9 It \,·as probably in- for a number of reasons. hence the names Asiarlc tEuro-
troduced with the ,\sian blue tick Boop/ii/us microp/us dur- peanJ and African redwarer, respectively, have come into
ing the latter pan of the nineteenth century (see Chapter I: conunon use. While 8. l,ovis is rhe more virulent oi the two
Vectors: Ticks}. parasites. 8. bigemina is probably more important In south·
Babes/a bige111i11a is principally transmitted by 8. mi- em Africa because of its wide distribution. In this review. we
crop/11s and by the common. indigenous African blue tick pay more anemion to B. bovis. main!) because it has been
(Boophilus decolorarus). This parasite would therefore 1,avc studied more intensive!}' throughout the world and is much
been endemic: !n the subcontinent for cenrudes. Indeed, bett!?r Underswod than B. bige111i11<1. Information obtained
redwater is said t.o have been the cause of high cattle mortal- from southern and eastern 1\frica is emphasized but refer-
ity in the present-day KwaZu lu-Natal Pro,ince of South Af· ence 10 relevant litef'i!nire from elsell'here is also made.
rlca prior 10 Europeim se!llement in the 1830s. '3
• Several accoums have been published of the history of
Aetiology and life cycle
redwater and other tick-borne disea,es in southern Africa, as
weli as of the people involved in the early battles against these The genu~ Babe#a belongs 10 the phylum .'lpicomplexa.
diseases. The5e reports are well worth reading. 6~· 8~ 137- 139. U 6 class Sporozoasida. order Eucoccidiorida, suborder Piro-
Redwater had a marked effect on the livestock industrie.~ plasmorina and family Babesiidae.3 88• 89 The Babe$ia spp.
in the subcontinent early In the twentieth cencury. espe- kno11n to infec1 canle ln southern Africa, and their pro\'en
cially in imported catde.117 • 1~9 However. its impact was vectors. are listed in Table 25.l .
406
 fJol'ino babe~iosis 407

Table 25.1 tr::w,ne Babesia spp. and their vecto1s in southern Africa

8A8ES1ASf'? VECTORS REFERENCES

Ba//esia bovis Boop/Illus mic,opt11s 119
Be//esia bigemina Boophl/us decolorarus 1ii, l19. 121
Baoph,lus m,c,oprus 11~
RhipicephJJl11s evensi evens. 22
82/Jesia «cu/tans Hyalomma marginerum c•
vJ
• •
1vfipes '
Unnamed 8a:;es1a sp. Hva!omma truncarum .::9

..
Babesia bo11i~ is dassically kno\,~1 ~s a 'small' Babesia
measuring up to 2 µm in diameter. while B. bigemina is
larger and can extend to the fl.Ill diameter of an el)'lhro-
cyte.1 tq However. the r.vo species both show considerable
morphological variatlon. making it difficult to Identify one
•
from the other.24 Large forms of B. boz•is are quite com-
mon.z4. 119 Figure 25.1 8a/Je.si2 bov,s 111 a blood smeat

Single R bovis organisms are round, o\·al or irregular In

shape while paired fom1s are piri(om1 or club-shaped. The
angle ber.veen the paired organisms is often, but not invari-
ably. obruse (Figure 25.1).71 • 119
Single forms of 8. bigemina arc elongated or amoeboid in
shape and contain fine cytoplasmic.- filament~. Paired forms
are typically pirifom1 with an acute angle between the mcro-
zoites (Figu.re 25.2).n. 119
Boophilz1$ microplus is the only known rick vecior of
8 . bc111is fn soltlhem Africa. 119 fron~m is.sion is transovarial,
-.,.
with engorging ad ult ticks ingesting the parasites and the
lar'\'al ticks of che ne:,,.'t genera tion transmitting the infection •
to rhe host" I I!? (Figure 25.3). Ensuing stages are not in-
•
fected.
Confirmed vectors of B. bigemi11a lncludeB. microp/us,zi
B. decolorauts 119• 121 and 11/rlpitep/ralus euercsi 1'11er1s/22
(Table 25. l}. Earlier reports incl uded R/1ipiceµJwlus app,m·
dic1dat11s as a ,·ecror, bu t this has not been confirmed. z~. 125
Transmission by Boophilusspp. is zranso\'arial, ,,;th engorg-
ing adulc ricks becoming infected in the same WO}' as
...
Figure 25.2
-
&/Je.s,a /Jigemma an a blood smear
8. bovls, but in this Instance the infection is transmitted to
canle by che n}mphal and adult stages of the nex1 generation Despite th£' fact that one parasite inoculated lmravenously
(Figure 25.3). There is some evidence of venical l!llnsmis· is capable of inducing lnfection.bO there is no e,idence for
sion in the tick. with infection pai.sing from one tick genera- iatrogenic or mechanical transm ission by biting flies. Intra·
tion to the next in the abse11cc or reinfec1ion.s;- °" In addition. uwrine infection has been reported. but is rarC/.
wandering male Boophilus are known 10 migrate 10 unin- IL appears 1ha1 isolates of Bt,besia spp. are m ixl:ures oi sub-
fested canle11l<) and ma)' pla~· a pa.rt in the transmission or populations differing genotypically and in biologic.-al charac-
B. bigemina.36 Transmission by H. e. ei:ertiii is transovarial. teristics.32 Selection of different subpopulaciom; or s1rains
and only the nymphal stage infects the bovine hosi.22 may occur during artificial passage in cattle or dun.ng growth
Hyalomma marginawm mfipe$ is 1he only known \'ector i11 t•irro causing some of these characterbtics 10 change. in·
of B. occ11lta11s: transmission is 1ranso\'atial with infection of eluding virulenc-e, transmissibilit)· by ticks and immunoge-
caule by nymphal and adult stages of the next generation. 65 nicil). These change:; induced by passaging or maintenance
Exrrac1s of infected ticks have been used to infect C"aule have panicular significance in the production and effect of
experimemally. 124 Both 8. boz•is and 8. big11111itw are easily live blood vaccines. 10· 37 Cloning c~perimems have sho1,~
transmiued by subinoculation of infected blood. a fact these subpopulations to be separable and to differ in 1iru-
which forms the basis of vaccination using lh·e parasites. lence, protein profiles and antigenic composition.60• 141
 C

C'\- ~ d ?a,recmero101tes

, , ,.v-::--~__v___ "7 ---

. ~
e Syngam1·

1
Mi OJ (luman o' gull

. G·
-•~ 0
Transo,1!tll!I :,a~S1111ssion

Fi gure 25.3 Lile cyc1e of oov,ne Ba!les1e spp.

a = s110·ozo,tes released !rom !lck saliva,v g!ands !merogonyl :o p,oduce merozoites
b ~ sw:ozc,tes invade erythrocytes g "' meicw t2s (spo,o, ,~etesl undergo secondary s:h,zcgor.y
c = soorozoites in ervthrocytes become tt0l)ilo,011es lmerogor·1! anc alro finu tlle,r way ,010 aeve.oomg o-.a lhl
d = limited mercgonv lsthizogony) whe1eb-, a pa,r of me,ozoites 1s i = ov,pes;r on cons: totes ,nlected eg9s11rans1)varia :taosmiss,onJ
ll!Oducell J • repllcat.on on ,n:ec.ed la",~ by SQ~izogonv ,n the g.,: oi :ne ,nfec:ed
e = s-109amy 'olklws in the lick gut afte1inges,100 or infet:ed aNa
e1vthroc,1es bv :ne eogo1gmg female ~ = sporc,ir.e:es e:en:ua Iv invade the sahvasy g!anos w~ere :~e;
I = ockioetes pe, evate gut epithehal cells anc ~n:lergo scl!,mgo~v undergo spcrogonv to pioduce 1porozo11es

Repeated serial passages of infected blood exen selec1lon different developmental Mages in the life cycle. al1ema1ives
pressures on the parasi1e populations. which may resuh in arc given belo,,. in parentheses.
reduced infectivity of B. /.Jovis for ticks ro1. IJJ and reduced l1a/.Jesia /.Jo1•is and B. l>igemina rollow ,;imilar de\·elop-
virulence of both species for tattle.33• •u mental pa Hems in adult Boophilus spi>.57 19 1~ Initial de-
The de\"elopmem of Ba/Jesia spp. in 1icks has been re- velopment 1akcs place in basophilic epithelial cells o( the
\iewed by Friedhoff.57 Despite many detailed studies. our gut wall 1 where, after reported fusion (syngamy\ of gametes
understanding of the life cycles or Bab~ia spp. is stilJ incorn- in rhe intestinal lumen, schizogony (multiple fission) occurs
plete.2' Some of the more recem investigations indicate that with the formation of distinctive. large merozoites cverrnic-
scx-ual reproduction does lake place. 10·1 To prewm confu- ules. sporokinetes)11 9 (Figures 26.4 and 25.5). Successil·e
sion about the terminology used in Lhe lilerature to describe L)'cles or schizogony then occur within a variet) of cefl types
 Bnvi,w babesiusl, 409

and 1issues, including Lhe ooq•1es. Tirns. transovarial 1ram,· are readily detected. bm dividing forms are less common.
mission occurs with fttriher development taking place in the Cells containing more than two parasites are mn.•. Parasi·
larval stage. In the case of B. Do1 1is, a final cycle of schi?Ogon~ 1aemias of up to I and 5 per cent respectl\'ely are common
(sporogonyJ 1akcs. place wichin cells in the salivary glands of for B. boL1ls and B. bigeminn. but can exceed 20 per cent at
feeding larvae leading to 1hc rormation of small merozoltes the peak of the reaction. Recovered catlle retain a latent in-
(sporoioites1 1zo (Figure 25.6 and 25.7). These small mero- fection varying from ~b, months to several years with de -
zoites are lnfec1in! for C3ttle (figure 25.3). In B. bigemilla. teccable recrudescences of parasitaemia occurring at
some den•lopment takes place in 1he feeding larvae. but irregular intervals. 92 The mechanism::, used bj 1he,-e para-
schizogonir In lhe ~alivary glands only occurs in the sites co e,·ade immun.: mechanisms of the ho~, have been
n)mphal and adulc srages 121 (Figure 25.3). reviewed.2
tn cau le, Baltesin spp. develop onl~ in the erythrocytes.
Each metoioite penetrates the cell membrane \\ith the aid
Epidentiology
of a specialized apical complex 122• 123 (Figu.re 25.8). Once
inside. it transforms into a tropho7.oite from \\'hich two Numerous factors influence the prevalence of redwater. In
new merozoites develop by a process of merogony57 1~2 - l?.l this discussion. the factor~ affecting the vector. host and
(Figure 25.9). In stained blood films of acute!).' affected parasite. as well as the concepts of endemic (enzootic) sta·
canle. single tropho:i:oices and typical paired mero1.oi1es bility and epidemic tepi;:ootic.) ~pread. will be considered .

•
..
, •
••

•
• ..
Figure 25.4 Large merozoites ,n a smear made 1mm the hael!'.alymph Figure 25.6 Small rr.eroro,tes ,n a smear marle ftom the
of a :,ck salivary gland of a :,cil

Figure 25.5 LOng.ruc,na section through a mature 1a,ge :ne.czo,te ol Figure 25.7 L~·,g,ii.a,eal se,110" tnrough a single rnerozo ,e o:
Babesis b()v1s in me !it< gul "~3 000 Babes,e !lo:es ,n a se1'·zon: ,n tle t,ck salivary glaM. • 48 ~00
 lnter-~pecific mmings may occur between 8. microplus
and 8. detolomws. resulting in I.he production or stNli<:
eggsuo and dic creation or a 'hybrid zone· where popu-
lation, of the 1wo species overlap_t35 This ,.one may have
prevemed B. microp/1is from fully exploiting all the climato-
logically favourable areas in the subcontinent. Its width and
stability are unkno\,i), but it is li kely lO have a significam ef-
fect on the epidemiology of B. bouis. Bttbesia big,!ll1i11a·s lack
ofrector specificity makes ii unlikely 1hat this species \\111 be
affected by competition be1ween tick species.
One infected larval tick is sufficient co transmit B. bo,•is.
but tick infection rates are usually low and the rate of trans-
mission to cattle is thercrorc low. Studies in ,\u~tralia found
on!) 0.04 per cenc of field ticks to be infected \\id1 B. bo1,is
Figure 2.5.8 Sec11on through typ1ca paired Be!JEs,s ba\r.s meroro,tes in where European canle breeds were i nvoh-ed. and C\·cn less in
an el)1hrocyte. x 50 000. Note ul1rastr1x:tural features o: ap:cal complex
the case of Zebu cau!c.94 9; Tick infection rate;, \\1th B. bfg-
emi11a are higher (0.23 per cen t in the Aus1rallan study) and
1hererore transmission rares of this species are al~o h 1gher
1.han tho~e or B. bo11is. As a result, 8. bigemina infection~ are
usually more prevafont In chose herds where b<>1h ~pecies 11re
presen1:•9 and le,s readily affected by factor~ such a,: climate
or management, which reduce tick number...
Babesia bo11is and B. bigem ina have high degrees of host
spccifirn.y curopean.Sanga and Zebu breeds are all suscep-
tlblt>, and all develop latent infection~ after reco\·ery which
pel'$ist for \"llrious lengths of time. European breeds of caule
can retain fl. i.lOl'is infections for life 111 and remain infective
for ticks for up 10 two years.92 while mos, cattle \\ith a ~ig-
11ifica111 Z;:,bu comem lose the lnfectJon wlthin rwo ycars.- 3
Ba!Jesia big<'l11i11a infections rnrely persist for more than a
year, regardles; of lhe hos,. and infened cattle remain
infective for tlck~ for 011ly four to seven weeks.'· 92 \frican
buffalo tSyncerus cajJer) and Asiatic buffalo (Buba/us bul>a-
Figure 25.9 Section tnrougn an intra·elV!lllccv:,c o,v c,rg 'o•m of
Ba//.;s·a t<:gem•na show,ng lhe developmeni of ,\'.1) 11.!ugh,er eel's lfs) can develop latem infections. 12 \ntibodie$ and transi-
fme,ozo tesl from :he moiller cell {trophozo,,el. x 37 500 Note d,viding tory infections have also been demonstrated ln other
nucleus aod l;\1Cpiasm1c d1fferent1a11on or 1r.e oe,e c:img merow1te animals, but there is liule evidence co suggest that non-
famlllal hobls .i,re imponam reservoir hosts 12 7l! even
The distribution of 8. bot1is and 8. bigemina is deter- lhough R. 1•. e11errsi infected wltl1 B. bigemillfl have been col-
mined by the distribution of the vector,. Bpbesia bo11is is lected off a sable antelope (Hippo1mg11s 111ger) with clinical
restricted to areas where 8. miuop/11, is prc-\'alcnr. usually bahesio~i~."~
the higher ralnfall areas in the eastern part, oi the subcon- Ro1h Ba/Jes/a ~pp. probably evoh·ed in cattle In lhe
tinent. including the Eastern Cape. KwaZulu-:'\atal and tropics. and well-de\'eloped relationships therefore exist
;\1pumalanga provinces or South •\frica. ''' SwwJland. bet,,ecn Zebu and Sanga caule and these parasite~. As a re-
,\'101,ambique.2" parts of easwrn Zimbabwe. 13 1.he nonh- sult, Babesia spp. are less important in tl1ese breeds while
easiem part~ of Zambia.; 5 and Mala\\i. 8ab<'sia bigemi11a is European breod.s are known to be highly susceptible. 12· 32
much more widespread In the ~ubcominent than 8. bovis Howe,t>r literalure on the resistance of tliese breed~ and
because of Its wider vector range (sec Chapter 1: Vectors: crossbred c.auJe i~ rather ambiguous. 11. 12 91 In a large con-
Ticks). fl is prcseru throughout most of the subcontinent. rrolled field trial in Aust ralia. 32 per cem of56 pure 80, indi-
~xrept ror the arid parts or the Western a!ld :,.;onhcm Cape r:us showed dewciable clinical signs after natural expo,ure
and the western Free State of South \frlca. :\amibia and to B. bol'is and 8. bigemina while 2 per cent (one animal) was
Botswana. as well as some high-lying parts or the Drakens- severely affected .. i In comparison. 87 per cent of 52 cross-
berg and Leso1ho.l.!1• ,s, 1~· 113 !n S0u1h ·\frica. ils dis1ribu- breds (hnlfbred Bo~ i,u/ir:us) were c:linically affected. 19 per
lion appeaNC 10 comcide wilh that ot B. decolom111s and c.:ent of lhem senirely. Typing of the para~it~ In $e\·erel)
8. microplus.·' 9 The role of the more drought-resistant affected canlcsuggested thnt 8. 1,011iswas the main cause of the
vect()r, /?. e. ,,,errsi.7" in the epidemiology i~ unclear. clinical reac1lons. In a parallel pen trial. Ba, i11dirus and
 Bovln~ babl'~losis .J 11

crossbred canle were more resistant to B. bigemim1 than Bos sex class rhat would be affected br dilTerent severities of the
11111rus caltle (none of sb: Bos i11dic11s and sh: crossbred cattle disease, and the inforniation is then convened lmo a herd
wasseverelyalfecred compared with six of the seven Bos wums mod!'[ to estimate the number of animals in each severity
caulc). 11 Howe\ er. su!lceptibility ofindivldual Bo$ i11dic11., and class. This model has been used 10 predict the potential im-
crossbred animals varied considerably. The mason for this is pact of Bal,esi(I spp. on some large pro pen ie.~ in Auslralia
unknown burlt may help to explain t11e amblguit} in the litera- and 1he cosl /benefit or control measures. t·1
ture \\ith regard to breed susceptibility to B. higemi1u1. Rabl!Sia bo1,i, is presumed to be absem or endemically
1'he virulence of 8abesi11 is not a stable characteristic. unstable in moscof southern ,\frica.~r.'rhis ismainlydue to a
Mild and ,irulem isola1es of both species have bet>n idemi- large part of the subcominem being ecologically unsuitable
fied from the field and are known 10 be capable oi rapid for 8. microplt,s. the large zone of hybrid imerference be-
change. A major factor in parasite virulence is a history of tween the two 8oophilt1s spp., and pre,·aillng tick control
recent tick transmission: long residence in a bo,ine reduces measures In the high rainfall areas.
virulence while irequem cvclical transmis!.ion in ticks mar Babesia bigemi11t1 h endemically stable In large pans of
increase the severity of the· ensuing reactions.i·• southern Africa but. whereca1tle are 1101 expo,ed al a young
l.solates and selected strains of 8. llovls and B. bigemina age, e.g. in situation~ as~11ciated with increasing aridity-19 or
also differ anrigenically32 but this appears to ha, e little epi· With rick contro1.J1 3 It may become important.
demiological significance. Cross-immunity experiments Se\'ere losses may occur when susceptible ca11le,
have shown that recovered cattle are more re~lstam to esµccinlly Bos rcmm~ breeds. are brought into an area where
challenge \\1th the same (homologous) isolates than 1,~th Boophilus spp. are pre\'alent. ~lor111lity rate~ of 5 to 10 per
dilferent (heterologousJ ones.32• ~2 Antigenic variation is ce11t are quite common under these conditions.39
also known to occur during 8. bovis and B. bigemi11a The distribution of the vectors of 8. l)m,f, and 8. big-
infections with l'ariants emerging at regular intervals during emin" is limited by ecological factors such as rainfall and
the latent phase of the infection as~ocialed with slight also by the extent of tick control measures. llemo\'al of these
recrudescences of para~itaemia. ~2 barriers may result in the ticks becoming established tem-
fl is possible to have both redwater and its vectors poraril~ or permanemly in previously uninfested areas. This
present in a cattle population without measurable eco- happened during a period of unusualJy high rainfall be·
noniic losses or clinical dL~ease. This situation is known as 1wct'n 1974 and 1976 in the normally arid 1'orthern Cape
endemic s1ability and defi ned as the srate where the rela- Province of SQutll Mrica, when B. decolorat 11, mOl'td sever.II
tionship berween host. agent. vector and en,iromnem is hundred kilometres in a westerly direction.l!• Similarly. a
such lhat clinical disease occurs rarely or not at all. 115 An im- breakdown in tick control measures during the pre-inde-
portant factor in the eMablishment of endemic stability is pendence war in Zimbab\\'e between 1973 and J 978 enabled
the age of first exposure. Calves have a nalUral resistance B. microplus to spread t'rom a few foci near the Mozambique
during lhe first sL, IO nine months of life and nuely show border to the centre of 7..imbabwe. 112• 113 Both tick migra-
clinical signs, yet develop ~olid, long-lasting immunity.32 • 42 tions caused large outbreaks or babesiosis in susceptible
The occurrence of redwater in a herd depends entirely on caule populations (see Chapter 1: Vectors: Ticks).
the frequency ,,it.h which the causative species is transmit- It is interesting that introduction of 8, microplus in lhc
ted. If there is frequent transmission, infection of all animals lauer half of the ninetee11th cenrury into Australia re,ulted
will lake place at a young age and immunity will develop in n major epidemic of redwatet2'1 paralleling lhe southern
without evidence of disease. 24 · 39· 9S ;\frican experience with East Coast fever. This raises the
Under conditions of endem ic instabilit}, some animals question whi the ,;an\e did not lrnppen when B. microplus
"ill fail to become infccwd for a considerable period after was introduced into southern Africa. Some possible expla-
birth and \\ill therefore be susceptible to clinical disease. !kt· nations are: 8. bigemina wa~ already present and therefore
hesia bigemina hns higher infection rales in ticks than.B. bo11/s only B. bo1•is\\1IS associaccd with the spread of B. microplus:
and endC'mic instability is lherefore less likely 10 de,·e!op to it competitive interference by B. deco/ora111s suppressed B.
than to 8. bo11is in regions where both are present.~6 microplus in some awas: the East Coast fever campaign in
A simple mathematical model has been used to predict the early pan of the twentieth century pre\'enced the
the level of stabillcy in a herd.9.' It utilizes che rate at which unchecked spread of B. microplus; and B. bigemina gave
infccc!on occurs In calves (determined serological!) ac a spe· partial protection against the introduced B, bo11is.
cific age). thereby providing a means of esrimacing the sra-
tus of B. bouis infections:1~· 9• It has proved useful in
lmmtu1ology
predicting stability levels in European breeds but less so in
those of other breeds. More recent!)". a disease prediction Cal\'es less than 1wc'l months old born to previous!} unex-
spreadsheet model was developed 10 more accurately calcu- posed cows arc susceptible to infection and the effects of the
late incidence risk from age-specific seroprc,alence. 1?6 The discase.24 while offspring of immune mothers are resistant.
model predicts the proporrio11 of animal, in each age and presumably because of a passive 1ransft'r or immuniry l'ia
412 ''-ClK'' mo: Protozonl diseai.es

Lhe colostrum. 1.1 1 r,4 Arrer the age of two months a natural. is mhogenic for bo,ine B lymphocytes and enhances imnm-
non-specific. innate resis!ance. which persists for at leas! a noglobu!in GI and G2 production by 8 lymphocy1es. i;
further four [O sL~ months and is not dependent on tile im- Specific and non-specific cel l-mediated mechanism~
nnme status of the cow. procects cal\'es/1· 14•1 involving T lymphocytes and natural killer (i'\1'1 cells ap-
Severe auacks of redwater ln cows at calving have been pear robe involved in laboratory Babesia infections: intra-
obsen·ed. and may be related to Lhe perinatal immunosup· cellular death or parasites occurs which is nor caused by
pression which occur,; in other ho,1-paraslte syscem~.~4 amibodie$ and which is non-specific with other para~lres.
i\ged animals may also suffer seyerely. presumably as a re- micro-organisms and even microbial extracts exerting 1he
sult of a general decline in immune competence. same effect on rapidly dividing Bnbesia orga11bms. A
:-.itost cattle develop a durable immunity after recovery. possible explanation of this phenomenon is the non-
Thjs immunity is nOl absolute but may last for life. even in specific ac1ion of ;,./K cells activated b}' either a specific
the absence of reinfection. Follo";ng recovery. the infection immune re~pom,e involving lymphocyte~. or by non-
pershts for ,·arying periods in a latem form without detri· specific agems. '·
mem to the host. The presenC'e of this lau~m infection. \lacrophages are irnponant for immunity to 8. bot1isbe-
known as prernunit)', was long considered tO be a pren•qul· cause they act as antigen-presenting cells (APCJ forT helper
she for protective immunity. tti It i~ now kno\m I.hat persis- [fh) lymphocytes and remove parasi1i1.ed erythroc;y1cs by
tence of infection is not necessary to ensure immunity. For phagocytosis. :-.1acrophages acti\'lHed by interferon-gamma
instance. caule which naturally eliminate die infection re- are though1 10 play a significant role in resi.tance 10 B. /1011/s
tain strong and lasting immunlty. 10· ;J, 110 while cattle cured by parasite suppression via secretory products rele.ised
of che infection by drugs show levels of resiscance more from macrophages. H? This has been demonstrated in 11i1ro
closely related 10 the degree and duration of antigenic where supernatant from macrophages s1imula1ed wid, 8.
stimulation than to the presence of parasltes..16 . !lO The con- /101,fs antigens inhibited the growth or 8, bovis. 1"' 11iese in-
cept of premunit:y in Bnbesia infections has dierefore been hibitory produces were nondialysable. resistant to heat and
abandoned. inact!\'ated by freezing and th;iwing; but no auempts t0
The qualiry and duration of the resistance to reinfection identify the inhibitory factors were made.
after recO\·ery is not affected by the age of the host at the The generation ofreactive nitrogen imennediates (R,'11).
time ofinitial infection. nor by the virulence of the strain in- such as nitric; oxide c;s:o) or reac1 ive 01>.,"gen Intermediates
volved.?4. <in. 98 Can le recovered from 8. bigeminn infections (e.g. H2 0,J. is a mechanism whereby acrh-ared macrophages
show some resis.1ance 10 B. bo,,.is challenge. 153 but the re- release microbicidal products. NO has been shown co be
verse is nor true. Furthermore. cattle harbouring latent 111· produced by bol"lne macrophages follo,\ing stimulation
fections of T//eileria bttffe/i. a benign parJslte in Australia. either with H. bovis mero.r.oites or\\ ith amigen-s1imulau:d
have increased resistance ro B. bouis but not ro 8. bigemina 8. bo1,is-irnmune ·t lymphoc)'tes. 111 :-.ro has been shown 10
infections.93 t\ similar effect of African fhi•ileriaspp. has not inhibit the growth of B. bo11is i11 vitro whilst H20 2 does not. 72
been demon~1ra1cd bu1 is worth exploring. Kon-$pccific The effect of NO appe.irs to be directly on the parasite and
s1imula1ion or immunity in cattle wid1 agems such as le- not on the e111hrocytes.
vamisole and BCG has been shown to moderate laborat01,· There is no direct evidence that 8. /Jol'is-spedfic T lym•
Babesia infec1ions;12 but docs nor appear to have a signifi- phoq,cs are pro1ec1ive because adc1ptive transfer studie!>
cant elTeet on field infec1ions. using T l)mphocy1es from immune animals are difficuh 10
The mechanisms of immunity in babesiosls are complex perform in cattle. Indirect evidence, howe\·er, suggests
and involve both amibody and ceH-mcdiatcd componc,ms. that T l}~11phocyu?s nrl' importarll for immunity. In manr
:\ml bodies devel,)p afier infection, but not all are pro1ec1ive protozoal infections. Tlymphoe)•tes are the prima11· effector
aod anempts to correlate antibody levels with immunit~' cells in protecth·e immunity. 54 • 112 Peripheral blood mono·
ha\'e falled. 2''· 93 However. some indirect e,idence for a pro- nuclear celb from immunized caule exhibit weak and
1ectl\·e role for antibodies is provided by the passive transfer short-lh·ed prolifcrath•e responses 10 a crude soluble
of maternal immunity w B. botli$ and 8. bigemina in calves parasite e~1rac1:i;o however. they may proliferate in
by colostrum intake. 12• L"4 response 10 rnero1.oi1e membrane antigen for up 10 1wo
Live B. bovis inoculated into cattle induces antibodies )'INlfS after il1fection.:ro Brown and co-workers have been
that remain at high levels for at least six momhs. 111 These stud}-ing T lymphoc)1es from caule immune 10 B. bo1iisin an
,Jntibodies are strain specific:96 and are or the lg.\l and lgG 1 effort 10 identify antigens of 8. bo11is which may Induce
isor:ype. 61 JgG2 antibodies 10 B. boi•i~ have not been dc- protccti\'e immunity.20 They have shown that (04·
1ec1ed.61 Specific lgCl antibodies reach high titres and are r lymphocytes (Th lymphoq>tes), bu1 not cos- T lyrnpho-
reacti,•e in a complemcm fixation test. an Indirect nuores- cy1es (T suppressor lymphocytes) or y.S T lymphocytes. arc
cent antibodr 1es1 and an antibody-dependenr cell-med!· µreferentially s1hnula1ed by crnde parasi1e antigens in
aced cytotoxichy assay.U2 In addition to 8. IJOvis antigens ir/tro. 18 Funher work analysing C}'tokine profiles of
being immunogenic forbO\'ine B lymphocytes. 8. botiis Di:\:,\ Th l)mphoC)ie clones derived from immune animals has
 llovinc bahc<io~is 4 l3

shown that interferon--,. interleukin-2. intcrleukin-4 or endothelial damage thus :iugmenting 1he effects of circula-
LUmour necrosis factor may be produced in response to iory stasis and h}'POtension. 311• 152
/11 vitro B. bovi.s stimulation.n l'rogressive haemolytic anaemia develops during the course
The cote or the spleen in fmmunicy to babcsiosis is well or B. bo11isinfec1ions. \\'hile this is 1101 a major factor during
known. 1' Splenecromy of latent carriers u.,,ually results in thl' acute phase of the disease. It will contribute to the dis-
fatal 8. bigemina relapses and in B. bo11is relapses that are e:ise process in more protracted cases.
some1imcs fatal. Infection of susceptible spleneccomi:i:ed
cattle results in fulminating infections ,,1th both parasite Babesia bigemina infection
species. Splenic lesions have been shown to reduce the abil· J>athogenesis is almost entirely related to rapid, somelimes
i1y of im:ict cah-es to contain anenua1ed B. /Jigemi1w infec- massive, intrarnscular haemolysis.2·1 Coagulation disorders
tions..~1 The spleen is not essential for 1he de,·etopmem of and 1he hypotensive state seen in acute 8. bovis infections
protection. but immunity is established more rapidly in in- arc not features of 8. bigemina infection.s. 3 ' 1'Ill
iact 1han in splenecmmized animals. sug_ge,sting chat local-
ization of immune cells fn the spleen and the splenic
Clinical signs
microcirculation facilitates the process.
The din ical signs which develop as a result of 8. boPis and
B. bigeml11a infections are similar. but 1he course and out·
Pathogenesis
come are often wry difl'erem.2'1 11,e incubation periods of
Babesia bovis infection nawral lnfoc1ions usu:illy ,-acy from 8 10 15 days.~;
The acute disease is characterized by a hypotcnsivc shock
syndrome \\1th vascular stasis and accumulation of para- Babe!;ia bovis infection
sitized red blood cells in che peripheral circul:i1ion. II is In acute infections. fover {>40'C) is usual!)• present for
accompanied by ac1ivati<>n of a coagulation/complemenc sever:il days before othcr sigJ\S become ob\fous. This is
cascade and the release ofvasoaclive compounds resulting followed b> i.nappetcnce. depression. weakness and a reluc•
in vasodilarion and circulatory stasis as well as generalized mnce to move. Haemoglobinwia is often pre,em, hence the
organ damage due 10 anoxia and toxic products of bo1h name redwater. Anaemia and ictcrus develop and are
parasites and damaged host tissue. 1.:;i espccially ob,ious in more protracted cases. Diarrhoea is com-
Parashe pro1eases cause hydrolysis of fibrinogen which mon and cows mu~ abort. Muscle wasting. tremors and re-
resul1s in the accumularion of large quantities of soluble cumbency develop ln advanced cases :md are follo\\'ed by
fibrin complexes which are not cross-linked. as well as tn coma in terminal case,. Affected animals may die from one
:iltered fibrinogen in the circulation. Thus. 1he co:igulabilit~ to se,·cral days after the onset of clinical signs. :-Son-faial
and ,1scosity ofblood increases bu1 inSoluble fibrin is C\~dently ca~es ma)' rake several weeks to regain condition bm
not produced, suggesting that classic.disseminated imravascu- recovery is usuall> complete.:!-I. ~, Surviving bulls mayshowre·
lar coagulopachy is 1101 a feature of 8. boPis infections. 150 duced fertility lasting six co eightw{?eks as a result ofthe fC\·er. 55
TI1e cause of cytoadherence129 in B. bo11is Infections may Cerebral babesiosis develops in some B. bor,is infections
be related to the formation of crythrophylic fibrinogen and and is manifested by a va riery of signs of cenrcal nervous sys-
soluble fibrin complexes adhering to infected eryihrocyte 1em involveme111. These include h>'J)i:>raes1hesia. nyscag-
membranes,.,., or due to the adhesive inceraclion between mus. circling. head pressing. aggression. convulsions and
infected red blood cells and receptors on endothelial cells paral>rsis. and may or may not accompany other signs of
caused by vascular cell adhesion molecules induced b}' in- ncute redwater. The course of the disease ls usually short
1lam111a1ory C)'IOkines.11 It has further beer\ suggested chat and the outcome is alrnosr invariably fa1al.
B. bo11is could express variable surface a mi gens functionally In suba.cute B. bm•i$ infections, clinical i;igns are less
similar 10 those expressed by Plasmodi11m falcipnrw11.3,1 pronounced and some1imes cilfficult 10 de1ect. Cal\'cs in-
Pamsite proceases also activate macrophages to release fce1ed before rhey reach cigt,1 momhs of age ofum develop
pham1acologkaUy active agents. such as histamine and an inapparem Corm ofthe disease.20
5-hydro:,.·ytryptamine. causing vasodila1ion. hypotension.
increased caplllary permeabillry, oedema and vascular col- Babesia bigemina infection
lapse. u9 This hypotensive shock state has much in common /\cutely affected cattle are usually not as severe!) affected as
\\1th endo1oxic shock, suggesting t.hat B. bm•i.< and endo- those with B. (Jor•is infcc1ions, but the disease can develop
toxin cause macrophages to activate 1hc same harmful very rapid!}' with sudden and severe anaemia. ic,ems and
mediators.2 9 · 30• MS In fact. the susceptibllit}• of laboratory death, which may occur "1th liule warning.~~ Haemoglobi·
animals to endotoxin increases dramatically afrer they have nuria is prcse111 earlier and morn consistently than in
been infected with BnbPsia spp.30 Laboratory evidence also B. bo11is Infections. Signs of cerebral lnvolvement are not
sugges1s t.hat Babesin spp. cause leukocytes to release free seen and recover) in non-fatal cases is usuallr rapid and
O>-'Ygen radicals.30 These radi~ls are toxic and contribute to complete.
414 '"~'~' l\,o: Prpto,.oal disca=

Pathology dark red discoloration of the urine. The spleen is commonly

enlarged. someumes to several rimes its normal size. The
Clinical path ology liver is swollen and may be yellowish-brown. with the gall
Babesia bovis infection Haemol~'tic anaemia. which is bladder containing copious amounts of thick granular bile,
charac[eristicall}' macrocytic and hypochromic. is a fearure The kidners and lymph nodes are also enlarged. Pulmonary
of 8. bot•is infection~. Packed cell ,•olumes (PCVJ may fall to oedema may be prest!llt but is uncommon. In the cerebral
less than 0.1 O. total ery1hroq1ecou111s 10 less than 3.0 x 1ov1 form of the di•ease. the grey matter of the cerebrum and
ml. and total haemoglobin to less thnn 50 g/l.2' 1• 147 Severe cerebellum has a characteristic cherry-pink colour
anaemia is particularly evident in protracted case~. while (Figure 25.J 0).24.l,.12, 1ss
very acute cases mar die with little evidence of anaemia. ~licroscopically. sequestration of parasitized red blood
In cattle which survive the infection. parasiiaemia levels cells In the peripheral circulation (Figure 25. LI ) and evi,
>tan 10 fall three 10 five days after the onset of patency (the dence of vascular stasis are striking in acute 8. bo11is infec-
first microscopic detection of parasites), and evidence of a tions. Large numbers of paras-llized red blood cells fill the
response mdicative or erythrocyric 'regeneration can be de- capillaries. sinuses and sinusoids of\'arious organs. and can
tected cwo to four days later. These signs include anisocyio- even pave the tmimn oflarger blood vessels. AccQmulations
sis. polychromasia, punctate basoph1lia. macrocytosis and or haemosiderin 1111d 1>hagocyto1.ed red blood cells. some-
reticulocytosis.24 Recovery is slow and it may cake several times containing parasites, are common in cells or the
weeks before nonnal red blood cell counts are restored. Leu- re1iculoendothelial system. especially in the spleen, liver
koc}'lic changes are variable. ranging from leukopenia to and lymph nodes."'; 127• i.:; Other lesions include: 24 127• 147
leukOC)'IOSis. l$7 • degeneration and necrosis of the epithelium of the con·
Changes in blood chemistry reflect the consequences of volu1ed tubule~ in the kidneys and an accumulation of
circulatory stasis and hypotension. including renal and liver hyaline or granular casts in the rubular lumens:
damage, and muscle degenerntion.' 3C\ Characteris1ically. • cemrilobular hydropic or fauy degeneration 10 extensive
the following occur:u 7• i:.o. i;i. i,;5• 1~: cemrilobular and mid1.onnl hepatic necro~is and bile
• significant increase.~ in blood urea nitrogen and plasma s1asis:
creacinine levels; • marked congestion of the sinusoids of the spleen and a
• marked increase, in unconjugated bilirubin levels. reduced ra1iooiwhite10 red pulp with thegenninal cen-
plasma creatlne kinase and lactate dehydrogenase levels; tres containing iew cells;
• the presence of haemoglobin in the semm 10 a level that • oedematous and congested sinuses in the lymph nodes
may be as high as 5 g/(; and deplerion of lympho1..ytes in the gem1inal centTes:
• increased plasma com·emrat!ons of fibrinogen and • oedema or the lungs in s<1me cases;
soluble fibrin: • marked distension of the capillaries of the brain by para-
• proteinuria during the acute phase of the infection or silized red blood cells (perivascular haemorrhages are
which l 5 10 20 per cent Is haemoglobin and 70 10 75 per uncommon);
cem albumin: • haemorrhages in the myocardium and hyallne degen -
• metabolic and respiratory alkalosis. as shown by elevated eration of some myocytes; and
bicarbonate, excess base levels and a lowered pCO::: and • degeneration of skeletal muscle fibres In the hind limbs.
• a terminal increase in lactate and pyruvace levels.
Ullrastruc1urally. capillaries In the brain are dilated and
Babesia bige111i11t1 infection Haemolytic anaemia is the filled \\ith closely packed red blood cells. most of which are
outsranding fearure and is very similar co that seen i.n B. parasiti.zed These red blood cells have scalloped edges with
1,ovis infet:tions. However, red blood cell destruction occurs fine strands apparent!}' connecting adjacent red blood ceUs,
more rapidly in severe cases ~d is accompanied by preclpi· as well as connec1ing erytl1roc}'tt?S and endothelial cells
cous falls in PCVs. red blood cell coums. and haemoglobin (Figure 25.12). Masses of ly,;ed red blood cells which have
values. o~mocic fragility of the red blood cells increase~ dur- undergone lysis but still contain intact parasites are
ing the acute phase of the infeaion.150 and serum haemo- l'requemly seen in capillaries. Changes in the capillar:•endo·
globin levels are high in acute cases. Evidence of kidney and thelium of the affected pans in the brain range from swelling
liver damage is similar 10 that seen in 8. bo11:s infections. 1H of the cytoplasm and nucleus to necrosis. Perivascular and
perineuronal spaces are enlarged. 1'18
Macroscopic and microscopic pathology In the kidneys. capillaries are not packed as tight!}' will,
B"besin bo11is infection In acme infections. congestion of red blood cells as in the brain and the para$itaemia doe~ not
most organs and tissues is intense with subserosal pecechiae exceed 50 per cent of red blood cells. Other changes are
and ecch}~nose-s occuning in many internal organs. Con- similar 10 those seen in die brain. Capillaries h1 the lungs are
versely, pro1racred cases show e\~dence of anaemia and ic1- packed with red blood cells. but only a small proportion or
erus. Flaemc:iglobinuria is often seen at necrop$y as a light to the cells are parasitized. 1$"
 Bovine bab~,iosi- -l 15

Figure 25.10 Cerebral babesiosis caused by Babes,a /;Ov1s Note mar~ed

congestion of th{l 9rev ma tier of the brain

Figure 25.12 Cm$S·Saci'on :h·~ugh a brain caprllary ~acked \\1tn

para51r ,zed ery.~roc,1es Note il>e pro,ettrons from nfeaec cell sur'al:es
and me rarge nters1 11al spares wh,th are orom,nem in :he •ei;•co,
, "
- however, anaemia dose!~ parallels 1he parasi1ae11,ia and
lhere is no evidence of auto-immune anaemia ha,ing a sig·

,. .. ~..•... •
• nl!icant pathogenic role, even though phagocyio,is of non -
parasit ized e~'throcy1cs is known co occur:'4
Immune complexes ha\·c been implicated as a cause of
1hc anaemia, splcnomegalr and glomeniloncphritis in Ba·
hes/a models in the laboratory. bur their contribu1ion to the
~
... .- . ' , ..-:,
......, , .., . '
J • ........
disease in ca11le has not been conclusively established.32 i •
Uabesin btwi, amigen has been detected in glomeruli and
Figure 25.11 Er,1hrocytes tiaraS1ti£ed by 8abes1a bovis ma bre,n sauash the endothelium of blood \'CSsels. and is known to be i11ti-
smear mmely associated wi1h ho,1 fibrlnogcn.61
Immune re,,ponsc~ to he1erologous antigens are sup-
Bn/Jesia /Jige111i11a infection l..csions seen a1 necropsy rc- pressed <luring Babesitt Infections. The most comincing e,,j.
~emblc Lhose In animals which have died after an anaemic dcnce of this immunosuppression comes from laboralo~·
crisi~. There is evidence of severe haemolysis. such as a pale Babesi(I models. h b manifested byan impaired ability of the
carcass. \\'a1ery blood and haemoglobinu.ria. Haemorrhages host to countt>r superimposed parasitic and ocher infec-
in Internal organs and splenomt.'galy are not as marked as in rions. as well as br decreased amibody respon~e" 10 inocu-
8. 1)01,i.~ infec1ions, bu1 pulmonary oedema is a common lated amige11s.J2 , 4~ fJtilX'Sill bo11is causes a prolonged
finding. lc1erus is evideni in protracmd cases. 24 • 2 r increase in the suscc:piibllily of calll<' 10 8. 111icrnp/11s infes-
I listological changes arc less pronounced than in B. /Jo1•is tation if rhe host is exposed to both parasites at 1hc same
infections, and sequesrratlon of infected red blood cells and 1ime.2 & Cortisol values also become mnrkcdl} suppressed
vascular srasis are not features of 1hc infecrion. Changes in during B. boi1is infoc1ions and may be followed by 1ransjen1
the kidneys and lh<er are similar to those caused b~ 8. i1011is. adrenoconical dysfunction.5"1
Ex1ensive necrosis of the red pulp of lhe spleen is common
and large thrombi may be pH'SCm.27 • 1• 7
Diagnosis
[mmunopathology History. clinical signs and macroscopical lesions may often
Evidence of auto-antibodies as mediators of anaemia in ba- be sugges1i\'c of redwater. yet are n01 absolutely characteris-
besiosis comes mainly from experimental snidies wi1h labo- lk of the disi:a~e Dlfforcmiation between 8. liot•is and
ratory infocrions of Babe.litt spp. In bO\inc babesiosis. B. bigemi11t1 infec1lons is abo lmpo~sible on these grounds
416 s,nK,, rwo: Proio,.oul dis~ases

but it is highly ad1·isable that i1 Is done becau,e of differ- of death. thus making ldentifkation imposllible on che basis
ences in pathogenesis, prognosiS and conrrol. The 1radi· of morpholo~'Y alone. However. va~1 accumulations of para-
tional method oiidentifying 1he agent in infected animals is sites in organ smears. or big differences in the parasltaemias
by microscopical examination or thick and rhin blood films of the general and peripheral circ11lallons. are indicative of
stalned with. for example. Giemsa's stain.r Details of these B. bo1,is infec1iom;. In B. bigemint, infections. parasites arc
techniques are recorded in the literature·; z, and the sensi· readily detected in the general clrculation. Accurate diag-
thil)' is such that par.;sitcs can be detected in mosr clinically noses are usually possible as long as intact erythrocrte, are
affected animals. Species clifferemiation is good in thin films recognizabll' in tht- preparativns. In cool weather this may
but poor in the more sensitive, thick films. This technique is be as long as three days after deatli. 2• · ·15• SIi
u~uaUy adequarc for detection of acure Infections but is 1101 For bistoparhological examination, specimens oi brain,
suitable for detecrion of carriers where the parasicaemias ~pleen. lh'er. lung and ~pleen should be submined. Babl!sia
•
are usuall} well below level~ detectable by this method. hot·is will be identifiable by virtlle uf its accumul3lion in the
Parasite identification and di!Tcrcmi;uion can be improved capillaries of organ:.. but n diagnosis of B. bige111i11a may be
by using a Ouoresccnt dye, such as acridine orange. instead diffic:ulc to confirm unless the parasitaemia i~ very SC\'ere.
of Giemsa's stain.•R A Quantimtive Buffy Coa1 method using Labelling of parasites with Uuorescein or hor:.c-radish
ac,idine orange 10 stain parasite, in capillary tubes was de- peroxidase conjugated ami-8. l>ovis and ami-8. bigeml11a
\'eloped co demonstrate Plam1odi11m in human blood and lgG is a sensiti\'e. specific laboratory tool to identify para-
has the potential 10 detect low Bobesia parasilaemias as sites in blood nnd organ preparations. provided adequate
well. bur differentiation is likely to be poor. 1" numbers of parasites are present.7'1· 83 Probes ha\'e also
For the best results, blood films should be prepared from been used to detect 01':A of bovine and other 8(ll!esin spp.
capillary blood collected. for instance, after pricking the tip bur are general!)' not more sensirive than direct micros-
of the tailor margin ofan ear. The temptation to use blood of copy and applicauon in routine diagnostic.,, is limited.tl. 58
the general circulation should he resisted as these speci- PCR assays have proven 10 be very sensitive particularly in
mens mny contain up to 20 times fewer B. /Jovis thnn capil- dcrecting B. bovis and B. bigt•mi11a in carrier cattle.zl. .l.'I Su
lary blood.i: In B. bigemina infections. parasidzed cells are Howe\'er. PC Rassays generally do not lend themselves well
e\'enly distributed 1hrough0ut the blood circulation. Mixed 10 large-scale tc>1ing and are unlikely to supplan1 sero-
infections 01 the rwo Babesi<1 spp. are uncommon. Babesia logical rests as the method of choice for epidemiological
bo1ds parasitaemiasare often low (<0.1 percent), even at the studies. PCR assays may be useful a~ confirmatory tests. in
peak of the reaction. while 8. /Jige111i11a parnsi1es are usually ~omc cases for regulatory testing. and also a.; markers for
more numerous and therefore easy to detecL The most vaccine strains D
common method of staining 1hese films is to apply Giemsa's r,, 11itro culture methods have been used to demonstrate
stain at 10 per cent in phosphate buffered saline fpH 7.4) for the presence of carrier infections of liabesia spp.bl' t 5A The
20 minutes after fixation in absolute methanol.r minimum parasitaemia dctectnble b)' this method \,ill. ro a
Thick blood films are cspcciallr useful for the detection large extent, depend on the facilities a\'ailable and the skills
oflo\,· le\'el B. boi,is infections:' These film5 differ from thin of the operator1" bm could be as low as 10- 10 • making it a
ones in tha1 1he blood is not spread O\'er a large area and is very sensitive method for che clemonstrarion of infection. 511
not fixed before staining (lh:ini: in acetone is optional), thus An added benefit is tha1 it Is 100 per cell! specific.
allowing lysis of rhe red blood c~lls and concentration of the A variet')' of ~erological tCSlb has heen used to demon-
parasites. These films are usunll)' stained \\1th a 3 to 10 per strntc lhe presence of antibodies to Babesia spp. Oi these,
cen1 Giemsa·s solution for 20 minutes. A high degree of the indirect fluorescent antibody (IFAl te~t is widely used to
cleanliness is necessary when preparing thick films.2 ' de1ec1 antibodies to Babesia 25 · 00• 11 ' The lf.\testis specific
Several factors may make it either difficult or impossible for B. bovis. bu1 cross-reactions witl1 antibodies to B. bovis in
to identify 8(,besia spp. in hlood lilms: prior tTeaunent of the the B. l;igemina Il'A arc a particular problem in areas where
animal with a habesiacidc will cause the parasites to degener- the rwo parasites coexist.' Unfonunatel)'. in the standard
a1e and disappear, often within 2.J hours; exposure of the IFA tesr the degree of serological cross-reaction that occurs
films to moisture or formalin \\~II make idemilicadon of the between the four Babesia spp. present in southern Africa is
parasites difficull; and din:y smears may mask the presenc<' of such rhat accurate differentiation is sometimes difficult.' 7
. .,.
parasnes."' The de1ailed methodology of chc IFA test for babesiosis is
• Diagnosis at nccrops}' is possible if decomposhion is nor available in Lhe literature.4 · 2 •
100 advanced. 111e specimens should include thin films of To overcume the disadvanrn,g~ of the If-A rest, such as
blood from the peripheral and genernl circulntiun. a squash low sample throughpur and subjccriviry, the de\·elopmcnt
preparation of a small p iece of cerebral cortex and impres- ofaltemathe tC!\tS has been intcnsivclyresearchcd. lntcma-
sion smears of. in order of preference. kidney. liver. lung and li<mally validated enzyme-linked immunosorbcmt assays
spleen. These are stained in the same manner as blood films. (EIJSAs) forthe diagnosis of B. bo11is infecrion ha\•e been de-
Babesia parasites assume a round shape \\ithin a few hours vcloped38· 10 " · in but. despite concened effons. there is still
 llov1nc bnbesiosis -I 17

no similarly validated ELISA for 8. blgemil1a. Typically, called minimum disease situation) is the most desirable,
£I.1SAs for antibodies 10 B. blgemina suffer from poor speci- permanent solution to the problem bul is rarely considered
ficiry52 but an assay recently developed in Australia shows practical or economical on either a national or locnl basis.8
considerable promise. 106 The alternative approach of allowing natural endemic s1a-
bilil}' 10 develop by practising limited or no tick control, is
similarly unrealist.ic in areas where R. appe11dic11la111s and
Differential diagnosis
t\111/J(1,omma spp. are well established.' 16 These regions are
The diagnosis of suspected clinical cases of babesiosis also endemic for Boophi/11$ spp. and essential control or
shov.oing fever. haemoglobinuria, anaemia and ictems. other tick species \\-ill ine\oitably affect the epidemiology of
should be con finned by demonstrating the parasites micro- redwater. It is recogni.i:ed. however. that the livest0ck
scopically In erythrocytes. Efforts should be made 10 differ- industries in rhe region can be guided along coSt ·effecti\'e
entiate between 8. bovis and B. bigemina infections. as pathways by ' managing' the ticks and diseases associated
specific diagnoses lead to a be11er understanding of the epi- with them. 8• 194 In the long term, this approach can be
demiology with regard 10 species distriburion. pre,-alence in achieved by Integrating the strategic use of acaricides. the
specific areas, and their respecrh·e importance in disease application of vaccines in endemically unstable conditions
outbreaks. Disease control strategies certainly benefit from and the use or rick-resistant breeds of c;mJe.8
such recorded information.
Species differentiation ls essential in handling redwater Tick control
outbreaks as it inOuences the choice of prophylactic treat· Control of ticks as a method of preventing babesiosis is
rnent to be used. and the recommendations for appropriate feasible in situations where ticks have made temporal')
follow-up vaccination programmes. incursions into em1ronmems which are normally unsuit·
Babeslosis should be differentiated from anaplasmosis able £or long-1erm sun'i\'al.~q Many farmers In regions
in animals showing anaemia and icten1s. especially in the endemic for Boopil/lusspp. also believe lhat very few or no
a.bsence of haemoglobinuria. Positive identification of para- ticks should be seen on cattle.50 lfowever. on the basis of
sites in blood smears. especially A11aplasma spp., may be cost and benefit. tick control in these regions for the purpose
difficult in advanced cases because of massive haemopoie· of prevcnri ng redwater can rarely be justified. 102
sis and low numbers of parasites at this time. lt ls common
practice in Somh Africa 10 treat such clinlcal cases wirh Vaccination
cimgs directed against both infections. Cattle develop a durable immunity after a single infection
Haemolysis, haemoglobinuria and fc1erus may also wirh B. 11igemi11a and B. bo11is. This feature has been ex-
occur in catde suffering from leptospirosis as well as from p loi1 ed in some countries with rhe use of live attenuated
non-lnfecrtous conditions such as chronic copper poison- vaccines to immunize cattle. lb, •J.I. 48· 117 Tradirionally.
ing and intoxicarions with Brassicaspp. andAl/ium spp. 81 th ese vaccines have been produced in government·
In southern Africa, cerebral babesiosis caused by 8. botiis supported laboratories as services to the livestock
can beconrused \\'Ith other nervous condltlonsln canle, such industries. amongst others in South Africa and Zimba·
16 138
as heanwater. cerebral theileriosis. sporadic bo,~ne encepha- bwc.' • so. (l lalawi also has the capabllity to produce
lomyelitis (cause<l by Chlamydophila pecomm), plam 143
vaccin.es. These vaccines are similar in composition ro
poisonings (such a~ those caused by Albizia spp., Cyru111c/111m those produced in lsrael 117 and Australla.44 • 45 • - 6 A sun•ey
spp., Sarcostemma r,imina/e) .81 chlorinated hydrocarbon of farmers in hearrwaterendemic regions of South Africa
pesticide poisoning. vitamin B1 deficiency responsible for showed that about 12 per cent used the vaccine.•0
cerebroconical necrosis. and bacterial meningit[s and F.arly redwater vaccines involved carrier-donor systems
menlngoencephalitis (such as infections with Haemophilus whereby blood of recovered carriers was used as vaccine.
sonm11s. Pasieurel/a spp. and Srrepcococc11,< spp.). These vaccines ,vere simple 10 produce but suffered from
ii is ofren speculated that new infections or ~tress might variable infectivity and poor quality concrol in general. From
'revive· a larem infection ofdilTerent haemo1ropic parasites, 1973 until 1998. a standarcliied chilled ,·accine was pro·
because mixed infecdons or Theileria, 8nbesia and Ana- duced a1 the Onderstepoon Veterinary Institute (0\11 ) in
plasma spp. are frequemly encoumered in blood smears of S0u1h Africa, using anenuared strains. which overcame the
clinically affected animals. However, such conditions are problem of poor infecth;ty bm not that of post-produc1!on
diffic~it 10 simulate in the laboratory and have not yet been quality comrol, including exclusion or contaminams. 128
S1Udied adequa1cJ~, under local condirlons. Since 1989. frozen 8. bigemlm1 and /J. bo1,is vaccines. \\'hich
are subjected to full quali1y cont rol procedures before
release, have also been produced at Ov1. These frozen
Control
vaccines steadily gained popularity and since I 998 are 1he
There are several options for conrrol of tick-borne diseases only vaccines produced. 5 :\ similar frozen 8. l,igeminn
in southern Africa. Eradication of rhe rick vectors (the SO· v:iccine is also produced in Zimbabwe. 80
-I 18 ,u:i.o~ nm: Pro1ozoal di.sea~es

~lore dttails on the use of these 1,-accines are provided The technical and biological shoncoming; of these live
below. Considerable progress has bl'en made in the setting of l'accines have pro1ided impetus 10 improve them and also to
international standards for the production and quali1y control develop inactivated vaccines. In uirro methods ha, c been
or &1be$iti vaccines. ;s. 118 111e Ondcrstepoort 8. /io11is and 8. used e~1>erimentally to grow organisms for preparation oflive
1Jigemiru1 vaccines consiSt of blood collected from animals in vaccine-r:-, 140 and alternative cryopre.,;ervati1•es have been
the acute stage ofinfecrionwith these parasites.$. 111The imen- srudied 10 impro,•e the viability of frozen products. 131 Re-
si1y oflhe infections is measured and blood is collected in an- search on inactivated vaccines in che past decade has
ticoagulant at the peak of the reactions. The blood is then focused mainly on three approache~ in idemfi)ing target
diluted to contain a standardized number of parasites per antigens. The first approach 1,,as to sequentially fractionate
dose. The ,~accine is packed in five-dose containers and merozoi1e amigens. 15•1 The second was to identify babesial
!,;sued 01, demand. It is dispatched on dry ice and each dose is molecules as p0tenrlal targets of the protecthe immune
1 m( injected intramuscular!} irrespective or the size of the response (such as the surface of sporozoites and extracellular
animal. It should be u511d as soon as possible after receipt and meroz:oites and the surface of parasitized erythrocytes). The
specific procedures are used when thawing the vaccine. third approach hn,. been based on the proposed iunction of
Thawed vaccine cannot be refrozen. Frozen vaccines are the babesial molecules (antigens) in ery1hrocytC, invasion
currently produced in monovalent form but indications are (such as apical complex organelles), followed by epitopc
that the production of mulLiva.lent vaccines is feasible. 9!I characteri7,a1ion and ability to induce protecti\'e Immu-
The strains of B. bigemi11a and B. bouis used in the vaccine nity. t 14 Experimental vaccines containing crude or purified
are of reduced ,~rulence, but are not entirely saJe. Reactions to Babesia antigens, extracted from infected cattle or cultures
B. 1Jig,m1i11a may be seen within seven day, and to those have been de1·eloped l;lu t the level and duration of protec-
8 . bovis \\ithin 10 10 14 days. A practical recommendation is tion against hcwrologous challenge have 1·arlcd.6 lZ •00
therefore to limit the use of vaccine IO cal\'eS when non-spe- Progress in tl1e development of subunit vaccines has been
cific immunity will minimize I.he risk of reactions. When older hampered by pol~'lllorphism among Isolates and species. and
animals have to be vaccinated. there is a risk of severe vaccine a lack of knO\\lcdgc of the immune c,ffcctc>r mechanism~
reaction~. The:se reactions occur infrequent!)", but Y,lluable responsible for protection against hacmoparasitic dis-
animals ob1iously warrant close attention. and should be ob- eases, 100, 114 b1n anugcns
• chat sumu
. Iate srrong anam ncsuc .
served daily for three weeks after vaccination. Be<:ause of the im mune responses h:1ve been identified. 1' 1 For any sub-
risk of abonions. vaccination of pregnant cows is rarely ad- unit vaccine to be successful it should produce an
vised. Ideally, rectal temperatures of 1-accinated canle should immunity that Is at least comparable 10 rhat or the live \'ac-
be taken and animals treated when significant fever de,·elops. cinesin degree of protection as well as the longmiryor the im-
Prophylactic treatment, includin_g use or low dosf.'s of babe· munity. :-:o killed or subunit vaccines were a,-ailable in the
&iacldcs, tecracyclines and aminoguanidine have also been year2000 for routine use against 8. bigemina and 8. bol'is.
used to reduce the prevalence of vaccine reactions.48• 59• 711
The main fea!\lres of the live redwater vaccines are as Treatment
follows; Attempt, at trea1i1)g redwater feature amongst the earliest
• Protective Immun ity develops in four to sLx weeks. studies on the contrnl of livesrock diseases in southern
• The immunity lasts for several years In the case of 8. bo- ,'\frica. A number of compounds ha,·c been used through
11is.~ but in the absence o f narural choJlenge, it may break tl1e years. some of which are shown in Table 25.2. Unfortu-
down it1 the case of B. bigemi11a_ll I llowever, the latter nately. none is Ideal for the purpose and while some of the
doe,- not seem to pose a problem under local con<iiflons diamidine compounds have proven to be both saie and ef-
and rel'attination is rherefore rarely ad\'ocated. fecth·e. problems with residues have affected availability in
• Red\\'ater l'accines can be gh·en at I.he same time as ana- some ~ountrie$, and potentially also export markets for
plasmosi$ and other vaccines, except hcam,•arer vac- beef and dairy products in others. To complicate mauers
cine. further, the supply by some manufacturers ha,, been er-
• Anenuated strains or B. bouis and B. bigemina impart a ratic in recent years. making it difficult at times 10 obtain
high degr.-e of imm tmity to other strains. even in other the drug of choice. Dose rates and instructions of rhe
countries and comi nents.40• 41 rnanufactnrers must be adhered to at all time,. The c:hemo-
• Factors influencing the development of protective im- therapy of babcslosis is comprehensively reviewed in the
. u w.as
munity include.~ potency or the vaccine tiiability), devel- ltterarnre.
opment of "breakthrough· field isolates. poor immuno- Drug resistance to bo,ine 8abesia spp. has not bucn con-
geniciry of the va.cci ne strain and poor responsiveness of fim1ed. Some tolerance by B. bo11i~ to fmidocarb has been re-
the vacciuated animals. m poned.3" but did :not aff£'ct the therapeutic value of the drug,
Recovery i~ the rule if specific treatment is given early in
Details on the use of redwater ,·accines arc prol'ided with the course oi the infection. llowever. lf treatment is delayed.
the vaccine. supportive therapy may be essential if the animal is to
 Bovine babesio~•• 419

Table 25.2 Chemotherapy of bovine babesios1s

DRUG TRADE NAME DOSAGEAND USES ANO DISADVANTAGES

RDtrTE OF ADVANTAGES
.'J,i)MINISTRATION"
DIAM!OINE OERIVATMS
Oiminaien'e Serenil 3.5mg/Kg1/m Rapid activity against both
Ganaseg 88/)esla spc.. well tolerated
Tl'{pazen
Veriban
Babezene
Dimisol
Am1ca1oalioe Oiampron 5-10 mg/kg s/c. i/m Rapid activity agams, both
Pirodia Babes1S spp.. wsll tolerateo
lm1docarb lmizol l.2-3.0 mg/1<g s/c Rapid ac11vi1Y a9a1ns1 bot11 Nephro- and hepa1010xic a; high
For,av 65 or 1m Babesia s'l)O, well totecated doses
Phenam1d1ne Pnenam1d1ne 12 mg/kg SIC Ot 1/m Greatest act1vt1Y agatl'.st Cnohnesteras.i nh1brl1on
8 b1gemina
QUINDUNE DERIVATIVES
Oumuromum sulphate Babesan 1 mg/'<g s/c Greatest ac1r:,tv against Slow effect on 8. bo,1s.
l udobal 8. bigemina Chollnesteiase inh1bll1on dose :ate
Acaprio should not be exceeded !atropine
Pirevan ccunteracts to~ic effec:si
ACRIDINE DERIVATIVES
Euilavme Gonacrine 2-4 mgl'<g 1/V Rapid awvny against both H1gn1v irritant 1f not given smci lv Vv
Euilavine Bobesiaspo
Trypan blua Tl'{pan blue 0.1 mgr~g 1/v Actrve against B. b,gem,r,a Little effect agamst B txms ,mtan!
11 not g,ven ,iv: a1sco1orauon of m, 1~
and carcass
Ao1ib1oucs
Tetracycline TerramytmLA 20mg/lcg1/m Mitigates Babes1a vaccine Doubilul efficacy in clin,cal disease
react,ons

• itm,. intramuscular. s/c = subcutaneous; ,/\' = intravenou,

survive. Xon-specific support includes the use or haema- a lso useful when susceptible canie are transponed through
tinics. vitamins. imravenous administration of fluids. good or temporarily reside in an endemic area, and when preg-
nutrition and provision of shade. Blood transfusions may be nam cows need temporary protcction.39
indicated in cauie with heavy parasilaernias and low PCVs The use of prophylactic compounds may be combined
(<0, l Ol; hisco-incompatibility is seldom seen at the first with an intensh·e dipping programme aimed at eradicating
transfusion. In acute B. bouis infections. use ofantio1'idants the vector ticks. or with subsequent vaccination to stabilize
such as vitamin E. and high doses of corticosteroids may the simation.39 Unfortunately, the prophylactic use of imi-
help to offset the hypotenslve and hypercoagulable state of dc,carb may interfere with the development ofimmuniry fol-
the animal. 14 In cases of cerebral babesiosis, lntra11enous lowing vaccination because the residual effect of the drug
use ofhypenonic solmions of ma nnitol or glucose may pro- may eliminate or suppress the infection.~9 · 43 The interval
vide temporary relief. between the use of imidocarb and vaccination should be at
least eight weeks if immunity to 8. /Jo11is is required and 16
Chemoprophylaxis weeks in the case of B. /Jigemi11a. tr diminazene is used. the
lmidocarb and diminazene are the only babesiacides with intervals for the 1wo parasites should be about four and
useful prophyiactlc properties for the shon-term control or eight weeks respectively.5
prevention ofbabesiosis. Treatment with imidocarb (3 mg/
kg) will prevent oven 8. bovis infections for at least four Genetic resistance
week& and 8. /Jigemina infections for at least eight weeks.5• 136 It is well known that Sanga and Zebu cartk• can} less
Diminazene (3,5 mg/ kg) will protect cattle against tl1e two ticks chan European breeds· and these breeds also have
d iseases for one and two weeks respectlvely.5 a degree of natural resistance to babesiosis. This trait has.
Shon-term comrol of outbreaks by chemoprophyla:<is no doubt, contributed 10 the popu larity of indigenous
may be valuable. particularly if the morbid icy races are high. breeds such as the Afrikaner, Nguni and Mashona in south·
Treatment of all affected and exposed cattle will be costlr em Africa and Zebu breeds generally here and in other
bm ,,~II bring about rapid. although temporary, relief.39 It ls endemic parts of the world. Unfonunatel)', the resistance of
lhese cattle b not absolute and significant losses c:an re..~ult Control of outbreaks
from exposure ro babesio~is. While lhe literature on the mpic Procedures to be followed during an outbreak will depend
is cor1flicting. it i$ clearrhat pure Bqs ind/cus cattle have a high largely on the number and manageability of the animals
degree ofnawral resis1ance to 8. bouis and 8. bii;emi11a infet· concerned. and the a,·ailability and coSt of labour, drugs,
lions while crossbred cattle are sufficieutly ~usceptible to vaccine and acaricides. One or more of the following actions
w.:irrant preventive measures.9 • 11 12 Using Ran1:suy's disease can be taken to limit losses:5
prediction model 126 on darn from Queensland.14 the benefit • Treat skk animals and separate them. if poss-ible. from
10 cost ra1io of vaccina1ion is obviously very favourable in the the rest of the herd .
case of Bos rr111rus cattle. II is also favourable for crossbr<•d • Have the diagnosh confirmed at a reputable laborat0iy.
cartle except at the 90 per ccm seroprevalence level. while. • Treat unaffected cattle for 1icks to prevem exposure.
in the case of &s lrul/c11s cauJe. It is only po.;.irive when 1hc • Consider immedia1e vaccination of all unaffected cattle.
seroprevalence is 30 to 50 per cent and only calves were • Consider use of a prophylactic tream1ent programme as
vaccit1ated. mentioned above.

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18 1uw,,,. w c " X""' , , ""'· 1~~ Haltom b<w.tr. bo,'Ulci helper T cull Hn~
" "-'U..~.. l ~ Ticl.. anti cick-bonn"' dL;c:isc ronnol A p.ruC1il'bJ rlcld re.Jcu, c \',1lh ~1ub?~ and mcmbmnc an11gen~ of mcro7.o1tcs
m.snu.11. \'ol.11. I id,,·borne dlsclbe conuol. Rom~ Food ,,nt.J T:.tpcrtm,·ma/ Pam,titofog;·. ,4, 1~8-199,
\s:rfculturnl Oq:.1nfza1ion ur 1he I Jn1100 Xation:S-
19 ntto,..~. \\ c.. \U''..U WAI~. T.I',, UOT'l.l4 , .. RUl.'tl• ._,,.• fUCE-FIC'II I ,\.C.. mo,.
:; A~o~. t998. R:tbc>fos,s. Onder\H:poort Ve1enna1y Rt·~3rch lnst1tut(I. n.w.. -..ut>11t7., c "L, 1;,n<. SM,t. k l'AI ,rnn. (;,u.. 1998. lmrmmudomimm1
Agncuhut3J Nl!\t!atch Council. south Afucn. µµ . l-8.
T-ccU •ntlg<-n.~.tnd cpuop<> or 8<1bu.,f11 /7<11•i$ ond &llh•~la bi11<mi11a.
ll llti,nW!. :.t.T, ,,o~"ftSfCiftO·JMJti"-, ~.• MM<», r .• A~~"ft,;, , •• ORREGO~. M,t. -~nn,,J,, of Tropical .\T1•dUin.-and /'rrrasltol?1;.>·, g2, -473--482.
&. .\Rf'\.\!>,J,.J •• 1~6. P«>duccion M:ml,indu,..ui.t.ldt \':ll;Un.A
-a.nl[·b:UW~ln:-i, f>rltttn euhr\'O In \1UO. \ ·,,rctrl,:.arw frr,p""'· 21. 59-741 20 '8ttm.. ,. \\',C. & Rlct·J-1e111. A,C,. 19!:14, lhr nf hcJp,l1rl cell, ~o ldtn1lfy
pnt<nttal v;,mne amtson~ or TJ/lb.,,</11 /1<wls. P1mi.;ttol1>&)' Todllf.
; ncs..~tsc.tcn. R."'$<'lJ10, ~,AN, J.u .• 1983. Serologkttl n-sponse or cmtll.: 10 10, 14i-M9
lnf~ti(lt1 with Bt:the,fin blg-~ml,u1 und Bab~ia bot•ts In ,outhrcrtAfrlca...
:.!I Htt.o\\':,;, \\., \\'(MIU,, \ 1,)1.. 00111111..\l'JU., n•.i\..E. 6' t,.Or..\,,, lrr..4i ~ 1~93
Ondl'r.SU'/J()Ot'I /IJlJIJttJ{ of Vl!lt1ri1wry• ll~.sc.'1rth, so. 115--l I 1
He1trogtncln In t;10,,no prolllrt$n( n,,i,,,./n /101•/1-.pcc:,Jic hn"nc
8 BlG..\l.J..'£ 1t.n , 191~. Thoughts on 1hc:, ru,urr C'Ontr0I 01 ucli;.j :md
et1-1• T·ccU clan"' a<1!1-.1l/d /11 "'"''· f11/...-1/nnt1n,I lmm1111/Q·. Iii.
d~k-b1>mo d,sc;i;e~ in '°nl~ in S0u1h Ainc•. Joumaf of,;,, S<iurh Afrfra,1 3273-JJSI
Biotoglro!Sod1•r.11• 17. 7- 17..
~2 uu!>-rtt11R. r.., 1988 Thr- infcict1on ot' ,,.uiou, Ock si,c,ctcs with /Jl1l,af1.,
~ B()Cl,., ft,.J·., Ub \()~. A,I, 5:JNG!-ro, I (,, & .\1CLEUAX, u., .. 1~97. l±fl'ect of IJ1,&t!mlnn it:f rra.n.,mb.SfDn am.I ldcnriticat1C)n. P-nrll~flfJ!ogy Rt1s.•drl'l1. i4 ,
brc:cc! of Cdttlc on lnr,nt~ re>L'{t:,rrc.: co fnt'cc1ft.1n wilb BtthtJi,a bo11U. 8 1
124-330.
Jng('*mi,;a and .-tuaplwmti mn"8,11J(lle.A1.,srra1fan trturlmr~· Jr,:,m(I/. 75.
:!-5. ~3 CALDt:Jt, 1..\.M,. let.DI)\". <,.ft.1 Cfllf.\'J.$, l.,, t'.OIUU -..:n·, C..U.. UlTfUi ll!.,
U\'l1''000n. •.11. ~O'R\'.\L. tt,,.,..1 •• :i.Mnu. r_ 1,, 1,u..,11., J-.11., l99b, ~tonito.ring
10 fLC)Ct;, JU .. 1H \'O,.), \,t .. LEW, \., ._lM0:,,.10.X, t.i., h fll.Ul1ll, l,fL 0 l~5. Siudit"
811/mla bcll'irinfoctions In canlo by using PCR-ba«:d 1,:,,1<.Jo11m11/ of
cin f.nJhtr(' o!T Airnm U\'c Bt1l:>P#11 llf'>t'IJ \11ccine . .Au.u11,11tm t·"ttrrilfmy
Clinicnl Mf(rol,tr,lo~·. :.,.i. 2748..2755.
Jo11r>111T, 72. 29&-300.
:e4 GAl!.O\\', LI-, 1~~~A11l1nt,J Hrallh fn A.u1trulln Vol- \'. PrmQ:JH1l mul
II t,.C)O.. n.E-.. ~,,(:S1C,~·. ·u; .. .sr.,so,:,\~ S.f. & DE VOS,., J•• t999. f.ff('(t of
nttk~ brc1..lt't on inn-ate r~ifotjne~ to fnO('uLuion or Bnl~'ltt hll{t'miutt Rltkeittu11 Dl,.=s. <:anherrn. ,\usn•li>n Huri·au of Animal Uc.tlth,
,\u,t:1lllan Go'<·mmcn1 l'ubli<hin~ Sc1'1ce.
Au...uuliun ~ 't•ftrinnn• /0tlfn(1/, 77 -465--166
l:i: PoCi,., K.k., -..iM1~10~. r,<:. fi o,, \'fJ..o,:, ,\.f., 1999. Efcct aftm..~ of caulc: on ~~ CAl.1.0\\', LL.,U'IU.\J. G....l..S. \'.\~UL,1J.1:Rt.JIL1 L, 1981. Scro!oglca.l
mm-sons.,,on rutL•and inmJ.ll" re!lo.i~1anc\• 10 mrection with Batwsm bo11u t.·<,mpa.rh:on of "\tram.~ of 8al>e.tfa lxwis ut;curring in .\us1mJia and
Md 8, hlg.!mlnn trAtl~miucd by H<x,phllus mfr:rop/1)$, t\us1mUa11 ~hr,.amblque fr())na1f A11imnf Hen/tit tmd 11todurtf,m, 1:i. t'l...S2.
# \'&'ftft1U'Jt'j' /ourun(, n. -161-46·1 !:& CAUO\\', I .t .. \11":(;RfOOft. W .. PAN-mt. R.f. k l)AL(;l.l.Jl,$11. k..J_ 19';'.\,
\'i 800:, R..k, uw A,L ,usr,m~. C., \I., f~.)TOS, v., f.-lOJU;ll~MiS, \\.J,: 3000. The immunil) of c~ttlc to Btt.btsln 1Jtgt·,11ou1 3.ftcr drug stcrlh.:lalt(MJ
.~pplka,ion of l'Ck '"'"')" to t(eienmn~ 1he j;el\Ol}")'le o! IJ/1/INfa b<wis of inftttlon~ of va.r;il1A duration_ .\usttalia11 l't1tl·rJt:no Jour,u,J,
p.tt11•1h~ i>olot<d from ""nlc \\lth clinical l>•be-<!o,i, ><>On n!tH so. ij-11.
\'ilcdnadon '1gnit-:-"t tftk fo\en ·\usrrnlian \ i'tmrmry· Joumaf. 18, J79- 2i CAI LO\\, L.L, 1'0<,(R.'\ n..1. ~ Of \'(h,, ,\,J,, 1.993, Tid;~bomc 014-fi)W~
181 Caulo,PU;thoJOjt\ und s..~rolt>g!,'. Ju: cutt~ll(. t-.\. & OAUU!!ol, r .J .. (ed.S).
1.1 a()C'.:, 11.f., 1999. Tick Fe\\"'1 Rt~.lteh ("...cnu'=', W1u:of. Quceiu;fanJ. Aunrnlim, ~uuulilfd I>fngrtostit Teclmlquc,•s fo.r .tltW)'UJl 1'lu1lJ•'J. C..O:,IRO
\t"trafa, unpublished do,o. tnro:matinn ~~"'""'' f..w \lclbournl!. pp. 1- 16.
 BtJvine babesin$iS 421

~8 CA.UOW. w.. fr.Stt\.,.AXi, N.P., 1978, rrnmunu>iuppression b~· Babes.ta bavis cp1dcmlology ofbo,ine b3b<s!lo,is. Ondtmepoorr /orm,al of\l,•r,ri1111ry·
agaJl'\)I itS 1:ck vector, Boophllus ullc'roplus.iVawr1111.c,ultmJ. 212. Hl!St1tm:lt. 50. 3-5.
818-819.
47 u, """'- o.,,..
1989 Vctenmuy ll111carch lnsthu1c. Ornlc1>1cpoon, south
~ Cl.ARK, r-A .• ,982. Correlnuon between su.."Ccptibflny co malarin and Africa. llnpublishcd dam.
b.ibcsio pamsht'S 011d 10 cndo1oxic11)', '/'ra11wc,in1110///1< Ht>)ril l,ot·:01;1• 0
46 Ot \'f,\.Al,. O,l,, 1996. \ ;\CCHHUiCJn ag.;1H'l'1~1 balw~ic.>!'iS.. /tr f>rt>CCl>dlng-. Oflht:
o/Trop/cal ,\f«l/cl11rt11ul Hyg4'11•, 76. 4-7.
VJll lntemationat Congre~ofParnsitology.Acw l'ornsltn!Qgfro rur,·itn,
30 Ct.Al(~. I.A .• HU~'T. ,.11. 4,COWOC."o:, w.n., 19H6. OX)'~tn•dvrl\t,:d free rndicaJ.) ZOSuppl.1, 487-l99.
In the pothogcncsi.\ ofp;,""!tlc di,caso. J\clv~11,w /11 P1trasi1alogy. ?S.
4>} IW \\'\.\.t.. D.T,, PQT(;lffffiJ\, f.t,, C".()MBIU~K., M,I•. & ,\\Al;O:,.;, T. t .. 19,.9'), The
1-t4.
lsolauon and ttun~»ion o( an un!domlllod lkt/Jesfa sp. to c~11le by
~1 tlXI~>. Q,\l, & co1•1•fl. k,l. l~SS, C:Yto:1dht$ion and ~aldporum .Woloriu1 ltyulumma rnm,·ar1w1 Koch 184-~ Omlurstep(X111Ja1m111lo/\"t1termrtrJ
Going wlth the J101,. Pnrtultology 1r,,luy, 11. 282-287 Ri!,f('(lft'h, ;»;
229-232.
~ UA1.Gur,sH. n.1 .. 1993. liabe~iOils. /n! \\'/\lt!U':r-:, X'.". (ed.). lmm1molQgJ1n11d 59 DU Pt.E»t,, l,l., DI! WMI. D,T. I< STOl.'tsY. w,,.,
,9,4. Asur.c)' oflhe
.\/ot.c«far Bfolog)' oj Purosiric T1ifec/io>1s Oxford. UK: Blad.,,~11 inddcnce and impon;.111ce of the: 11Clt•bomc diseases heam,-ater.
pp. 352-383. r~dwat~r and ,unrplatr-t'l'losis in the hc;mwo1c,~cndcm11; regtQm, of South
33 DM,CUESH, ILi,, C.\U.OW. LJ..., .MliLLORS, LT. tc. ~iC<m,tGOR. W., 1981, Afrieo. 011d,w~poort Joum(I/ o/\c1erin<1Ty H<'$1J11,;/1, 61, 295-301
~ve!opm•nt of a highly lnfcc:th·c lkl/JMio blg,•minn vucdne ot reduced 51 OUlCT. \\',J, !'10tCilltTUt, JI.-Y., tJ.£\\'r\\l., U-T.. V.\>'\ Omt LU(:T. J,f.41 \ ' \~ t>Efl
vfrule.ncc. A11suttlln11 V~r".rinflt)'}aur,wJ. Si. 8-1 J vw1fR, F.H .. 1s61. Chronic splomt,s os a posslb!e compticotlng factor in
3A Oi\LGUtsU, R.J.. MOLLOY, f,8., 80Q... R.H. 41 JORGP..~S.L"I:. \\.$~. 19'9,S 00 cal~:i- Inoculated ,\1th an .:meounted rcdwoter vaccinc.Jm,n:11/ OJ rite
pam.site nntigt>.us on ~rytbrOC)"les deternunc host•pnrrurit.e relotionsh.ips South A./ricm1 \ ~tenna,y ASsuefclfrafl. S8, t 31-131.
in !Jabc.-.ia infectlons,fn caute-1 Lessons from :v1alarln. /n:coos~ L.<. ROTH• !,Z f:L•C;\\'Slf, A.. SVNl)Ql,11.t.'T, U., Of>tJ~l "-!S$0N, u,,,.. 111~"\U, M fr ~A..,'S-L.k•.,.:,1.,
(eds). Pl't)C('l!d/r1gtoftht211d 1111~null/011ttl Co1>j<Tffiteo11
CIUU>, " - 1996, Ob..,"1ttio1is on 1ho use orf.US,o\ for ~«~Ion orBn!N$ia
Tick,bome Pathogen~a, 1hl! Host•J 1tc10, /11ttrjhet: •.\ C,'fobtll 1xmt/Ji"Clfl._,.
IJlg~mlna~peeitlc omibodfes. \fell'rmt11), Pom$/tnlog)'> 62, Si-61
Urtiled Utho.
&:. c ~ Q.$,. \\'AC~'IJC. c.c•• CM tr., r.M .. I l~SI\U>h \l,JI N HOWU...
35 DALC.U~H. J\.J, - srr.WART1 s.p.. Jgn. ·rolt'ffl.llC~ to 1m1docarb induced 1983- Hypocholffierolemla.u1d l1)'11C><'onls.olernin ln "~~m,Nid 1,:minill
cxpertmentilly in tick•ua11smiued Oabesia.ar~.tntir,a. :'\u.uralian (J(1l;,..,alxi1-ftln(tctioru \i'111rlnaryP/lmsllo/i1i;y. 12.1-1 I
VNarillOt)' foumnl. 53, 176-USO.
5,1 l'\U,10·, l'-1 198l. ,\dopdw: 1ransfcrofhnnnmlly to lnfccdon \\ith
J6 OALC'd.tr~tt. P..J.. STE!\\'ART, '\',P.. ~\.Ali.ow. 1..1... 19;8, 1rransm,s.s1on or rJwllt"rlt11,arit1 11::.a_,t C:o.a..'t Fe\·t:r1 httwcen caule twin>- ~s.,•a,,h m
8'1/x,yia b1f!,cmiun by mun:fe:r of ;,.duh mnle 8oqplritu1 miuovlu$. Vttorinary SciMel', 30, 364-$;'
rlusl1'alttt11 l'•rtri,wryJo11mnl 5,1, i<)S-2~
ss t.l\"T\VlffLL r .. 199.2. Effecis orht3t sire,~ on reproduc,h-c runcdor. or
3; w. ...: .. 01:: \'Cl:,••,.,." \\·fuatn. (,Ci .. 191)2',
lMl.k\'Mf'U.. fl.Ii .. JC.~>.(:t!.Ni-t'S1
hlill~. l>I: IIOLKOI u, K.<l, (ed.,. Jf11/l l'~rtll/ty, Con(on•ncc ond \\'orl..,fi,;p
An'olym of the composition of S<>mples 1>f (J(1/>,,sla bo1'1sand the $Crie, QC93008. D~por.mun; o( Primn,y lndu,Lm'>, Qll<!en.,ll!nd.
mrluence of d,ffercnr en\rironmenmJ condition,. on geo.e~1c:tlly <h!.tinct \11J11railr~
>Ubpopulauon..,. lmeruarfq,ul/ Joumo/ for ParasitQ/ogy. 22. 131-737.
56 1-IG\U'AUA, t,\'.. CHlf\1b, L.P, tOHSW>-, G,$,A- HUl...'-"lNG 1 G..'-1., 1S9:t.
38 DE P.C!J-1,\.lt>li. S. t ., f'..C:twDf... 1.e., Gr\200, A,U .. :Ol.AXCOLO, A,J, 1..1Jtti\1t£$f. C.I,,
Oeiectlon of Bnbe..fa bfgrmfna,infectocJ cniricr~ by polymora.<c chnin
,·.n. ,., c:;ucUEL,10NU. A-'\,, 1995. Evi;11utuion of an cn.ie)rne·lln.ked
\'A.."1::r,?SI
reaction onipllficntlon, Journal o/Cli11f<11f M/trobloll>g)·,:SU, 2576-2582
hnmunosorlicm ossa)' kit to detect Bttw,ta bom ontlbodies In cnttle.
Pnwemfre Vetenna:ry Medicine, 24: 277-2.83. Si 1:An 1)1-!ttJ t . -..?,. 1~. TranstniS$i01\ Of Btr/J(!J/11. In: AJfflC. M.. (ed•. ,
/Jobl!sfosis ojDom/JJtic Anlmtt/s amt MaJI. llc>C:1 Room. Florida: GRC
39 t>E ,o, . A.J., 1979, Epldcmiol!)l!Y on(! conuol of bo,1nc babesio,l> In
Pre.<$, In<.
South Afr!c.1 /n,mwl 0/1/i, SQ111/u\/r/ca11 Ve11ri11ary• .4,>S<1Clmla11, 5(),
3Si-362 S8 rnntouorr. ,.-. .a..aose.11 .. 199..t, Ret'cnt dt•,·elopmc.nlf. in diagno~1ks r,f
some tfck·borne df~~llSt'S. Im u1u..\'onnc. c .• PO'lMt:-:. A. $. tl,\,."SE~ J.\\".
40 m: \'OS. A,J .. B~.SI..NGF,H. "·' FOVI\I~ C.t;.., 1982. ViruJcmcc and (od>J U;g ofapplfr11'11t• blot,'Chnologicnl nmh"'U for diagnosl11g
hc1erologous strain immunity of Sou01 Afric,,n and Au11mlian Bttwtln ha8moµ.arasltt,s. J>r0t:ct-ding.s O(lh~ l:.'\'.pcn C:ansultad~n. Merida.
b<>vl, ,trains 1Sith reduced pa1hogcnki1y. ()11d~me11oorr /ournnl of Me.~ko. 4~ Octo~ 1993. FAO. Roino, Ital)'. Jljl. ~6-li7.
l'c1ermt1f)' Rrsearc/1, .;9, 133-13(>.
S9 C:At,I:~ K.M .. \\',\LTIS.fmHL r>.l .• H.OWOES. f.M .• J0flG"4~St:~. \\',K',, ~I\TJruo,-:. J••
~· 01: \'05. A.I .. C:O)!RR.tr-:Jr:, M.1,.~ flF.SSf.NGtJt, n., 198.2.. Ha.Wiia btgemi,w t.A~·r. t.J .. 1..\):.Rl.£\\'~li.l. 11. ~ U:.A TCH, r. .• 1998. Amelioration nf vrru.Jt'nt
\'lltelnc: Comp.,rlson o! lite effi""cy ,ind safc,y of •\ u<trnllan ;ind South /Job<,lt1 b<>vism(cction in clil•'l'S by 3dmlnlstnul01\ of dte nltne o'1de
Afrfran ,tritln.s. under cxpe:rlmcmal COndllioru fn South Afric.a.
s~•nth.1su inhibitor amj11ogw1n.1dino. Ptuas/11., l,mmmulogj'. ::?O. -141-4-i:S.
()ndrrsc~poort Jounu,/ o/Vtterlna,y Rem,n:lt, 49, 155-1;;8.
.;~ OE.. \'I)~. A,J.• OALGUESH, R,J. & C \ LLO\\ , LI_, 1987, &JJ.wsin. In: SOU.1....<'ll' ll,f,L,
~o mu. A,C.. COW>L"· "·'" , ...., . . .... ~.,.......... D.j, ll<Tl>IM&. P.. 198:
Ccd.J. lmmwtf Rtspon.i:cs fn 11nmsilic /.Jlfi"Ctiont.' lm11wnufQ6Y,
H<1bfnit1 bot·is: ~1ol«uh\r ~nd biologic.rt! clmrntteri~tfC$ or clo~Nt
J>'ll".Sitc line,. Exp.-rimr111nl Pa",slrologi•, 63, 180-188.
Jmmu,1oparholot,· anti lmmuuopropit)'faxis. Vol. 1l 1. l'1otc,wt1. BMa
Raton. Aorldo: CRC Press. Inc. 61 <!Oa,,' f. w..i.. t. ,..,.,.r..stft;. cu;., 1982 Tht boi.11w lmn1une respon!te to
uck,dcrived Hnb,·.<fa l,ovl,m(ec1lonaotol0Ric..'ll studies or lrola1Nl
43 Dt VOS. ,\,I .. DAl..GUfTSH. A.I. & MCGR.£00A. w.. 1986.. Efft'clof imtdocarb
il'nrnunoglobuirns-. Veu,rinar)' PmttsitDTogy, 11. 109-1'?0.
d1propion3<t prophyl:ixis on the lnfoe1Mty ond lmmunogcnicin• or 3
Blllwsiti bouis,·.acclnt" In c-ttule. 1\usrral!on vl,u:rimuy Jounu,J. 63. 62 cor-r. w,t. 1r w.,c~Ell. c..G.. 1984 n,c rolt or specific immunoglobuiio, In
17,l-lii amibody·dcp<'nd•ni ceU·mediaiod crrotoxlcily assa,-s dunn~ &1~1<n
bov,sinfec::.ion Vererl1ttlT)•Pa111si10Iogy. 14.117-128.
44 OE \'nS. A,J • .it.JOHC~SES. \Y,).., 19~:t. Protection of cuttle ag_3il1S.t
b•bt>•iosl> in troµ)col and subtrnpit'11 coun1rfcs \\1th au,·~. frozen 63 t,;QQD<lf.Ri 8,\',., RODt•SRA.\L.\~lS, Ko If \\'IUC111', J.C.. C987. ProroaguJam
,-acdnt. In: fl\'J.7... a.H•• ratr.-£\', t.~ & 110-St:A.:. 1.c. C~dsl. Tick \'i•ttor activity o( Babrsin bai1is•inf&rtd C'C)1hrocytcs.Journnl ofParnsirol~,
Blo!Qg;,, .W>?<l/ool mid V.tMnaryA~fH'<'IS. Berlin, Springer Vl'rl:,g. 73, 1002·1053,
pp. IS!l-1 ;4, 64 (;()Qf)(;H.R, 8..\'., \'."NGHT u:;.. ~1.\liOSti\'. n.r. St MCKE~~A. R~\'. l98o, 1Jabt1,;(J
""5 OEVQS.,\.J JORGr:~S-!!.S, W,J.,1(.\101.1,0)', J.t)•• 2000, 8o\1DC' babcsi()si:s. hJ: IJ0:1ts (or.gemin((): StudK-':' on ihe <'ompQ~ition and loan ion of anncum
M<muot ofS1a11tinrds for Ditl/(nostic Tci1.s ami Vaocints. Third Edftlon1 nssociated , ..;m in!ect.td t-1)1hroc) 1es. lmerntJ1imwl JcmmrJ/for
Office lr.wmatlonal des Epl1..ootie. in 11ress. Parn#roll)g)< 10. J:$-36.
t6 gt vo!!-,A, f, t:i Pm1turnr.1t:, r.T•• 1983. ·11iecJfcctoftick.comro.! on the 6S (OM\', 1.~ & Pf'\ \'OS. -\.J .• t98t. ~tudil'~on a bcninr Jkwr.""Sfa tn\Jl."-mirted by
4.22 ~~""' f"\,u; Protozoa! diseases

l~ralomma margi11n111111111f111,s Koch. 18;4 01ul,wi,J)Ollt1 /oumt1l of 11<1br#osu o/Dom..itr ,lnfm~ls n111f Ml!tt fl<K•a Rolon. r1orlda, CRC
\'t'terft:(11)• R...,Ylrclr, ~8. 215-223, l'ross Inc. Xew Yori.
66 GRA> 1,., 1,, 1'01'CIET>R, •.r.. 1981 The retention o( &1bc$/a blgm1f11n 85 surn.l!R, .,,. ,.. AUU. \ o.. 1984 Chcmolhmp)· of bab<!$loil>. !11 i,mc. ><..
lnf<ctlon b) Boopiri/,u deoo/orm,u txpchod 10 lmidrn:.trb dlproplonu1c '"BROH·HI0\1'~. P. t ~llfl>k, , .. Jell•). Mnltuin "1:d /1(//,f;/osls
durlr,g •ngorgemem. 011dtr,iepoor1 Joumnl oft "t:crl1111r; R,nmcl,, ~8. r>ordrochc \1"nlnu, '-i)ho( Publbhe,...,
225-227. es l.\\\R~~CI:. 1--"·· fOGGJS, L)I & SOR\1AI-. tt..-\,1., 1980. The t..•ft('l(1_< oh,-ar on
67 HE..'l:SISG, M.w.. 1956, ,,t.oimnl Dls,•ttrc•si,t Sc1utl: A{rft.'?:1 3rd t..-dn. Pr~tori~: 1ho ,.,n1tol of di,o.,= In lht'Ste>ck in Rh1>ll.,..i• ll'.lmbah\w). n:,•
Ccnital 'ie\\'~ Agency Lid. I ~tlru:11,y ll<Y1>m, 10,, a:?-8,.
.fi:8 HOU~..... P,J •• FR.EltfCHS. W..M .. r.J11£VES. L.£ \\'AC~-nt. O.G.. l993, Culture a- uwRr,;ct. , __\ "~nR\'AL 1L\.1 .. 197g_ A hl!i10~· of rick) and iicl,bome
conllrmalion of came, $1a1us or Btlws/11 caba/li,infeo1«1 hors~s.Joumal dJ~ises of tnult u1 RJ.1.odMi:a... Rlu.xt<11frm Vi.•li..·r:'no~·Jot1m(1I, 10. 2~0.
ofC/fnf<ill .\llcroh:o!nio•- 31 . 698-701.
88 u,1st. s.0.. 19;1 Taxonomsoi 1http1rop1:s~ms.. Trrrns11ctfo,~o/rht
6g HQ\-*£.1 .. stn\ou, s .• M\,lsopz.,...-A, o, i, \L~\AA. 5., l998, hola1ion and ,1m,ricn11 Microsr:op,cnl StJCiol)•, 90. 2·33.
cltaracrorilrndon of a /IQbesfll spec,e,; from Rhfpfe,phalus ,..,,m ~,,,.ml
!!c.s p1c'ud oil• sable nn,elopo tllipJ)(Jtri<gu.s nfg,,r. which di,-d of
89 u;,w,. , o.. 1985. l'i'tori11<1f)' Proro..wt~·. 1\mc,. low1t: low;i St••~
Unin!'l'~1ty Prcu
ocme babeslosi,. 011d,T$WJJOOrt /Oumal of I\>r~rlll/JI)' R~ro~Ji. 65.
75-80. 90 t.oHR, J;. •J'.. 19".'~ lmmun!t)'Of 8nOOft1 b!g..m11111 fnex-ptm.rmmtall~
Infected conic. Joumal of Ptou,:00/11~·. 19. 653-1i60.
70 HOWEU- C.J .. \\AU".ER, ..... ~ X"b'\'llJ.. E..M ... 19:,a, Tltkl:. ,\fittsm:d lnfl,"Cl"S of
Domwic.A11imal.s III South .Afr,a,. Por, I. V,.sc:r1prio11 a,uf 8ro/ogy. 91 LOIi~. -.-F.. 19:-3. Su...:ep1ibtl!cy of non-,p!enectomb;td •nd
Scientific pamphlet No, 393, Deponmrm or,\gneulrur4J Te<hnfcol ,ptrnmom11~ (;.thiwul c.ttdr. 10 ~·rfmcn1ol &1l>i•f(tl b:gifml1:r1
Scr,ic.,,;. Republic orsoui.h Africa. lnft'Cllon. 7.eru,11/h/1111 fur l'tt,•rinarmet/1::in. 20. ;2-56.
71 HOlTI!.. u.,t.b.. 1~:-6.. The titk.-(cvcr pornshcs or c.lale:. Pr««dir.gs ()/11',• ~u.-dy of faC1Qf". l'Ot\(Cmi!d tn
92 \l,\H()!\:0, U..f,, 1969, 8()\"UlC hJ\)~itJ-"-i~ A

Royal Soc:fc1yofQua,11slmul. R7, v-,;ii. uanmili<lon . .A•wnlsof r~1>ict1/Mlvfif/11,1111d /lara,/to/QIO·. 63. 1-14.
-:2 JOlf.,~o~-. w.c.. CUIFP, c.w., (;()Pf. \\',L & \WATT, C.A., 1996. Rt"at:ti\'C 91 ~1A110~·£\. u.1, .. t9':'2.. lrnmunc rt)~fmnse to hat:mopro1om.;t. JI. &liu~ftt
O~ll•n and nll,og•n inicnm.-dlo1e< and produc~, from polramino :,pp. /n: SOUUO.l, E-J.l.. , red...,. lmmtmiry10A11lmOI Par~Uttt. Xe-..,·\'ork:
degrododon are babesluldal 111 ufrro. .A11nals of1lu.\'11,· York,lca1temy \Q.d~ic Pn.•s,
of&:i,mca, .91, 13&-J.1,. ~ ~WtO:.:E~1 O.fl.;. MIMI.. G.B.. l91l.. Bo\ine b3bttsios!s: E-s.timation of
·3 10,ia, r.:<.. 19;8. The dur.atlon oiL1u·n1
101<.,sroi,;, 1.,.v.. U!.\TCII, c;. L lnf<e1lon n>te> Ill the 11c\,,·ec1or Boophl/u, mlrropl11s 1Ca,.,.smn1,.
lnt'e~tion •nd func1ionaf lmmunicy or Diough1mas1et and Hertford ..\tWblJ IJfliOJJ!cO.( M~lcir,~ a,ut PtlfllJilOlogy. 65. 309-317.
caidc following natural infection 1\'11h 8ob.!11il ntg,nr/na ,ind BnlNs/11 9:; M4HO~tv. o.J "Re»-~. n.n.• 19n. Epboot1ologir:d fnc1ors 111, the conrrol or
bigwnfnt1. Au.s.uah'm: Vtttrlnn?' /1Jurnal. s.;, 14-18. hovint babe~,01o1;;. .tlutrt11i,11t \ ·(,Mtinllry./oumal, -48. 292-2.98
7,1 JOH'-'STOX, LA.\',, TRUF...>M.S, li:.F. t. l"f;AR$()!\', R.n. 19.; Bn\im: b;ihc~,(>~i~: 90 'tAlto:-.n. n,f., ,,"R1C..KT. 1,G. & c;ooD(,flft, o.,·., 1919. lmmun1ty1ncnnle to
Comp3ri:<on of Ouornscen, nntlbody and C!emso Stoln1ng In posl Sal,esta bo11J, airer single mfe~uons \\1th 1>o.rasite< ofr.uiou, Qng,n.
mortem diagnosis of Jnf,,c1lon. A1wraU1111 va,~rli:a,:,• /01minL SS. 1\usru,U,111 \'s.'remwry/oumo.l. 55. 1~12.
2?2- 226.
91 fi.lAltONl\', t>,f .• h1U01n 1,c;.. 1.oOOOGl:.R. 8, \.'., \UI\AE, C.ll.., MITHFR$T. R.W, &
;5 ,osC[IA,.X. r., 111:!RltY. {1..0,, MOOJttrOUSt! P4D.~•• \fUN.St. F.L.. pf.(;RA..\ l. H.G. " VTf('.11, k k..\\l., 1981. The trtU11>ml~~lon uf lln/;..•fla l10v/1in htrds ur
'""c:,ct.",
>1 .• 1988. Epidemiology ofbo,1ne bobe<!osls and nnaplo,mc,si! European ~~d Z~bu x F.uropoan cnnlq 1nte.1cd wilh the uck, iJ,xJpltllus
In 7.l!mbl•. Tropk"I ,111/11111/ Henf1h 111,d Pt11t/11r:lon, 20. 23-1-2-12. m,,ropln.s.,\1utrallan VN'1rinary)o11mtfl, 57, 461-469.
;6 JOIIClt.-<StX. w.-..
0€\'0S..,,,. & OALGI.IUII. R.J_ 19a9.1nu,c1M1yof 98 \L\ltO~t\'. 0.P.. \ \~j(jHf l,G.. &~muu-.c;,,t,.,, 19;3. SO\in-c bnbestoi.JS:Th('
cryopre~rved Babesin boui.t, Bnl~~u, i)jg.,;mlnn andAJiaptasma ctntmlc pe.r~~U:nC'e of 1mmurtit~· co Ba-1Jt1,1/t1 nrgt•mtn,,;,nd 8. bigt•mtnn In C'31\'t'f.
for cattl~ ~i1er th~,"'1li;., dilution and l11cubotlon 0130 'C. 11cturi1111ry (/lo$ tnc,11is. u(m nn1ur.llly a<:quirod infection . .Annn/5 ofTropfwl
Pnrrultoloi:;.•. 3. 2·13-251. .Wedicln,·nnd l'rlmsttoloJD'· 67. 1g1.2QS.
'7 l0Jt(';~S-riN', W.~•• \\'AU)RO!I:. s .1•• MC.Ct-A.ATtl, J,. ROM,~. R.J., Pf VOS. A,I, & 99 Mr\jl:00Ln, A.I,, \'Al\ZJM, \'.It,, t:X'IL\11>1!, I .ti:., DE ESCll\lOf., $..T \ '01..POG-!1,1,
WILU~\.'fti, Ji:.E.,t99·2, Gro\vth or Bot,.:sla 1Jlg11mI11a pa~i1t:-s ln SU•f>t.•nJ.ton
"·" .- r.ucum \IOM. ~.• .. 1996. \~abnlty after 1ha1,ing und dllu1!on of
culmres for \"3cC!inc prodhctlon. Pnrtt~ftnfogy ResMFrh. :'8, 423-426. ,1t1111!1ancou,I)' ot}',UJ'l<>rrved ,accln3l lla~tn bo10J •nd l/n/,,>sin
78 JOHGE~~fi:O:, W.~.. BO<.'K, R-E., KJSGSTQ.S, T .G., J:>E \'OS.~\.J. S.\\'A10R0"'· $,J., b,gm1/nn >tflllll> cul1ur<d In ,i1ro. V•rulnmy Porndto/(©\ 61. 34:,-3-;8.
1993.A»essme:ni o( tetrJc.ycJinl" .ind Ba~&i(I culture ~upemtt10n1 a.~ 10<> ""~'· i: ,.,. ,.,.,.Ai.•.,.,..
1981. n,e m,n,fcrof Brxipl:flrz• m/(roplzu
prophylactics (or modm1ing rie:tc:dons in uule to tivf!I (Jqbi?Jfa and (Acarina· lxadidao from lnl,.,.od 10 umnr...i.tl ca1d~ under fitld
.;unp/asmo Wt.Cti,1rs.A1urralltm Vt•r,rinnl)·)oumnl. 70. 3$-36. condlrfon~. \'tlmm•I)' Pt11,;,/1olog;·, 8. 1115-188.
;9 JO~f..11. LP.• .OO:<:<tus. 1.. 19;g. The epidom1oloi;y of b:1besiol IOI >t>WS, T L, l'OTOllmR, T J . "v.i, R.ll>'SBURC. , •• t9ll6. Too lnablll~· of•
infecricms..-\d1-ance;t in P,uaslwlOg)•, 17, 113-141). '\ou1h Mrican &,i,..,,,, bol•I,, ,'!lcclnc ~11.1in (o lnftct /loQ11lii11L1 m:<rap/us.
So l>:AT'SA,'01., T.c. ~· nmTO,-. J.A., 1$5. Progre,; on the productlon offro,,cn 011d,m;poort /ounwl of\'r:,rtnaf)' /l-1r(h, S3. 1;3-H&.
.4nap/a.<ma,md 8ab,•s/awholo blood rncclru,, ai Cenlml Veterinn') L02 '1CCOS$.:.Ut.. P.f 1981 'fhc global importance of b.n.bdto.sls fo; 1\l.'l.TtC. .\l t,
L.lbomory. Harore. l.tmbabwe \'c1erlniuy Assodarion roogres>, ktt.uo., ,., .. !!dsl. Ba1N1l0Ji1. New York: Ac:a<femtc Pic.>s.
JU!iasd4le. ll-lSSep1emb<;r 1995.
103 MtHUl'l. o. & CUIU:1'. 1t.. l.97J, St•rologic;il im·fitllr-\UOMOn ;he
81 lr..1-..LLfA\tA..~. T.S.,COl=T/.lifl:, l •A,\\:, t,, ,.,l1Dl. T.,,., 198a.
P/Qm Poiumt11~ pre\-aloncr ofa.n:spln:,-mosl!tand piropl~mosis tn canlc In Boawv.ne
1111d .11.,.,,,rOJ<i(Ofes ofU1,a1,x/; in Somlr,rn Afrl(Q Capt Town: O.\lorcl rroptnnwd.i:iti und 1Jar~iro/uglu. ?S, 3-10
Unl,..:rsity Pr""-<Southcm ,\lrlc;i.
10.t ,\U:CKL'\::STt.DT. u.• G.,\O(R. '.\I., ruc:m;.. P.. 7.APF. fl•• SC:HU~. ,. • )lt.lCUtORX,
..82 KUMA1t. ~... GOOD, M.f'.. t><)NTI R.uD. r .. \'l~(T°t.. I
,t. $. 'ULU.lt. l..J-C,, 1969 H.. 19%-D:XA me=r~m~n1s re,....rdlffvronces In th~ lif~C)'Cl~ of
lnttrdeptndcncr afCD-1+ Tccll; •nd malonlll ,pl«>n rn lmmunlcy 10 lk:lmib big11mina and B. Mnis. l\\O rypit:t1 m~mbt.'t.. of,h~ &l!'nu.<
P/asm(>(//:,m ,1,rrk<'I wzckel. /o<1r11DI oflmmutto/ogy.1~3. 2017-20"..3. &,i,.,,111. Pnm.•11ololl)· lleJM1rtil, 81. &9!Hi04
8J .._i.lxc·u, ,i. w. t. ('.()OlX',1 A, n,\',. 1990.A :.Udt" cm•.ym\.4 lin.k~d .t\tt'l)' 10:; MOLLO\" I 8-., 80\\US, P.~L. 80CJ\, R.t., tlJftTI>'-, l,A.. K\T'S.~''t)E, i.C..
(SWSAI for th~ diagnosis al Bni•-,!n bo1·ls infcctio11s and for th~ ~\fES'Dl 1,M., MAllJ>:AtHt:CHE. L.~.. WAl.1)1\()!-., S..J,. 8UC:.tn. G,\\', lc
<Qroening of &lws/11-sp«l6c mono~lom,J an1ibodle,, ln11mat/011nl 0;.\LGUES:H. ft.J .. 1998. E,~luntfo1) or an F..USA for dc1cctidn of 3.ntJbod!c...""t
/ournol for Parasitology. 20. 3H-34$. 10 1/<>i.*sill /JOL"iJ u, c~nie In A11s1r.1l!~ and 2.imbabw,:. 1'r?1,•mi1•r
Sa •'VlTID. "-L. \938. Chemolhorap) of b•bc<lo,i<. /n: •1,11e. ,,. ll'<II \ i!:eri,ut')' .I/Minn~ 33. 59-6,
 Bovine babcsic,sl~ -123

106 ,,ou°' 1-!I, lfflWU~• ......'t. JD.'1'0:V,?,J•• 8RU\"f.rti.."t•.\.{;.~lffl\\1)f~. J.M .. it-& Ki.\:\I~"·· G.<" 199;-. SC.UmGAnlm.11 Jle4f&h l'ri,ufties:1;\ \"Cttrin;iry and
BOC~ Jt.l !UM.GI ~,rs. \\',1'. UUGHT.0,\\ ,., UAlt.Uf~n.",. 1q98 l:f."Otlorntc J.tl31ysh ,..1:h \pci;ii\l rt•forence ro 1he conuoi nt Hahnia l,oz:U
Dcn•IQpm~nt nr .an t.'111.),'tTle-llnked imrnunosot~m ~~41\' for dl!1~tit1n In e.>nirJ.IQueon<l.111d PhU fh,sh. Unl,·c;;ltl ofQu..ns!o.rtd.
01 ,tntlb<><l!.. ,o llnl.,,lt1 )>Jg;,n,f,,a In ca1tk> Pntr<•w>l"f!:!' R=t1ft/i. &4, 121' ROOQtS. R.I .. 1971- QM!!f\~tlna!',Of the p3lbOlD$:)'rOr BQbi•S!t: nrg~UitUf
ij5l--6SG. 111:fe,ction.s fn c.otdt :\1u:ttll1tm l'P11.·r11wry Joun:al• .;i. 2·t2-?4-
10; Mo!'., ti.Au:(,Rt _•· u \"\', ,1.c... m$11c..,1. & ,.,\1£.'i. !'It.I\•• t98s. Grt)\\i.h
1:?8 H()('ilmS. Jq. J>J'1MVllti, C.~•• Uk \'IJ.~. A..I, ¥c AOOWUJ... l).J ... 1988 l\0\111C
lnhlbjtion of &JL"'.~stn J.xmic in ..:ulture bysec~tions. fr"orn bavinu leuC0$1S \irns conu1minonion of a \'accme produced in 11u'O ~a.inst
mononu1..~lc.•ar phagocytes. /11/ection tmd Jm,mmil)', 50. S23-S26, bovio<: b:.1be,-1<>>i> ;;ult! :lno.pliumo>i$./\11.{tttclto11 Ve1.-r111t10• Jomnnl, 05.
108 \tO.S rlSt:loRO•IA>St:.S. S. lORt), ,\1,, UlOS, t... ClUILll.'' ,\.1' LOPlZ, R. l!l 285-28;
LOPEZ.\\ .. 1992- Immuniwion of <'attJt- With .m ina,uH-.t~ pcil~"\·DlNn 129 ~fUITTERS, T ,P•• Jr.'1£USl'L\~. , .• 'Sf Jf\)1 rr~ N .. r-..onr.!fflOT, \ .• 19--98. Pomsue
,iacc.ine n~t1u, an:tpl.tsmosisnnd bi1l~1os.¾i.•-\J11mfs. ofth," .V1-"1t' l'i'.i,1.: localu.auon and di,><.-mina1ion in !he llabez'ift•lnfecwd ho,,1,.-ln11nl, of
.4,"t1dMWOf&J~11tr. 6$3. 112-121.
Tnmrcal M1•tl1r11l Pma.<itol<>~·. 9Z. 513-518.
109 \\V~IJ~I .\.I,. 1•,\Ut(J\, G,U .• >,1(£,\.WAI!\'., T.f., ,~!\t, \ 1 \10J.t:\ t.R D, 1996.
1Jo s-Ptcnn. ,.,.1 f.· ·,1,1.A.,. J,k. 1978. Crl'nL'~k inrompadb1Htybel\\-etm
•

Pro'l)ecis !or subunit \'accmes. :1g_aill$1 ttd::,borm.• dl'-.t.'115,l>s. British lloaphr/11, dua/oram, .!;oth, 18'\4 ~nd Booplt/111, mir,np/u,
\·,rm,111,;• /Qum11/, 15c2. «?!~. (C.mesuini. !888. and hybr.d stcr!lit)· ofAus1robnn ond South.tjnc:m
uo ,,.,,.,, \\.O. 19~1 The occumn~ or &rbt-$111 ho1•/,, In S11u1h \!rl...,, Boopiulus uur,op!ui{l\cann~: h.odid.leJ, OndersrcJ)(>IJttJm,rnal of
Jo11mo/ t}fthe South Afrlam l'a1eri11nry Mtdlcnl .-1u<mnt1on. IZ. oZ-llo. V,1crttu,ry• R(fStarrl1 1 H. 1}g....f 5J
ui ,,ur7. w.o.. 191,g, Obs<.'l'Vlltion~ on t.he durarlon of pr•mun!\y lollo"i"1l 13t n ..,st,r.\.,T. :o;,f. " ,ut«.11:-:~1 ~. "'. ~., tm. C:ornp3rls.on c,fthe: anf\•c:u,i1>· uf
auminhtrntlan of th~ bl\'alcnt redw;u,..-,.,cclne. Journ11/ of th< Sc>111/r Bni>tsJn bn1·is.. &d,au1 t,~ufn'1 and .:lll(rp/asma atllml~for c.:iUJ(' .s:(icr
African \leterl11ar)' Mrd/rul ,1$10eintiur>, 40. •\ 19-IZO, n,·opreS<:r.-a11on In e!thcrdlm,thylsulphoxido ,o,-tSO/ or
112 NOR\~\L. EtA.t.~ 1979. T[tl lnfosra.tfon~ and tfck-born<' d.l;;;t1h1..~ Jn p0lyvi11;1p}tolfdonl.' P\ V! r1usrralitr11 \ 'tt~rinary l<mmttl. i5. i;;t..63.
llmh.:ib\\"t Rhod~it,-/ouma/ ofthe South Aftira.tt t ·,m,rimJ.l'J,' J\jjt:Jd,ufo,J. t32 ~"Tli\\',\KT, s.f>., 1978. Dlf<tn:nct•-" in 1he: Ufc t.'}f('-Jt btto,<.oen :s ,·4rrinc .. truln
50.2811--.292. and an unmodifiNl ,1m!n ol /Jltb<.,/11 /J,wls (Bnbt'<. 181191 In ili• u,:.
U3 SORVAL. R..\.I., HVJ\Z. S..H., lA\\'ltl::.~. I.A."" OAJUtC(>URT T., 1983 B(Jc;phHw m(UQp/us {Cnhi:... t(int /011m11I ofP1p1a.ux,lqg;.15, .;g·-501
EpldemfOll>g)· of tick-borne disclls~or caul~ tn 7.lmh•bwe. I l;J..l q'fCH, ft,\\ •• ,ur.m.\ I.. L. 01\Lf\\'l;.S, )?.. \l')U:fl, a.. f\lS(.,I.
r.\\' A. URM\':\'
B:tbcsi~i~. Tro11i(1J/ .~uimnl Hmllh mzd Pmtfur1inn. 15. 87-9.J. ..,.< 1~~ 5Umufationoi nJini:oxfd~ produe1ion ln macroph-a~1..·1, ~
n..; r.r.. 1995, ~l()le(:u~ ba~ti for \'.SC'C:lnl.'
PAL\.1tlt, G.H. & MCU'\'Al:O.". &,bl,l/a bot'ls. /11/,'<'rwrt n-id lmmmury, ~ 4130-l 136.
de\'tlopmenc agamsc anaphtsm.osi!.and b3bcs.1u~1,. \'tt~rinnr; 134 s1mnsU,f A.w.• 1CJS.l '.\tnn1tg.t:mvnt o:' anhtopnd pMitsilk.m in li\·e1,ioc:k.
Pm11MroloJ;>o 57. :?33-2S3. In: ou~:r..10n.L. t.D. trd. Tropln,J PMrlSJl(J;!,(!S nnd PhTIM!tic Znmu,u•,..
115 1•t•ro u,P., 199&. Ep1dt<niolog1cal lnd1cu1ors und their •Prlicutinn 10 1h• Ptnh: World \sso<iatfo>I ror tho Ad,'3ncem•nt o(Vctcrin,111
conuol of U<""k•boml' dis-c.i~. fir. Mnmwl 011 Tick cmd Tlfk·lJOrtt•· P;mhitotug~:
t;,Js,.,,,,. Comrof, food ~nd Agrlcuhural Org,nitiauon llome 1,~ ~UTH.Etm. 8.\\., tCJBi Tht! dyru1mk~ o(bybrtd2ones bctWl'<"tl tid. (.\earl
llb tJIJtP\", U.D '.\111:-ISt, f-.l,,. l'l'GAAM, M.C:,-" .SCll.l!L,°" l.t..f. l985, .\.,.)('~$mCllt or $f>tth:.,. llttr'h:tlll1.J1t(lf /01,rnalfor Pt1tti!irot,,g,·. 17, 921-92.6
enderntcstabilttyto tlcl,-borne diseases, Worl,tt111/mal Hen,u·, ;<,, 136 TA\'l()Jl. M,J. 6 Mf':U \t\bY. , . , 19;9, PrcllmlJ\31)' ob$Cr'\'31it')nS oi, tht
24-3? 1.·ombln~d thi' ot lm1do<"arb ;1:nd Bl1betia blood vsctirt~ in t:mlL' Journal
ur rip., "1!1;,, t~,. i~t. Fiait•n van:int:l!t against Utk. f,,vth Qt coulv. t,,: ,,t, L~ £.. oftltt• South ,\,frltttn \ ',•t1•rJna')' .·~odtttrou. SO. 326,-.329..
(ed. El,.,..,,1111 lmm:01/onal Co11gre,; 011 0/s..-.ires ,,rc:,111/r. Haifa. hrncl·
Br<gm;u, r,~,. 13-; 111aLER• .\ .• ago> Rcd•,,a1cr. Tra111n~nlAg(l(uiltuof J<mmat 3.4:EM%.

138 nn1u-1t. , .• 1912 Gwl,.;Jc.knes...,ofimporttd canle IUld 1hc protc1:tm~

ua vw,~o. t., 1995. Live vacdn~ .-ig.iinst hemop4m.,ftk dl"'eas.c.... o(
moculauon ~gains1 thJ.lt df~ase-. ,¾:ritt,lrmal /oumnl, 3. l-21
Lh e-,iod•. tlttt>rl11m:v Para..-rirolqgy, s:. 213-:!ll
13~ nttra Ut o., t'J-:-> Pa.;.t worl;e"'on tick illnd rltk·humc dl\taS.,..,, m
tt9 l'OTGltt~"· ~.r 19:-;-, rhc H(e ci cle ol &1b.•,in bo1·/s1111</ &ll>;sw
big,mirr11 In Uclr., and ln onl• m South .\flico. Ph.D. Rund Th,~,, southern Africa Joumnl C1{titr, Sa111i1 :'frl"cm, Vr1qilu,ry· As.santttton. -'46.
l-8.
·\fn.l<anu, Uni\"tf'lt}·
,:rtJ r,orntLnlt r.1. a. as. tt.1 .. t976. l.ight and dL:"tm>n tn!tfO'.\c."'Opfc
1.40 TlM\1$ P •• IMU.r.lMII, R.J fl,\RKY. o.~ •• D1MMOC.lo.. t.J; ,. ROll\\1,U .. ,,., ..
1983, IJOJ;l>,rin hm"i>-: comparl.son of ndturc,deri\'-t."Ci parn>ite1, nun·li\ing
qti'!lcn·,1tlon.: on the dM't'lopHH:nl or~rna11 rnero7.Qlt~ or fk1W~tu /Jfwf.,.
antigen a.ud cun\'t<lllion,11 vaccine In 1ht" protectlun or can!-: ¢:un.51
in floQp/1//w mlrroplt,..IJ\!\'8C, 011damqpoo11 /a11mnl ,1f\e1,•rlnnry·
hctcroroguu"" chw.lcng,·, :\ustmllon Vell!riunry1fozmmJ, 60. 75\ 77.
ReJMl'l'lt 43, 123-128,
1-11 Tt,1\1,, P .. ~m,"m N, 4. tJl \'I» ,q., 19<)0.$1Udy ohirulenc,~ and ,·c.-c\ar
1~1 a,o-r<,1t'rt..R. r.T.,. ns..11.r .. 1977. IJght ;md el«tmn n11c.rn:orc:o;,te
,ran,.'1ll•slon ot 8n/,.,tn lxwlt b) U-"' of clonoed paro,hc ~ne. /11/,,·11011
Gme-n·:itlons on the dt'\,:lopment of lla«s.ia big,~nhm In lar,.;nc,.
tmd lmmunil)r1 58. 2l 'il--2l;O,
ll)m}'lbac :.nd non-replet~ females of Boop/1i/11s d«n/1>r,111...
O,u1,•r,tt•1N10n Jt111mnl of l 't'fc!n-,uu,· R~,rrh. -M ~ 13--212 Utt 'M)tM< , .• Sil\\ \Ul,' r ,t()t)\\l!U.• 0.1 "MM\'. D.S .• 198.,. lmmunt
1:?2 ,o~sY.nJL F,r., u., u.,., 19n. The fim_~'" trunure o!inrr:a.•rl)ihrocytfr n:,pc,ru1e,of cottlr foUo\\1ng ,·nccinatlon \\11.h 11,in, and non-lt,ing
~tilgt!~ nf Rab<'Jln Mg~minh- 01tdf'rtttspt>0n J(lu,,u,I of \','l.i.'flll(lf}"
Bal,e,;fa lt01•u anttgen-,... \~ritrlna1/' P<arMtolog, 16. :?·:3-1i?.
ilill«Ud1, .;_.;, 157-168. 1-tl TJ()ft,\"I ll<'IJ,).... L\\\'Jlf\Cf,, 1-... MJ,lJ\\ -\, 11,1'., \\'1UU1\'.\D. A.P,._ CJl.n•11 !<A

c;,3 N)-rt.nTI R r.1.,. rr...,. 11.,., 1979. An c ll"C!mtt mft:Urt1copic t.tudv l.lf n.,.11., 199; lmmuni<minn of <mallholdct dalrycatde n~ln<t
inUa·N)'lhrocytl~,uig,•~ or /krbv,111/xwt!ln the br•ln caplllar!e,of anapl.,.m11,r, and bb1"',1n,!, In ~lolowi. Tropirt1/ ..immnl fl,·t1Trlt anti
lnftoctt-d "'J>lrnectomizcd cal\'EtS. Ondeh'lepoort Joumnl nf\'t1terifUU)' Pr11<l11<llort 29. ;;-112.
Rl'1<'t1rtlt. 4b, 41-19, 1-1,4 TitutM-.\:-., ._,,• .t, auc,Hr, c,.w.. 19?8. The c.·n'ccu. o( .'l~t' nn r~f~i.wcc nf
au l'O IC.Jl'. tl R, f~ t. &\'AN \'\Jlllt:E.~. A.S.. 101.i. Tht transn\1~'Slc)n or &ll.Nsfn c:nnl~ 10 Bt1/Ji:s/11 /101'/t.. .•\tLtllnl/ml v,1tri1U1t)'Jouma}, 5-1, 301-ao;,
bot•is LblnJ:i frot1'n infccri,·e mmerinl ob1ahtMI front Boophll111 145 WAU&SflUHL. D.J • t001)(..i~. ll.\', \\lUGHT, I.GA c.o~i~ux,. \t..\ ,. \UJIO~l '1
mittophL\ l:H'\1lt:. <.Jndtrlftep1)()tt Joun,(lf ofl',u<•1i1:.ttty R(•srart·IJ_.: I, D.F-.. 198;. An c1~1tH~ Un;:.i!d tmmuno.sorbcnt as~}· to dia~no...:e 8ab.·:1!a
;9-ao l•H,1nr,,.t1on In cA1tl• PMmiw/ogyll,,,..t1rrJ1. 73.12G-131.
l25 l'OR:\El.L l\..l, a,v,:-,.·.\G.\'S", 0.. &. Qt(C)\\11:, {.,G,.0.. 1970. J\H~mp,t:d 1.ss woo11.,rr. s.a. .. 1913, Bo\.ine pitaplu.'1l"ioUS.. h>: 110Aat. t.w •ed.••i
tr•n.m1~s1011 nf Millie p,roplasms b)' thlp)e.,phahd 1!cl:.,. rro,11,111 Sys.11-•m of\ tler-inoo ,\ft!iflcint._ \·nt I. Mu.·1oiJial Di&M:$M. London •
•-lnimlll Ht-t1ltl: nnd /lr6ducrion. 2. 1,16-1.iO. B.!IDl.-re. nnd,111 & Co•
424 sECl'los 'l\,o; Proto:roal diseases

u; wmG>n I c.• 19,:i. Srudles on lhc pMhogcnesb of Babtlia llf8'11n:f,w.•nd & \\'\LTISBUHL 0.1 .. 1987 Prottct1on of BIJbt•sl.n. l>fgemino-imm.unc:
Bab,sio bfgomltra infocalon• !nspleneccomt<Cd ca!,·es. ~l:sthrift fill anlmnls :rg:tln.~1 <ub><:quonc rhalleng~ with ,irulcn, Ba~u i>oiis.
Parrufmnktmb, 39, a;-102. lnfi-ct/011 011d fmmun/ry, ar5. ~&1-368.
s.ca ,,1e1<an, LG.. 1912. An ch.-otron micc.oscopic smdy ofinuu,\'aSc:ulnr 15,t \\ RIC.HT, 1,0., C,\SU, ~ .• CO~fMl~S. ~t.A.. OA.LH'\'MPU!. lL.P,, GA~ ~ .k ,,
tlgglunnallon in the c:crchrnl cortex due co &,/,.;ill nrgentlna infection. f,UCUXill.N, n.v.• ,uouu,-.. ,,.,,•. WALns:num.. o.r .• Mll!T1,. 1.. ft.ERJUE, D,.A, rt
lnr,•mar/011111 Joumalfar Parosl1olog;1 2. 209-2! 5. al.. 1992. Th• dc,•elopment of.i tl:l'Omblnant &1bt'>la,'llC.."Ul~.
149 w1uG1rr. 1.G., 19;9. The k.tlllkrcin•lnnin ~1..,.itm :snd it> rote in tht l'etmnor, Pams/10/ogy. •14. ~13.
hypo1ensh·e shoe~$) ndrome o!3mmal5 '"focml "ith th, u.v.. \\C.Kl:.X'l:f... ll.\', &- ),lAJIOMlY. U,f_ 19?9, Acute
1(.5 WlU(;}ff.. l,G.- GOOIJGHC,
hacmoprotoio;m pa{ashes Bnbe:sio, Pt'.asn1odi«m and Tr,ptmosoma.
Babes/" /Jtmt ,nfccclon: Astu~·orchc vascul.11 lesions ,n l:idney and
,,.11,ml Pirarmaco/Qgy, to, Jl!!-325, Jung. Z,msd1rif: fur Par,1$f1Ct:kw1du, 00, IS-2i.
150 WRICtl't, r.G .. 1981. tliochcmlent characc•ristic. of /lab;,,/" and
physkochemtcat rcocc!oos In ch• hem, /tr. iume. ,,. ",...,,x, 1.P.. (<'<l•J. I~ WR(GHT. 1.(L. \\'AlTJ(IJUIU, 0 .J.. MAIIOSEL l>.L&-COOOC."'E.R1 B..\·•• 19~.Al'Utc

Babt-;Jo#s. ~,w \'ork: ,\c;,drinlc l'rc>s, IJt,Jx,,ia oo,,/s Infections: Metabolic :,nd blood gas cho.'lgeS du.ring
lnf..,don. !Jrillsh v,1,•m111ry/011mnl, 138, 61-69.
151 \.\!fltc,n, r.G. • CO()Or.r.n. fl,\'., 1988 Vathog(mcsl< of babesiosb.. /n: 1us11c.
M., !\\Cl). ll<lbMot/$ ofVOmt4tlc J\nlmnls t1fld ,\Inn Boca Racon. Flonda, 157 z..•,'.urr. n. '- hi'OCkUiSB'II'• v.w.. 1979. BabcslC'i,h Xonspecitk :-esistanc.e,
CRC Press Inc. :-1ew York: Academic Pms, immunologkol ioccor.. ond pachogene,1$. Ad,'<lne-.tm Pal"/JJ/rolog)· I7.
lmmunop:.tl.hophys-Iology
t.S2 \\,ua1rr.1.c-.. (;(.,Of)Gt!ft, B.\'." cu.KK. 1.., .. 1938. ~9-113.
of BalJ<'sla /Jovis •nd P/as111od/umft1/cipnrw» infe<'Uons. Parnsltology
Tcdcty,4,214-218.
c;.. JQST,
15a Z\\'6W,;.,urn1, 1;,.. VA'S MEKERK. ,,.c..
"UI:! WMJ.., D,T., 199S- ,,, t4tro
culuva,lon ofo 8ah.,,1<1sp. from c::inlc tn 5Quth 1\lrica OndIJrs«poon
t.53 \\'HIGHT, l1G.. GOODGDR, It,,\',. U:ATCH. C ....\Yt.W,\J-Q.),J.H .• ROD~BkA:.USIS..~, Journal o{Vr1,n11(1ry' R,'Srorch, 62, l:r.l-142.
 26
Equine piroplasmosis
Synonyms: Biliary fe\·er, equine babesiosis, Babesia equi infection, Theileria eq11i lnfecrion.
Babesia caballi infection, galkoors van perde CAfrik.)

D T DE \ VAAL ANO J VAN HEEI\DE:-1

Introduction Furthennore. ;>;uuail and StricklanctRl showed, by means

of cross-immunity tests. that horses which had recovered
Equine piroplasmosis. more commonly kno\\il as biliary from infections with 8. cnballi were ~till susceptible to 8. equi.
fever. is an acute, subacute or chmnic tick·bame disease of No published record of the first cases of 8. caba/li infec-
equids caused b}' the intra·eryth1:ocytic prorozoa 8t1besia tion diagnosed in outh Africa can he found. apart from the
equi and BalJesia caballi. It is characreri7.ed by fever (which casual reference by De Kock 11 in 1920 to tl1e presence of
is sometimes imerminenr). progressive anaemla. icterus, 8. cabal/i in the blood of some experimental horses.
hepatomegaly and splenomegaly. Bilirubinuria and haemo- It is dlflic11lt 10 assess tl1e economic importance of
globinuria ma)' be present in the later $tages of the disease. equine piroplasmosis in south1m1 r\frica. as a ~uitable re-
Pregnant animals may abon as a result of these infections. ponlng ~ystom is not in use. but ill one survey ir was round
In 1883, Wiltshire. as cited by Henning,3~ described a that the number or cases of piroplasmosis created exceeded
fever of horses. which he referred to as ·amhrax fe\'er'. Th is thar of every other iniectious disease of horses. including in-
is apparent!}' the first record of equine piroplasmosis in fectious respira10ry diseases a nd African horse sickness.66 11
South Africa. At about the sa me time. Hutcheon·15• •16 ob- is also an important cause of wasrage (days lost by race-
served a similar condition in the th,m Cape of Good Hope horses due to not training or being withdrawn from a race)
Colony, which he named 'biliary fever·. Nunn 81 first re- in Thoroughbreds tn South Africa.a.~
garded it as the 'bilious form' of African ho rse sickness, but In earlier repons on equine piroplasmosls little effort had
17 6
Hutcheon ' ~ insisted thar biliary fever should be regarded been made to determine whether the infeclion was caused
as a separ-.ire disease enrity. In West Africa, the disease by /J. equi or 8. caba/li; the infections were consequently de-
was described tmder the name of equine malaria. 35 scribed as a single disease. In time. howc,•er, distinct differ-
Theiler103· 104 demonsrrated tha t biliary fever is noL idemical ences between them became evident. particularly with
to African horse sickness, b ut pointed out rhai a hor$e may regard 10 their morphology. life cycle, vectors and suscepli-
suffer from the two diseases co ncurrently. biUty to chemotherapy. It is therefore important to differen-
At Lhe rurn of the cennu:y. Lavcran,~ having examined tiate berween B. equi and 8. cabalii infections.
blood smears fr<>m South African horses received from This chapter deals mainly \\~th R. eq,li because almost all
Theiler. classified the organ is ms as Plroplasma ec111i. Some cli nical cases of piroplasmosis In equids in South Africa are
comroversy has accompanied the classification of this or· caused b~ this parasite. B. cabal/i infections being only
ganism over the years,:!5. en. 80• 115 and more re<:emly it was rarely associated ,,irh clinical ~ii,,ns. 811/Jesia caballi Is sel-
placed in the genus T'1eileria, 73 based on elucidation of the dom diagnosed because it causes an exrremely low paras1-
life cycle. However. general acceptance of these changes by taemia which is difficult to dett:Cl even in blood smears of
the soiemific community still needs 10 be assessed and clinically affected animals. 12 Equine piroplasmosiS is one of
therefore the name 8. equi 61 will be retained in this chapter. lhe mnjor reasons why many coumries ornside the region
In 1904 Koch 56 identified two morphologically distinct prohibit the imponarion of horses from ~out hem :\frica.
equine Babesia spp. in Zimbabwe. Later Nuttall and Strick-
land82· 83 clearly demonstrated that either one of two dis-
Aetiology and life cycle
tinct species of parasites occurs in horses suffering from
bili~· fever. They suggested the name Piroplasma cabal/i The genus 8abesia belongs LO the phylum •\picom-
for the larger parasite. plexa, class Sporozoasida, subclass Coccidiasina. order
425
426 <OCc!Os rwn: l'ro101.oal di<ea><cs

Eucoccidiorida. suborder Piroplasmorina. and family Babe- Thedcvelopmenrof 8. equlin its vector ticks {Rhipiceph·
sildae.61 However. the recent discovery of apparent exo· aluseL-errsi el!ertsi. Hyalomma detr/t4m, Hyalomma a11a10/i-
erythrocytic schi;,.-ogony by B. equi has cast some doubt on cum l!.ttavanim. Rhipicepha/us rura11/c11s and Boophl/us
its taxonomic position9R and it has therefore recently been micropl11s) has only been studied in detail In their salivary
reclassifie<l as 111eileria equi73 (see above;. glands.3-1. ~. 78 · llGOn the eighth da)•afcer nymphal tkl<sde-
The occurrence of possible sexual stages in the life t·ycle tach. sporonts of 8. eq11f are found in the salivary glands.
of Babesia spp. in thei r tick vectors was first considered by They persist unchanged in these gltmds during the nymphal
Koch in 1906.57 Recent work indicates that a sexual stage to adult moult. The mau,u·ation of sporozoites does not
probably does occur in ticks: it has been reported to occur in begin until the infected adult rick attaches to a verrebrate
a number of Babesia spp., such as 8. bigemi11a.32 · 113 B. mi· host. after which the process is completed within five days.
crori.94·s5 8. ca11is7~ and 8. equi.73 Toe sporoz.oltes of B. eq11i are3 to3.'I ~un in length.n Sporo-
zolte development of B. equl in the saliYary gland is very
Bal,e.sia equi similar to that of se,·eral Theiler/a spp. 90• 9 •
Holbrook and his colleagues described the lntra-erythrocyt.ic
developmental stages of B. et111i.~z Recent studies further indi- Bt1besia caba/li
cate that, unlike mosr other Babesia spp .. 8, equi apparently Babesia caballi develops in the venebrate hosr exclusively in
also has astage ofschizogony in l}mphocytes in the venebrate erythroC)'!es. 29• J 2 The merozohes are pear-shaped (pyri-
host.77 9ij :'-Jo schlzonts have, as ye1, been observed in blood form). pointed at their posterior ends, vary in size by be·
smears or tissues of B. equi-infected horses in South f\Irica. 12 tween 2 and 5 µm in length and by J and 1,5 µm !n width,
Merozoite fo1mation in the lympho~'!.es takes at least nine and often form pairs. The angle between these pal rs is usu-
day~ after inoculaUon of sporoioites into lymphocyte cultures ally acute.36 The trophozoites are polymorpt1ic. but round,
before it il> complete, or 12 to 14 days after infected ticks first oval or eliptical fom1s varying between 1.5 and 3 µm ln
attach to susceptible animals before merozoltes are released diameter predominate (Figure 26.2).36
from l~mphocytes to invade erythrOC}tes.1• The imra-erythro- The fine structure of the trophozoites and merozoires is
cytic s111ges probably include trophozoites, dividing forms, and essentially similar to those for other large Babes1'<1 spp.36• 99
merowites. These different developmental stages are diffic;ult Red blood cells rarely contain more than two parasites in
to dilferemiate by light microscopy. ·n1e trophozoites of B. i>q11i B. caballi infections. unlike the situation in B. equi infections.
asswne a \'aricty of forms (ovnl, round, elliptical. or spindle) The parasiraemia fn the case of B. caballi iniections is invari-
and are up to 3 µmin diameter. 99 TI1e merozoltesoccur either ably low - llSUally less than 0, 1 per cent of e1ythroC)1es a.re
as two or four pyriform parasites within a red blood cell {Fig- infected even in splt:nectomi2ed horses12and. even though the
ure 26.1). ln the latter cnso. they are arranged in the Conn of a parasitaemia caused br B. equi may exceed 20 per cenr. levels
:Vlaltese cmss. 111e parasites .ire smaller than B. caballi. being between f and 5 per cent are more common.
onlyc. l.5 µm long. The finesm1crureofthe erythrocyticstages The initial development of 8. taballi In Dermace11101·
of B. et/ui resembles that of other Babesia spp. and has been 11itens takes place in epithelial cells of the gut in the replete
describe<! by ROlan.36 female lick. Successive C)'dcs of schi:c:ogony occur and a

•
-
.. •
•
..
•
•
•• ••••

• ••
• • •

••
•
•
.. •

Figure 26.1 arood smear Babesia equi Figure 26.2 Blood smear: Ballesia caba//i
 Equine piroplasmosh .J2,

variel)' of cell types. including Ollll. become infec1ed. Babesia considerable period (probably more than 12 months) alter
tY1balli ooki11e1es (vermiculesl enter the salivary glands at binh and are !hen fully susceptible. 1 1 For more inforn1ation
about the time oflarval moult (610 10 daysafteranachment) on endemic stabillcy and instability. see Chapter 25: Bovine
and develop 10 become sporoioites (the infecrh·e forms) babesiosis.
shonlyafter nymphal ricks begin feeding. The-se sporozoite.s In pregnant mares. infecrion of the foetus at any stage of
are liberated into the lumen of the saliva~· gland and are gestation. panicularly \\ilh 8.. equi, is quite common24• 76· ~. ~
oval or pyrifonn bodies. 2,5 to 3 µm long. Tuey are thought and 11 per cent of reproducrlve failures in Thoroughbred
to be injected while the nymphs are feeding. There are also mares in South Africa have been found to result from intra·
indications 1ha1 the infection may be carried over from the uterine infections. n Infection usually resuils in abortion
nymphal to the adult stage ai1d, transovarially. to 1he follow- with the foems showing the lesions characterisric of equine
ing generacioo. 38 piroplasmosis. Infected foals. however. may be born alh·e.
No e,idence has been published that either confirms or showing $igns of the disease, or they may develop signs
conuadicts the e~stence of a sexual cycle in 1he develop- \\~thin a few days. Due tO !he persistent namre of the infec-
mem of 8. caballi. which has been studied m the tropical 1io11, a carrier mare can produce more than one infected foe.
horse tick (Dermace,1ror 11i1e11s) and in other ticks ~uch as 1us which occurs at random in her life-time. In a recem
Dermaceiuor rnargi11a111..~. D1trmr1ce111or ph·ms. Dermacemor study It was shown thaL foetuses can be infected with B. equi
siluarum and Hyalomma plwnbewn.33 ,,~thout lheir de,·cloping disease and 1hat they can be born
as carriers of the parasite.63
After being infected \\'ith B. mballi, animals mar remain
Epidenliology
carrie~ of the parasite for up to four years. while the carrier
Both .Babesia spi:cies occur in Portugal, Spain. France. Italy, scare and immunity is probably life-long after infection b}
and in large pans of the former USSR. Equine piroplasmosis 8. equi. 12• 30 Carrier horses. donkeys. mules and zebras are
probably does no.t ocem in the UK although Dermacemor infective to tides. Batie.sin cabal/i infections may persist for
reticularus does occur in parts of that country..)() :\u 1ochtho- up to live generations in RlzipiceJ)hn/11s sn11gui11e11-S (prob·
nous lnfecrions have not been observed in Germany. $\,11- ably R. 111ranic11s as I.his !'pecies could ha,•e been incorrectly
zer!and and Austria. although the potential vectors a.lso Identified as R. S(l11g11ineus by earlier workersJ.22 'There is no
occur in those counrries.114 Equine piroplasmosis is prob- cross-immunity berween B. equi and B. cabnlli.
ably indigenous to Asja. 30 Australia was free of this disease TI1e occurrence or equine piroplasmosis coincides with
until 1976 when B. equi was brought into the country the seasonal actl\~ty of the adult stages of its \'CCtor ticks and
through the importation ofSpani~h horses, but because of therefore clinical ca:,e~ are encountered more frequent!)'
the Jack of a suitable tick vector, it has remained localized.'1 during the summer months.12 Both parasites frequently
111e disease occurs throughout the African conrinent and share the same \"C4!tors in a given region and are therefore
;\,Jadagascar. \\'bile South and Central America are also en- close!)' associated, although B. equi is generally distribu1ed
demic regions. Both Babe,Sia spp. were incrodu('ed into more \,idely throughout the \\'Orld than B. calx1lli."1
Florida in rhe USA in about 1958 when Cuban horses were Rhi11iceph(ll11s e1•ertsi euerr.1i. a rwo-host tick, is the only
imported, but. with minor exeeptions, the disease ha~ re- confirmed tick vector of B. equi in South Africa. M, l<MI rrans-
mained resuicted 10 this area mission only occurs transstadiruly. Two tick species have
The distribution of the disease in southern Africa has not been identified as n,ctorsof B. caba/liin South Africa. namely
been smdied: h probably coincides ,,ith the distribution of II. <'- evertsi and Hralomma mmcm,1111. Bnbesia caballi is
the respective rick vectors. For thi$ reason 8. cabnlli i~ tcansmined minso\·ariall} by II. trwu:amm. also a two-host
probably more wi.despread in the subcontinent than lick.1 3 a nd transstadiallyby R. e. e,,ercsi.16 Both these tkk spe-
8. equi 10;, 108 (see Chapter l : Vectors: Ticks). cies are widely distribuLed in South Africa. R/1ipicepha/11.<
Horses. donkeys. mules and zebras arc 5usceptible to everrsi e11errsf is present lhroughom the Limpopo, >:onh
both B. equi and B. caballi infections.110• 96• 116 Foals born in West. Gauteng. .\lpumalangaand KwaZulu-Naral provmces.
endemic areas usually receive maternal anribodies in the the nor!.hem part of the Free State, and along the coru.t of the
colostrum against B. equi and 8. caballi. Colostral antibod- Eastern Cape and Wes1urn Cape. 11 is present in Swaziland.
ies can be detecled for up to fi\'e momhs of age in foaJs. 14• 19 Zimbabwe and eastern Botswana. bordering South Africa. 107
,\l. in the case of calves. foals are prorected from Babesia in· Hyfllomma mmmmm occurs tlu()ughou1 the western and
fectlons by passively acquire_d and non-specific factors dur- the northern pans of South Africa. It is absent from the higher
ing the first six to nine months of life. 14· 19• 30 Infection and wetter pans of the midlands and the Eastern Cape. Free
occurring during this Lime will cause immunity without any State and l\lpumalanga pro\'ince, and most part, of Kwa·
overt signs of disease developing. and an endemicallrstable Zulu->:atal and Swaziland. It is not present in Lesotho. In
situation will develop. 1~· 30 On the other hand. an endemi- Zululand (the north-enstem part of KwaZulu-1'atal 1 and the
cal.ly unstable disease situation develops when, due to infre- adjoining lower lying areas of SwarJland. It occurs in pockets
quent exposure. some animals fail io become infected for a where drier climalic conditions prevail In \•alleys.1/yalomma
428 >l£110" ,.,-,,: Promzoal diseases

m111ca111m is present throughout :'iamibia {including the and death in hor~es have been associated with pulmonary
Caprh1 Suip). BotS\v.ina, Zimbabwe and Zambia. It has oedema.67
also been recorded In Mozambique. 108 The development in some cast?s of Jaminitis with
Tran~mission of B. eq11i and 8. caballi b~· R. sa11guine11s gas1roimes1inal ,iasis may well be due to the absorption of
(probably R. 111ra11ic11s - see above) and B. equi by R. t11- endotOxins.110
r1111icus and Boophi/us microplus has been reponed.2 J. 22. Experimental infection of horses with B. equi or B. cal1alli
J.l . ~,. T. 30 Infected blood inocu lated into a susceptible ani- often resul1s in a ~ubclinical form of the disease. 17• 53 Parasi-
mal via the Intravenous (i/vJ, Intramuscular (i/m). or subcu- taemins in 1hescanimals are usually accompanied by a drop
taneous (s/cJ routes is capable of inducing 111fec1io11. in packed cell volume and platelet count.
Iatrogenic :;pread of 8. equi by the use oi contaminated hy-
podermic needles has been repcmed: 8 but there Is no evi-
Clinical signs
dence of mechanical traJ1smission by haemiuophagous
insectS. Most clinical case!> of equine piro1>lasmosis in soutllern ,\f.
rica are caused by B. er,11i. Babe,(ia l't1balli infection is usually
not clink-ally apparent and is only rnrel}• responsible for se-
Pathogenesis
vere anaemia 1;1nd otller signs typical of piroplasmosis. It oc-
Very linle is known abou1 1J1e pathogenesis of B. 1tq11/ and casional!~· causes chronic inappe1ance, poor performance.
B. caballi infections in horses. These parasites, however, are loss of body weight. pale-pink mucous membranes and mild
probably able 10 evoke tl1e same pathophysiological events tachycardia. Splenomegaly is usually present in 1he~e cases.
as described for the oilier Bnbesia spp. (see Chapter 25: Bo- The incubation period of equine piroplasmosis varies
vine babesiosis). This may well explain the dh•erse clinical from 5 10 21 days.63 · 11 9 1 93 • cos;\ rare. peracute form of the
manifestacions of disease. disease resulting in horses being found either dead or mori-
The progres~ive development of anaemia is I) pica! of bund. has been rcporrnd in South 1\frica.63 Acute cases are
disease c--,rnscd by Req11iinfoc1ions: 1 Peaks in 1>arasiwemia characterized by a fover. usually exceeding -~O •c, V3l')'ing de·
are accompanied by varying degrees of thrombocytopenia. grees of anorexia and malaise. and elevated respiratory and
hypophosphatacmia. h)'J)ofcrronaemia as well as bilirubin- pulse rates. Fever is often accompanied by sweaung. con-
aemia.17 gestion of mucous membranes. 1uchycardia and palpilating
Although 1he pathogenesis of rhe anaemia has not been cardiac sounds. fhe faecal balls are smaller and drier than
elucidated. hypophosphataemia may play a rolr. Bnbesin normal and are yellowish-grelln.
parasites depend on red blood cells for their energy supply. Subacute cases show val)ing degrees ofanorc~aand mal·
and 1he increased uptake of phosphonis by red blood cells ai~e, weight los~. an elevated or nonnal rectal temperature.
may well be responsible fore he development ofa hypophos- and increased pulse and respiratory rates. Hean ratts of 60
phataemic state. This may contribute to increased fragility beats per minute may be accompanied by medium-grade
of parasiti,:ed red blood cells.27 Hypophospha1aemia in hu- 1,ystollc murmurs. The fever is somi:times more in1ermi11ent
mans has resulted In ATP depletion as a result of the me- in 8. equi than in 8. mba/1/ infcctions.6 ; The mucous
tabolism of glyceraldehydc-3-phosphate being inhibited. A membranes mar vary from pale-pink or pale-rellow 10 brighl
deficiency of ATP predispo~es 10 haemolysis. The same yellow. Petechiae or ecc.hyrnoses may also be ,isible on the
mechanism seems to be responsible for postpanurient hae- mucous membranes. Normal bowel movements are some-
moglobinuria in daily ~c,ws.M Uypophosphataernia has also times slightly depressed, and the animal may show signs of
been reported in malaria in human,.6'2 The haemolysis may collc. such as pawing. looking at lhc nanks and l~'ing down.
result in haemoglobinaemk ,1ephrosis and uraemia in se- nit' faecal balls arc ,..mailer and firmer in consistency than
verely affec1ed horses. normal and are coated with a chin layer of mucus. Constipa-
Disseminated intravas<:u liir coagulopath}' may develop tion may be followed by diarrhoea. The urine is orten dark-
in cases of equine piroplasmosis complica1ed by 01her in- yellow 10 orange or evell brown. but in some cases it is
fections. such as African horse sickness or worm infesta· reddish-brown as a result of haemoglobin and bile pig-
35
dons. In vitro studies have indicated disturbances in tl1e mems.1?. Rectal examination usual!) reveals an enlarged
clorring mechanism but uncomplicated piroplasmosis In spleen. Untreated or neglected cases become severely
horses has not been associated wilh disseminated imravas- anaemic and show signs of general weakness. such as droop-
cvlar coagulopath)'. 109 Thrombo~es have been observed in ing of the head and neck, stumbling, muscle trembling. sway-
me 11\·er and lungs or a splenectomized horse.71 ing and lying d0\\11. There may be a mild oedematous
The pathogenesis of 1he disease in horses infe,ted with swelling of 1he distal pares of limbs and marked praeputial
B. a1b11/li has been related 10 an accumularion of parasiti1.ed oedema.11 0 but oedema of the head. limbs and ventl"dl abdo·
red blood cells in small b lood vessels and capillarles.3; II is men has not been observed in horses suffering only from
possible that the development or an acute hypotensivc state piroplasmosis in South Africa allhough this has been
mar be tl1e cause of peractlle den1h~. Hyperacu1e infections reponed in other countries. ss. 112 African horse ~icknesi, or
 Equine piroplasmosi> 429

infestations ofhelminths occurring concurremlywith equine is charac1eTi1.ed by low sodium and chloride concentrations.
piroplasmosis may explain arypical clinical signs, sucl1 as and elevated urea and creatlnine concemrarions There is a
oedema of the supraorbital fossae. and oedema of the ventral drop in the specific gravity of the urine. 1to
abdomen and 1hora.,_, sometimes associated with the disease.
Clinical signs are apparently less severe at ambiem tempera- Macro- and microscopical pathology
tures below 15 •c. tot Gross pathological findings include anaemia and iccerns:
Chronic cases usually present with a histol') of non· oedema or subcutaneou~ and subscrosal tissues: varying
specific clinical signs including mild inappetance. poor degrees of emaciation; hepatomegaly and splenomegaly;
performance and a drop in body weight. Some cases reveal enlarged pale or. in haemoglobinuric cases, red-brown
pale-pink mucous membranes and mild tachycardia. The kidneys with subcapsular and cortical haemorrhages;
spleen is usually enlarged. asciws. hydrothorax and hydropericardium \,ith epi· and
Neonatal piroplasmosis in foals is characrerized by endocardial haemorrhage$; congestion and oedema of the
weakness a, birth or the rapid onset of l_istles~ness and the lungs; and enlargement of the lymph nodes. Catarrhal
development of anaemia. severe icterus and malaise soon emeri1is, myocardial degeneration, peri- or e1>icarditil, and
thereafter, often prior to the ingestion of colos,rum.24 pneumonia hnve also bc('n recorded.Gs
Affected foals become prowessivcly lethargic and are HistopatllOlogical findings include congestion and
ultimately unable 10 stand on their own or to suck. There is oedema of 1he lungs, and fatty changes and bile sia~is in the
usually a fever and pecechiae ma~· be evident in the mucous liver. The nephrosis is charac1eri1.ed by hydroplc nncl fatty
membranes. Some foal$. however. are appareml) normol a t degeneration of the tubular epithelium. and by protein and
birth. but develop signs of the disease two to three days later. haemoglobin casts in man) tubules in the! conex and
Nervous signs are rarely seen in equine piropiasmosis.40 medulla. Congestion. haemosidero~is, and a depletion of
Marked ataxia, generalized trembling and mild to moderate lymphocytes are observed in tho spleen. 71· 93 In addition to
ronic-clonic spasms have been ob~erved in young foals these changes, marked proliferation of reticulo-endothelial
associated with high rectal temperatures a11d positive cells In 1he liver. kidneys, lungs and lymph nodes is e\'ldent.
B. equi smears. 110 Thrombi in the blood vessels of tho liver and lungs have
A \·ariety of complications have been described in equine been reponcd in splenectomized horses. 71
piroplasmosis. Acme renal failure causing marked eleva- Foetuses aborted rela1ively early in gestation are usually
llons in semm urea and crcatininc conccntra1ions haw moderately 10 severely autolysed. The lesions in aborted
been observed. 110 Some horses may show signs of colic and foetuses and neonatal foals include anaemia, moderate to
enterilis.3 " · 110 The prognosis is poor when the latter occurs severe icrerus (Figure 26.3), pecechiae on visceral and sero-
together with laminitis. Pneumonia maydevelop.101 in stal- sal surfaces. h}•drothora.x. congestion and oedema of the
lions. panial or complete loss of fenility. temporary or per- lungs. splenomegaly and hcpatomcgaly (Figure 26.4). The
manent. has been reported.65 Mares mar abort at any stage micros<:opicttl change.s are vel')' similar to cho~e in adult an!·
of ptegnancy as a result of systemic disease and pyrexia, but mals. While the parasitacmia in adult animals lhat die of the
often there is no history of previous illness In mares that disease is usually low it may often be more than 50 per cent
abon; abortion occurs most commonly in 1he last three In aborted foetuses and nec>nnral foals (Figure 26.5). The
months of gestation after intra-uterine infcction:>o. EIS and is placenta is often oedematous and contains scattered ~mall
generally accompanied by simultaneous expulsion of the haemorrhage$ or large haemorrhagic parches. ~Ian} of the
placenta.. Uterine i11volution is normal and subs·equent con- red blood cells. in the placcnt.1 arc par;isitized.
ception is not affected.
Diagnosis
Pathology
Clinical signs. examination of blood smears. serology. sub·
Clinical pathology inocularion of blood imo a susceptible animal. and xeno·
Reduction in the haemoglobin concentration and in red diagnosis are all useful In making a diagnosis of equine
blood cell and platelet counts are 1}1)ical of B. eq11i and piroplasmosis. It is not possible 10 dllieremiate between dis-
B. ca/Jalli infecrions. Acute Infections are characterized by ease caused by B. equi and 8. calm/Ii infections on clinical
lymphopenia and neutrope11ia.~~ Ncmrophilia and lym- signs alone.
phopenia ha,·e been reported in splencctomized donkeys.sr. The parasites can best be demonstrated by staining
Decreased plasma fibri nogen concenmuions and place- blood smears \\ith Romanovsky-rypc dye~. such as Giemsa. 3
let counts, 1 as well as elevated bilirubin concentrations. In peracute. acute and subacute plroplasmosis It is often dif-
have been reported in horses suffering from plroplasmosis. ficult to dcmonsrrate the parasites in 1h!n blood ,mears.
Lowered serum phosphonis and iron concemratioi1s 1; and Even in acute clinical cases of B. ca/Jal/I infection. 1he para-
varying degrees of haemoglobinuria are also present. The sirncmia is VCl')dow (<0,1 per cent) and parasites may be dlf·
serum of horses sufforing from haemoglobin uric ncphrosis flcult to find In a thin blood smear. The thick blood smear
430 ><<:nos T\">' Prot0zt>al diseases

technique-0 is useful in detecting parasites. panitularly in

those cases where the para...itaemia is very low.
Identification of carrier animals by means of blood
smear examination is not only difficult but also inaccurate.
Serological techniques such as the complement fixation
[CF)~· 28 · 39 and the indirect nuorescem anribodv (IFAJ 9 • 69• ; 9
tests are preferred. especially for the purpose of screening
horses before exportat ion to countries which are rree of the
disease.3- 101
In recent years. the enzyme-ilnked immunosorbem assa}'
(ELISA! has also been used to detect antibodies to both
species of parasites in experimemaJJr iniecced horses.33• ;.
Serological cross-reactions between B. equi and B. caba/li in
Figure 26.3 Aborted foetus showing severe iclerus
the El.lSA33• 75• 114 and false-negati\'e or false-positive sero-
logical reactions in CF and !FA tests, may be encountered. 18•
23• 102 To overcome the problems associated \\1th cross-

reactions and antigen purity, ELISAs based on recombinant

amigens and specific monoclonal amibodies have been
developed for B. equi,53 · 54 and B. caballi? Both appear 10 be
more sensitive than the CF test, bm further field validarion is
necessary before these tests 1\lill become generally accepted.
.'\lthough a cumbersome and expensi\·e procedure. the
subinocuiation of large quantities of blood (c. 500 mil imo a
susceptible (preferably splcnectom!zed) animal can be used
to idemi!Ycarriers. The subsequent close obsenration of the
recipiem animal for cliniclll signs of disease and 1he .
.'·. , . ~-
presence of parasites in the red blood cells is essential. ,~
.~..... .~..z: . ..
'-1
D:--A probes for both 8. equi and 8. cab,1/Ii may lead to the Figure 26.4 Aoorieo foetus note severe hepatomegal,•, spleno;r.egaly
development of a rapid, sensitive and specific diagnostic and icteias
1es1.2 117, 118 Such rests "~ll be invaluable in the screening or
animals for export, bu1, to date. no such tesrs are routinely
available for de1ecrion of either B. equi or B. caballi infection.
Carrier animals can also be identified br xenodiagnosis.
In this technique. a specific vector is red on the suspected
•
carrier, and the organism is thereaf1er identified either in
the vector or in a susceptible animal after the vector has
•
been allowed to feed o.n it.
Recent advances in tl1e ir, 1,irro culture method of equine • •
•
ptroplasmosis has enabled researchers/inves1igators 10
identify carrier animals of either B. equi or B. cabal/i infec-
tion. some of which were microscopil!allr and serologlcally •
negarlve.44 119 Depend ing on the skills of the operator and
the facilities available, 5 parasitaemias as low as 10- 10 may be
detected. 31 making this a very sensitiye technique for the Figure 26.5 Spleen smear from an al)oned roerus as a rasuJt af Ba/Jesia
demonstration of Infection with the added benefit chat it is equi iniecucn- 80 to 00 oer cem parasitaemia
I 00 per cem specific.
An accurate diagnosis of the specific parasite involved in
equine piroplasmosis is essential because of the difference Differential diagnosis
in drug sensiti~icy of the two parasites. Thorough screening
of animals for che presence of parasites or antibodies is re- The clinical s[gns of equine pi roplasmosis are often non-
quired before they can be exported from countries where speclfic, and the disease may be confused with a variety of
the disease is endemic to coumries free of it. Epidemic conditions.
spread of the disease may occur if carrier animals are intro- ln the febrile stage of equine inlluenza before typical
duced into countries free of the disease and in which the tick clinical signs have developed, it may be difficult 10 differ-
vector is prevalent. emiate from piroplasmosis.
 t:quinc piroplasmo,iS -131

A low percemage of horses suftering from equine en- For chis reason. and because more eliective drugs have been
cephalosis may exhibit fever and mild ictenis. which should developed , it is now rarely used.
be differentiated from those occurring as a result of equine Tetracyclines such as chlone1racycline hydrochloride
piroplasmosis.23 The development of an..'lemia and icteru~ (Aureomycin, Lederle Laboratories) and O,\ytetracycline
in horses showing classical signs of African hor~e sickness hydrochloride (Terramycin. Pfizer) are effective only agah\St
should prompt the examination of blood smears for Bnbesia B. equi when given i/v dally for two or more days at a dosage
parashes. since concurrent infeL,dons do occur.35 Equine in- rate of at lease 5.5 mg/kg body weight.49
fectious anaemia. whith has not been reported in somhern Preliminary results indicate that parvaquone, an anti-
Africa, may be very difficult to differentiate clinicaUy from theilerial dnig, can be effective in the treatment of acure
equine piroplasmosis. 8. equi infectio11s in horses.39 but it-docs not eliminate infec-
Tn the ne\\ · bom foal. piroplasmosis should be differenti· tion} 17
ated from both equid herpesVirus-1 infection and iso-eryth· Depending on the severity of the disease, supportive
rolytic neonatal icterus Ciso-immune erythrnlysisJ. The therapy is oflen necessary and may include blood and/or
lauer occurs within a rew day~ of the ingestion of colostnnn Ouid infu~ions and i/v administration of essential phospho-
and is often afebrile. lipids and antibiotics. Polyionic electrolyte infusions mar be
Abortions as a result of piroplasmosis mosc often considered ror severely dehydrated horses showing anor-
occur when pregnant mares are affected after the fifth or exia or sufferlng from diarrhoea. Oral laxatives may also be
sixth monrh of gestation, and should be differentiated Indicated.
from other potential causes of abortion such as equid Chemosterilization of 8al1esia spp. infections is rarely
herpesvirus-1. rec<>ml'nended. but may be indicated when horses are co be
moved from an endemic country ro one free of the disease. t\
fe" drugs "~ll effectively sceriliie 8. cahalli infections. buc
Control
elimination of B. equf is very difficult. 100 Orngs u;ed for che
A \,idc variery of drugs has been used to treat equine piro- elimination of B. mballi include diminaze11e diaceruratc ad-
plasmosis with varying success. ministered i/m at a daily dose of 11 mg/kg body weight on
The small (B. eq11i l and large (/J. calla/Ii l babesias of cwo consecutive days: 51• I.\O amicarbaJide ,tdministered iJ m
equids follow a pattern of drug susceplibility similar to lhar at a daily dose of 8.8 mg/kg body weight for c:wo consecutive
of rhe small CB. bovis) and. large (B. bigemi11a) babesias of days: and imidocarb given i/m ac a daily dose of2.2 mg/kg
cattle: the small babesias being generally more re~istant 10 body weight on rwo consecutivtl days. Diminazenediacetu-
rreaunenr than the large. Acute il1feccions respond to ra1c causes swelling and necrosis at the injection site~. Toxic
many of the same drugs used in cattle. but higher dosages doses result in respiratory distress. depression and other
are usually required.. signs of imo:dcation.100
Aromatic diamidines and related compounds including t\micarbalide fails to sterilize B. equi infections at dose
diminazene diaceturate (l3erenil, Hoechst) at l l mg/kg body rates of 8.8 mg/kg ru1d 11.0 mg/kg given i/m daily on four
weight administered deeply i/m: amicarbalide diisethion- consecutive davs. 10· 100 Tn one Lrial, four i/m treatments of
ate (Diampron. 1'·1ay Baker) at 9 mg/ kg body weight i/m and lmidocarb at a dosage race of 10 mgtkg at 72-hour interval$
imidocarb diproprionate (lmidocarb. WellcomeJ at 2 mg/kg eliminated 8. equi from 5 of JO ponies, but severe side-
body weight sic or i/m, are effective against both organisms. effects (colic. diarrhoea. and anorexia) and even death, were
Repeated 11ea1mems. however, may be required co control also reported. 10 In another experiment. four individual
B. equi infections. 100 treatments of imidocarb i2 hours apart. given at a rate of
The acridine dyes (euOavine and 01.hersJ at a recom- 4 mg/kg. eliminated B. equi infection in l3 or 14 horses.?6
mended dose of 4 to 8 ml/ 100 kg body weight of a 5 per cem However. eight similarly treated donkeys dled from drug
soluti.o n whh a maximum dose of 20 ml, administered i/Y toxicosis.26 Yet. in aJ101her study. imidocarb given at t11ese
very care.fully.8 are effective against both B. equi an.d B. dosage~ 1v-.1s ineffecti\·e in eliminating 8. equi-carrier srate
cabn//i. Subcutaneous administrations of these dyes cause infections in nine mamre ge!dings.!,q Adams and Corrier. as
severe irritation of the tissues at the site of injection. cited by Kutler et til..59 showed that for horses. the LO., for
Resistance of 8. equi 10 eu.flavine has been reported. 1~ The imidocarb administered rwice at an interval of 48 hour~ is
simultaneous l/m administration of acropine is recom- l6mgtkg.
mended to coumernct the potential adverse parasympatho- 8allesia equi-carrier geldings have been treated with the
mimetic effects (primarily abdominal pain) of imidocarb anti-theilerial dnig. parvaquone. at a dosage race of eirher IO
diproprionate and euflavine. 111 or 20 mgfkg for two or [our rreaunen1s seven days apart,
The bisazo dye. trypan blue. administered in a single without sterilizmg the infeclion.59
dose of 150 co 200 ml i/v of a2 per cent sollnion. only has an In summary. it would appear that none of the existing
effect on 8. caballi.11 This drug also has the disadvantage of babesiacidal drugs is satisfacrory in che s1erlliza1ion of
producing a discoloration of many of the animal's tissues. B. equi infections.
432 m:'rro." r,,'O: Protozoa! diseases

In highly intensive management sys1ems where animals vectors effectively. Strategic tick control will expose foals to
receive dally individual care, i1 is probably feasible to control ticks and allow them 10 develop immunity earl)' in IJfe,
lhe disease by eliminating the contact be1ween lhe tick vec- which \\111 result in endemic stability.
tor and equine host by means of the regular application of No vaccine is as yet available. and therefore fully sus·
an acaricide. However, under free-ranging systems ii is ceptible animals introduced into endemic areas should be
difficult. if not impossible. to conuol these multJ-host tick care.full}' watched and treated when necessary.

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A.Oali.FtTso:-.:, G.!lf., 1979. ll\'aluatlo:1 of an indirc<:1 nuort"Sccm anr[bf>dy ~6 FRERIC.US. w.~r.. AU.£.'\;', r.c-. & HOLBROOK. A..I\., 1913, Equine ptroplasmosi.1o
test ti'\ dl;&g.no,se, Bnbesifl f"flm'infeet.lons In ho~e~. A11t.trnlin11 Vtleri11af)' (Ba/*"iaeq11f):Thorapw1ie uials or imidooarb dihyd(ocblolidt in
Jo11mal. 55, S5s-5;i9. horses nnd donkeys.. n:e \'e(erI,mry R,.1cord. 93. 73-75.
10 C.4.kRRF.\', F-A., A\') kV. R.J .. .:x()WLE.& .R.C:.. II Si\$U, s:.c.. 1971~ Chemothtrapy 27 •••••en~. 1,.,q, HOIJIH<><.11:, "-'·· •!17~- Pc!ed[ng mcchanlsl)1$0( Bal,..,/,t
or eq_ui.ne babeslo.siS-./oumaJ <>J"lltt".,lmrricnn Veterinary MediMI «/tii. Joomal o/Prour...oolor;y. 2. 707-709.
A.srocia1fo11. IS9, 1538-1545. 28 FAJlklCJIS:, \\'.)S. , HOLBROOK, AJ.. ~ l(')H!",;SQX, \.J., 1969. Equlnt"
n DE. ..:oe'K. c.o.w.. l92(). Drug Ln:atment in num.llliosi!i.. i1h & 81/t RcptJrl of piroplasmosls: Complement lixlltion rltre.<o!hor<,s 1nr~1od "ith
the D/r,ctor o/\ff!leri11ary Res,;nre/1, Unio11 o[Somlt .'lfri<x>. 6S9•675. Ba/lttfa cabal/£, Amrrir1111 To11mnl o/Ve1erfMry Rttettrch, 30, 697-;02.

c:, D£ w.v.1., o.T.. 1963-19$8, i>3rasitology 01\i&ior,, Onder.1epoon 29 •'••eOHOH. ~.T.. .IIJ81. M~rpholugle ospcc:tJrnf Bau;o.sia ln the tfck. /11:
tusnc. M, & Jdtatk, J.P., (eds). &ibe1.l0$k. N(!w York: Academic Pr~""·
\ 'eterinary lns1it111e, Ondersl"!'OOn. lJ1\jmblisl\ed dot•.
so FRIEDF!Orf, ••T.. 1982. Ole piroplasmen der Equldcn-Bedeutungl\lr den
13 oe WML, o T•• 1990. Tht:.lran.so\•arial transmlss!on of Bam!smcalxtlliby ln1cma11onalen P!etdm•en:ebr, &r/111er im d Mfi<rche11tr Tltrdm://(/1•
Hyplomma rnmcamm, Ondcr,steJJ()Crr Journal o/Vt1enn1Jry Jlc.,s,earch. 57.
lV«li,11,w/,rift. 95. 363-374.
99-100.
31 FJUEDHOF1'. ~- & BOSF. "-· 1994 Recen1 dewlopmenis In diagnostic$ or
1.i DE WA.At..o T .. 19f>S. Oisffibulion, tnmsml$Slon and serodfagno;;tS of :,0111«,? rick•bomc di$t'a:Sc~ Im VILF.N8J;R~ (;., Plli\.\11}(. A.. & IL-\.,"-:·5 ~. I,\\'.,
Babes(a eq,11 and Bulws,n wba/11 in South Africa. PhD ,hr.HS. Facuhy of (eds), Us,;o/Apfl//c,1b/e 8/011'1:h110/ogia1/ Mtrluxl,for Dl11g,,an11g
\~eic-rinarySience, UnivecsiLy oft>reiomi... Sot.1th Afnc::a. Hnumoparafit,s. Prqa,ed/11g, 0/1/re Expm Co11..,,J1nnu11, lftrfilo, ,\lf!Xlco,
1S DE.WML, D.T.. HO~'<. n." JOSCMA,Ss. ;\.J.• 1~99.
F.p!demtolog)·of cqufne 4-6 0<:101/,·r /993, F.,\O. Rome. Judy, 4o-S7.
plroplasmosls In So111h Africa: De1cc1lon nnd duradon of coiosltal S2 FRJSDHOFf, >-..T "HUSCHER, c., \.9;U. nedl!l~\'CJ'}' ofJ.:'.oeh 's
~n11bodle;and che lncldeM< or new infec1ion.s m fo.t<. Prrx,edi11ss of S1mhlenkorpcrof Habe,ia bfg.•m/1111. 7.£it,<cltri/1 /fir Pnrtulm1k1111de, so,
• tl,u 8th F.i1um• /11/ocrrnu.s D¼,a=Co11/vrtnL.:, Dubai. UrutedArab 34S-347.
Emira1es. 23-?6 March 1998.
33 oon, r,., 1982. Untcr,,'Uchungen ubet die Brouchbru:kei1 von EU5'\. !FA,
16 vt \\'Ml, o.T. & P01'Gllf1'1U1, ,.T., 1987. The tmnsst1d!al rransm!sslon of lllA under KBR zum naohweisvon Rnbesia cq111 lniecdonen, l~ugural
Ha/xosfa <aballl by l/llip/ri!p/ln/1,se•'f!l'tsf ..-~mi. Ond1•mtp(1()rt )Qumal of Ofsscr<ation ou,dem 11.sutut ror vergl•khende T,openmudiz!n und
v,,e,inary Res:imrth, 54~ 65>-oSG. Porll$itologfe dcr Ti•rllmlioh«n l'al:ullU1 dc1 Unl\'er!:itd1 MOnchen, 1-:0.
17 DE WM.1. t).T.. v~-.: HF,J!.RD6N, J. 4( IIIQTCICTER. F.T•. 198;.A prclimlmuy 3,i om;o-1.Aki\Ji..-c;,. "· M.• u""'· J. o. "RJ6llftO, ~,~ r-.. 1.998. Sporogon~· and
lrwe."!lgatlon imo tho cllnic;al pnthologienl changes nnd serologicnl e~nn:corn.l trans.m1.ss1on of 81Jbesifl equiby IJ.wphilus 1mcropl11.r.
respon~ ln horses experlmomnllr ,n!eeted \\11h Ba~sla equi nnd PafiL<itoltY,:y Re.wrrJr, 84, 323-327,
 Equine pi1opla,mosJs 433

Yi rn ,~'1sr.. ~t.\\ 195ft. :lni,nnl Dlst.<Ut'J In .~,t,rlr ,.\frl<«. !lrd cdn South 57 ~U<.H. n,. 1,00. Be!rr.tt,:<' ror f.11twlc'klun~gl•\Cl11chh~d1,•1 Plrop}.!.'>mt'n,
Africa: Ctntrol :\t~w~ :\~ency. Ltd. 7.elt;Uhrt/r fl,r lfrgtrn«· ,md h1/«tfonskrrml..'htrti~n. S.I, 1-9.
3n Ho&.\., M., 19113. \ ·c~tkh1:1;de Untor,uchun;wn d~ FeJJ't\tru~mr \·on 5lt -.on-J foR.. J.,L, (,IP*>',(...\ .• (,Orr, \ .. ,1.. A JOll~!<io~. L\\,. 198(. ExpcrfrthmUl
/Jnwn11 ,nba/11 ~'(un~IL 19101 und lllrbc,rn """' J.;,,eran. 1901 H4tltt"Sin t•qu/inlccthm In matuf~ hu~1..~. \uwtf,·,m /otth:nl of\ j,•uw,mry
htHUJWitd Oi!-M'nau-cm 1u1..Uem ln\lhu1 ru, \·t.'rgf\'Jtht:nd~ R,·l~t«h.4-i, l6ml,,,lt;;~O.
Tro1,.an1,-dlzjn und Pa,~sitolopu lier 1let~rzliichrn fw.ultJc dN S-l f,.:lJTTUJI 1.., ... ~.Auc;c1. J.1 ",,ui~'.\:. C.A .• t98":". lmidocarb ;md p;in·.1qur.1n1:
Um,~r>l1J1 ~llln,hcn. l-~~- in thl' trcJ..ummt of pJropl~mo~is t&lb...':)'/(1 r11w) 1r, equid,. .\tr1N·1rcm
.17 uouittoox .u., Nh.i, l!qulnc r,irn-plafi:111f!l"i1' un,t it-. rlt3jtn0'"~ /l)u1ut1/ uf V1Ht'rinnt1• Rt•Jinrrh,48. 1613,,,tt;tG
Pr,J<"tttllt:qs oj 1/11• l'lti·,•ftlh .\,rn111,1J C:t,m1mtitm of tilt• lm1·t/((m fio L\\'11M'\ ,., 1901, Con1rlbu1ion ,f re,ude du Plravlth·ma t,1111. C:ttmple>
't.»lxlllliot1 oj f.'1JJ111w Pr11rrlrfrmtf'I, ~ti~mt (h.~.sc:h. Flondu rcndu~dh s~anres. ~1.·t,, dt Bioioii~ ,,, ''" m· ri/U1le& ol. 38.5-J.88.
38 11outROol.. ,M,.. \...,·rno,\'. o.\', .. & JOllSSO:-.. ,.,.• ,~t;a. Ob:ser,..atfun~on ah~ Ol 11vr,t "< .o 1985. \ tlnfoctr) ProW:l1<>ftJK\\ ,\m~- fQ\,.,l S1.atr- lJnn'L1hh~·
<h.•\t•lopmt=m ot' Bnl~·,fnt.Wbt)lli ,.'lu1m1l In 1h,• ttopit".il hot"l' tkh 1•n.~\.
lA.•111"1rtntfJr nlt.tt~ ~ewnJnn. Jrwrunl OJ Proto:nhlo~· 1S,, 391..J9t.',
G=.! u \\1:,,, o.J.• 19$7. 1l}p~plt~ph111..u\.mfn: A fe:icun• ()f m.Uarfa. Brrtislt
J~ llOJ UftOO~. \.A .. l"OCWf.(:.JUOl:JC, O.F~, R011l. 'r,O. ~UO•:JH\t, 'J.W, 6- 'IAl'lll' .\1.-,/fr,i/ Juumal. 29.. ~ 11....i I 7
w,u .• 1971. PrQlo(~11 tor the.• (Omplemem fi-<,Hlnn ,est tor equine
6J ll\\1$. a.n.. 1998, "f:.in~,1an•n1.t1I HM.>mi,'"1011 ,,1 llal1t'!!lo <'QIII m hu:~..t~
p1ropla.<"mo>ii, Pro..-,·,·tlitJR~ oftlte 75th ,\,umn{ .\lf~tln;tol thl! lJnilc1f
,i.mJ chl•tttt>tltCt;,f)\' of ~;.II &1loai11 spc<.:i,>ir. PhD lhf'Sh. Facull'\ ul
StlJ/dAt:inwl H~1!IIJ .'U40tintlon, Okl.ahom.1 Ut\" Oklahnm.:i
\\•tt'rittiil)' Sdetrr Um,•cr).lt:-' of Prttt1rh1. S<>ulh \fiica.
40 um Dfloo;; \,\, .. 1-111 ,uq1s. ,, .r-i.. 1968. l'.qulm:.•plrop~mo.,,. ..
/Jroc..wting., cf:Irr :2,ut ·\nm,al .Wn:mgo/ rltt· u,111«1 ~:mis Ammal
6-,1 11t111i(:>: ••,. • Mll'l \'.\, ,,.j.,
\9~. zum \ 'nrkornmcu und 1.ur •.-ei1l,m,•lhm
Bed{fllwngdc.r/.cd,c:n f)l:rma~mtJr nuugtmnu.< lM1tl.er, tj'7l) \lnd
Jftyi/Jh i\s;W<i11rion. ~cw Orleans, l.o.s ,\ngdc-..
ru,n,wt't'mor r,rlculnm1 1-Jbrtciua. l'i9·J 1m 1.kut.i;.dtJrmd,
u m,1 n.Mt'W)~ ••u. rm"i<:u ..., \\',M. •\I":-., v.c 19~ L.1bor.1tM)' cli,igno'il-. 'l'h1Jx-1w1i!dfcfoL• um/ Para,.uology. 2.-. 3!U-IO~
nf tqwne p1rnp1U..••.mosu•• l'rorMilng"uf rJw flllrd /,m.•r,illlJOunl
65 I ITTT.£(OH:-., .\•• J9(~3- aa\>e~~OSI~ hr. fl(>.",:(;. f,i , CM\ tin. LI, <~H.. \ \,
Ctmfen.,t~?uu £quin~l,rftttfou~ l)/Jc'JL~ Puri, "t Karg~t. 8.).,vt
1011,,a,:-.. , J ~ MO w,t-. h:di.,. E<1111th.. \f;,v/fr-111,•nntf~:Jrg,ry ~.:tnta
,1:: KOUR<)(.'JX. ,\. ,_ jOJaC.,-..:o~ ••q. ltl)lAUl>L'- 1•.\. 1,1,1&. £qumc
8.irhn.m. Ca.lifom1..i· A:m:m·an Vctonm1ry J•uhf1ra11CJO.'
plmp!:uinv:i ..i~: lnu~c.·~1h:'O<.')ttk dt.-,•dopnttmt o( &l~Jifl ttJbfl/11
~~ I IT"l I l.JOUS .\., ~'fill rl,rl\UI( u. "·"-,. ll
uou:,..' l>.1 IU';'"T, Pan,n,, ttf
(~\l\Hdl) ,md Bul~ln et/UI. Aml'rlta11 Jnurntil 11/\,~tt'.rlmu-:,.• RM1,,r,·Jr, 29,
29,-30.l.
equim.• dl,1:t»C in -<outhvrn Africa. 1n 197:t Viwrlnmy<:t1111(ltm, ?:i, 3-11
:lrtd 26.15~0.
,:t k01 ...,: \S. Jt.J .. 8[CU. I .. &L.\~OS. L, l'OUJ:.IJCJ. t':., CJUll. D. $t \\ \(,\l If, c••r, ..
67 MAUfJH mt .. tu;.. lillU ..... tc,,i. w !)1:\'\.'.;\..:.0I. ):.,. 195;. !>.3.tholot,'rttl
1~8. Bal~it: fVIUi fidd i,olut~ ruhurt.-d fMm ho:sc• blood 0~1111! n
mlc,ocentrifuj;e rn,r.hod Jo11mr,/ of !•am;/ralog;, 8-l. 6'36-699.
vrr1er'.'-,n in &lttsit, fJlll:'t, 1nf~fu11t.. Tm1Huumrdi:i11 u,:d Partu.ito[og1,·
8.~~-1-1
44 IIOI \1,\~, 1•.J.• FRI RIOlS, \\' ,M,, ('I lmvts, I t- WMiS.fR. c;.c., 19'!'3, (..uhurc-
t'ill M \f'~l'..\'.~Jl:.ll l, U,. r. \U(ft, \1,, ltU('ll•111 11., ~APf. 11,. ,1 111 I~. r...,. \UH ~IOli'.;.
ron11rm1UJOrl.Ol the ,~rner SitO.lUS ul Tf(l/,L·.~,t., tdlNt/H•ttUc,m:d hohch,
n .. 199l- D~A mt.l,,:Urt•mt-·nl!'i rt!\t-a1 \llffor~nn°' in the ltfo ryrlc-, o!
Ju,tr110/ 11fCl/nio1/ .',ltrrobitJIOI{)\ 31. 6~11-;01
flll/lt'$lft blgl!mifu, on1J ll w11#. two 11,,'f)kOI mombcl'\ nflh1~ ~~nu~
-l5 flUTt.H.tQ"\ 1 tft.83 adJar,· fc:1.Tr Rc!/ltJ(l ,JfrJ,;,· <.:oloruat \ ,rr,,nary &J!JcsU.1 Pnrcuttl.J!ogi· Re~,·a,ch. 81. 3,9~~
~urg1ton fo:- tlu }far l&l,3. 32-Si
()9 MAOnr.;, ft.A..., UOUJROOJ,., \.,\ :95&. J:.quln<." ptropla.sm'>SS~; lndlr«I
4t1, llU tc'HH>~. t>.. 1A85,,Rdi.iry fcv\.'r al 1he> hor'M!. R~·uart OJ tl:t' (A!fh-tlit,J 11uor~nt andbe>!J,. h.•..; (or Babts.ln adJ11/ll, Am.:rh:an/our,ml of
\·,tNin,m S1,rg<"'11 {Cir,,.,. r...:,, 1118.'i. 21.-28. \"'1lCrirU,f)' Ri!tt.•md,, 2.9, I Ji-123.
J7 11u1c.m:.ID., u.. 18~0 HIIIM\· fc\•cr or Jltundlcc in thr hoN· ..rtgrirultm,,r ;o :.,.rno,n. n.f."' ,\.\I .. ,.n.. 1~1, Ru\'irwh:.b~~fei\h: Thtd. bl()Qd mm~ fot
Jrmmal ti/lht· Ci'lfW o/ C4.mtf llt,111~. 2,361 ""'"'lfi2. 1hc tktenion ot pnrn,itnrmliJ AwrmJirm VNm1rary•Journal. 1;, .;.;-,;.
,11$ uu1t11w,. u . . :s~n. 811inry(c.;..:r.A;:flcu/r,1r11l.loumnlo/1h1·Ca:N•uf -, ,w,o,t-\·, D.F•• \',"JllGHT &.c.; .. 1 fttRWH'4i, \\ ,.\I., Utnl M ..SD\l., ~ , 0 'OriUll J\".\,"'\
G0<,d f!q;,r, IJ. 1~-I ij. u. ,1. ,11:.21,1::f('N )tc:. • \\ ,nm.u. -u t .. Ht77- hlcmlfimutt>n or &1lwsm cwu
.,y IA'-M•\'. . a.c... 19>3 The r,anii.i1k1di.1.l cffc.'<I nfaurcomrdn 1l.c.'dedt~1 <m ;,, ,\uMrali:tt ...-\1u11t1lu,n Vt-t,,rlnal)' /1mrm1l. 5'3, -161-164.
11'11N,ulf l!f/UI .l..i,trnn I 6991fn ,plenoctoml«'d donl:c), 011t1,,,u,•p1JWI 1va.a. The r,lmpl3'-Jn1to: IJfto t'\Cl~~nd ~L·xun!
,.., ,,ru111t>K'.\. u ,. ,<.:Jrt1:rir..1
Jr,umal o/\tu ruwn 11\7{.Ylrth. 26. l75 ltsl. <i.igco, Tn: ~tUllUL Jt ., tt.A~•Jt1 l,R., Nio,1. ,l.lftwuc.-1,1 /Jormitnf11<0··
G,o 10\ ,f1t. &.. ,,•• oo:-..~·HJ.\' ,. • uut:r, R.."\., t~h Comp1l•mt"nt ~tintt h°""-b l.fmdun: Acadt-mtc Ptt'......_ 2l 3; 103
for equiM plro;,l,.,mn~h ,llab,ec/11 '1/llt and /1. ,111~1/11 prrlom1t!<f 1n dtt 7'J ,.1ltU llflft~ 11 .. " ~of.I,. r 19...q8. Redcsrrtpunn or &1Uffw n1ui l.;a\·or-1n.
UK during J9;r,co 1!179. /',/11/lr, \ ei.rlMr; /011mttl. 13. 103-IUG 1901 a< TT1ctl1·na ,q11/\ltblhom. ~chein 19Sas, /'t11t1sl1ol,,g;· l<Mr,11rh 8>.
51 ~mK1M,1, \.\',\\ .. 111<)~. The tNurmcm or &."I.JUlne b;1b\!~0:-1s.. Jtmmnf of ri:t ,)67 ·175.
~·\m,•ffcm, \'ilcri11t11)' .UeiUu,1 A.Uorltu/(m, I SS. 45;-t6t. 7,1 M111uu.u,s, tt.. :,,<:1111, r 1,. \nl(.,t \\,P, 1980. I :ght .m<,l de-ccron
52 t:11:,,.:a;:,v.,,. r;,. 19tH ~pom1.ohtn-.1~1bilate vor. &1111."SJn 1YJU: .1u~ mrc-roi-.ec,pt.c ...,ud~- QO dcv~lnpm\"1H,tl ... i.i~c· of &:,JA~«tt<(Ulb within th~
11,wtummu a(l(lfQ/lc:m: a,wmllrum und RltlplcrpJ:allo r.,raruc,u. gu1 of1h" tick D..~rnm1 -,,morn·rfr·r,lams. Journrll of /1ora~i1t1lqg:,. t~
lmaugural OJS:i\:rtulum ;au,."I. dcm ln\tiiu, fur P-.in.1.1iiok,Wt."d,·r 2?0-?28.
llor!,rt<bchon Horh«:huk liar111owr. t\-1',, 73 ,11'f1~ r. LJ t'llJJ. Untc-hu1.:hun1t.tn UbN· dfc Br,tuchb:irk.t!'h \·on KUR
l,.,l J,;SO\\U !\o, t>1P. 1ft. ~\l'l'MM'1 R. Loe;~, ._TIIU H.. n.. lll~XA.Otlt, ..,t., ,._. IF\'I und CUZ..\ ~um n.s,:hwr.h drr 811ltt.~i""''"'III lnrt;.,11011 \I~" Ptrrdh-
Pl!.llft\"'"'~· t~r .1992- .\ntibod)· co.i t\.'i:bnthim.un 1nL~rc)1.oih: prcu-:,m lnnui;ural o;,,crouion iJU:,odc111 tn,titut {Ur\'crgh:lchcndc
cpicopc- ¼d~ntdlt~ hor,t~ int°t.'"C:t1.od ,,i1h &,t>t..,1« ,qm, /01,nml uJ fl,nfc~,, rmpcnmodi,Jn und P.tr.,i1ologr., d,,rTiurumllch,'11 Fal.'Ult•t do,
\/1<1oblolOR,·. ;JQ,.ll2?-3t2b. Unht.'Nittt MUn,'1-1\•n, t ~-t,;
SJ ~M)\\U!<- U,P. JR, PFJl..8\\l-\.~. LL. );.\Pfl'l>.\Mc. L"'- .!.. lJLS\,\~fA "G .• !'.,91, ~ Ml.\i\·AJW. 1 .\. 1\J5\ rlropta.. mos~ congCmtJl~ rhcz leCh("..ul Rto.:ut•l ,,~
1)~1~uon <,f l"qUint" anubod~1 10 ~,,.~,, 1.'t/ui momtnH.c prot,•ln, by n ,\ffti,"tlrie \·'1.·tt·rl1wJr-.., 1.27'. 340..3.i2.
rnonoclon•I 11n<tl>od~·ba><d compolftlvt lnhlhltlun ,·nlt) mc•linl.~~, 77 MOI r~tA~~ u.a. ,o IIUIORS, H.• SCHI IS. r:.. ftl llkUS, (j,. ,·nrr.1·. ~\ ,p, ~
1mmun;norbcm o,s;,y. Jmm111/ o/C//11/r,i/ \11 rabio/o,:i,. 29. 20¼-2058. ~\\'l'Ytr\KtH. r.•• 1983, fmt· ..1rutturt 01 1111/x'#a ,.,,,,; ~ernn. 1901 \\ ithm
ll k-'O\\ I ;._ot., fLC. ~V."-1~· • UJ\'D. R., 1()33. F.qulru: piropl~u·mOSl( (B;ab.csfchtJS> l)mph0c~1~ and et)'thnK",·11:,- or ho~: ,,n w ,,wo nnd m t•::rn•,utdy
of th~ Hab.•Jin adttdli type. El/tlil/4.~ l 1mtrfu. 3. 18-:.!:!. /ru,r1mi o/PamMttJitJ}{o', "9.111-120.
36 ~OCH~ R., 190,i Khod~i•m 1m l.....;,flgatlon).. <.npr Agnn,l111m1 Jo11mal. 24, 78 \IOI r,t\X'.\ u,c;, "\LUIUIOJ\.'.;, H .. -,cnu~. L, \'Cll{".f. \\ .r. '-' l·'RU:UJff•f"f, ,c.1..
l.l-1', 1933 lJl1ra~rrunur.il ,,udi on 1h,· dcy,.-lopmcm ol Bn/Je,,in ,·q111
43'1 ,ici '" two: Proto?.Oal diseases

~Cuccidi:a.: Piropl:3.,.mla:l In 1he sali\'Ory gl:mds or H;: \'t'"<"lnt tlck~Jnun,n/ 99 StMP.sox. c.1- 1rr.rn1m.A~1. \\·.w." n1~<.:. J.~, .. 1gG;. Compmh,·e
ofPro10.oology. 30, 218-22S. morpholog1~ fe.ttur<'>' of Bab,s/u cnixl/11 ,nd Bobesia <lful.Amc,iron
,,._p,, SROCJ,.'t~\-. IJ,W, "- fli\Rk-WlSt- Q.t.. 19:-; £\'aJU3lfOn Qf
~ }fOJtl,..\JU\. /tJur,uzl of\ t•:.·nntJ')' Stttnr~. 28, 1693--1097
1he1ndlrect 1luorescen1 antibody 1e,11or 8aliesi11 m(l,'t>r and T//ril,rin 100 T,\'i LOlt. w.M .• 1972, Cht.!mothernp)"of equinc pir'oplasn1osts. Pt{'1Ct.>ttlin-~
muutns in en,atn The t i!wnnrr1:y Ru.ard, 100• .;,;i;-i87. 0/1/ti! rl1itd /11:4.•rmutonn.l (:Onfr11•n(C ou Equimt /nfi"trfous Di.sta.set.
ao \\'.O .. 1956. <l•1>11lca1io11, tran,,mluion and biology ul plropl:i.sm,
;s'Effl:. Paris, s. M4GCI. 8"5.<-L
of dom~uc animab.. Am1als n/ llm ,\tw \'Ork .irademyofSc1~11w, M, 101 f,\\1.0R, W.)j., ffkY,,.,"f. J,E..A.."1:l)lJtS(')N, r,H.. & \\1u..EM, ~.M.~•• 196-9, Equine
S6-lll pir<>p1Mmoii) ln th~ Unitt-d sra.tcs-- a fl'\iC'.\·, Jounral oftli•• .'4.mtrltan
81 'CU"' r •.. 189,. Sp{'Cific (cv<,r< of rtt31art:il origin 1n l.'<!ulncs. l 'e1,ri11a')' \ ~1c'rinnl'} )1,Ylitttl .J.Jsocintiott, 15~j 91S.-Sl9.
/our,wl. 39, 40i-l05. 102. T'l::.'"TU, "·''· N FR.)£nHoff. s;.r., 1986. SC'rod.1ngnosis of 1!xptrtnrnm'11 and
8'2: strn'.,11, <.,.u.r. 1. srmr:sa.Asn. c; .. 1910. Oi~ P.1.ras!um d('s natun\l &1W.sfr> ,·,Jlli ,1ndBabc.(iff rolwlli infeccton.$. \ ·,mrrina.,:.·
Pfcrd•pi,opla.,niosc resp. der ' Slllut)• rc,·tt'. ~nrratbt,m/fir Pt1fl<<ltolol(>', 20. ;9•61.
BaJ:wriologii', Parasiu:nkwutP wul J11fek:ton.d:,1a11kh.e,1t•11 AbttiJung I. 103 TIIEJ.LER, 1\,. mo, Ole Pierde-Malarta. :)r/m~t:er Arrhivp,r TlrrhrWmntk.,
56. 5'?4-S2S, ~3. ~"3-280.
81 ~UTl-'U, C.H,I•, & S11'tlCK1.ASO, C.. t(Jl2. Qr~ the OC'C'U!ftDCC" Of tWO Sp«'1C'5
io, nnu.F.a. , .. 1902. Equine malaria /or1m11t ofCompo,01/i,· Pmlro/og.·m1d
oi parJl$itCS In equine p1ropl11s111os,s or bll!\U} fever. Paraslrolog,·. S. T/1ompe,11irs 15. 40-~5.
&o-96.
10; 111£11.Ut, ., .. 1905, furth<r not<:> on plropl..,mosh of the hone. mule and
8~ OG.-\W,\, ~ ).'t)J11\\"AStll. x., ICO"ltUU~. "·· }1U)i..\t, , .• 198; 80\.ine
donkey, /ouma/ ofC:On:pamrfr~ />01//0/o~·and Tliem/Hl:,trc,, I B,
p<»;\J)artuat'nc hamnog!obinacnua: Hypophosphat:st-m1,1 ;md mcm1Q01ic
229-235.
dl;ordor ln 1ed blood (ells. ·\merfcu,r /011r,1al nf1·,w1nar, H-rd,, 48.
1300-UOJ. 1()0 TU.EU.fR, ..... 19®. Tran~mf~Jan of equlm.• ptropla:,:no~~ br tic-~ In SUuth
Mnra Jout,:al ~/Compmo,/1,v, Jlmlu1IOl/)•n11II T/rmt/J<J/llic.<. 19. :!113-292.
as ()1.J\,FA. J\,. SURl"(')N, l,f", +.· GUTHl,Uf"i. i\.j •• tS9;'• .\n epl1-..oologfc:d ,,udy (1(
w:m~c In 1.ho1oujlhbred roc~hOl"S~ In Gouieng, South Mrica.Jo11m11/ nf 107 TIW.LER, r. .. 195<1. 7.uologicahw,·,,y of the Union ol Sou1l1.~lrlcn Tick
1haSo111h Afrl,m1 v,,1cri1tc1J}'A.sSO<t«1ion. 68. 115-12!), ,ul'\'<), port V. Di>trlbu?ion ofRhlµlrepl1'1ltt.1rtwctl. the red tick.
A6 P>.NDrT, s,11• ,...._,~GH, ft,, 1982. Babe-sin ~qui infoction an ~tM\nrashtrn: Ondt!rste.Pf)(Jrt Joi,mal of Vttirln.myS</f·nc.., nnd .-tntmnl JrrdtJJ'flj', 24
Pra'l\lcn<e and dlnicopatho!oi;Jc•I chnngoslnduu-J b)' 33-36.
splonec1omlzalion In clonkC)'$. /111tl1111 Jo,mrnl of' 'tl(riru,ry .\/tdlc/11c, 2. 108 ntEll.1\11, c .. 1931;. looloi;ical,urwy or 1ho Union ofSou,h Mnca Tkk
~9. >\ti'\'<') -pt1n lX. Th.,dlmlburion ofllw 1hr<.'<·S0u,hAfr1am
8; POS~f..TI. c...s•. 1988. Veterinary lh~!'urch lnstim1e. Ondt'tstl•poon. 1l>'1llomma, or honip001s. 0111/erstfpt>Orr Joumal ofV,1<>r/11ory Rl!St'arc/1,
lJnpubli<hed data. 2,. ~9-269.

88 ~.,rrr. f->," .\M&Ao~,o• .R.t .• 1989

O~.i\ probes fordeu.-cuon 01 Bahtsin lt>9 \'"-~ ,\M,ml-. ...,,.. UU\1ER. c..e:., tuivt-M. 1.. 198;, Gtd,,u.,nhttcmie
¥"qm ••\lDl«utur,md BuX'hcmical Porr,srtulag)', 34, rs-;'8, ln1ra,·a,,ul~cc ,rnlling In perdt ·n v,.,,log ,..,1 $<>,g•vrul, '""""'' 0/111~
$01,1!, ,!fr/can ieNn11ary •.\st()(/11Tio11. 58. 100-112.
8~ PURCtl-A.~1. H.), .• 19;4. Pirop1n.c.mo~t in kt.al at binh. n1~ \,~11.'rinmy
RcC'Ord, 59, +19, uo \'A., 1n,i:ao~, , .• 1987-1998. ~ICdic.al Unh,~r.sltyorSoudwrn J\!rica,
90 PUASCU. ft.r. & 1or.i.at. LP., 1-.968. rhl• de\'elopmt"nt of ni,•il1·rla p1m,a fn 199(H998. Prl\Jte\'c1crinal') Pr:ictke. 16 t)alham Road.!(imbcrl~')',
the sa:Ji\'ru'\' ~un,h ofth~ rick Hl1i11h'i·µhalus11p1>tmll.culnrus. Su\lth Afrlco. UnpublL<hcd obs~n·otiOn"-
POrflJilolu;:,·. 58, 7.!.5-732,. Ht \.'AS HUJt.01:--:. f,. 1996 EqulnL· bab(loslosis in South Afrle3: a ,~pon ort\\'O
91 iUmu. <;.P•• 196.1 A compnnson of cquinf;' p1roplasmD$l~ m Somh Afn~ ~b. Et7ui11,, \/1:tdrznry.Edumtlo,,, & a,...s,
and ,he Unite<! S1a1es. Jou ma/ of1hr .4morieon \'i'1,rlna')' .\'ltt/11:nl u.: w.v.:-.:s,1, .,• .t 'H)klll~. o.o.. 19&7. l racnwlyttc tm:se.mlas. /n~ Rotos,o.'I., ~ .r...
,lssocwr1on. 145.1112·916. (td,). Cumm /Jr<Va/J)' ,n F.qul11c Ml'dlt/11,,. Phll.ldclphla· \\ 8. ~•under;
S2: Pl~TJC- M., 191$S, f.quinc bab~..-iosls :i.nd nypanosoml:s.:.i-,... /:in1 Compan)
Jmer11auon11I S)'mpoJium on Eqwuc Httt/mawlog, MichJ~n. Mlfhlgan u,,1 \\'LnrR,G. & FIUtnHon. -.;.r 197:. Prclimlnt11)' c>b~en·.adon" on 1hc
Stifle Unhtrshy.
ulrrostructur., of suppo5t'd $e:<11•I ~'·'S'"'
of Sabet/,; /Jlg,1111,;n
93 M)BERN. E.t>.• MO'liU101JS~ 1..J•• (';,\ISi.ft, ,.H.&· Mt.D,"11-:.L. "·'·· 1962.. (;qmnc tPiroplosm,o,. Y.1:lur/1riJift1rParnsitr11k11111/q, Sl. 83-!12.
pfroplasmn'Si<./aurn"J o/rhe J\mtriton l/c1,1rfnal')' .\lrditr1J .~/nrion.
11~ \\TILA~J;>. c., igS-&.. Spcch,..,-spcdfle ,~\radfagnuft1i: of cquin1.1 plropl:ism:?
UI. !323-1329.
Infection.. b) mc>nl of complomi,nt lb-;ition 1~1 (CFTI
9-t RUD1.JN~"t;A., ~'--"·~Sf,'U!.L.,tA,..;, A., JHIJ(. ~.F.. U\\'EX(-l{Uft. S,I , PIESM>.:-., I" hnmunofluur~n.t•c fFt\1 and en:cymc·linked immuncuorbonc a~foaJ
1979. lntra-trylhrocytfc: ·garnctoq,es· 01 8ab?.sia micrott and thC1r 1eUS;\I, V,ttr/11ary• P11rttSf10/o,r,•. 20, IJ.-18.
m1uur3uon In ttds. Owndlfm Journal ti/7.Dof.og.•, s;. -1:?ii-434
tt5 wr..,"\Os. c.~ .. 1926 PfotOUJOlO[G', tt ,\ftmunlfor .\U1dfcttl .\/tu.
9"5 Runt.r~~" M.A. 41 r~<·•m•. w.. am. Fdrmntfon o! mcru;c-0itt:) in Vnm,u,r/11r1, ,wd7.1><>/ogw,. Vol. II. l.1>nt!un: 8alll!~tt Tindl\l & Co.
mtra--eT)1.hro~1:ic Bahcs!JJ mferoti: An ultrutructura! siudy. C.mwdlnn
/oumal o/7.<l(J/ogy. 55, 91ll-933. 116 \"OtlSG. "-,s, x. PUA.'-:'CU. AJ:. l9i3-1 Obscrvotir,ns onBalx,sio ('l,u1 in the
satl,·azvgloml< o( RJ11pir,,µltnl1u ,i,w,L /J111/eli11 of Ep,:,001/c Dl$MS& of
96 S.U..\HMUUi\, f·.f.. GOl>StU:.-w. ~ •• t-(RJUlJt, , .• fl\U~U.. r.• \'lU.\LU/\, G.;. tOJ\CC.
,lfrlrn. 21. 3,-383.
1.. 1981. 8"11,•,la rN11tr(Jllin) ~/fli y llnl/<'$1<> tabo,//1 •n cebra< Equus
.:,,bll1 . Oiagno,1lco niorfolaglto 1· wrologlt,1. ,qel'uro c:11bana dt Hi :,..,uGG. 1.Lk L\:'1:"I.. \",!1-1.. 199~. Efficary o( bup:rl'\"l.\Guone .\Sa 1herap.~uti<'
Ciendas VrttrlmmtlS. 2. t 71-176. ~nd Clcarlns •Rt~• of Rnv,,s/n Cl{fll QfEurc>pe:u, origin In horses
Amtriton Journal of\1l'1Rrltu1.ry Rl.'$i.''1trlt, 53, 1396--1399.
q; SCH.us. t.. ~ FRn m 10 1,J1, K.T.• 19;8. IJthtmikl'C>skopikhc Unr4..•r-:uchungeu
HS Z.Wt\*Gi\inll, ll,, JUSf. M,C~ & DE \\'Ml.. n. , .• 1993. Conunuous Ju ('Jt,(J
, Ober dt• Emw!cklung ,..,,, 11,el/er/a 111111ulmn (Dschunkowsl:y & Lulu.
l.90.1, in H_t't1lOmma anmolicum t'.trat1t11wn ;:oeh. 18-t~J. 0.'fts(ltrififilr C'..:.ltl\'tuon ofl'J}'throcytic ~<~~of &bnia equl. Paraaiu,to,:y Rt!Sauth.
Pt1rtU(tfll~'U1td111. ~ 28i-303. 81. 35;;..JSB.
9.3 sanJ.S, I?.-. kHthfJX, G.., \'OICT, \V,Jt. ~ l\vt:.\'C..\ltT1-I, (., 198J. &ti>tr.Jtn t(Jlti 1r9 7.WE\l.,.\kl"lt, 1 , rusi. ,,.c. • OJ \\'AA.L. I\. 1•• 1997, In :•!rm cuiti\·.:nton of
0.,,Vcrnn 1901 u.....1opnie111 an horses rmd in J1111phoe),e•ul1ure. Babes.in eqm: dctcc:uon of carrier animals and i$0ha.lion of i,ar~h""--s.
Tr<ll)t111mtfdI:in ,md fln.rasi101tJgl,:, 32. 2i3-22i. Ondurstepoort Jtmrrml o/Veterlutu)' ues..•nrth. &;. 51-56.
 27
Porcine babesiosis
Synonyms: Porcine plroplasmosis, Bnbe!iia rrawma1111i infection,
Babesia perroncitoi infection

D T DE WAAL

Introduction measuring 0,7 to 2 µmin diameter. A variery of other fom1s.

such as o,,al. quadrangular. and pyriform, also occur. These
Porcine babesiosis is a lick-transmitted, protozoa! disease vary in size from 1.2 to 2.6 µmin length and 0,i to 1.9 µm
caused by either Babesit11rawma1111i or Bnbesia perro11titoi in width. 1
and characterized b~, fever, anaemia, icterus and haemo- The life cydes of the organisms in the tick vector are as
globinur.ia. Bnbesia perronciroi does not occur in southern yei unknO\\'n.
Africa.
The disease was first reported in Russia in 1911 by De-
Epidemiology
memjew.'° According 10 Knuth and Du Toit the first reliable
description of the disease was given by Trautmann who Clinical cases of babesiosis caused by 8. rra111r11mwi have
studied the condition in Tanzania in 1914.7 The protozoan only been observed ill domestic pigs in southern Africa. The
concerned was lacer named Babesia 1rautma,111i.' In 1989. bushpig (Poramochoen,s porc11s) has been shown to be ca-
Cerruti identified another protozoan parasite in the red pable of harbouring the parasite for at least 24 days after ar-
blood cells of pigs in Italy which were exhibiting clinical tificial infection, but \\i thout apparent disease occurrlng. 1;
signs ofbabesiosis. and he named this parasite Babe.~ia per- The disease could potentially occur in those pans of
ronciroi.' Babesia rra111ma1111i also occurs ill pigs in Zaire9 southern Africa where the tick vecror(s) and bushpigs are
and :"Jigeria.3 • 14 :-rore recently. 8, perronciroi has been iden- found. Bushpigs are particularly associated with forests.
tified in pigs in Egyprs and Senegal. 19 thickets, riparian undercover. reed beds or hea\'y cover of
In southern Africa. porcine babesiosis was described in tall grass where there is water. 18 Agricultural developments
1948 in pigs near the Pongola River in the south-eastern re· have favoured them. In South Africa, bushpigs occur in the
gion of the Limpopo Province, South Africa. 6 and in Zimba- Limpopo Province, eastern parts of Mpumalanga. central
bwe.8 It was also reported in 1958 in the Soutpansberg and eastern KwaZulu-Natal. and along the coast in the East·
district in the Llmp.o po Pro\ince. 12 In all three outbreaks the ern Cii\pe Pro\1nce. They also occur in Zimbabwe. Botswana
causative organism was B. rrmuma1111i. and Mozambique south of the Zambezi Rh·er. They are
Although the disease is probably endemic in large parts widespread but do 1101 occur in the more arid west of Zim-
of southern Africa, it ls seldom reported and is not consid· bab\,•e, while in Botswana they are mainly confined 10 the
ered economically impona:nr. Okavango Swamps and adjacent river systems.18 They have
not been recorded in ="amibia.111
Infection is probably endemic in domestic pigs kept
Aetiology and life cycle
under free-ranging ~')'Stems of manageme111 in endemic
Babesin tra11rnumni is a large Babes/a found in the red blood areas. but clinical disease is rare. Although intensely fanned
cells. of Infected pigs. The merozoites (which are usually pigs are usually not exposed to the parasites, clitiical cases of
paired) measure 2.5-4 x 1.5-2 µm and are oval or pyriform in babesiosis may oc<!:ur when they are turned out 10 pasture,
shape. Single fonns. probably trophozoltes, are amoeboid or when infected ticks are introduced into pigsties ,ia gra!.s
or round. The host cell usually contains between one and used for bedding.
four parasites. Circums1amial evidence suggests that the cicks Boophilus
Babl/$ia perroncitoi is smaller than 8. rraurmnm1i. the rlecolomnlS' and Rhipicepha/11.s mranictts13 are the \'ectors of
majority of parasites being pleomorphtc. annuiar and 8. rraum1a1111i in Africa. bu1. it was only relati\'ely recently
435
that R. 1mi11man11i was succesfull) transmitted \\ith Rhipi- Babesit1 para~ites are demonstrated in red blood cells. Blood
cep/u1/11s sim11~-.2 and Rhipicevhalm 111m11ic11$. rn The nymph smears. preferably stained with Romano\·sJ..·y-type stains,
and adult progeny of experimentally infected remale are therefore essential. The percemage of parasitl;,;ecl
R. simus and R 111ra11icus ticks 1ransmiued the infection ro erythrocytes may be as high as 60 per cent.
pigs. ·n,e prepatent period varied from 6 10 15 d,1)<s after tick
infestation. Attempts at rra11~miuing 1he disease with 1-f,I'·
Differential diagnosis
alomma margi11ac11m. Hyalomma 1rc1m·mw11, tlyalommt1
mnrgi11arum rrifipes. R/1ipicephallls S(111gctf11e1111. Rhipi- Porcine babc,iosb should be differentiated from cpcrythro-
ceplwlus macultl/lts. Rhlpicephalus rosaic11s. Amblyomma ioonosls (yellow belly) and leptospirosis.
hebrae11m a1\d Dermace,uor silt1arum ha,·e failed. 10· 13 Of the two species of cperythrozoon which occur in
The pos-.ibility of mechanical transmission by haemato- pigs, on!) F.pery1/iro:;0011 suis is pathogenic. These rkketl·
phagou~ flies. such as tabanids and Sromo.,)~ spp .. has been sia-like parasites can easily be distinguished from Ba/,~sifl
suggested. :i spp. by blood smear e'.l:amination. Eperythrozoonosis is an
Although not proven, R. scmg11i11e11s. Oermaccmor rerirn- acute disease of fattenmg pigs characterir.cd b) anorexia
lmus and Hyalomma aegjplium have been suggested as and pyrcxia in the earl)' staie of 1he dbea~e. and by
vectors of B. pen-o,1c/1oi In Europe.' anaemia nnd lctcrus in the later stages. ll ha, a low
morbidit} and mortality. The distribution of porcine
Clinical signs and pathology babeslosis In ,outhrrn .-\frica is restricted ro ,1reas where
bushpigs occur.
Babesiosis caused by 8. 1rm1Cmm111i i.s a mild disease4 11 ,a Acme leptospirosis In pig-s Is manifested by septicaemia.
which. in endemic simations. usuall, only results in severe fc1·er. anorexia. haemolytic anaemia and ietcrtK .\ppro-
clinical signs 1vhe1\ che animals are uncltr5tres,. such as tha1 priatc labormory procedures miisr be taken in order to
caused by malnutrition or helmintl\ infestalion5. Healthy exclude it.
adult pigs expc>sed to infection for the first time. howe1t'r,
may contracl a severe or even fatal disen,e.
Contr ol
The disease is less severe in younger pigs than in adults.
Clinical signs indude listlessness. fon?r. anorexia and Paremeral adminibtnnion of diminazene leads to a rapid re-
anaemia. Jcwrus and haemogloblnu.ria may appear during gression of dinicill signs. ~ Treatment with phenam,dine
the later stages of the disease and abortions may result. isothjonate i$ effective against both Bobe;ia .spp.. while uy-
Xecrc,psy findings include anaemia, iCtC'ru,-, a ~light in- pan blue b. onl) effeetive against 8. 1m11mu111T1i.
crease in volume of the lluids in the body ca1ities. pulmo- It seems that animal~ \,:hich have recovered from the di~-
nary congestion and oedema, petechiae on the serous ease ,,;11 ha,e a life-long immunity. but there is a chance
mcmbrant,s, epi- and endocarclial haemorrhag~. spleno- that relapse~ mar orcur. 9 It has been claimed that B. 1mur-
mcgaly. and hC'patomegaly with pigmentation. mm111f can per5ist in pigs for up to eight months after
infection.1"
Control oi the di~ea~e should be aimed at tick control a~
Diagnosis
well as the prcvemion of contact between domestic pigs and
The history and clinical signs may raise suspicion of porcine bush pigs.
babesio~;,, but a specific diagnosi~ can only be made when No rnccine b available.

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 Porcine babesiosis 437

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Powmo,tlw,·ms 110n.·uJ.. ,,ws.."Jwm1. l..lnnbc1.g. 10 mr~cuon \\itb BuMzo
of dumi:-;t:c ;,nim4l., .•"''''u,Jso/tlll',Vt'W )'ork .~rnnrmyofSrumc-r. 64
trnututmwi Jo:mu.rl oft/ll' :Wm Tr Afrfc·n.n \'t•ft!ritf(lf'.I' .\l,~lit11l ,.\.<AA':a1Jon,
S6-ll 1.
31.,151-15)
a.i moo,. t.o., l~:"6. Dloud pann,ites. ol loc31 Pl~ in lbad1U1. Tr1Jp;,c:.al Ammat il3 H)SJ. 11:t< Miwmwl.s ;,J 1/w ~urhtm A/rlrlll: SuiJr.-gicn~
,~m 1-11- n,, ft.It..,,
llrultl1 nud l'rodtt(flO,,, u.. ,u. Ptcu,n.i. PuhH\h...,J h}' 1h1t Unin•Nt) tJI PrNorfa.
IS 1'1:.CU ,: \ .. 10 MU1.10, F ... ftl,\ '-"Ul)JI o, r;., 1!).l,11.
Efflcaci~ W!l '$erem1' J9 \TKC:R-U'l'Sf J, tr l'\1\1;,I, f{., 19-81. 00'\t'l'\';ltmn d'une CfU)'.()OUt, di!'
111.~la 0Jt1r c,h•Hu p1ropi~,mo..-1 suh,~ da P1ropltl.$ma :rurumannl ed~ b.ib1.-•,fo,c: p<>rc,nt o 8(,1,,,,.u, J)('rtmtCJIQ1, Cerruti. 1939 au Scneg.if
Babtsfdltt p.lrro,itiltJr. \f!terinnria ltnluum, q_ 611-616. .•\mu,(,;., d.- ta "I~ ,h..; BdF.i!•fl." ,\NdrrJm,"' Tmµfaif;:. 61. 1z.;...1. l,
 28
Ovine babesiosis
Synonyms: Babesia 011is infection. Babesin motas/ infection

l YER U HAM AND A HADAN J

Introduction che more pathogenic. It appears that some isolates of 8. mo-

1asi are infective for sheep and goars while orhers only infect
Ovine babesiosis is a tick-borne disease caused by the intra· sheep.6 Three other Babesia specie~ have been reported to
ery1.hro~1ic proto1,0a11 parasites Babe.sin 01,i! and Babesia occur in sheep- B. !!ergemi.90 B.foliarn73 and B. crassa35 (in
morasi.62 The acure rorm or the disease is characterized by Algeria. India and !ran respectively). These ha1·e a restricted
fever. progressive anaemia. ictems and bilirubinaemia. local importance and their q11idiry is still debated.
Haemoglobinuria is rare bm may occur in the late stage or
the disease. Pregnant animals may abon. Susceptible nocks
Aetiology and life cycle
may suffer a high rate of mqrtality. Recovered animals that
are latently infected are usualli•immune for a cenain period. 8(1besi<J ovis Is a small parasite of 1,0 to 2,5 µmin length. The
There is no cross-immunity between the two parasites. majority of parasites are oval or pyriform. tending to occur
Ovine babesiosis caused by 8. ouis is of co11siderable at the periphery of the erythrocyres.3 2- s.. g; The angle be·
economic importance in the southern pan of the Pa.lae- tween the paired p)'Tiform parasites is obmse. Cross-forms
arctic zone (Mediterranean basin. l3alkans. Kazakhstan. are uncommon93 (figure 28.J). Babesia crassa. described in
Kirghizstan, Turkmenistan. Iraq and Iran) 01·erlapping the iran. 35 has a similar morphology.
distribution area of the vector tick. Rhipicephallt$ b11m1. M Bnbesit1 morasi is a relatively large parasite measuring 2,2
The pre~ence of B. ouis in sheep in Madagascar86 is ques· to 3,8 µm in length by about 2.0 µm in \,~dlh. It commonly
tionable. appears as a pair of pear-shaped organisms. separated by an
Rhipiq1pltalus burs" is widely dist:ributed between lhe acute angle, occupying rwo-thirds or more of the affected
31 and 45° parallels nonh, including the :\lediterranean erythrocyte.' k7 Ring. oval, elongated and budding forms
basin. 3a. 40 · <8. so. 79 · 92 the Balkans. 22 · 61 • 69 southern former are less common. 8abesin motasi often possesses a double
USSR. 66 lraqH and lran:11 II has not been found in the nucleus. which is invari11bly single in 8. ovis. The intra-
southern hemisphere. erythrocytic stages include single rropozoiles. dividing
The geographical distribution of 8. morasi is very wide forms and paired merozoites. The fine structure of the
and includes western a nd northern Europe. the ;\1editerra- tropozoites and merozoi1es are apparently similar to t.hose
nean basin.Asia (Turkey, India, Chilia) and Africa (Somalia, described. for other species of Babesia.26
Nigeria, i\ ladagascar and southern Africa) and is transmit· Rhiµicepha/us b11rsn, a two-hose tick, 1°0 is the only
ted probably by H<1emaphysalis spp. ticks. Bahesia momsi known vector of 8. ouis. 97 Haemaphysalis puncrma, a three-
was first reported !n southern Africa in 1902,42 and sub· host tick. is the only proven vector of B. motasl in Europe7
sequemly oVine ba besiosis was diagnosed in other c.oumries while H. bi;pinosa and H. imermedia are the only proven
in the African continent (Somali~. !l:igeria, Madagas- ones in tndia. 3• 51
=J.18· 24 ·47.:;.i;. 7u,86 s1nce the reportofHurcheon and Robert· ;\larkov (c!lcd by Erashov 21 and Warnecke~ reported
son in 1902.42 there are no other records on the occurrence on lhe transmission of B. momsiby E-Jaemaphysali..~orophila.
of ovine babesiosis in sourhem Africa. Babesia morasi causes a a three-host tick. The possibility chat R. bursa may be a ,·ec-
se-.·ere disease and mortality in sheep in A.,ia 11• st and Africa. 18 tor or 8. morasiis e>."lremely doubtfu!.87 0pasina68 suggested
!tis regarded as a benign parasite in western Europe.19• 49. 67• 87 tl)atAmhlyomma uariegawm. a three-host tick, is a vector of
bur occasionally causes anaemia ill lambs. 1s B. morasi in goats in Nigeria.
Babesiosis caused by 8. 011is and 8. motll$i affects all Bnbesin 0111"$_ and 8. morasi develop only in erythroqTtes
breeds of sheep and. more rarely. goats. with B. ouis being and multiply by binary fission into merozoires (merogony).
438
 Ovlne babesiosis 439

Figure 28.1 Cross-form of

8.3bes1a ovis in a blood smear

The life cycle of B. ovis in 1he vector R. bursa ha:; been 11lor- result in a decrease in the degree of parashaemia and appar-
oughly studied I. 12• ?S. 27• 59• 60 and consists of gametogony em ancnuation of 1he parasite: this feature has been em-
(sexual C)'cle) in rhe gut of engorged female ticks, which re- ployed in the production of a blood-based vaccine ...- 88· 98
sults in the formation of oo!dnetes (zygo1es). A primary lntra-merine infection of the foetus in pregnant sheep by
schizogon)' (= sporogony) produces i.porokinetes. A number 8. ovis also occurs. 83
of sch.izogonic cycles occur in the gut and various tissues.
Schizoms have been observed in the haemolymph, eggs and
Epidetniology
salivary glands. During the fi nal sc:J1izogony in the salh•ary
glands, the infective forms (meroz.oires) are produced. The larvae, nymphs and adults of R. bursa. a two-host tick.
Young kinetes of B. ovisin haemolymph and eggs of R. IJU/'$a parasitize the same animal species. and their namral hosts
are round in shape, but subsequently become elongated are wild sheep and goats.' 1 This tick is one of the most im-
and cigar-shaped (Figures 28.2 and 28.3). porrant species in endemkaUy infected coumries (Cyprus.
Transmission of 8. ovis by R. burs«'l9 and of 8. motasi by Syria, !sraei, the Balkans. Turkey)2• 11· 44 · 48· 69· n. ~. 98 and
H. puncmca is transovarial as well as transsiadial.7 12• 97 All favours a habitat characterized by rendzinas and terra rossa
instars of H. puncrma are capable of transmitting B. mocasi soils. low average precipimtion and a long, dry summer.39 lt
parasites. 7 has a strictly seasonal life cycle with one generation annu-
Adult R. bursa ticks become infected with B. ouis during a ally. Larvae are found on the pasture. and larvae and
blood meal. Transmission to a susceptible host occurs by all nymphs on the animals during the winter period (October to
instars of the succeeding generation. 97 Rhipicephaius bursa February) in the northern hemisphere, with a peak preva-
can be infected with B. ovis both alimemarily and venically. lence in No\·ember co December. The adult ticks first appear
Alimentary cransmlssion gives rise to a higher ra1e and in April. their numbers reaching a peak in May, and persist
degree of tick infection than that following vertical trans- in the field and on the animals until July.96
mission. Eggs infected \~ith B. ovis appear earlier in horizon- There is a striking seasonal incidence of O\'ine babesiosis
tally (alimentary) infected ticks than In venically infecred caused by B. 01,is. The disease QCcurs annually In R. bursa-
ones. 101 infested areas during the acti\'ity period of the adult ticks.~.
lnfecrion taies in ricks can be esrimaied b~ exami ning The prevalence of babesiosis reaches a peak in May, corre·
their haemolymph 7 to 12 days after their detachment, and sponding to a peak in the infestation rate of R. bursa on the
those of eggs 5 to lO days after the commencement of ovipo- an imals. It thereafter decreases and. by the end of July. dis•
sition.9s In the salivary gla:nds. infective particles can be ob- appears.2 · io. .:-i. 50, 6 1• 69· so. 9; Ovine babesiosfs occurs two
served and are transmirred to the host during a blood meal. weeks after the first appearance of the R. bursa adult ticks on
Babesia ouis infection in sheep has been observed 11 to pasture.9 4 The disease may reappear annual!)' in the same
l2 days post-infestation by the ti.ck vector.57 It is readily flocks in endemic areas. This epidemiological pattern differs
transmitted by inocula1ion of infected blood.98 Successive from the state of enzootic stability known to develop in bo-
serial passages of infected blood in splenect0mi1.ed lambs vine babesiosis.s.i Sporadic clinical cases of ovine babesiosis
-1-10 stcn~, mu: Protozonl di~ea«:s

Figure 28.2 Kinetes oi Ba/Jesia

ovis in a smear of ,he haemolymph
oi a t,tk

•lit

•
Figure 28.3 Kinetes of Babes1a
C'ns in a :1t<·egg smea·

may occur outside the main season or adult R b11rs,1 activity. regular e:1:po,ure 10 the parasite. and is influt'm.:e:d by clf-
concurrenlly with lite rnre appearance or few adult ticks. nn matk condition,. animal managemem, and dipping. It ap-
Relapses of chronic lmem cases due to stress factors. suth as pears 1hat en1.001ic siabilit) in ovine babesiosi< in R. bursa-
heat, food shonage and helminthosis. may ah,o occur in the Infested area.s 1s rare!} e-,tablished. In general. unstable
autumn months. erv.ootic situation:. accompanied b} outbrenks of O\inc
The inc-idence ofbabesiosis in 6· 10 12-momh-old sheep babesiosis depend to n large extent on tour facto~: tran,...
in their first grating ~eason may be almost double r l O per humance (animal~ are moved 10 endemic area,1 health
cent) that or ewes (5, 1 per ceno?~· <>; The average rate or management (tick control 3nd chemotherapy . espec-ially
t)arasitaemia in lambs and hoggets (i,5 per cem mar be uttfavourable climatic conditions, and chemoprophytaxis.
some\\'hat higher than in ewes (5.6 per ceml.93 These rate,; lntensiYe tick c,i111rol should be avoided to maintain
are rather low when compared 10 those recorded in endemic cnz<>Otic stability.""
a reas of bovine babesiosis. 16· 65 Recovered animals remain patcnc or lmem carriers of
It appear.; that a certain pcrcenmge of sheep are noT B. 01 •is for \.u)Wg periods. ·1be parasites persist in the blood of
exposed annually 10 infection and consequently remain sus- infected sheep fnriwoyears/ 1 and 1fcl:sfeedingon the animals
ceptible. Enzootic stability in bovine babesiosis depends on during this period may become infected. The presence: of
 Ovin~ habcsiosl$ -14 I

para$ilc,, in the circulating blood maintainl> a state of premu- In chc absence or non-o\ine susceptible host;,, the tick
t1it).78 might serve as a reservoir of 8. ouisP6 Babesia 01·is and
High anrlUal rates of seroconversion have been obsef\ed LI. motasi ha1·e been n'ported to occur in unusual hosts20 but
in both hogger:. and ewe~ (84.5 per cent and 89 per cent. re- the role of these. as a~ymptomatic carriers. ls unk110\,11.
spccti\'ely . The ·autumn-winter rise in ,crological reac-
tions observed in young animals might be cau:,ed by larval
Pathogenesis
and nymphal infe~1a1ion6.97
in young animals. immunity to babesiosis may be con- Upon infection ofa susceptible sheep with B. ouis or B. mo-
rcrrecl hr culosrral antibodies and t'"xposuw 10 prc-lmaginal tasi. muhiplication of the organisms occurs in erythrocytes
a11d/or adult R. burs(I ricks. Morbidity in nocks gra1.ing in with the development of a clinically detectable anaemia
endemic areas can be anribu1ed to the fact that only 75 per which results in hypoxia of tissues that may cause the dea1h
cent ot rhe colostrum samples of the hogger~ and u..S per of the hose. Organs. panicularl~· those with high metabolic
cem oi tho~e of the ewe, may contain maternal antibodies activit). such as the lil·er and kidneys. r. are the mos1 se-
to 8. Ql'i$, and these may per$ist for onh three months in vcrC'ly affected in 01·inc bahesiosis.n ·,., 4 "· " 5 q1
lambs.JI 9 : ,is compared co the longer period of resistance Severe anaemia may occur in sheep harbouring a 101,
known in 8. bo1•is in caule. 75 It 1herefori- appear~ that l.imbs 1>11~icaemia, which implies that addirional factors are in-
must be exposed to B. m•is infection up to the age of three voh•cd in ii« pa1hogenesi,. Erythrophagocyto,is of both
months in order to maintain an immune state. Lambs in• parasiriied and non-parasitized el.)1hroc}'t<..'S has been ob·
fe:;,ed with R. bursa lar\lae may develop amihod~ titre, after served in case, of 8 o,,;s infection. A similar finding has been
a ro2la1ively long post-infest.uion period 139 days on average) recorded in Infection, \\ith other species of Babesfa. 3; ••1
and ma, suffer from only mild clini~·al renrtions when sub- The anaemia follo\\ing il1foctions with Babesia spp. is
sequemlr challenged 11ith infected R bursa adult Llck,. 91 ron~idered 10 be caused by:
The delay in the serological reaction and the fact that only 85 • mechanical damage of red blood cells by im'ading organ-
per cem of the lambs may become serologically po6ith·e are is1ns;
probably due to the lo\\' 8. oui:, infection rate o( the pre· • au1oimmune phenomena: and
imaginal stages of the tick.9~ • adsorption of ;mrigens omo nom1al erythrocytes fol-
fhe relationship beLween antibudy levels and the degree of lowed by the formation or immu ne ~omp!exes and thr
immunity to babe<iosis is 1101 clear. Serologically po<itll'e ani- aclivation or the cl.i,$ical complement pa1hway resulting
mals. when challenged. may develop clinical babcsiosis. A in their destruction."
similar ,JtuaLion has been reported in bo1ine babcsiosis.13· 29
In .iddition to the climatic and husband!") facto~ mentioned The pathogenic mechanisms or B. 011is are broadly similar
above. the precarious enzoollc stability in 01·ine babesiosis to lhOSI! of B. l101•i;-, pro,•oking lesions characteristic of a
caused b}' 8. ovismight be due to the fact that the actil·eperiod hypovolaemic and cndotoxic shock proces~. The lauer b
of R. lmr.<a adults is short, ~ulting in low levels of colostral .in- probably the rt!,uh of a massive rdease ol catabolite, from
tibodie.~. a,; well as the short lifespan of these antibodies in the lysis of erytliroc)'ti:S and necrosis of li~sues.3'3 fhe le·
l.imbs. The transmission of B. 01!is tw lhe pre-imaginal stages sions observed generttlly arise from 1•ascular aherarions.
d.uring the aummn and 1\inter months, causing <eroconver- ~uch as \'asodilaiion and rnscular ~Iasis. leading Lo hypoxia
sion. might play a major role in pre-immunizing and boosting of the tissues.J~ \pan from the changes in the brain. other
the immunity in shc,ep against infections which occur during findings are similar lO those reported in 8. boz,is mfecrion
ih~· spring to ,ummer mon ths.97 of canleY 1
01ine babesiosis caused by 8. mowsi has not been studied
in detall. IL has been suggeswd that :,;igerian sheep are resis-
Clinical signs
1am to infection \,~th 8. morasi::• Furthermore, C1?rtain iso-
lates of B. morasi might be of lower pathogenicitJ. while Ovine babesiosis. In general. is an acute disease particularly
others are knmm co cause severe disease and e\·en death il1 in susceptible animals. :'\•1orbidityand monality due 10 ovine
sheep in lndias• and In Somalla. 18 It ha~ been proposed that babt>siosis in flocks in rndemic area~ occur annually. Balle·
the virulence or different Isolates of B. morasi and the se,·eriry si(I or•is infec1ions may cause 10 per ccm morbidity with
ofth!' disease might begrt>ntly affecred by the tickvector. 51 • 16 Z per cent mortahrr.93 but a monalirv rate as high as 25 per
Ahheugh 8. momsima,· be a significant pathogen on Ill> own. cem has been reponed.-l
it ~eems more probable that disease in cenain regions ma~ be The clinical signs of o\1ne babesiosis caused b~ either 8.
complicared by ocher tick-home pathogens. such as Theile.rin 01,i.~or 8. mo1as/arcsimilor. In gencml. tliesigns obserl'cd are
,pp.. Eltrlfcliia spp ..~2 and Nairobi sheep disease ,1rus. 18 typk-al of those due 10 an anaemia. An Incubation period, gen,
In goat$. clinical babesiosls cases are rare.''1 It <eems that erall)' ranging from ~e\'en 10 nine days. is followed by a febrile
the local breeds posses, a certain dcgrcl' or natural resisl period of several day-s dunng which inappe1enri.> and depres-
ance w babesiosis. sion are t?\idem. The febrile resporu.t· in babt.'Slosis caused
442 ~=io.~ 1wo: Protozoa! diseases

by 8. 011is precedes a de1ec1able parasitaemia by a1 least within a few hours of death. thus making their morphologi-
2.J hours.99 cal idemificauon impossible. Thin blood smean; from lhe
Initially, visible mucous membranes are reddened a.nd peripheral circulacion. and impression smears of the spleen.
the urine is normal in colour. bul as the disease progresses liver and kidner$ should be made.
visible mucous membranes become pale and lcreric. Hae- A variety of serological wsts has been used 10 demonsrrate
moglobinuriu is uncommon. Constipation may occur. a11d thepresenceofantibodiesto B. ovisandB. morasi. 517• 13 31 45-
7
faecal pellets may co111ai11 blood and be co\·ered by mucus. 49· 63• o. 8 • · 9\1 Cross-reactions have recently been reponed
Respiration and pulse rates are increased. Weakness and within the genus Bnbesia. A high correlation ts found between
p rosualion develop together with muscular uemors, lachry- B. morrui and B.crassa, and a lower one between these t\\'O and
mation and salivation. The parasilaemia in 8. ovis infections B. ouis. 63 ~o croS!.-reactions have been found between 8. Ol'is
is usually low (less than I per centJ.99 antigens and B. mot(Jsf, B. cras,;n or 111eilerin ovis.~ Serological
Death may occur at any time after the onset of the clini- tests are used mainly in epidemiological sun·e}"S. The indirec1
cal signs depending on the virulence of the 8a/Je.1in strain immunofluorescence test (IFATJ is one of the simplest. most
and the susceptibility of the animal.' sensitJve and effecth·e techniques used for thls purpose. How-
ever. as in the case with other serological techniques. the
specificity of the method is nol always satisfr.ictory.~3
Pathology
Infection with either B. ovis or B. mo1asi leads ro a macro-
Differential diagnosis
C}'lic hypochromic anaemia which coincides ,~ith the peak
ofparasltaemia. Rericulocytosis. poik.ilocytosisand anisocy- Ovine babesiosis should be d ifferentiated from infections
tosis. together \,1th hypochromasia. indicate that haemopo- caused br other tick-borne blood parasites such as Ann-
etic centres anempt to compensate for the anaemia. There p/(lsmn 01,is, Theileri(I ovis. C,•ioeceres phagocyrop/lila and
i-s an initial leukocytosis. largely due lO neutrophilia,6 and J;peryrhro.0011 011is. Copper poisoning resulting in an acute
increased total and conjugated bilirubin levels. haemolytic crisis with icrerus. should also be considered.
The clinico-pa1hological findings in babesiosis due to
B. moiasi infection are considerabl} milder than those
Control
observed in 8. oi>is infections.6
Affected animals show a decrease in the packed cell vol- The comrol ofbabesiosis in sheep is s imilar 10 that used for
ume (PC\/), which ranges berween 30 and 40 per cent. 99 the comroi of bovine babesiosis (see Chapter 25: Bovin e
There is an increase in plasma transaminases. particularly babesiosis) and is based <>n a combination of s1ra1egic cick
~-panate aminotransferase and alanine aminotransferase, control. vaccination. chemoprophylax!s. when necessary.
dehydrogenase, blood urea nllrogen and creatine, and a de- -and chemotherapy.
crease in sorbitol dehydrogenase. alkalir\e phosphatase. lmmun!zarion can be achieved by inoculation of a whole
total proleins and albumin.30· 56 ,\ proteinuria occurs in the blood vaccine containing auenuated parasites.~ 6- ••• 88
course of the disease.30 A11enuatlon i$ achieved by serial passage of the parasites In
ln experimental B. ovis infection in sheep33 · 99 e\idence ~plenectomized lambs. A crude exrracc of B. 011is parasites in
of anaemia, marked icterus. splenomeg-dly. a grossly en- Freund'~ complete adjU\-alll has been also used as a vac-
larged dark,brown degenerated liver, and nephrosis are cine.• There is. however. no 0\1nc babesiosis vaccine avail-
noced. Ecchymo1ic haemorrhages are present in the epicar- able commercially. An effective method for the control of
dium. endocardium and subserosa of the viscera. 1hedisease in endemic areas might be the exposure of sheep
Erythrocytes parasitized by B. o,,L~ have not been ob- 10 pre-imaginal s1agc:s of R. b11rsn in October 10 February
served 10 block capillaries in the brain, 9~ as recorded in B. before the main adult tick period in April lo July.
bo11is infections in carde. This explains the absence of neu- Animals suffering from clinical babesiosis sho1tld be
rological signs in acute ovine babesiosis caused by 8. 01,is. A treated \,ith the babesiacidal dnigs used in bo,1ne babesiosis.
disseminated in1ravasculat coagulalion syndrome has been Diminazene acemrare rserenil®) is used at a dosage tate of
obsen·ed in sheep suffering from B. oi,isinfection.33· 99 3 to 5 mg/kg body weight subcutaneously.8 1. ,i; Dim!nazene
can also be used in the chemoprophylaxis of ovine babesiosis
Diagnosis witl1 a "ithdrawal period as recommended b}· the ma.nufac-
turers.28 lmidocarb dipropionate (lmizol®> is used at a
Acme ovine babesiosis can be confirmed by microscopic ex- dosage rate of 1.2 mg/kg body weight inrramuscularly. Asec-
aminarion of stained blood smears made from capillary ond treaanem may be necessary in case of recrudescence of
blood. The parasitaemia in B. ovisinfection is. however, low. the infection.li• Amicarbalide iselhionate (Pirodia®, D!a.m-
Diagnosis of ovine babesiosis al necropsy ls possible ff de- pron"") at 5 mg/kg body weight subcutaneously has also been
composition is not 100 advanced. Babe-sin parasites deform used with success.117
 Ovine bnbe~io~I~ 443

References
ABR\.\-?0\, J.\•. OY,U.0:-,,.'(I\'. L..P ..-~"tEl'.,NOV.A. -.:.L 19:"3 The uJ~lnJC{Uft .23 J!tiR.1t1iR~ D., CA~LL\. ,. N CUTTUIRl!.Z. J,F.. l998. Scropt~ltn(r of Bal>tsin
and c!t...-vt-!opmtn; or Bt1bt1.sla ods in Utks and in :.he \\'3f'ffl•blooded host. ou!tln sheep In C:uulonl:1. nol'lheas1em Spain. n11,rim,n· Par.1stro/ox;.
Pmg,,'# i11 P,010~/ogy••~Mtmct orP•PNS read at the 4th ln,~mntional ';9, 27S-281.
Congrc,,1 of Proto:wololl)·, Ckrmont•Ferrnnd, S•p10mt><r 2-9. ISi'l. 2J ll(l~li.R$. c. • w1L n .. 196;. Blood parasuesin conic. ,hc~p and go.au m
3: ,\80-~Ht:HADA. '.\l ..."l.. MUWAlJ.A, M.)t. & Ti\\\r'f(D:. F., 1.988. 0,.1nt bobcsto~rs northern Xtgerln. Br<IIN/11 ofEpl.eoo1lolo11J• Dis(l{<m iJJ r,frlcn. t&.
Jn Jorduni3.n sbttp mistakenly dingnosed tt5 ph'\n1 por.sorunf,. Pt~Jfl1nt/n... 121-123.
\l,ter/,,ar; .\l,:dlrin•. 6. 235-238. 25 >R!nt>KOF>\ !(.T ,969, Uchunilaos.kopis~he Umen;ucl1w1gcn llb,-r die
3 ,\CHAH, ~o. « !liRIKA..'TIAu. G.s.. 1934. A note on Babomn morasi, \ftn~oo Entwid:lung ,..,n Bafx,s/a ovu (Plroplasm,deaJ In Rlrlpretplia/rc< bursa
(19:?tH m ~cop in Xysorc1wte, J,rdian Vt"rainnry)oumfll, 10. 2i0-2:73. (lsodoldeal. I, Die FJll\Vidlung in Wcib~d,en 7.e<ktn nllch der
4 \1..-\81\\' M.. OU1.GU~. ~. CltRCJ, H., \\'RIGtJT, 1.<i.. \\.U..tf5JlUHI D,J, • Roplction. Z.•iu(/,rlftffJr PnrasirMJ.:untle. 3Z. l!!l-21S.
GOOD<,"EM, ti.\ .• iga;. o,me babesiosi,;: Induction of a pro1cc1Jec lmmuni! "6 FRW>ttOfl', ,. , .. 1s81 Morphologk aspt>etsof B(,be,in la the tick. In:
rc,pon.,c with cn,dc <Xtr.t<'u.ofoi1hcr Ba!µisia 1/oiis or 8 01·i~. R.is<0r.h
ir. \'twr1nary·~em:~. 43. 401-402.
rumc . .., • ti;MUi.Jt. J "· itdi). Balwiic,.ds. New York; A~adem1c Pres.!..

s <LANI, ,\.I., ~()NMI>, I," 1'EflH.aR'I", J,\'., l98;. Serodiagno,,:, or Bhbtsih n FJUEDl:fOFf'. J. .l. & ~;\tm1.ao•• 1981, Tra.nsmi~fon o( 8..'llwslabr U(k. In
RJSi1C. ,1. ~ OWEA. f.J'. ttd, 1. Baut.rios!s. Ne-.·.- York: Academic Press..
mow: (WalC$J. 11i,i/r.rfa r,'N>mlira (l\'ale•J and Q.·tlJttltl411hago-
cy1opl11/aln!,:etlon in Sheep, Rtkur</1 in Vtter/,:a,;•Scfon,~. ,:3, 104-106, 28 <:ro•cu.«:u. 1.." TW'm:A. , .• 1966 Prevention of BnlH!si•l!n 1111£, •nd
s ALA......:t. A.t.tc Hl!.P.OERT. t.v.. 1968. The p3th()logy or Bnbl!Sfh mown {\Vnlt:.,l Baw~ln nrornsf m(ce1lo.n,ln ,hctp by lnje,:1ion o!Berenll L11crnrifc
in!ec1Jon lnsht"<lp. \l•terim"y Paras/10/0,0•, ?,, zos-220. l11stitut11lut d<' C~n-etan Vt11.crfn(l~ si 8iQprc:paratcPas.ttur. 5. ~2';' -43,8

1 ,u..As-1, A.f. ~ HER.81!1IT, r,v., \988.. 1'he morphon1,1rics o( lt(ilwsm motasi :!9 eoLDMM(, ;\1."' R01'f..,1UiR<1, ,.$... 19;4 hnmun<,OuoreSc:cncc .1otud1b of th4:
(Wales) on.d its transmission by H<1emaphysalls 1mnc,arn (Cancitrini ~mall 8ub<!li1(! 'l'We$ of canle fmm dUTorent_geograph!cul area..
and FllJU•ltO· 1877110.shecp. Ve1'rb11<ry Parn,,10/ogy. 30, 87·95. Tk~,v,h In Vtttrlnmy Sl.'lunu. 16, 351-35-h
8 A~Gi1.0,~, T., PJ!ftl0\'1C. -,.. &: To.,10(:0\'A.. 196a, Piroµl~moJ.f, in ~beep 3n lt.W£1A. M.A. IUJ!'o~. o .. ~.\\'AJUU:n:. 1• • HJ-MSA.~Ut'ZAODJUGUU. s.• 19-8;-..
in SR Macedonia, Vetennors41 G/a,nlk. 17, 86Hlf;; A11en,1lan, of 1he rtnol function In the nv!ne e.<J)<'riinen1ol b,1bl'si0>l<
9 e.uiu. ,,. 1892. L'erlologle d'une enzootl• des rnou101>.<, dN1on1m'ce
n,trd Mfdlttrrm:ttm Co,1fe.rvnteo11 Pamsltolot:,•. 2.;...2; A\J.b'U$1
carceag en Roumanlo. Compr,s Rcndus. MtUfcmltdts Sdm«J Porisl. Jemsalem. brae!
115. 35!1. Jl l~\8£L\. M-'<, IU!IM, D .. '<,\\"AJ\Rl'TI!, 1., 19~. :lntlbody .,,pcm.>< ,~d
10 IHDtSC'U, C roP0\'1L'"l. I. fl ~IIHI. s,...1968. Comribtnlc,n to thl!~tud) of 1he durouon of lawnt lnfcct,on in ~ht'ep follo11ing <.'Perlmemol ln/ocOon
oc.ologrof the ixoclids <m species) of the ~"llhal s,.,·a
paswre of the "ith 8<>/.lr.s/11 ov/J. I ',•rtrlr1arrParas/1ol"I/Y. 35. 1-10,
district of Giur¢u (Rumanfa). lucmr//eSl',r./11111/ S11/nriji<,e. lnsrtwrul 32 HA.kF.IA, ,1..,,., ~m:-u... D.• :;U,TO.C.J. ~,W,\RRcrt. 1.. 1990tt, lsr,Ja.tlon and
.-1gt01101>,i~ ,Wrolae /kJlmcu /C), 11. 311-3~5. ldentific3tfon or Bn1N>il101,fs In t:.,;11emadu1'3 (Spolnl, l',NT:/UJI')
11 BIL\S~\M t{A(}. P.. SUI\C....:DKA....... N.S,,& lV.MA(:H.A.:oJOk,\"k"O· \989. !,tudyon Pnra#ro/ogy. 35, 233-2;18,
outbr~k,ofbabosiosis inshc;,p In llndhm Pradesh. Jndilm V~tl!Tinal')• 33 RABEL\, "1..\., KEIS"A, o.. SA\'A~nrrr:. 1,, fll!:DO~UO, 6. a. H61l.~AXOE7,, :, 1990,
]Ullhllll, 66. 343-351.
)ilsiop•thological change, In ,hoep c.pcrimemullr inf,..:ted with
\~ KUSCHe:fl,v.. FR.UiPHQ>'F. J..'T., A: Et.•Al.U.WV. T.A.A.. 1988, Qu~ntllative Babes,a 01,,s. \'eunnnf)· Parn1itoley, 38. t-12.
c!l'Scnp11on or 1htde,elo11men1 of Bn/x>no o,11 ,n Rl11p1<epi:n/11s /111rsn 34 19,r. Hhtopathalogfcal change.sin
HAu.eun·A. ~1 . .,.,. 1•AuLO,·. ,,
(h,u·molymph. o,·•l)·, egg,). Pt1r()>irul1>gj• R,srarch. • ~. 331-33!1.
,plencctomiic<hhc<p lnfcc1cd \\ith 8nl>(>sin oi,/i. 1·mrlna,y Se/en,-.•. 14,
l3 CAJ.M'I\\ LL. ~ CGJtf.<',Oft. \\',, PA.RJ.'l!lt, R.J. -6. DArr.uc.su. R,I,. 1974. The 50-56.
lmmunh~· of cu,tlt ,o &,~a (l,gtntmnaf1e: drug sienll,uuon of
JS IW!Hf.\H·Ft;$11ARI.~. f •• u1u.,ar11C, G.. 1981. Bnbt,,12 CfllSlf(I n.5p.
infections or ,,ming dura1lon. A11Stralln1: 1,urinq,;,· /or.rnt1/. so.
6-11
(Sporowa. B~slld11l'i ofdoru~tlo~heep in Iran 'n:e lt11rint11)'
1.i c.........-snu~a. c., SA~t.A(.O, r... 1811. lntomoa.gli .icart tu1lianl. .-ttti /11irmo Quarur!)•, 3, 1--ll,
Vimc.'th d, 'S<feuu. uu,r.:.
("ti .l\m, 4, 69-206.
36 >LA.~KJl\1Mf.t..,"'lll-\M¥.J. 11.. 1~91 , Q\1n.c and c:aprinc babeslosr.i fn Iran:
15 Ck1HSTESSSOS, o. & THUXtCAflD, f... 1981.
BafJe..ti(I momslin sheep on the ttoatmunt with 1midocarb nr, l'ercrl11t1ry flecord, 119, 383-384.
i>lnnd of Go1lond In Sweden. Vi<1M111,ry PnrM/10/og;y. 9. 99·106.
37 HIU»U11A<r>r, n .. 1981. The organ and va·s cutar pathology of b3bt!<losl~
10 Cll.R.'<O\\', , .... 19n. Srud,cs on the cplzoorioloil)· ofbo,ine b~beslosis m 1,r. RISTIC, M," ~MllfR. r P (eds). Bnwifo1l1. Nr.wYork:Acodomk Pr~,.
11Qrthc.tstem ?\:ew South W.ates.Australirm Vtt"erin.t~I)· Journal. 49.
284-297
38 HOFOIA."',~x .. G•. HOKCJl~E», f ... SCllEI~. !i. C'&RH~, fC,C... 1971,, Seuonl11
ocaJrrc:nrc of t!W and plroplasms in domLloStk anim:t!s in thr 3.Siauc
I:' fJU7..(iU"• .\., \\'IIIGHT l,G.. WAl.-TI-SaVJIL. 0.f .. GALS. IUt. C.Q~OOU, I,\',
pro\;fnth or Turko:,. &,rl/nrr und .llur1el111erTlmlr:1/lcli,
"'"'<>IE.1,0., AIAB.V, " · ~ Cfillct, If., 1991, An FJJSA for 1h• diagnosis of
Wl><IIMS(/trift. 84. !02-156.
Ba/1,sla 01'1Slnfcc11on utlllzlngo synthc,rc, B-1b.•,rn wds der!\'e<l
:rn1igen. I ctmnm,· Pnrasirolog;•, 39. 225-231 39 tick,, /11: " TU)E 1.>:.11. («!.) Tc(t•®""
HOGStl!MI, II., 19;d. B10Jogr Ol

18 rnu.srt~. "·" .. l.9i3- The drs1rib111lon Md pre,·alonc" or :,/~1tobbhcep Dlµas~and rhw \'«rQr;-. Prl)(ecdlngs<>f an !mem•tlomtl r.on!ercncc
di;eo" ond other l!tk·borne Infection., ofshe<ip ~nd go>l' in Xonhem Univcrmy o( Edmburgh, C..orre for Trop,cnl Vewrinal} Medicine,
Sorn•lia. Tmplrnl ,tmmal /iooltlt11nd Prodmlmr, i, 29-34. Ea5rnm Bush, Ro~Un. M1dlo1hian. Ulo..
19 Ellie~.~. 19.;6. D!e Sthofplroplasmose In Dcutschland. Oi111x1~ .,o The epft)ennolugy or lick borno Crtmtan·COngo
H OOGSJR'<AI 11., 19,9

T/erlJr.tllrh,• l\'tx;lr,nschrift, 63, 161-162. hemorrhagic lever In Aslll. Europe and .lfrica./o«ma/ u/Mw./rol
Emomol(®·, 1S. 30i"-4 l7
20 1:XIGK, ,: .. fltlfl>HOfr:. 1:.1. & \\'lMM-1.AOlRllDIA, s.. 19&.4, Zur ,,irts.spedfllnl
\'OIi Ba/Jes/a »wtasiw,d Babllslo ovis U1ropl~smrdcal, 7.rlru:lmft /hr ;If IIOOGSl'RAM- 11, ~ ,·~to,;z, 11 .. 1980, Tick$ (l~odoldeul from "11d sheep and
Porasitcnkundt•. ~4.309-318. goo is in Iron and medical and ,·otcrlnat)' hnpl!callons Fi<k/ia,w:
2ool{®~Xi'"1l'S,..,nf!4. i-16,
:.?t t:k.\:,HOV. v,..... 1956. PamsltoJogyrmd Pnril...llir t)iJ«t{(':1 o/Ll1'<'1tc>ei:.
ls:oel Program for ScienlfficTmnslallons.1960, mmslotion of original ,~ IIUTC/tto,, o. /. llll~tiaTSO~. w .. 1902. ~1olariol ca1onhol (C\·tr of ,J,«'P
Ru..ian ediuon of 19>61. Tlrr V,r,.,mary F/m,11/, 14, 6:!9
Z2 FlJOlJt 7... IIIAUCUI.\Oi, C. •~Ufl.O.XfSCU, 1... 1.958. Thr ;(cir.) ofRunuuU:t. .,3 ~O<'.lf, u.. 1968. Elnls• kllnt:<ch• Bolundo%Ur &,b<>~fo, 01·is- lnfe~11on d<>
Qtl:()$/0,..,11,i,11 Parnsiro/ogi~. 5, 71-87, ~haf~. !n•ugural Ois.ertulo11. T1tramlkhe Hoduchule, Hannover.
.:4 ~C'llrUR, <• .,. HQFT<;\tA..,, G,. f ASllS.CHXl Jr,; .y \\;U't['\.Ulfl''ll.JI, 1 .. t'!lfi-;'. 1-9,0. Plropfo-;.ms of
t,,; Jl'.,P.\t.KH•omo:-. A MQ:;).ut.D. ,1. • Pf;.lltt !\ ,1
Srnd1('S lO\\;trds imowh.-dg(.• of lhe u..;~,;. found m .Syn,1. 7.1111flmf1 fl,r dom~w. ;1111m.1J... in the \l:.ccdonja rt.~1rm orCir,•«1.·. 3, Paop14S-m.:t 01
Trr1f)(:mu«ti:fn ruul f'11msimtr,grn. 18. J:'S-381 ~mq1J ru1n.nrnm, ~ 1.•1,•nmuy Pflmsirolog1•, 63. b:"-7-l,

.:s ,;,·unTov. ~. 19;6. Use m 1hc 11nnnmo1fuoresccnce n.~cuon m lhe 65 ""'"''· at u.• '\IU':'>1,t. Lt. l'&i.,.':t<AM, ru.•. lt'ICHf-1~, tt L l~5 .\\~'llnent of
dr.t,:.t1o<l" of Bal.w.1:i11 <111is inft:cilion. \'ut.rrinnmo .\fni{rsin~ki .Vm,ki. i3. onzoudc'1ahllit)· 10 tkk borne dbc.i,,e,. Wnrt,/ ·\11/mnl R,'tM~. M. 2-l-JI.
61J,t;, 66 l'O,'tER."rn, "·' 1q,o, fauna of 1hc, US.SR .\c,chnWll I\ \:o.1. l,odfd
.16 l.-\WJtI.~cr ,.., •. :99· f;(:)1,\'entionill \'na:!nt, for l1cj.,butn~ 1"kl-.. •bodidar, ••·\md,•,11,\•o/St:lt•11us \lo.k'VU USSR
h,tt•nuJp3.r:tiltCt dl~:i~l"S or .\heep .ind ,;n:u, Parr.sito!Ol(id. 39. 119-12 l . 6; ?UJl:~111.1-1.f t I WI ... n .. lltlL\t.\S. '.\I.A- k \'OU Sf•• I .... l~ftt. Jm·es.ns:.arion,.
on• &,/.,.,.1~ i,olat('J from Scuttishshccp l'oras/10/tr-0· 83.3~7-35l;.
r I H·f"l.\'\:t,;, , .. 1m. Ilck-bomtrdlS4.ra ...c-,; ol domt-.s.Ucantmaf~ ,n nonhem
XJ,:erfn. I. I ii:!olorit..".11 fl...,,it.•\v 1•12.4,. 19Gb rtvpfrnl •.tmm11l Httilth mu/ 68 ov,\,;.'""·'· t, .., .. l'83 Ctt•nfll'\!µiu1Tull'S.etml~ 1n(«oon ~( Jt~t'-. T1npie,1l
Jtrodur1101t~9, t..;-:• 151 \uimnt lli•,,lth ,m,I P11x/ucl:on. I~. 106,

.i8 t J Rlt"Ur. 1•..o .• ,\LT.\:0.:, Y,.C.\.\lrm 11. J.ll 61" Ef'i~\TIIIOU .. o.r ., 197.f- ·nc.:L.~ t,;i osW.\LI>, • . 19.19, Con Yup.q..la,ian 18alk~n 1id,, h~~oldcd. Ptlwsltn!og;,
'lxock>1dM!. old~rnc,1lc: a.nlm:1loti in C)·pn~. Bct/Wu'u ofF.nromO!OJ:fr.ol ·u , 2:r1-:!so,
R"'"'t1tcl1, 1w 53-63. 70 o.e1.u.., o ... l9 ..9, '."llud11.•-. tn ,he )trulogicn.l dl,tgnh~'-"' <,! 8i1l1tsta ,wb
19 1 f\'.'t..-', t)., HOU,~\~, ,1.M,. ~HNU I,. ft.[. 4 YO(hr,, l!..K., J!,81. ht\'t<'"tl~.tl(H'I
infl'<'tlnn In ,h,.p h, the lluor,:,i:cnt arnlbod)' 1,'(l\nlquo. Nntllk
~'tu,,m,., ,\ flkro/11\'"foji /)1•rgis,. 1l. i'0-83
on IJ,nMj'/a murr.sl t~lnce-d from\\ all.' .. R~:-.ti1f(/t i11 t'-•ttrilwry .'icl••nu.
,1. 239-243. -;, o:r.:.oc:., u., n, '"' 1 A. rK>(,flU, c., 1~g2..An _J.Jl\.''"'ll(Wlinn an !hC' rrJ:u.i ono!
SC<l'.f:()~t:il actl\"ll} of Rhf1;t,.,_.pfwlus Imm, (1~·1h!Ohfo:1: w::h IIIJAM1,1 o:#
so uf111sc.u• ..._., i-uMm, ,1,, 1970. Btolt.11th:~d .indtco1ogic:nl ,tud?c, .,,, 1h, ~...
lnfoctinn 1n •h«•p In th< \lonnaru region. Pt·m//1. I r1,rl11,r
oilhc-gcnera Boopillltu. Rhi11ifi•plml1ts"nd lf.,"!omuu1 in ~}'fi..i.. /11.
l/ikral>J;o/01/ lJ,·rii1s,, 14. H-52.
wltm. J.1r1..n. ted,i. Tlc:k-1,ome Di.~•VM•~ mut rlwu t n.·rnrs Procttdlng-ol
:\U lntcrnation:U Confcrcocc. Un1\'('r,l~· o!Ed1ob~rgh. U:n1rc-tor
fl: AAfl \." '\L\.ttt..\.,u. c..• ,966, 111; R~putt oj'FAOJOTI!. fmt•moUrmu.l

Tropic:;il\'eiorinnry \lcdlc11w. l!11<,un1 ll~<h, Rqslln \1ld101h1on tJ~. (.Jmfi•rt•rt,-c uf~ltt!t•f) D/JNISl•S._ fAO. lMtt'Hr 1966,
7J 1u,. 11.s. ., IV\CUU\ ,n1,H.1. lri .. 19 11, Obscrv;iuon~ on lktlxt.\!(; f,,llmn n. s;:,.
51 J,\O\~~Alll, M..~ •• JlCC.OL.LS.. Sllf\ \fe\"il. K, A ..:AG.\Ai\J.\. x,· .. ,9:;: /\11
ou1brcat.. or" b:1besmw,. in shc.'(•p and ~oo,... and Ii~ control .'.l_\'1ort
rrum 2: ~h(tCfl /1:dinn Jrmmnl of\ wurimll)' ,.,<'ivuu. l J 2!9--:!.·l~

/01mwJ nf,\llrieufruml &t'1/U'$. 8. 4·1 l-+13 ; ~ no,~su,., non a. '·". • \L• \\,\TIA.'n'll), 1 •• ,968 rid;.... i.,odo1dt:aJof
don:e~dr .1mm.:s.J,;, 111 Iraq: Ptin 2.. Summer inft":.'it.ttion'.\ jn tht"" hw~, nf
S2 :\HPH,W ,. J,.,.R.;.\., 1.966. ~mr o~n.·nnonton .Bnba.w mu:a.<i jnftttion m
I!Iii>. Korb:li;. n,wnm)'> ~nd :-a,il'l}O JQumnf n/ 1/11 1/,•lf/rnl
.shctp fn.:\ndhm Prndc:<h. Tttdftut ' JL·rinllf) Jo:,mal• .;). 76,S,-169.
\'t_
Emomot,,,r. ,'l. :?1-31 .
SJ ,v.t.GMAnH, 1.. c.u..us, u,,1 tcl'J>HA..,UL ~. 1957. Pathological proccssc,;1n iii no~&.. J.fl',J,4' COHR, t,..r .. l9i'O.: Ubenrngung und \'<:twcJld:mrr,on
Rnlli'!S:a r.nnu Jnfrctm1b. /.A!ftithrlfrJUr fmµcm,:.1df;l11 umf
K<itostr.U \'"' uml·n;,n Ba~w l)11wmi11a und t\1111plns11w mnrgfnnle
flnmslwlogir. 8 48:,--&t i
Amiknrpl~rn /t1h'('l1rt'lt ft,r tmf1('1Wh!tbzm 1111d J'nnL'l(Off>'~iP. 2:1.
.s~ !•!.\UO~f\, n,v. .._ ROS$, 1,.1,.. 1972. Thi! <:pf:t.(\Oliolo~cal filnof'. In 1he 101 H'
ct11uro1 ofbo\·lne ba.httbio,i~J"\Ustmflo,, \ 'l!Urlnnry/011mnl. ,18. 2'92-298.. 7(, TilO\bO\:, J,U & JL\LL, G,X .. l!tl.1 The nccummct- o( &Jbt-;.111 mnmSI
S5 ~,.,t,nn. o,,.,. & niPt.Ut u. o.u.. 198-t\. Spomanl."OU!> infet'tlon" nf litk·hornl~ \\"cn:on 19:!6, in ..,h,"C"J\ ln northern ~igt:ri11. ,,ith J diwu ..,io-n on the
cfa..,~1mannn 01 the pm:,pla!'m,., Joum(tl uJ ('rm11>0m1u·r- Prul.oiog,:, 4'.6,
blood p.u,1:sit{\'- tn ~ple:ncctoruit(.·d indiJ;l•nou~.sl!~p antl~al~ In
Jbldan. ~'iRt.oria. Rt.•t-·,w,t'l!l~l ,ti:'' .\li·tlrt•in«' wurl,mir.- l'1~"1 lm11r'trt\',
18'1-191.
.l'~ , ,
218-231 .
r,..(,1,1.~, ,. l \ .. l&ik) fmmunilAltiQn nf $1n,-ep a~:dn~ ov:n~ mthc,"lus1,
sa \\.Ut.,_o, . ,. \. &. .\DnAMO\', i.,• .• 1y;,o. tlt~uh~o! t\-.•ent)· )'f".lro. obi..etvJtion~ \ /'(('TUfcJt)'fll, ,\ (l>kOll', ;li• J J.
on rcp<-c1tcl11ifc C'\~l-..... of Babtt~:'a ut•U"in ~.; ~cnt'r;1tfon~ of Rhipl~1,lutlus -s -,.1,.1tc-,..~,. f .•• OC)X\rlt, "·· 1•,11H01, 1... l.N~"'fOQU.\l1fj, r ... l'tA:-.:n1t1F.U1>, f,,
J,una, Trudy l-'.u'to)';'ntlj;;,' l1Utiw111 JJ:.s,wrlmt'ftlnr1mi \.!Wrin11ri,. 38. ,~;. Ecu~1t.""' '-~Pt!llmrruaJellj, ,ur-lt."'i PiroJ,1,l~l'J'IL»-l'' R<;vlni:" u· .\lgerie.
5-1-1 (Rcfi t ·,·uri,mry r,ullttlu, 1gn, .;z. 4). ..\J11mh?1/w·un11Pmwur rf>:m~•.11. 72.1-7U,
:.~ ,ufCJ, ,... .KO~ I.A .• X'A\'Amtn'I., t." tMtll:U. >1. , .. 1995- ~.\pon,1c-tt1~ :1:I :'9 ~tkC.1''-l t.: .. 110:\:,llt', \_,V.\llNll,L.&!LL~ltQU.\tlll,f.. 1g.;;.hu.tdes1u1
e:onotJn1~nul 1:pidt-rniOhlJ;:.kO di! l,1 b.i.ht"lffh.is O\i?'IA &lbniti cm In <1t11, fc-s Piroplil.smoih Uo\'in-.~ Jnstlfut Pasteur d'JllJ:tvrit•, •\ltet
p1min1,·i3 de, 8-ad:lj-o..t St.·1o;m:\"t1lr1,di1 \ tt1immbfon ~pcrimrmnJ por 60 $HDtl\()\·, \L.> U•k.\Hlt \'. a~OX..~)U,l.., 19i6. \'°Ur\C\'OfpollilSHesb!ood
J\odtdo, \/ttlicinn Vt1rrittntl11. I:?. -lS0--:70. dht•.a"'i.: • rte;.. Hor,,~ Fl"\'e( hl dorllC'!tllC' nnim,U, In lotdJn. /!f)JU /01,maJ
;El ~HUJC. D.. 196.!. Eih Bc--ht.iy dcr K,•nncnli.dt•t \'t-randcn.inhtn ltl th,lt ur.d of\';•wri11ruy .'idn;a_ 13. :?'J-35.
dtm (.C\,.:t'hc.1n b(i,i den nn dt!r Virorl,1,mo~t- ctl.ranktcn "C'h~fcn ·\tm 8J ~lMll (;U, t:.,P., '\(·\'[,'\lf \' f.:- oii1QA.1 IC:.. S .. T956, l.4"' tt31h'Inf:nt dl.· J:a
l"._·ttrinurit,. &'ograd, 14.21?-2:~ p1ropia~mu~ O\'inr <<t I.a piroplaSnlQSc htt\inl" par 'bcrcnil' .•o\cro
~9 ,101.n.t.\S\. u.,;., Ml:.i1ut<Jtl:\. H. ,._ 11t110110.r·, ~.t.. 1982, llcctron H:u11unritt. Bt·<,gmr11 6. 3-l 3.
m1cru<ropfc '-ll.ld~ on th~ cl,o.vdopmem 01 &W<1•ltt 011~ Plro;pJ,t~,mla) iu tra ~,11 u1. M,n,. 1983 &1btst,d.,on->: C:c1mpu1c, ,111ndaucm of thu rd·.stiunlihip
thr ,.1!iva11 Fland... of rh~ \·cctor ,i,•k Hllipu.·.,111Jtnl11t buf't(j A, m J'tOJJ/rlJ b~twt'-''" ,h~ ll<'k \'CttQr p.lrJ.siw. and bo\'ln\• hm,J. £'(1)('rt1m•11Ull
J9.J9 -10. P:1rastrtJI~•. :'>U. 27 ~10.
WI "101T\HSX. U.G., ~MtUt(JN'(, II~ LI f\11..Pllnn.. fl.,T•• 1~:: Ullr11.~1rutlut,'II l\3 -suu1,usn\ , , ,,, 1!P'3, Jntrau1enne-infectlon of ,;hccp iems wotl\ &rPe'Sla
..uid) of the dt\ e1opmc.m uf Raf"-',s.ia ,wi> Puopla~m•al n1 the ov~ nr o;•JsOage>t.an M~d Turkmcnia Tnlfty V.M•soyu..;rit>.;,.'O hu:trucn
1ht\~tordd Rhipfr.tip/111111.1 hwso Juu,,w/ ofl>roro;o#!o~ W, .3()..J8. t::k.,.1n.·r1m,•nr.:/'nm t ,•tt.·r,m1rii. 31. 307-308.
61 \.10\'.0\', M,. r.t.'Tt\Q\' u. ~ :\111 OUli.HfV. I , lf){J7,
'l'hc O(t'Urrc.~nw. '!lf)f"('IC') 84 >UU1),.,.,o,·., , .. 1!','.'B. Dn"Clupm<,n, nnd 111orphol9lr.,' of &,/...,f1>01•Js
f>mpu"hlrJ.n 1.nd wa.son1ll al·U\'hy or lxodfda~ fanuJy u(:k.\ 1n narthW<:$1 \8:tht::,. 189:.n ln &he- blond on tht! inwnt1-brate host l.iJ1,•,1twmlt~ Stwkt,
BuIR;tria, t ,•:.•rittnnw Mcdltsins.kl ·Vmll.:.I f:,<1fi.1J 14 t8-54 ?.114··00.
~2 \.1l)f ,\S,. C .. 1903. 1.n ptroplu,nll')$(! O\'h\l-" '('3.r((!I.I~ Cumplr1 n, .mlu1rl,~
1 85 <trn.u. i \·,,t·tn.: , co~,,.,.•'-• l\'.'ISC'\• .• tto1.,nv. o. kGOnv.1 .. ~c..w 1,,s.
~mrc•J SorJrwll~ Blt,/~11«1 ~1,lt•S1,.~ F:Ua/~~1.-1.u«I'-'-'' :Pan~,. :H. dat.t .and nb..l!'l'\':tUmh ,m hrtbcslosb In "'h-t."i.'p In I r.ln\~l\':tnfa... Rulh'rin
U:!2-1.52.; /11>t1/llt1tl ,1,.wr;r1or11/r 29. I 07-1 ()9.

63 V,\.P\DOJtO.Ul.0-', .a.. l'UUI ~.,1. • UUI\:IJER(,. G.. ttt96. Pitopl.bt'r.,i uf 8Ei-- r.. 1,1ft1. ~OIL"' Mir J'"'- liumalMoa.lrcs. 1.•s tlque1, d~ o111irn;mx
uuv..111 ,.f.
dnmc.._uc anim11~in th.I! ~f.JC\."<ltanin r<-gion o!Grc«c. 1 ~1olog1rnl domc,tiqu,... a \1.lllugn,..;or. Rrl'l1tlfe Eftl'(f/1>' \INl-<i>i, 1·,·1,•rmh/rt Pfl}~
Cffl'!'i""'·tt•a.i..'110JtS. Vtll'rinmy P11u~ltt1/ogy. 63. -it,...~t'i. tropiuu. 17 3.37-:{59
 Ovine bube,,io;.1$ 4-15

37 tlllJ XUltK<i. O.. JU>\tu>.r:fl, ,1.e.., l'I kll, ',,M. I. .'.\\.\Kl. f>., lft80. Blood 95 v~u,,,,,1. 1,, :~9.5- Utibc(io,i\ in ~heep! 't"he ~..:olo~· of the- drk vcctt>r
parnhe; or ,h..,•p in lhe :--etlierl~n<ls. II 811/x,,/11 morasl 1,pomioo. ll/1/plrt/Jltolru /Jt1($(1 1mn,ml$$lon of Ral.>cs/<1 o,,Js, lmmunr,logl..-Jl,
l!obeslld,1~). Tlie VN1•rmflryQ11t1rrcrl;•, Z, 3-14 clinical ;md chnlco·p;Uh1>lngfcal "''ll<?Cts. Phi) The-I, The H<bre"
88 \'lQ-tfRJ,.:JS'. s.--... 1~.si. lmntuni1.a.tlon of,;hce.p-liROinst Bal'i'Jitt odJ
Unl\·cf")lty o( lcnt}lulcm..
ln.Ccetlon, Trudy tft!$0y1w1ogo lttttftuttJ tk$j)l•rinUwtalnof v,~:,-r!narli, 21. .UA \'l1HUI-IAM, I .. 1um,1 ..\ .. (".\UJ,R. • & flOSL.~. "· 1996 lh.c .scasonlll
2;5-2119, OCCtlrteneeof 1kb (,\Cart: lxod,d1lrl on $heep nnd in 1hc field in
Rg \\'ARS£Cf(f. ,s .. 19:8. A~ur\'ey of 111e fnf.cstotlon ~te or Lu:idid ucks with ;h,• )udc>n arc• n( h:-,u:I. l-:XJWlm,11111/ 1111d -~fl/!lll!il At:<1ra/ol.{\·. 20,
piropl~ms.in furi..1!\. hr. w1uu•• J.>:~11 .. tcd.J. Uck•bort:e D#t'h.MS a,:tf 47-SG.
1/t<rlr \ 'ertO/'$'; Proctl.f.•dlng: or an Jnn:motfon;,ll Ccinfort..ncc. F.dinburgh. 97 \1:RUHi\;\l, I. tt.\UA.'\'.L \ ~ G.\IJ.ifiM., f'.. 1,g133. :Wnw cplUWJtiologh:.al and
19,G. Edinburgh Unh'<lrslrJ I•ress cllnftnJ :u.:pett.> o( ovme bab~~lns1~ caustd hy 84best'<> ,,,,;, - ;1, revi~W.
50 wl!.,"YOl<,<:,M., t926.Pro10:011logy, Vol. II, London· 8•illle«. Tmd.U and \141t,·rin11ry f1nr1Mitol<11v', i,;, 1SJ-l63.
C.ot. 93 .a.
n-nuHi\M, 1.. ltz\DA'\I , °'"'-"~M..,.,.
1998. Aucnunlfun or &ri'PM.h miJ,. 1,~
y1 wrucan. u•... U>OOOt-M. B \.. • \1At10s-n. o.t-.. 1981. \'arulcnt 11nd a"lmlen1 rapid ,ucce"'1w pas<at<'> m ~pltnec1ornize\l Jamb~ Troplca/i\11/mt1/
~uuin or Rttba/11 bQ11u. The rt!l.ufonshlps bt(\\'~"(.lrt p.;sr.u,hc pro1e1h·~ IJM//11 m1</ /1rodllt"tiUII. 30. .>3-S5.
content ,nd p;u.hophysiolog!c;tl ellce1 of the <train. /mlmal of 9g vmtUH.\M, 1•• 1t.tD,\i\1, ~. c;,ut:n, L, i\\'lrMJt. v. & tl<X:.t"' , ... t\198 Oinica.l,
P1t>ro,.oofoa:,•. 28. 118-120.
cllnlco·pa1hologkal .1ntl wrnlogltal stud!"' or 811/t<>,ir> m·u in
92 YloRUIIA.'1, 1., IIAOA~I.A, GAi.>.,•• I •• M,'IUlf>M, e. IIUKI'\. ''·"HQ;>,.,, ""J><firn1•ni11lly lnfet:lc:d >hccp. /1J11r1111/ o/W1en111.u;- M,'<llclt11• ~,·r B. 45,
,~as. The g<'<>i;rnphlc.11 dbuibutlon :>nd •nlm•l ho,u ur l/l11picupl111/11, 38S-3~1.
bmm {Canc'Slrinl and Fanmgo. 1877) In lsrael.11,iwr d'E/;mgr ,,
100 V.UUII.\M, I.. H.\ll,\"· A. O< .......... ~•• 200(), The lire cy<le o( RhlpiNJJlw/11s
.,re,fednr r'e'trrinare P(Q'S tropl(YlU:t;, Jtl. 173-179.
/mm, (:,,nt,1r!nl •nd Fai11.ag1>, 1877 1Acari11•: lxodlda, untler
93 '1'1:.RUHAM, J., lt,\()ASI, "·· Gi\ti:t',k, J•• ftO~f'.\. ~ d, ~Hui,., .. 1g.J2. ,\ field lnboro101) condllllln, l'ewi11my P,rm{ftol<>i:y. 89. 1OiH lfi
,nudy orh~tmnp;lnlsit~ In'""' flnck., of<heep m 1,rael. J,rud /ow1111/ oj'
tOI \ 1 J\IHlA'l, , .• 11.\l)i\~I ,\. l,f.,\1$1'.JI , ... 2()00, The clfect c,f lhl: O\!nC ltc»I
Ve1,ri11aryMtdlcl~. -li, 10,-111.
pM\sitaemlnun thedeveluprn•ni or &1/Jes/1101•/s (Babe<. 1892. In 1hc
94' l"'f1tUHA}1, t•• l!Al>i\.~1, .\,. GAtKFk, r.. HOSF..~, S. & SCHC.U1\, J,. 1995- ASIUd}' c,f tfek R11ipirc-pl1{1/USbt11*1 fCsm.~uint And Fiin1:ngo, 1671 ,In
an ~nzoouc fot11$orshe~p babesrosi, CIJl1/x's/11nv/s.1!;1b\h 189~1. Preparation)
\ erari1tt1.f)' Pnm.<fro!ogJ•, 60. 349--35-$.
1
 THEILERIOSES
Synonym: Theileriose (Arrik.)

7111Jilerft1 is a genus comprisi ng tick-transmitted parasitic called Tlieilerin tiuffelilorienralisgroup, i~ \\~dely distributed
protozoa in the family Theileriidae. order Piroplasmida. around the world in cattle. but rarely causes clinical disease.
subclass Piroplasmia. phylum Apicomplexa. 6 It includes except in eastern t\sia. More recently, two other species of
parasites pre\1ously allocated to the genera CJ1t111xzoo11, Theileria hnve been described In eattle, both usually non-
Gonderia and Haemaro.,·e1111,t. The taxonomy or the genu~ at
species level is controversial.
. pathogenic, Thei/eritr (Cyw1Lrzoo11) 1111trotragi ' and Tlieife-
ria (Haemmoxe,ws) ueli{Pm: 5
Of the specie$ of 111ei/erin that infect cattle, the most im - For the purposes of lhis book, East Coast fever in its clas-
portant in sub-Saharan Africa is Theileria parz,a.13 Originally sical form is described as the type example of theilerioses in
described as a single species, it was for a period divided into cattle in southE!rn and eastern Africa. Other diseases caused
three subspecies: T. par11a par11a (the cause of classical East by members of the T. parva complel' are considered in com-
Coast fever): T. pan,a lall'rimcei (the cause of Corridor dis- parison with East Coast fever. while the other species of
ease}; and T. parva bouis (the cause of Zirn babwe theilerio- Thei/eria are discussed only to the extent that their presence
sis).5· 16 This subdhision has now been abandoned. 1 11 affects the diagnosis and control of 1.h e 7: pnrua complex.
1'/ieileria annulari 1 is the on ly other major cause ofbo,'ine 711eileria <11111ullun theileriosis is described as a separate
theileriosis and occurs in North Africa, soutl1em Europe and en tity.
Asia. In the course of investigation into the development The extensive literamre on the theilerial parasites and
and implementation of control measures for East Coast the diseases that they cause in caule have been re,iewed
fever and Theilorta tll/1111/ma theilerlosis. a number of other comprehensively.:?. 4 • a, IO. 12
theilerial parasites have been identified in canle (see Chap- The ta~onomy of the specfes of Theileria that infect small
ter 29: East Coast rever: Table29. l ). The first such organism ruminants is very confused. but only one species. Theilerin
was Thei/eric, 111wa11s:• a species which is usually non- /estoqu(lrdf CT. hirci J is of economic significance. It occurs
pathogenic. although it 11111y cause severe or fatal disease in in the Mediterranean basin. :S:orthAfricn and Asia. The other
East Africa. Theileria bujfe/i was originally identified in Asi- species are general!) regarded as being benign and are
atic buffalo.9 It. or species closeJy related to it 111 the so- widely distributed in Africa. Asia and Europe.

References
""os.. 1~119. TheUeno,;,; In E0>1,•m. Conrr•I ~!Id S<lutht'm Afnca hr. ; \SJ\RTIX. u. t.--8ROCkUS9\. o.w 1900-. A ne-w parasite oftht- t1and. 11:e
t>OIA..~. T.T. ltd., J'lroc~Y'fli11pof1t Wark:1.ltt11>nll Ea.nC/JiUI Ftl'i"f l',•rrrl,UV)· R«ord, '.'2. 33! ~,32.
lmmunl..<ario,1 hdd in l.flongu.•c, .\Ja/1m.1l. 20-22 Stpt1.:mbt•r. 1988. 8 ~"'EITZ, \\ c,., t~i- Thellerios1s. gonderioses .snd q1n~.oonoses:A
NalrobL !.:on}-. lnttrnolioruJ Lnbonttory !or R<'>i.':lrch on Animal rt,·1t\.... 01.<frr.m:poorr Joumnl o/Veunnol)· Rest·tu'rl:. ~-;. ?i5-130
9 ~fi\'CU•lL\tYAn. ~1 .. 1912. P.'1m.!itnlogft!d1.."$J\nlmm1,.1 DomfstltJUl'l, Pan!.
2 a.\ItNm. ~.F.• 1968. ThtU~_n:ssh,. Ju; wE,~MA'-, o. k.rusr1e. )t•..e~J.
Lamarre
lnfec1/011s Blood Ditetue.< of,\fa1J and Attlmn/s. Vql.2. ~cw York.1.ondon:
Ac-ademft Pt~,. ,o "·-" ,·ou~c. A,S. {C'ds.). 1992. Th:i: Epidemiolog,·of
SOJ\\'At. R..A.t., PtflR\', n
n/l'ilcrlru!s in ,lfriC11. London: Mademic Pres.>.
3 nSCHUr\'.XOws~-..·. ,. " LIJHS:, 1.. 190,1. or~ Plroplnsmosen drr Rine.for.
Ccmrnl/J/otf fiir BnJ:1,•rit1log1,, 3S. 48&-192. u 11 1iHR'r. u.u. fi ,·m,,~c. ,\.$,. 1993. rhe naming s,arne! I.he changing fonune$
or J:;,,,t COct>I r.- Cl ond Tlu'i/rrta /J(lrl•/1. f11U Vt1rri11t>1)' RMord, 133,
4 ut,·,s, ,,.o."' ,10RJu,-O:"I., w.1., 1.987. hnmunopmhology, unmunology ~nd
6[3-616.
1mmunoprophrl•liis orT11ellerit1 inr~i:ilons-/1:: souLSBY. E.1.1... <t4l.
12 PVJt~fU.. n,,-., 19.: Ea~t CoMt fo\ler: Samv n-ctnt rc,;earch 1n E~st ,.\frh:-a,
Im mun~ Rrspontts m Pamsill! /11fcctlons: Jmmunolbgy~
lmmm1opalirol08)' and J111nmnoµrophyln;tis. VOl Ill. ProuJ:t)(J, A(lilflnn"I in Ptun>ltaloJD•, 15.&3-132,
R<;ca Ra1on. Florldu: CRC rreM. Inc. 13 THIJ.IUR,,\ 190, E4$t (o.,,., fo\'er. Tra11,,f!v>nl Agrlcul11irtJJJ011111aJ. 2,
·12t-l38
> 19;9. lno dUl',remlal dlagno,i< or •h• bo,ine 1heUertis or
lAWRE..'<CE. /.,\..
,outhcm Afric.,./oumal of1hc So111/1 ,>.friroo t ettnr.1uy M<Kwtw11. so. " 111mUJ1. A.. 1904; Plrop:,ismn mutnn.< tn. •pee.) o!So~rh African eatll•.
311,-313. /oumal oj'0,111varn1i1v Pmliolug)'llnd TJ:emp,utir.,. 19, 29?-300.
G 1.t\1St.:,. r, , C01\US$. ,,o... co., . P.t.G , Dl'.M.OU!t. t, OM1S, , .. no~u;uaG. l5 u1u-~ut:RG. c .. 1t16.J. Htwmaro:<Nrni t•.•l/forrlt. n.g.. n.~p.. parasile inanne
f& .)1. u:.10.,u.. G..F... Wl8UCJt UI, A.R., L0)1, J.. LVNS. D .• ,11:1:urno. E.G.. ~tis du s."U1.g de bb\"in,- i ~IJd.iga..;car Rr:•w:d'GIP'&..Vlgi: tt di!:\l!ldicir.t
tA,<;;t. r.c.. 1"01.IA.:"'~lr.'1', c.. ,piu.nut, v.• \'\\'RA.,.&<- W,\1.L\Q, f.C. t98o,. A \'l/irt,wlr, d,s l'mY TropktJllX. 17. 655-{;62.
newly misl!d c1..,.,1l(aUon oftlte Pt~to1.oo. /111ir11olofParasltologJ•, 27, 16 u,u.~hEk!i, ~ .. ,,,s.
llt~·bome 11,•.. tock dlse....,s and the-tr ,i:ctor5. !.
J;-sa. Epi1.0o1!0Jogy of totk·bornc d1;ca.es t\'orltf A11Frnnl R•l'icw. 17 8-15.

447
 29
East Coast fever
Synonyms: Cattle-derived J11tileric1 pari,,1 infection. Thellerin ptirva parr,"
infection. Ooskuskoor~ {,\IriU

J A LA\·VRE:-SCE, B O PERRY /\:>:OS ~l Wl LLIAMSON

Introduction East C.oasi fewr has long been endemic in ea~tern ,\frica.
The paril!>iu, probably originared l'rom ..\frican buffalo Syn-
East Coast fever in its classical form is a usually fo1al discasl.' 1:eru; t:t1ffi•ri popul:itions in ea~tcrn Africn and became
of cattle caused by Theileria 11ar11a. It is transmiued prlnci- adapted to caule following their introduction and dissemi-
pally b} the brown ear rick Rhi/iicepha/11$ a11p,•11dir11/t1t1LS nation in the region. Mpeciall)· In the Lake \'ictoria Basin
and is characterized by the proliferation of l}mphoblas1s and :ilong the coa~tal strip. 12•• 11 spread widelv through che
infected with theilerial schizonts throughout the body, par· region during the early part of the nineteenth centUI') 11$ a
licularl) in the lymph nodes, lymphoid a__i:gregates. spit-en. result of Eumpean .;eulemcnt. which hl\'olwd changt'~ in
kidneys. liver and lungs. The disease nccurs "idely through pauerns of movement. extensive use of ox-drnwn transport.
the range of !Is main vector in eastern. central, and southern and importation of ~usceptible cattle breeds from over~ea~.
Africa north of the Zambr:d Rfrer tsee Chap ter l: Vectors: The l'ir~1 World \\';u (19 1.\ 10 1918), during which the cam-
1'icks). It was introduced into the region south of the Zam- paign in whot was then German l.!.ist Africa. now Tanzania.
bezi River in 190 l /02 and spread through most of 1he range took place. also resulted in unusual catLle mo,ements and
of its vector but was subsequently eradicauul (Figure 29.1 ). furthc•r dl~-,;cminarion of the disease.
The classical disease is seen in cauJe of European origin rast Coast fever was introduced !nm the area south of the
which have been exposed to infected ticks. Cante of \fri<:mn Zambezi River in the period 1901 1<> 1903 by cattle which
origin ha,·e a very variable response tO infection and the di~- were importe~I from Kllnya and Tanzania for the purpose of
ease may be insignificant or subclinical in Zebu cal\'e~ born restocking the region after 1he ravages or 1he rindPrpest cpi·
from immune dams and raised in endemically infected areas. dtm1ic of 1896 nnd the Suuth African War (1699 to 1902i.-
The disease was first de.scribed as an unusually virulent Thc disease initially appeal'ed ln 7.imhabwl<', ~ourhern
form ofbabesiosis byGraya1\d Hobertson 1' 1 in 1902. follow- l'vlozambiquc and the northern pans of South ..\frica. and
ing its introducLion into Zimbabwe. It was initially named rhcn spread •outhward along the cast coast ol $outhem
Rhodesian redwater. T<och 73 identified the parasitewi1h one Africa. through Swaziland. KwaZulu-Natal, and the Eastern
that he had seen in 1897 in cattle on the coa<t of Tanzania. Cape Province. In the period up 10 1914 ii is c,timatcd 10
I-le named the disease African Coast fe\'er. the namr being have killed one and a quorter million of the four million
subsf?quemly modified to East Coast fc,·er in mo,1 countries cattle that were present in the atTecrcd territories at the
in the region ocher than 7Jmbabwe. The causal organism beginning oi the outbreak. The southward adYance was
was named Piroplasma kochi b} Stephens and Christophcrs halted in the region of 1,ast London in the Eastern Cape
i11 1903. but the name was changed 10 Piroplasma parv11111 Province and the disease was subscqm•ntly eradicated in a
by Theiler•:.: and sub.sequemly to Tl1l'ilerit1 pnn,a by Betten- prolonged campaign consisting of mO\'Cment control. tick
court. Fran<;a and Borges in 190i (a, quoted b) llenningi.1). control destocking or infected pasture\\, and <laughter. It
Uilenberg 161 proposed tha1 the organism be named 1111:ill!- was eradicated irom southern ~lozambiq111: by 1917, from
Jia pnrvll pt1rvt1 to distinguish it from other subspecies of T. 7Jmbab\\'e by 1954. and from South \Irie-a by 1955. The last
pan,a which cause diseases that can bl' dlffC'rentiatcd from case somh ol the Zarnhczi River occurred in Swazlland in
East Coast fe\·er on epidemiological grounds. However. it 1960. TI,e disea~e has persisted in the region north of the
has slm:e been agreed that classical F.a,;t Coast fever l>l' Zambezi Ri"er to the present dnv and is a major consnaim
known as ca1de-derived theileriosls and that it is caused b) on the de.-clopmc-m of cattle production.
the same para~itc a!. that causing Corridor disease and Janu- Eru.t Coast fe, .:r if uncontrolled, may cau~e O\'Cr 90 per
ary di,,ease. 1 · 128 cent mortalil}' of the susceptible canlc following its
 •'

Uganda

•.. /
~)
'\
\
0
u
Mozaml>}Qua
•

Figure 29.1 T~e approx11naU1 dlstnbutton cl cartle-assac,aie~ Thei!eua {Jilrva tnlec11on namely Eas1 Coast rever and Zimbabwe thelienos1s. "' eas;ern
aflll s ~ther~ A!11ca. trom Lessard er al. (1990). B~ffalo-assoc,a,ed 1nfett1on. ccmmonly refe;reo ,o as CQ1·,oor ,:J,saase. occurs ,n areas.~ wit ch ca,re
and Alncan :iultalo JSyncerus coffer} have contact. It nas a s,m,lar out ',\oder d1stnl1ut1on that incluaes Scuth Atr ca. Soil1wana and Angola
INSET Tre aPllroximatc d1Sir10Ut1on of tire Ahcan burfa,o. 'nod1f e<l irom Lessard ill al (19SOI

incroducrion into a region. In an area where ic is endemic. rick moult>. a club-shaped motile kinetc forms within the 7-)'·
mortalil) among locall~ adapted Zebu-typc cattle mar be gote. This is llberatcd into the body cavity and migrates 10 the
negligible but there isevidencethm rhe disease causcsasig- salh11ry glands via the haemolvmph. The kinere im'lldcs an
nificant reduction in growth a1,d producti\ic) . Su~ceptible epithelial cell of the salh ary gland. almost exclusi\"ely F C'ells
cat Lie introduced into an endemic area are very ,·tdnerable of type Ill ,lclni. ;,- and develops into a large si11c-~1ial sporo-
ro infection and the cost of the control measures required is blast within which many thousands of minure. eiongnLod
a continuous financial burden on both live5tock owners and sporozoit~ deHtlop during ,he early pan of 1he next feeding
state veterinary sen;ces.51 · toa stage of 1he tick. :\cini ol' ialivnry glands containing mature
sporozoices can be identified easil)' in whole mollm prepara-
tions of tick sall\'al') glands Mained wi1h methyl green P)TO·
Aetiology and life cycle
nin. t<,<, A poli,nerase chain reaction CPCR) and D;'\A probes
71,elleria J)(J11•a. the causal organism ofEas1 CoaS[ fever. aher· have also been used 10 Jecect T parvn in 1icks:i;;, IGR and to
nates between ox and tkk in its life cycle (Figure 29.2). The discriminate 1: parva from r. 1<111ro1mgi. 18
~exllal ,tage of dc\·eiopmc.m oe¢urs In the gut of the 1ick. 11 The sporozoiws are lib,•raced imo che sallva and lnocu·
Piroplasms m 1he erythrocytes of the ox are ingested \\1th the lated through the skin of the ox as che cick reeds. Tiler cmer
blood meal of the tick and develop into thread-like micro- lymphocyte:. and deV(·lop within rhem. becoming mulri-
gameres and globular macrogame,cs. These fuse 10 fonn nucleaic .>chi.zoms after a period of about three day,;. 1411
zygotes which invade the epil helial cells of the gut of I.he tick. Sporozoite entf) b rapid and enerm• depcndem. and is
Complerion of de\·elopmem occurs in the immature stage, of effected by a zippl.'ring proces:. between sporo1.oite and
the tick only after they ha\'e dC'tached from the host. ,\!. the lympho~,e plasma membranes?;· J.l'.i. 169 Difl'11remla1ion
-130 "°'"' rwo: Prorozoal d1S·ea~es

Gut
Saliva()' Garr.e1ccy1es
gland

Gamonts

Figure 29.2 Ltfe cycle of Theireria parva

into the schizont sli.mulatcs the host lymphocy1c to rrans- blasts and the piroplasm in erythrocytes. The schizont is a
form into a l)~nphoblast and thereafter the schizont dh~des in mass of protoplasm contained \,tjthin a thin membrane
S}11chronywith the host cell as it undergoesmitosis.65 ll is be- which is situated imracytoplasmically in the infected lym-
lieved that schlionts have the ability to transfer from onehost phoblas1 (Figures 29.3 to 29.7). It varies In diameter from
cell to another, but the mechanism by which they do so is about I to 16 pm 1,ith a mean diamctcrof5 µm and contains
not kno\,11 and the frequency with which transfer occurs within it between I and 85 chromatin bodies. \\itb a mean of
is probably limiwd. 10" The kinetics of schizom replication eight. 13 In macroschizonts the chromatin granules average
have been described in detail.1°2 Schizont·infectcd cells l µm in diameter. while in microschizoms they average
proliferate exponentially. becoming distributed rhroughout O.S µm. 118 The size of the schiioms and the number of chro-
the lymphoid system. with metastasis of infected cells to matin bodies that they contain Increase as the disease
non-lymphoid tissues a lso occurring. Tnitially the schizonts progresses. 13 With Romanowsky stains, e.g. Giernsa. the cy-
have large chromatin particles and are called macro- toplasm stains pale blue, cypically of a lighter shade than
schizoms. Later, a generation of schizonts -called that of the host cell. and the chromatin granules stain red
microschizonts develops with small chromatin particles. (Figures 29.3 to 29.7). In haema,oxylin and eosin stained
Merozoites liberated from the microschizoms invade the histological preparations the chromatin is basophilic and
erythrocytes, in which they are frequently referred to as piro- the cytoplasm neutral. The macroschizont was first de·
.p]asms, thus completing the life cycle. Merozoites have the scribed by Koch in 190375 and is often known as a ' Koch's
ability to dMde \,1thin eorthroc~'les co produce four daughter body' or a 'Koc-h's blue body', the laner name emphasi.Zing
merozoitesJII but intra..erythrocytic division is uncommon. its appearance when ,1ained by a Romanowsk·y technique.
Recent studies on the fine scrucnire of the developmental Schizonts a.re r.ound imracellularly in lymphoblasts in
forms of the organism have been reviewed.93 lymphoid tissue., throughoul the body (including lymph
Two stages in the developmem of T. parva are readily nodes. spleen. th~'111US, Peyer"s patches and the lymphoid
identified in the infected ox. namely tl1e schizomin lympho· aggregations in parenchymatous organs which are a feature
 East CoaSt fever 451

.
••
.••.
,
,t

· ·~, ..

•
•

-·---
-~
..
••
,
-·
•
"lo •
~

•
....t! •
.. .
'
.

Figure 29.5 Blood smear drawing of lymphocytes and lympl\oolas;

containing macro· and m1crosch12onis and red blood cells rnfette~ v. ilh
piroplasms 1By courtesy of the Editor of ,he Onders1epoorr Joumal of
Ve1erina,y Research. Printed in Volu'lle 27, Number 3. 19571

•
,~ ~
...
'
. .•
; •
• .•. .,.
• •

•
,.

..
• ••
•
' •
• •
••
_., .,
Figures 29.3 (top) and 29.4 (bottom) Impression smears irom
Figure 29.6 Blood smear of a bovine Infected with Theileria paNa
prescapular lymph noce: drawingsof lymphoblasts and lymphocytes
containing macro· aoa m1croschizonts. Note several schizonts are
extracellular after tiis:niegration oi lymphccyt1c cells (By counesv of the
Et!itor of tile Onders1epoori Joum1J/ of Veterinary Rese21ch Pruned in ,,
Volume 27. Number 3. 1957)

of thellerial infections) as well as in the blood. They

frequently appear extracellularly in smears as a result of
•
rupture of the host cell. The number of lymphoblasts in-
fected \\~th schizonis is variable and ranges from less than I
per cent to over 70 per cent (i n fatal cases).13 The infection
rate increases as the disease progresses. There may be more
1han oneschizont in a single infected cell.
The intra-erythroc:ytic pu:oplasms were first described by
Koch74 in 1898. although he mistook them for a develop-
mental form of Babesia. Th e majority are rod -shaped, but
round or oval forms are also seen (Figures 29.5 to 29.SJ.
They measure l to 2 µm at tl1eir greatest dimensi.o n. wilh a Figure 29.7 Spleen smear of a bov1ne1nfected with Theife11a {laMI
neutral or p(IIC blue cy1oplas01 and a red chromatin granule
at one pole in Romanow~ky-~talncd blood smears.Although
usuall, singlt'. them mav bl' several parasites In one ery1hro-
c~1e in hea,·) infections. \1umber;; vary con;idcrably and~
many a~ 80 per cent or the crythro,11es may he infecied. 118
Following experlmemal infection. schi:r.oms ma) be dc-
tecred between :i and l.5 dayi;after anachmem ofrhe tick, 118
and ther persi~t ror a 1'i11'iablc lcng1h ol' time. Plropla~ms are
likely ro appear four 10 five days later and usually disappear
abou1 l Oda)'$ ofter fir1al recovc11 of the animal. but man1er-
sls1 indefinitely.

>
The ~chizom of r. porM wa~ first cultivated i11 11irm b)
Hulliger \\'ilde. Brown and Tu mer r~ in l964 and was subse- .
.
quently established In bovine lymphoblastoid cell lin,:,\ b)
;1;faJmquist. 11:yindo and 8ro\\'n11: in 1970. Lymphoid 1issue
or clrculaLing lymphocyte, from init.>cted canle can be
cultured dire,:tl) or seeded onto a feeder la} .-r ofbO\ine em-
bryo spleen cells. Schizont-infected lymphoblasts dhide
and become established ah a setr-perpernatlng line of cells
growing in su~pcn~ion. The ,chi1.ont~ $eparaw Imo daugh-
1er schizonts in association with the mno1ic apparams of 1he
hoS"t cell during divi:<ion and are d1stribu1ed one 10 each
daughter cell, wilh the result 1ha1 almost all the cells in 1he
culture ore infected. Clllmrcs can ebo be established by 1.he
Inoculation of uninfected lymphOC)1e~ with sporo1.oites dc-
rh·cd from inf1•c100 lick~:1' 1 S<:hizon1 cu.liurcs have proved
e,1remely useful as anlit,ens for use in serological 1es1.s to
dctet·t scrum amihodies in ~:attlt'. ~3 a, antigens for mono·
clonal amihOd)· characteri1.ation of isolates of the para-
site. u" as a source of D:'\A for molecular studies.3'' as targe1
material for ~treening chemotherapeutic agents, 811 and as a
•
source of Infective marerial for tran~mlssion 10 cattle.:,J
Techniqu<!s for the propagation of tlieilerial parasirc, in cul-
ture are describcd in detail by BrO\m.23 · r
In the field. 1ransmission of F.a-,,,t Coa~t fever takes place
only through 1hc medium of1he tick\ ector. The natural vec-
tor is R. apprndicul<!ru~. a l·ommm1 para;,ite principal!) in-
fescing the ear~ of caule ;md other herbi\'ore~ in 1he mun:
humid axeas of i.outhern and eastern Africa (sec Chapter I:
Vectors: Ticks).82 Other members of the genus Rhipiceph-
a/r1., and se\·eral Hy11lommn spp. have bt'Cn !>h0\\1l tO be ca-
pable of transmiuing "/: pari•a in laboratory conditions
(Table 29. 1).~1- 2~- 78· 118 There i~. however. no e,idence 10
~uggest that any tick other 1han R npp,•ndiculnrru plars an)
signlficam rolo in transmission of the disease in nature.
Rhipicephalus appendic11/n111s i~ a three-host tick, and
transmission of T. pan,o is achie,-ed only by n)mphs in-
fttted during the preceding larval stage or by adults infected
<luring tl1e preceding nymphal stage. Transovarial transmis-
sion doe,. not occur. nor is 1.here transmission between larva
and adult ii the intervening nymph feeds on a non-suscep-
tible host. The tick acquires infection when it feed,. on an O)i.
at a stage of clinical disca~r or recovery when piroplasms are
prcsem in the blood. The succeeding stage transmits Infec- Figure 29.8 Li~e era·,•, ogs of p1roplasm; ot T/le1te11a spi;
tion as ii feed~ \\'hen 1.he ~porozoites m 11,. salivary glirnds oilier than ·hose which ct,-elop veils
 F.ast Coas1 fuver 453

h:l\'c ma1urcd. At moderate 10 low ambicn1 1c111p1:rature~ site of Alric-an buffalo whkh ha,. become adapted Ill cattle ft
ma1ura1ion ohporo7.oi1e~ in adult tides ts 1101 complete until has been found m infect wawrbuck Xo/Jus dt•.f'<,sstl) t iK under
three to four cJar~ af1er feeding commenct'l>:' 2" n; matura- natural tondition;.. :md the i\slatic buffalo ,Btt/l{l/1ts lmbali.s1
uon m nymphs is more rapid.' 33 At ambient 1e111peramres of undt'r experimental conditions.97 h is not infective to other
3, C or ,1bo\'e, !.porozoite maniration can occur In the ungulates. nor 10 any ,p<'Cie, ofluboratory animal. There is
unfed tick.· 77 and Infection mar be trnn~miucd ns soon as 110 e>\·idenc<: thac \\'Jltllif.:, play" any role in the epidemiology

the tick bc~ins IO feed. of the dhease.

Aniticfal 1ronsmisslon tJf '/. part'<I can be achieved in Theileria par11(1 depends on Its ,•ecror Lick R, fl/Jpemlin ,.
two ways. The first is by 1he inoculation nfa $Uspenston of Imus for 1ran~mi.,~ion from ho~t 10 host. The potential distri·
sporo-.oites prepared from homogenized tick ~alivary bution of Ea~t Coast fever is thus res1ricted 10 those areas 01
glands or whole ticks, the so-called GUTS (ground-up rick ea.stem, ccmral and southern \fries whert- ox and ti~k t<ll.'x
superna1an11 prepara1ion.~ 2 15~ This method mimics is1.su In eastern and cemral •\frica. this Includes much of
natural rransmi~sion and obYiates possible \'nriations in Ken}~· Ugand:.i, Rwanda. Burundi, the eastern µan of rhc
tick auachmem and £ceding behaviour. By pooling ,1 mnn- Democratic Rt•public or Congo. area,. of .. outhem Sudan
ber of ticks before homogenization ii i~ also po:;sible 10 bordering Uganda and much of Tanzania. In ~ot11hcrn
eliminate the variation in infec1ivity between batches or :\frica i1s range is more limited. and it is confined to the
ticks. The mc1hod ls widely used £or 1hc charactcri1.a1ion of nonhem and central rt.>gion~ or \lalawi, 1be nonlwrn. east-
isolaws of T. pflr11t1 In caule and for the infcc:tion and treat· ern and cm11ral 1·egions of Zambia, and the Tete Province of
mem method of immunization (see Control: Vaccinalion, ;'\1owmbiquc. all lying 10 the north of the Zambezi Ri\'er
below). S1abila1es or infective spomzoite, can be stored in (Figure 29.1) The disease cau,cd b~ T 11arut1 in the South·
liquid nitrogen and will maintain their viability and bio- cm Pro\ incc of 7-ambia has b,!en hiswrically rl.!ferroo to a~
logical characteristics for long periods. 43 Corridor disease hut it seems l1kel\" 1ha11t is predominaml~
The second method is b)' the inoculation of ~chfzonb. catrle-a,,ociatl:'d, ratherthan buffaio-as'>Odated.
Su:,pension, of bChi7.0nt$ in ,plccn and lymph node~ from The tick wcror R. <1111M1dic11law$ h widclr clistribuwd
infect«::d animals were admirtisrered intra\enously by th.roughom 1he wanner, marl' humid areas of east cm. C'en-
Theiler 1' 6 · i;; to minsmit T. parl'a as a method of immuni- trai and ~ou1hern -\frica (see Chapter I: Vectors: Ticks).
Wtion againq East C:0~1 fever, but the 1ech11i4ue was crude Within it, range it, abundance \arles considerably. being
and the results erratic. Suspensions of infected l}111pho· governed b) ~oil moi,ture and temperature, presence of
blasts in culture prm<idea far safer and m()re rcliahle source suitable habitat and avuilability ofho,,1~. Till} distribution of
or ~<:hizont~. Infection can be transmiucd rr.,gularl~ to re- C'limatic $Uitabllhy has been pinned on 1he basis of ecocli·
cipient caule when 108 iniecred donor cells are admini~- matic ind ice~.• w. 111· 1•• 1 Interesting!) areas oi ecoclima1ic
tered ,ubcutancously.24 HoweYer, when lower (."ell dose;. are suitability l!xist where the presence of the tick ha~ yet 10 be
u.wd. result!> are lnconsis1en1 and some redpiem, do no1 recordC>d, u, illu,trating that other factor, aJ,o play a role in
become infected. although an inoculum as small as I~ cells determining di~1rib111ion . .-\niong these is \'Cge1ation co\·cr,
will produce infection in the recipient animal if the rccipi· which influences the microclimate of the habi1a1 of 1he cick;
ems own celb infected in 1•i1ro are used. 101 'I he critical fac- overgra7ing ha, a murked adverse effect on !he tick popula·
tor appear. 10 be the ability of the inoculated cells 10 sunive tion. 121 · i·; Also lmponam is the livestock production sys•
for a sufficient period for the parasite to transfer 10 1he tem employed. panfcularly with respect 10 how manage-
recipient hoSt cell~ - this dcpcn~ on the degree of ment affect; cxpoMtre 10 ticks. Thus in Kenya 1here b a
historompatibility between donor and rocipion1. considerable difference in tick infes1aclon le\•els and corre•
lransmisslon of T. pan•a by the inocuiauon o.fblood can spondmg infection prevalence b,nween open grazing amt
be achie\'ed hut the rcsulis are \'Cry crm1ic 1:;9 as succes~ de- stall-feeding (1.ero gra1ing) management ,;y;,iems.;,i l ran,-.
pends on the presence of sufficient number, oi infected misslon of l:ast Con~t fever may be expected 10 otctir freely
lymphoblasts in the circulation. Techniques ha\'e been in area, where ecological factors pem1i1 an a\"ernge total
de,cribcd whereby blood rrom infected anirnah can be population of 1101 le:.:- than IO to 15 adult ticks per ox. t:>J. · .,
transferred to nymphal ticks as a means of isolating the It is imponant 10 note 1hat on the margins of R. npp,mdicu-
parasite from blood. This can be achieved eithl!r by inocula· lams dis1rib1.11ion. the distribmion may expand and commct
1ion imo a rabbit on which the ticks are feeding. 1:i-1 or by \,irh seasonal or secular change~ in climatic suitability. 12;,
direc1 inoculation into engorged nymphs. 14i The seasonality of Eas1 Coast fcl'er varies throughout ih
distribution, but is generallr dependem on rainf:tli and the
abundance of adult stage» ol the tick \·ector. u~ lo the high·
Epidemiology
land :ind coastal region& of eastern Africa, \\ith its two raim·
The epidemiology of East Coasr £ever and the other 1heilerial seasons. seasonality is not ,·cf) marked, and m area~ where
iniections has been re\iewed exumsh•ely by :'\on-at. Perry all ~age,. occur simultaneouily throughout th<' year. ,o does
and Young. 125 'fhef/eria pan1n is probably original!) a para- East Coast le\•er. In southern Africa from sou1hem Tanzania
454 ,ocno., l\,o: Protozoa! diseases

Table 29.1 The the,Jerioses of hves,ock in eas;ern. Central and southern Afnca

PAAASrTE VECTORS DISEASE SPECIES AFFECTED PATHOGEN!CliY DISTRIBUTION

inei/e(la oarva .'lhtpi&epnalus Eas, Coasi ie\-er Oi. As1auc <:1Jmest1c Usually fatal !benign Endemic thtO'Jghout
eppeM!cularus· • buffalo te~penmen;elJ in resistant Zeou most of the <ange of
R di,rront' ealvesl R. append;cu 'aws
nor.h of Zambezi Rille·
R. l8mbez1ensis· Afncan buffalo Benign
R cepens;s Wate:bticl< Benign
R carniV()(8/is
R compos,ws
R. evens, evens,
R, koch, Corridcr c:sease Ox Usually fatal localiied. in
R. pravus African bt:ffalo Benign associa1ion with
R. pulchel/us African buffalo
R s,mus
Hy11/omma anatolicum Zimbabwe theileriosis Ox I Vatiable -ofi~n !a,a1 Zimba!l't-Je higllvel:
H dromedarii
H im{}lessum
Thei/ena raurorragi R eppend,culaws· • o~ Usually benign- Widespread 1eoc'emic1
R evetrsi eve.as, raielv fatal
R putchellus Eland Sometimes 'a;al
'1 zambe11ens1s
TJieilena nwums Amb/,;omma Ox Usually henig" - Widespieal! 1enoemicl
hetne~m·· somesuains
A 1•enegarum· • pathogenic
A. as1rion African buffalo Benign
A cohaerens
A gemma
Thei!efla velifere A hebr11eum· • Ox 8en19n Widespread (endemic)
A variegarum' • African buffaro Bemgn
A asuion
A lep,dum
Theile~ sp near H;1emaphyse/1s spp, Ox Benign Widesoread
T. ooffelt
Tile1fer,a sepa,at.a R 6'1w1si everrs, Shei;p Benign Widespread tenaem,eJ

Important local vec:c:s

Pnnctpal natural vec;o1s

southwards, seasonality Is much mo.re marked. with clinical East Coast fever is maintained by a continuous cycle of
disease occurring predominantly during Lhe wet months of transmission between cattle and ticks. \·\qthin the infected
January tO March, when adult ticks are most active. 1- 2 areas. tha incidenc:c of the disease can vary ,,~dely depending
Ticks will transmit infection if, during lhe preceding on mm1erous factors. including t.he abundance of t.he vector
stage of development. they have fed on an ox with circulat- and the susceptibility of the host. There is a marked 1'llriation in
ing piroplasms. Infective cattle may be elinically ill. recently the susceptibility of cattle to infection. Taurine breeds are gen-
recovered. or persistent carriers. In general terms. the infec- eral!}•more susceptible than Zebu and Sanga breeds. 1n addi·
tive dose the:: licks receive will be higher from clinlcally tion. there are variations 1vithin breeds depending on t.he
affected a nimals than frqm carriers. for many years it was epidemiological state of Easr Coast fever. Under certain spe·
considered that cattle recovering from infection had a solid. cific circumstances. F.ast Coast fe\'er can exist in an endemic
sterile immunity and that ticks could only acquire infection form in which it has li11le or no influence on morbidity and
from recently infected animals. There was thought ro be no mortalit) in the herd.9. qg This situati.o n was first recognized
carrier scale. and maimenance of infection was thought 10 by Bnice in Uganda.31 was characterized by Mettam and
depend on a cycle of rapid transmission. This is certainly Cannlchael.~s and is now commonly referred 10 as endemic
Jrue of Lhe classical laboratory stocks studied intensively in stabilit:y. 12.:1, t:19 However, endernic stability 10 T. paroa infec-
Kenya tknown as ·;,.tuguga') and in South Africa (known as tion appears 10 be relatively limited in its diStriblltiOn and may
'Schoonspruit '), and appears to have been true of Lhe dis- not be achienicl easily. The more common situarionseen in the
ease as it occurred south of the Zambez.i Ri\'er in Lhe first half region is that of endemic instability, in which varying degrees
oft.he rwentiethcentury."2 However, investigations in Kenya of clinical disease are experienced.
have demonstrated that t.he carrier state appears to be Three major epidemiological fonns or states of East
1\idely prevalent in endemic East Coast fever areas. 7 1. 176 Coast fever are recognized.iz,,. i:is These are endemic
 East Coast rever 455

stability. endemic instabi!icy and epidemic. A State of en- liferation ofbotl1 infected and non-infected lymphocytes, pos-
demic stability implies an ecological balance between ox. tick siblybysrimularing production ofa T-ceU growth factor similar
and parasite in which regular transmission ofT. pan:a occurs to interleukin-2.28 Lyn1phot'}'le proliferation commences ar
in all age groups of the canle population and high levels of the site of inoculation. but from five days after infection para-
population Immunity are achieved, with minimal effects in silized cells begin 10 appear in the draining l)•mph node.
terms of morbidity and mortality. This is most commonly The appearance of parasltlzed cells in tl1e lymph node
achieved in areas \ ~here indigenous Zebu cattle are kept on coincides with the onset of fever and stimulates active lym-
pasture in areas highly suitable for the vector, and where phoid proliferation and the release of large nwnbers of
major seasonal fluctuations in vector abundance do not infected and non-infected l)mphoblastS into the efferent
occur. Infection pre\'alence in cicks may be in the region of lymph.102 After a further two to three days, more distant
I or 2 per cent,79 infection intensity in infected ticks is low. lymph nodes in the chain become infected and hyperplastic
infection of calves takes place early in life. and reinfection and paraslrized and non-parasitlzed lymphoblasts enter the
throughout life is common. Endemic instal;>iliry is fou11d in peripheral circulation. These establish in other lymph nodes
cwo forms, low incidence and high it1cidence. Low incidence and l~111phoid 1issue (thymus. spleen). as well as in many
instability is found in areas of very low infection challenge, parenchymatous organs (notably liver. lddneys. lungs. myo-
either in areas marginally suitable fonhe vector or on proper- cardium. adrenals), bone marro\,, gastrointestinal tract mu-
ties where acar!cides are applied Intensively. High incidence cosaandsometimes the brain. The distribution of para$ites In
instability is characterized br an infection challenge I.hat is non-lymphoid orgnns 1s variable and may be associa1ed witl1
insufficient 10 induce population immunity. This can be as a incidental pre·existing lymphocyte infiltration.
result of ineffective tick control. or intermediate levels of Four to five days after the Initial appearance of schf7.onts.
infection challenge. Epidemic East Coast fever occurs when che process of lymphocyte prollfera.tion is succeeded b)• a
the disease is introduced to areas previously free of the dis· process of lymphocyte destruction, and lymphoid organs
ease, and often occurs on a ·easonal or secular basis at the and lymphoid foci In parenchymatous organs may show
margins of R. appendiculamsdlsrrib\ltion. 173• 174 evidence of necrosis and depletion of lymphocytes. Both
The distribution of endemic stability and instabilil)' is parasitized and non-parasitized lymphocytes are destroyed
affected by geographical and production system influences. and this appears to oe the result of activation of specific pro-
Endemic stability is thus most common in Lhe smaJlholder tective cy1omxicT-lymphocyrns (CTll and non-specific CTL
systems prevalent in the humid art:!as surrounding the Lake ('natural killer' cells). 5'1 Lymphocyte destruction causes a
Victoria basin, where the presence of endemic T. par,>a fall in serum immunoglobulin leveisHb and immunosup-
infection in cattle prohibirs or severely llntitS the keeping of prcssion.165 This may enhance che progress of the disease
more productive taurine breeds of cattle. Low incidence en- and facilitate ~econdary respiratory infections.
demic instabilil)• is found on the large-scale commercial In non-fatal cases, development of specific protective
beei ranches of the entire region. and in the small-scale immune responses 1erminares schizont prollfera1ion.
dairy systems of Kenya.5il 59 High incidence instability is Recovery from infection is though, to depend on the sunival
characteristic of newer commercial \•muures. in which an of a sufficient number of effector cells 01·er the first 1-l days
upgrading ofindigenous caule is undertaken through cross- and on their capacity t0 achieve a specific protective
breedJng. and tick comrol is ineffective. Under such circum- response.:,.; A proportion of recovered animals remains un-
s1ances. and in epidemic East Coast fever. the case facaliry thriftyand unproductive, and foci of infected lymphoid cells
may he extremely high. may be found persisting in a \'l1rie1y o( organs: 8 tmmuno-
suppression may predispose such animals lo secondary
infections which ma) eventually prove fatal.
Pathogenesis
Lymphocyte proliferation and des1ruction are respon-
Theileria pnrva primarily affoc1s lymphoid cells and the sible in a variety of ways for the lesions which characterize
course of infection ln rhe ox may be divided into three East Coast fever. Foca l or diffuse infiltracion of lympho-
stages: a prepatent stage: a stage of lymphoid proliferation; cytes and lymphoblMts may cause degenera1ive changes
and a stage of lymphoid disorganization and depletion.~ in liver. kidneys and brain. while focal infiltration in the
The prepatent sta_ge covers the period between the mucosa of the gastrointestinal tract b responsible for ne-
inoculation of sporozoites by the tick and the appearance of crosis and erosion of the overl}1ng tissue. Accivation of the
schi7.onts in the drainage lymph node. The sporozoites invade coagulation cascade leading to an accumulation of fibrin
l}~nphocytes and develop into recogni:r.able schizonts after degradation products85 • 1 H suggests that dis~eminated in-
two to three days. There is evidence to suggest that. travascular coagulation may be an imporcam component
although sporozoites can infect a similar proportion of both T of the cerminal srages of 1he disease. There is also acth·a-
and 13 lymphocytes.1 it is the subsequent rapid multiplication tion of the complement cascade. t+< and va~0<1Ctive com po·
of infected T cells that is responsible for the patho- nents are released as a result of lymphocyte disintegration
genicity of the parasite. t04 The schizoms stimulate pro- in the lungs. These may be responsible for the pulmonary
45-6 ,s·c 11u., nw: Prorozo3J disca~es

oedema which is often [he final cause of dcuth. individual anunals h,b been success full) us('C! to predict thC' re·
The invasion ofel)·thro~1es br piroplasm, i~ a ,;triking fea- suh~ ob~l·rved when these cattle were subjl'CIC.'d to hetcrolo -
ture of the disease. but the parasite in thh fom1 doc, not ha1,e-c gous challengc. 1·;i l Jowcver. asa result of the work done in 1hi,
any conslstem pathogenic effect. !':on-rt•genemtire anaemia area. !here arc a ntimber of molecular Loob .ivuilable for char-
and ictenis:ire sometimes described in i:a~t Const fever, but no acterizing s1rain~ which can be applied in other importam
evidence of a haemolytic process has bccm prci;cmcd. areas, for i:-~mplt,. in vaccine production for idcnt~ing \'llC·
Immunity to r. p(ln.1<1 is considered to be principally cell cine sto,ks, in mOL'litonng changes in epidemiolot,•y post-im-
mediate<l, 12 ' the most important effec1or celb re~pom,ible muni1.arion. :md in distinguishing possible vacdnc brcakdo\\11
for clearance of infection and recover)' being CTL. which (failure 10 immunize) from breakthrough (immuni;,.a1ion suc-
belong to the cos· subse, of r cells."2 91 These r<K·ognfae cessful but not protec:tiw against a field strai n!.15
specific Theileri<t antigen combined with major hiSt0\'.'om-
pntability complel\ ,).JHC) antigens on the surface of
Clin.ical signs
schizom-infected lymphoblasts and their cnotoxic activity
is thu~ parasite specific and genetically re,tricted.";· 68· 104 East Coa.1 fe\'lff in the classical fonn as first described b)
There is evidence to suggest that the cytotoxicity is also Gray and Robenson"-' in 1902, is characccri7.t'<l by P)TC\ia,
srrainspecilic, this property correlating with the variable de- enlargemem of 1he superficial lymph nodes, severe pulmo-
gree of protection to he1erologous challenge observed in nary oedema and wasting. ft usually tem1inates in death.
cross-immunitI trials in caulc.' 5i Other cellular responses The incubation period i,; generally about 15 days from
arc likely rn contribute. For e.xample. C:D4 • helper T cells the rime of rmachmem of the iniected uck. hut m:l\' range
mar be involved in the induction of parasite-specific CrL. from 8 t<> 2.5 days. The period and £he c<mrse of the disea,e
but their role appears w be 5econdary 10 the main CTL im- become shorter as che challenge i$ increased."'' The fir,t
mu11ity.6 Re-exposure of recovered animals 10 homologous clinical signs are foyer and increases in pulse and respirauon
challenge stimulates activation or memo!) I lymphocytes rates. n1erc may be a sharp decline in milk production_ The
which, in rurn. stimulate acth11tfon ofCTL, \\hich eliminate parotid lymph nodes, which drain the car 10 which the in-
the infection. although some development of scl1i1oms and fecced rick has anached, an~ enlarged. After a few dny·< the
piroplasms mai• occur and cause a mild clinical re~ponse. animal becomes depres$ed and lethargic. The temperature
Humoral responses to schizonts are not thought to be comlnucs to rise, often to 41 or 42 •c. Anorexia may develop
imporrant in the immunity of recovered animals. but but is not inevhable. Lachrvmaiion com111011ly occur« to-
circulating ;.mlibodies which de\'elop against schi;:onts do gethcn,ith oedema of the eyelids, and may be accompanied
pro\idc a useful indication or expo~ure and ure the basis for by photophobin. The animal is often constipated. Therl.' ls a
serological diagnostic tests for T. pnn•a \See Diagnosis, generalized enlnrgemem oi che superficial lymph nod<!~: thc
below). Cattle repeatedly e.'(l)Osed to sp<>ro7.oites. as ocrurs prcscapular and prccrural nodes become VCI)' prominent.
In the field in areas of high T. pnrl'(I challenge, develop anti- The dis,•asc \l.!;Ua!ly progre,~es O\'er a period of about 15
sporo1.oite antibodies capable of neutmli1Jng ~poro:toite in- day~. but may temlinau,, after live days c,r be prolonged to 25
fcctili[) in 11/rro.111 In these situa1ions. ami•sporozoite days. The fever remains high, although in a small proportion
immunitr may play a role in protection. Significantly. the of case:; chem: may be a 1emporarJ re111ission for one or 1wo
ncmralizing acrlvity of anri-spnro1.oitc antibodies ls active day~. \ppctitc nnd rumination hecome incrcasingl~ de-
agau1sl a wide range of T. pcm!a isolatC!i, unlike dw anti- pressed and there is a SC\'CW loss of body condition. increa;,·
schizom immunity described ahove. 11 ;: Ing wealmes!- nnd ataxia. and frequent recumbenci,
Strain differences exist ,,ithin caulc-derived r. parr•a which Con$liparlon i~ ,uccecdecl by diarrhoea and there ma~ be
are manifost ru; variations in the degree of cross-prote<:tion af- blood and mucus 1n the faeces. Opadty of Lhe cornea rnai
forded 10 field challenge or between differe111 i,olate!- of the de\ clop and petechiae rnay be detected in the mucous mem-
parasite. TI1e topic has been revi<>wed b)· lrvin. 0: Smuns hal."e brane<;, e~pecinlly those beneath the tongue aad in the vul\·a.
been idencified on the basis of differcnres in pathogenicity. 11 AnaC!mia and iccerus ha\'e been repon:ed bul are i nconsistem.
crt,ss-irnmunity, 139 antigenic characteristics oi the schizoms Xodular ~kin le~ioru. have been described In a small number
in 11icro as detected hr monoclonal antibodies%· l:i:? and geno- of c,-perimenral ca~es. 1"' Prcgnanr cows ma} ahon.
typic variation decectecl \\ilh DN:\ prohes.1• 1,1 Ji;. 1' 1 !he anti- ln che rerminal ~,ages of the disease dyspnoea develops.
genic diversity Is complicated br the fact thac mo$t field wiU1 ru1 increased res1>iratory race. a watel)· cough and the
i~olates are composed of mLxed parasite population~. Consid- discharge of l'rothy fluid from the nosrrlls. rhc di~charge is
erable effort has gone into £he de\·elopment of i11 l'irro methods copious both when the animal I, recumbeni and immedi-
oi charac1eri1-ing T. parm parasite~. particularly in an arrempc alel} before death. Evidence of pulmonary ()edema and
10 find a method ghing re$lllts which correliuc \\ith cros,-im- hydropc>ricardium ma)' be detected on nusculuttion. 'itcmal
muni[)· so as 10 a\'oid !he need forche use-()f canlei n e:qiensive and submandibular oedema may be present. The enlarged
and laboriou~ i1111i1-o trials. ;'\one of the method~ studied ro superficial lymph nodes b~n to rcgre;i., the rotctai
date has achieved thi$ aim. although qrain ,;pecifici() of CTL in temperature fall~ to ~ubnormaJ ie1els, and 1he animal
becomes recumbent and die,- in a coma. sions may occur in the inrc~tinal mucosa, panlcularl) in the
,\ smaU number of animals. usually abour5 per cem. may Peyer"~ patches of the terminal ileum.
recover. bm co11valc~ccm·e I, prolonged and the animab Opacil) of the cornea of the ere is often noted. especially
013.) remain umac-iaicd and unproducth'e for month'-. at the lhnbu•. and re~ult$ from Infiltrations of lymphocy1.1c
The disease may assume a le~s severe fom1 in animal~ n?lls and from oedema..
with panlal immunit> or inherent resbtance. but pyrexia Otlwr chMges may include emaciation: subcutaneous and
and enlariemcnt uf superlkial lymph nodes remain con· lm mmt1sntlar oecfomn: asciic.,: sero11$ (!trophy of fut: pallor of
stant features and in calve~ may be accompanied by persis- the liver as a n.-,,ult of mild degeneration: pctechiae in l.he mu·
tent unrhrirlinc~s.' 00 \!lid dbra5c has also b111/n reported cous membrane~ of the oml cavity, SC!rosal membranes, and
occasionally following infection of fully susceptible animals the epi- and the enclOCllrdium: :mn.cmia: !cterus; and gre)ish-
with strains of T. f)Cfft'« of reduced virulence. 1' An atypical hrown foci of hyaline dcgenermion and necrosis of muscles.
cerebral form of T. pnri•n infection i, dl!~crlbed in Chapter particularly in the larger muscle groups of the hindquaners.
32: Turning s ickness. Hyperplasia ol lymph<>id tissue in organs and tissues in
in the ,\siatic buffalo 1he discru;e resembles that in the which it is normally prcscm. or hmphoid infilmnion of
ox?; but in t he African buffalo infection is invariably sub· other organs. ~uch as the kidneys (Flgure 29.1 I ). the lh·er
clinical or mild. 11 and lhe eye;, may be recognized macroscopically and con·
finned histologically. ti;o Microscopic examina1ion will often
reveal lymphoid infiltration in other parenchymatous or-
Pathology
gans. notably In the lnter.;titium end aln,oll of the lungs. the
East Coast fever is characterized by a severe panleukopenia portal tracts of the liver. the myocardium. 1he adrenals. the
which coincide;. with the on~et of the febrile reactlon. 1.0, i-o bone marro"· and the perhascular spac~ in thr brain. In
In fatal ca.'>es. leukoL-ytc counr, fall pr<'cipitous!y until the ani- the t?arly stage~ of the disease. the lymphoid tissu<' contains
mal dies. Non-regcneratin•. nonnocytic. normochromic numerou< lymphoblasts which are characterized by their
anaemia, in which erythroq1c counts may foll to 50 per cent lnrgl' ~i1<,. a large h}'Pen;hrnmatic nucleus, and rclati\1!1)1
of normal. together \\~th thrmnbocytopenia and !e1erus. have extensl\'e hyperchromntJc crtoplasm. :-litotic figures are
also been reported on occasion as prnminem features of the conunon and germinal centres may be prominent. 10• •11
d isease. There ls a moderate hypoprotcinnen1 ia, and in the Schiwms may he seen in some of the cell~. the characteristic
rcnninal stage~ t.here may be dinkal pathological changes in feature in a h;1ematC>:\1'1in-and-eosin-staineci section being
scrum indicative of renal and hepmic dysfunction. a duster 01 bnsophiiic gmnules within n lightly• staining
S.:hizonb may be pre~ent in lymphoblasts and pi ro· vacuole in the t")'t<lplasm. In the terminal Mages of infection
plasm, in erythrocytl"i ln ~mear, made from peripheral necro~is of lymphocytt'!i \"\1pervcnes and the l)'mphoid
blood. depending on thes1age of the disease. tissue become, rather dcplNcd of cell,. 1,ith ceUutar debris
At necropsy the most prominent feature is o[ten sc\•ere being scattered throui:h Lhc tbsuc.
pulmonary oedema."' There is a copious amount c)ffroth in Piroplru.m~ in erythr<icyw~ and ,chizontS In lympho·
the bronchi and trache-a and often also In the nasal pas$ages. blash ma~ be rerngni:r.cd with the rransmis$ion electron mi·
Emphy,-cma of the lungs may develop. and a moderate hy- croscope. Uhrastructuml pntholog1cal chan~es In lymph
drothor-dli and hycl ropericardium may also be prc-.em. node'> ha\e bt•en described br De \lartini and :Vfoulton. ' 4
There is a variable degree of enlargement of periphernl and
\i~ceral lymph nodes. which results from hyperplasia and
necrosis of cells of 1he lymphocyte series, as well as oedema.
gi\'ing the lymph nodes a greyish-whim, ne.~hy appearance.
The nodes arc sometimes discoloured hy hyperaemia and
haemorrhage. Splenomcgal~ ma) be pre~ent hut is inconsi~t·
ent. In young animals the th)11lus may be enlarged.
l11c kidneys frequently exhibit one or more raised red or
whillsh foci or nodules up to 20 mm in diameter within the
cortex (Figure 29.9 ). Tradllionally. but incorrec1l). these are
known as 'infarcts' but they are ,imply aggregations ot large
lymphoid cell~.
The mucosa of the aboma~um l~ generally thickened.
due co lymphoid hyperplasia. and congested. Pecechiae or
ecch) moses ma) be present and a few ulcers or superficial
erosions up to 10 mm in dlamt?ter may be round. panlcu-
larly in the pyloric region. The ulcers arc associated with the
mucosa! lymphoid hyperplasia (Figure 29.10). Similar te- Figure 29.9 \lu,t•fotal lvmpho1d nyoe1ples1a ;n :he re~al torte~
458 s.cno" f\1·0: Protozoa! diScases

Diagnosis

The diagnosis of classical East Coast fever ls based on the

characreristic clinical signs and lesions, and mar be con-
finned by demonstration of schizonts and. in the later stages
of the disease, piroplasms (Figures 29.3 to 29.8). The pres-
ence of R. appendiculatus on the animal provides support-
ive evidence. but tick infestation may be very light and the
ticks may have detached by the time the animal is examined.
Theilerial parasites are generally numerous. lo the live
animal they can be demonstrated in smears prepared from
peripheral b lood and fh;1e-needle aspira,es of superficial
lymph nodes (Figures 29.3 and 29.4). Parorid. prescapular
•
or precrural lymph nodes may be sampled. In the dead ani-
mal, smears prepared from the cut surface or the spleen and
an)' enlarged lymph nodes should be examined. Smears
""
Figure 29.10 Ulcerations and lympnc11d hyparp!asia in the mutosa of the
must be of good quality and should be stained with Giemsa abomasum
or any olher Romano\vsky srnin. The presence of large hy-
perplastic Jyrnphoblasrs should be an incentive 10 concinue
the search for schizoms if they are scanry. Care must be
taken to differentiate between schizonts and normal azuro-
philic granules. \<\'hen smears have not been taken, the
diagnosis mayscill be confirmed if infiltration "ith infected
lymphoid cells can be demonsrratecl In routine histological
preparations. The kidneys, liver. lungs and brain are par-
ckularly rewarding tissues for examination.
lotercurrent infection \\ith other haemoparasites may
complicate the clinical and pathological picrure. and these
parasites may be. detected microscopically. In those areas
where ir occurs, East Coast fever ls generally the most virn-
lemof the haemoparasitic diseases of cattle, and its possible
presence should be given priority over any other disease
\\'hen making a diagnosis.
Retrospective confirmation of the diagnosis of clinical
Easr Coast fever can be made by demonstration of a rising
antibody titre to -r. parua using serological tests. :--lore usu- Figure 29.11 lymphoid
ally, serology is used in cross-secrional surveys or longimdi- inf1l1.1a1,on 111 the kidney
nal srudies ro derect the presence of r. parua infection and
to detemtine the prevalence, inciden.c e and seasonality of
infection and disease. Ci rculating antibodies \\'ere first antibody of IgG I isotype and ancibody levels remain posi-
demonstrated i.n recovered animals by the complement tive after a single infection with T. parua for longer than with
fixation cesr11 and since then various techniques have been the!FAT.6·3-l
de\'eloped using schizonts or piroplasms as antigen.40 The An alternative means of retrospective diagnosis is the de-
serological test of choice is a T. parua-speci.fk antibody de- tection of infection in recovered carrier cattle (see Epidemi-
tection enzyme-linked immunosorbent assay (ELISA) based ology, above). Morphologically. the piroplai;ms of different
on the use of a recombinant polymorphic immunodomi- Tlleileria spp. are difficult 10 differentiate especially when
nant molecule (PHvl)."3 • 160 This has superseded an indirect presem in small numbers. Thus. tl,e examination of Glemsa-
fluorescent antibody test (IFAn, which employs cultured stained thin blood smears to detecrpiroptasmslacksspecific-
:r. parua schizonr-infecced lymphoblasrs as the amigen. 6• 33· iry and is also relatively insensitive. Tick transmission
st The lFAT, although extremely usefu l and still widely used, (xenodiagnosis) is not practical on a large scale. The need for
suffers from subjecrivity in interpretation and lack of an improved method of detection of 'f. pa.rua carrier animals
speci.ficiry due to cross-reacci\~ty wi(h some other Theileria has led to the development of a number of PCR techniques
spp.. most importantly T. raurotragi.32 · Ml The ELISA does using T. paruri.-specific primers based 011 single copy genes
not cross-react with T. raurotragi, T. 11mta11$. T. 1m1mlata (pl04 and PIM antigeti genes)66 · 160 ur a repetitive gene se-
or T. bujfeli and shows high sensitivity. It detects quence (TpRJ. 17 The besc of these, ba;;ed .on the sporozolce
 F.ast Coast fever 459

p I 0-! antigen gene. is sensitive and specific; T. pt1rva, and not Serological examination of recovered animals may per·
T. ta11rorragi. D.Nr\ in the blood oi Infected caule is amplified mit a retrospective differentiation at the species level \\ith
co a detectable level. This assay is currently undergoing specles-spedfic EIJSAs a\•ailable for T. pnrva'3 and T. mu-
validation and comparison with conventional methods of rans-Z and standard IF,\1-G for others. Biological transmis-
detecting long-Len n persistem Infections in cattle at the sion of the parasile may also provide a means of
International Livestock Research Institute in Nairobi. differenciatlon at 1he species le\"el, bUL is expensive and re·
quires experimental caule. Theilerin murt111s and T. t1<!lifera
;ire rcadil}' transmitted by lmravenous inoculation of hep·
Differential ruagnosis
arinlzed blood 10 a susceptible ox - an inoculum of 10 ml
Clinical!~•. East Coast fever must be dilferemiated from other should be adequate. Theiler/a wurorragi can sometimes be
febrile condi tions which usuaUy terminate fatally after a transmiued in the same wa)'bUt may require a larger inocu-
course of one to two weeks and which feature progressive lw11 (100 to 20() ml). while T. parva cannot readily be trans-
emaciation. respiratory distress. diarrhoea, corneal opacity. mined by blood inoculation. Tick tra11smission may also be
and lymphoid hyperplasia. Rinderpest.' bovine virus diar- helpful: T. m11w11s and T. 1relifera are transmitted by Am·
rhoea/ mucosa! d isease and bovine ma!ignam catarrhal b/yommn spp. bu1 not by Rllif)icephn/us spp.. while the con-
fever may all be d istinguished by the presei1ce of ulcerating verse is true ior T. parua and T. 1a11rorragi.
or necrotizing inflammatory reactions in buccal and nasal A number of molecular. biochemical and immunological
mucosae, and conjunctivitis. Contagious bo,ine pleuro· tools ~ available that enables differemiarion between
pneumonia and other pneumonic conditions arc character- 111eilerin spp. l'ol~ mcrasc chain reacrion (PCR) combined
ized by coughing and evidence of pain o n respiration, Acme wilh oligonucleotide DNA probes detecting ribosomal RNA
trypanosomosis does not exhibit diarrhoea or severe respi· sequences can be applied 10 !)'mph node. blood and culture
rator:y distress. In babesiosis and anaplasmosis. anaemia material. even when only small quantities are available.2
and ictcrus are the major presenting signs. The clinical pic- \<\'here larger amounts of 0:--1A can be purified from
ture in East Coast fever may be complicated. however, by schizom-infected cultures or piroplasm-infected blood. two
secondary respiratory infcc1ions or by concurrent infection additional methods are useful: r6triction fragmem length
wirh other haemoparasites. It is imponam for the im·estiga• polymorphism (RFlJ>J patterns obtained when an extra·
tor to look beyond the immediate presenting signs for the chromosomal fragmen t is used as the DNA probe: 70• 106 and
primary lesions of lymphoid hyperplasia and subsequent glucose phosphate isomerase isoenzymc panerns.3 • 94 Spe-
depletion. cies-specific monoclonal antibodies used in the !FAT are
A definitive diagnosis of East Coast fever is possible if also valuable for recogni;dng piroplasms60· 72 and, where
microscopic examination of blood and 1}'11lphoid tissul's re- available, schizonts in cell culrures. 20 96· 132 More recent.ly.
veals largl' numbers of theilerial parasites. If only small species-specific PCR assays have been developed 10 distin-
munbers of schi:i;onts and piroplasms are present in asso- r.
guish p(lrva, T. 1mirorragi, T.111111a11sand T. bujfeli. based
ciation \\ith R. nppendiculawJ ticks and severe clinical ill· on primers from their respective major merozoite/pi ro-
ness. rhe possibillty exlSts that the disease under plasm surface antigen (mM PS:\) genes. These assays a re
investigation is not classical East Coast fever bur is caused more suitable for routine use on large numbers of samples
by infecrion with buffalo-derived 7: parva (s ee Chapter 30: from the field than the methods described above and are
Corridor disease). Where schizonts are few and a variable undergoing validation at 1he International Livestock
number of piroplasms are encountered in mildly affected Research l.nstilme in Nairobi at presem.
animals. the presence of another species of Tlteileria should The differenriation be1ween the \'arious disease manifes-
be considered, or it may be that the T. parvll infection Is in· tations <:allsed by T. p(1rl'll ls 1.Jased on epidemiological.
ciclental. For example. Zebu-type calves in areas endemic clin.ical. parhological and parasitological factors. and on the
for East Coast fever tend to undergo moderate, mild or inap- lrnplica1ions of the diagnosis in relation ro 1he animal health
parem reacrions on infection which do not require treat· control policy ()f the cou ntty. Veterinarians in cotmrries that
mem. The presence of piroplasms in association with have eradicated (or are on the ,·erge of eradicating) East
anaemia. minimal lymph node enlargement and fe1,·. lfany, Coast fc,·cr are far more careful to make a distinction be-
schizoms is Ukely co indicate 1: muums infection. tween the forms of dlsease tl1an those in regions where East
To distinguish between the differenr species of Theileria Coast fever is endemic.
microscopically may be extremely difficult when relad,·el)
mild infections are occurring. panicularly as mL~ed infec-
Control
tions are frequent in the field.f" the most common of these
being combinations ofT. par11a. T. 1nurotragi and T. mma11s. Trea tme nt
Morphologically th e piroplasms are difficult 10 differentiate, Progress In the chemotherapy of East Coast fever has been
except £or those ofT. 11elifera. and the schizoms are indistin· reviewed by Dolan.4 ; The disease proved rcfracrory to treat-
guishable. except for those of T. mraans. ment for many years umil '.l:eitz. 11 • in 1953. found that
-!60 'RIil>~ "'": l'ro101.-0aJ tlist3$CS

certain tctracy<:lines had a marked suppressive effect on the lnfectlon.ll 8 Immunity wlll develop 011ly in animals in
schizoucs if administered during che incubation period. Tet- which iu!ectlon with schizoms has been established. Effec-
racyclines have been widely used in 1he treatnientof clinic.ii tive methods of lmmuni7.ation arc achieved by controlled
disease. sometimes to good effect, but 1hey have not proved Infection of cattle using ~pQro1.oltes derived from ticks. or
co be completely reliable therapeutic agents. They have schizont-infccted cells. '!'he infecti\·ematetial isvirulem and
been used succe~sfull), however. in immunization against lhc method ofadmini$tration must be carefully regulated to
East Coast fever b) an lnfoction-treatmen1 technique (sec avoid a fatal outcome.
Vaccination, below). In the early years of the t:\\'enticth centul)', before the
.-\n important breakthrough in the control of East Coast introduction of acaricides. crude techniques of controlled
fever was achieved In the late 1970s by the developmem of infection were developed for immunization. In Kenya,
two highly effective therapeucic agents. Par\'aquonc. a animals from endemic areas which were thought to be im-
napthoquinone. known during Its developmem trials as mune were e)(posed 10 naturd! infection on heavily infected
993C, proved 10 be a \'aluable therapeutic agent when ad- pastures. The majority survi\ ed and could be used there-
•
ministered by intramuscular injection at a rate oi 10 mg/kg, after as mmspon oxen 1hrough infected areas.' 25 In 5outh
repeated after 48 hou~.88 RE>covery rates of 90 per cent or 1\frica. cattle were immunized by tlie intravenous inocula-
better \,•ere recorded in field trials.Jt• 11~1The drug Is effective tion ofpreparations of spleen and lymph node from infected
against both scbizoms and pirnplasms. bur treatment does animals. and up to 70 per cent were protected against natu-
not achieve a parasitological cure. and recovered animals rn.1 c:halknge. 157 Theie was a signiflcant mortality from
may remain intermittent carriers and ~ome may ta\(e sevcrnl lhcilerial infection or from bacterial contamination of lhe
months to retun1 co normal produc1ivit)',4H.fin analogue of inocuium as a direct result of the immunization procedure.
par"aquone. named buparvaquone, has been de\'eloped91 but the technique was used 10 comrol the first onslaught of
and has replaced parvaquonc on the commercial market. It the di~ease in the Transkei region of Lhe Eastern Cape
is recommended at a dose of 2,5 mg/kg for the treatment Pro,·ince. A total of 275 000 canle was inocularnd and it l,
of clinical theileriosis. 8 ' although, under field conditions estimated that 100 000 $unin•d. 1,1,
where clinical caS"es are often detected late in the course of '-o funher progress W".iS made until 1953, wht>n Neitz dis-
the disease, a cwo-dose regimen is advised with the doses covered that the administration ofchlorcerraq•cHnc (/\ureo-
being given -18 hours apan.49 mycinJ during thE> incubation period had a marked
o\ second dr11g, haJofuginone, a qui11a1.oline derived from suppressh·e effect on the develop1mmt ofinfection and per-
febrifugine, is effective when administered orally as the mitted the estahlishmem of a solid inununity. 11 ~ His obser-
hydrobromide or preferably the lactate salt at a rate of variun was followed up by a team of scien1[sts working in
1.2 mg/kg, repeated after48hours. 110 Ir is active ngainst the Kenya under rhe auspices of the Food and Agriculture Orga-
schizonts but not the piroplosms. The therapeutic index is nization 'f-AOJ of the United Xntions, and nn infection and
relatively low and overdosing may cause severe diarrhoe.a. treatment method of ilnrulmization was developed. 136 The
but the drug has achic\·cd recovery rares or 80 per cent or method is described in detail in F-\0 and Office Interna-
higher in rhe ficld.3 ,; tional des Epizootles (OIE) manuals_t. 6 ,\ crude suspension
These drugs bavc rhe disadvantage of being relatively ofsporozoitesis prepared from adult ticks whkh ban? fed as
expensh·e and this reduces their applkation in the field. n}1nph,. on infec1ed cattle. have moulted, and ha,e been
but they can be used ns a back-up to prnvent mortality if pre-fed on rabbits for three 10 four days ro allo"· the sporo-
other forms of control prove incffecilve. Experimental :wices tO maiure. The suspension is mixed with glycerol as a
work suggests that bupar\'aquone could bc used chemo· tryoprotectant and stored, deep-frozen at-70°C or in liquid
prophyiacticiilly in areas of inrense r: pnn,u challenge. nitrogen. Aliquots are titrated In susceptible cattle 10 deter-
nie administration of a single dose of bupar:aquone m mine an optimum dose for immu nization. The stabilate is
5 mg/kg seven days after field exposure has commenced inoculated subcutaneous!} into die canlc to be immunized
may protect susceptible cattle from disease \\'bile they be- and simultaneou~ly an intramuscular injection of20 mg/kg
come immune. t;i of a long-acting oxytetracycline preparation is admir1is-
To prevent t.hc danger of carrier animals. no affected ani- 1ered. Ahemath·ely, two injections oflO mg/kg of a standard
mal may be treated chemotherapeutically in South Africa oxytetracycline preparation on days 1ero .ind tl1rcc or four
withom the wrinen aurhorizaiion of the National Director of may be used. Infection develops to a mild or subclinical level
Animnl Health and Production. bm there are no such re- and the animals arc thereafter resistant to challenge. both
strictions ln other countries in the region. experimental and natural. although infection ma) persist
and rhe animals may remain asymptomatic carriers. lbe
Vaccination duration of immunity appears 10 be comparable to that
The dc,·elopment of methods of immunization against l:ast \\'hich follow~ natural recover). Suparvaquone is also effec-
Coast rcver has been reviewed by Bnm11.26 Protective im- tive as a prophylactic and may be adminislered at the time
munity to T. par11a persbts for at least five years after natural of stabilate inoctilation.90 However, tht>re is a risk that this
 Ea,t C:oa.t fever 461

drug may bt.> so effective that infection fail~ to esrnbli;:h and The possibilities ofimmuniation against r. /Tfln•o by in-
the animal is left partially or fulh suscepcible.12' oculation of schizont•infected lvmphoblas1 cultures. which
The infection and treatment technique provides good im· ho:- pro, ed so el!ectlvl' with T. r1111111/nta inl'ec1ion. have-
munity against the homologoub immunizing strain. bm is been ,tudied int<>nsivdy.~ 1 The lechnique i& possible but
not consist!'ntl)' successful against wuclatcd strains. There, imprac1ical, as VCI) large numbers of cells arc requited 10
fore effort, have been directed towards finding a master' establish lnfoct!on consistentl). \'\11ereas \1~,h ·1: ammima
<;tock or combination ofstock!. which, whrn used as the vac· inocula of Io•· cells 11ill regular!) es1ablish infection, \\ilh
cine. gi1·c protection against n 1\ide rat111e oI :tntigenically T. porl'a it requires I o1' cells in 100 ml of culture fluid to
distinct T. 1)(//'llfl rarMitcs. On the bnsis of C)(tensive cross• establish lnfect1on reliably in allogeneic animals.27 30
immunity 1rialh, three isolate~. identified as cattle-derived ,\ number of 01her approaches lO immunlr..ation is cur·
T. p11ri•11 '.lluguga anti T. pt1rl'/1 Kiamhu 5 and buffalo-de- rentl)' under im·esriga1ion. Tiw presence ofsporozoire neu-
rived 7'. p<m·a Serengeti 1ransfom1ed, have been combined rrntix1ng amibodil'S in canle repeatedlr exposed 10 field
in the so-called Muguga cock1ail. u; Th!s mixture provide:. infection. or to ,porol.Oites br ,tabilate inocularion,
protection against most i~ola1es frot11 outbreaks of East providt•d 1lw rationale for the development of a potential
Coast fever ,n ca.stern \frica and ,s used in Tanzania. vaccine based on a recombinnnt sporo7,oi1e surface antigen
Uganda and Malawi. The u:se of a 1·accine in areas other than or T. pnn'(I. knc,wn ,1~ rerombinant p6;.1t 3 This protect,
those from which lhecomponcms \\'ereisolmed carrieswich 1wo-1hirds of immunized animals in laboratOI)' trials 1to and
it the potenrial danger of imroduciug parasites oi an arui- preliminary c,idenc·e indicate, that it may afford a broad
gcnic type against \\'h!Ch local caule may have no inm,uniry. ,pectrum of immunit) again>t dilforent 1'. pan•<, isolates. 111
To n1·oid this it may be desirable 10 isolate local strains to use Field trials arc In pmgrc,,. file definitil'e ~ubunit \'Steine
in lhe technique, as has been the practice in the Eastern should ideally contain antigens from both rhe sporozoite
Pro1ince of Zambia. where a local Isolate kno\\'n as r. pnn.•a and $Chi1.0nt Mages, es1,eciall~ as lhe lattt'r is known to
Katete is used!11 In Kenya. an i,;olatc from Coast Pro1ince, s1imulatc the main e1Tector mcchanbm. Identification of the
T. 11nri•a Marikebuni. which has been shown to afford a antigens recognized by CIL is ihe main fot·us of research
broad spccrrum of immunity 10 C'<1ttle-derivcd r. 11an'<1, has towards a \'accine based 011 tht> schiwnt <-tage. 1 H
bt:cn selected for countl)"liide use. 1i:; The \.luguga compo-
nent of 1hc ''.\luguga cocktail' doc>s 1101 induce carrier st ams Other preventive m e a sures
and the capadryol' 1he mh1•r t\1·0 components to doso is not In 1he ab~c.mcc of an effective 1·nccine. control of East Coast
known. while both ·1: ptm•,1 '.\larlkcbuni and r. pan·a Katete fever ha, dc;wndt.'Cl in the past, and continues to depend. on
do cause carrier srntu!> to develop in immunized animals. the prevention of' info~tation of susceptible caltle with in
'.\lost of the vaccine stoc~ in use have been fully character· fec11.'<i tick!>. There Mc t.lircc po>.~iblc approaches:
!zed. although unique marker; are. so far. only a\'a!lable for • lsolallon Susceptible cattle main1ained in a closed
T. pnrm '.\larlkchuni and T. pamn Kiambu 5. herd on properly kneed pascures {which exclude all pos-
111e infection and trcntmem technique I:, relative!}' e~-per.,,il•e sible ct>ntact 11ith otilt'r cattle) will be protected from
and cumber;ome, but h is bt?ing implcmL·ntccl on an increas- l:ast Coa,t fol'er. a~ infection c.:an only be introduced by
ing scale in a number oi countries and has generall1· pro\·ed ticks which ha,·c, dropped irom infectl'd cattle during che
10:, t~ 1:ill 1"-' Ii .d .
~ucce.s fu.1 ,.\ qui nnrogcn co11
c cI1ain
. .• . I
tscssenua preceding stage of the me cycle. \\lwre the pastures bor-
in order 10 maintain para~ite \iability and. compared toot.her der an area or high rbk. ,uch as a publi(' road on which
veterinal) immutli7.ation., . more Lraining and expertise arc re- infected catLle may pass. 1wo tcnccs 5 merres apart \1ill
quired fonhc vuccine to he delil'ered safely and effectl\·ely. ,\n· prmidc on effcclive harrier. Ha} and straw from poten-
other important limitatltm b t.hc ri~k of occurrence of clil1ic:aJ tially infected areas ma)' introduce unfed 1lck~ mechan·
reactions afterimmun.ization."" Incorrect dntg or vaccine dos- ically to the property and must be excluded. Other
ing and immunii.ation of cattle wider stress from poor nutri· animals. sud1 as horses, sheep and goats, ma)' transport
lion or intercurrent di~easc are predisposing fac.01-:. for infected 1ick., mechank·all) and musl be treated 11ith an
animals to suffer clinicul theilcriosis as a result of immuniza- acaricidc if they ,ire introduced. Wildlife ma)•also 1ram-
tion. '.llonitoring of animals in I.he month aftL'r immunil.alion pon ticks nnd should be excluded. In situa1ions where
i, neces.sal) 10 allow 1hl'ir idl•ntification and treatment if the} fencing i\ not practitahll!, caule ma) he hottsed and fod-
are not to die. Various monitoring strategies are pracrised in tler tran~poru:Ll ro them !'7ero-grazing'J. Isolation as a
I.he ceuntries using dw infection and 1rca1mem melhod. and te\:hnique ,~ too prone to brc2kdown tO be relied llpon as
in Tanzania. 1he use of higher chcmoprophylactic doses of 1,
the ~ole method of <·0111rol ar\d normallr u~ed In con-
oxytetracycline (30 10 .JO mg1kgl to reduce the reac1or rate is junction with lick control.
ptO\ing promising. 16 Serious problems with chronic and cere- • Tick control Susceptible ca ult• can be maintained in an
bral tl1eilcriosis in immunized caule maimained under hea11 infected environment pro\·idccl that they are kept com-
challenge ln traditionally managed herd» have been reported pletely free of R. app1mdil'ulnt11s. Dipping of catr!e in
in one area of Kenya.1H arsenical prepur·a1ions at three·, five- or seven-day
462 Stcrno, iw!I: Protozoa! dise~ses

intervals was widely used in the first half of the cemury to able success 10 control tht> epidemic in southern Africa in
comrol East Coast fever. In conjunction \\ith clipping of the first half oflhis century. 16 They \\'ere supplemented by a
hair from ears and tail-brushes to expose ticks at these programme ofintensive ~'1.lr\'eillance, whid1 invoh:ed the ex-
sices. and local application ('hand-dressing") of acaricide amination of blood and spleen smears from e\'ery bovine
to cars and the perinea! region. shon-interval dipping animal which died or was slaug!uercd othCI than ac a recog-
provided very good control of R. appe11dic1tlaws and pre- nized abanoir throughout the infected area. in order to
vented ticks feeding 10 engorgement on infected cattle identify residual foci of infection. Infected premises were
al)d thus infecting the pasture. 167 Control was not suffi- quarantined for up co two years after the last confirmed or
cienLly complete 10 prevent a proportion of susceptible suspected case, and persistent foci of infection \\'ere finally
caule contracting infection when grazed on infected pas- cleared by slaughrer. Implementation of measures such as
tures.~3 Arsen ic has been superseded by organochJo- the~e requires a comprehensive infrastrucrure including
rlnes, organophosphares, carbaniates. amidines and elJeccive legislation. adequate and competem veterinary
pyrethroids. and techniques of.application with spray staff. transport. good communications. fencing, dip tanks or
races, hand-operated spr.a1ys. pour-on formulations and spray race~. acaricid<' and, above all, a stable politi.caJ envi-
impregnated eanags have been developed as alterna- ronment. Failure co achieve rapid control of che epidemic
tives to the dip tank. Cattle must be treated two or three may well result in the disease establishing permanently in
times a week where che challenge i~ heavy. and even this an endemic form.
ma~· not be sufficient to provide absolute protection. If In an endemic area the greater pan of rhe cartle popula-
used in conjunction with isolation, it does reduce the risk tion sun~ve~ in equilibrium with the disease and there is
of infection 10 a minimum. little incenrive for the implementation of vigorous and ex-
Oestocking If infected pastures are kepc free ·of cattle pensi,..e control measures fort he population as a whole. The
for 15 to 18 months, infection \,~II disappear. This period dist>ase may well cause an increased level of calf mortalit)•
is greater than the limit of S11rYivaJ of the parasite in in- and a reduction in productivity, and where intensive re-
fecced adult ticks in che field . 119· 155 Ticks on the pasture gional dipping programmes are introduced, there may be an
will contin ue to feed on ocher species buc wi.ll not pick up improvemem in the geoeml health of tl1e cattle. However.
parasires for transmission. Oest0cking was used effec- such programmes are very demanding in tem,s of finance,
tively In che control of Ease Coast fever in the early parr of organization and effon on the pan of owners and veterinary
the twentieth century. Callie were moved from infected sen~ces alike, and in the long run they usually collapse and
to clean pascure through a series of'cemperature camps', che cacrlc revert co their previous scable form. Control can
animals being checked for evidence of fever daily, and only be justified when East Coast fever is found co be a con-
those showing a rise in temperature being culled as po- straint on attempts to improve production and establish a
1emlal early cases of infecdon.u• The infected paslUre ca1tle indusrry on a commercial basis. particularly where the
was fenced and kept cattle-free for 18 months. The tech- introduction of improved breeds of cattle is involved. Large
nique fell into disuse as the cattle population grew and produce ion units, pro\ided the~· are well managed and have
vacant clean pasture became more difficult 10 locate. De- adequace finandal resource$, can control the disease effec-
stocking may also be achieved by the removal of aJI cattle tively by isolation and intensive acaricide treaanent, al-
for slaughter and prohibition of reintroduction of cattle though tl1e costs of shon-imerval dipping or spraying are
for 18 mo11ths. The pasture can be used for sheep orgoacs very high, both in terms of acaricide and labour and of re-
in the interim. duced production resulting from frequent mustering and
1rea1mem. The effects of the occasional breakdowns in con-
Practical approaches to control trol which inevitably occur can be minimized with chemo-
Selection of the methods for control or East Coast fever in a thecapy. Small-scale producers. anempting 10 run small
gh·en situation will depend largely on 1heepidemiologyofthe herds of highly susceptible cattle in the midst of an environ-
disease, i.e. whether It is newly esmblished in the ar.ea and me111 of uncorurolled ticks and iufection, have little chance
spreading in an epidemic form, or whether it is endemic. of proccccing their animals from the disease by convemional
Where East Coast fever appears in an area for the first zoosanhary methods.
cime. e\'ery effort should be made to eradicate it. This entails Control of Ease Coast fe\'er in endemic areas should
the vigorous application of conventional zoosanitary mea- ideally be based on an integrated programme aimed at redu-
sures. which include an embargo on all cattle mO\'emen1.s cing the challenge and increasing che resistance of the cattle.
imo. within, or out of the area; enumeration and identifica- The use of Zebu or Zebu-cross cattle, which have an in·
tion of all canle in the area; and regular inspection 10 detect creased capacity for 1he development of resistance to ticks,
evidence ofinfeccion. In addition. the enforcemenc of short· should be encouraged: they will be subjected to reduced dis-
interval acaricide treatment of all cattle is essential to sup· ease challenge and less rick worry. Exposure of calves to mod-
press the tick population and reduce che opporrunities for erate tick chaJlenge in early life should be aJlowed to faclli1ace
transmission. These measures were applied with consider- i.be developmen1 of tick resistance and to permit infection
 East Coast rever 463

,,ith tick-borne parasites of all types while lhc animals are In considering any control programme for .East Coast
partially resistant. The use of acariddes should be limiled ,o fever fl is essemfal to remember tha, the disea.~e does not
the minimum required co prevent excessive ,ick loads: this occur in isolation but forms pa.rt of a complex of tick-borne
\\ill reduce production costs substantially. Chemod1erapy diseases and tick worry which may all have an adverse effect
may be used to treat clinical cases. Where such measures fall on the health and productivity of canle, especially im-
to limit the prevalence of clinical disease to an acceptable proved, lligh-producing breeds. Ullenberg 162 h,is reviewed
level, immunization by the infection and rreatmenr method the results of field immunization trials in Tanzania and
should be implemented. The 'Mugug;i cocktail' of isolates has Malawi in which. once East Coast fever was controlled.
been well tried and is readily a,·ailable bm may not protect heamva,er, babesiosis. anaplasmosis. T. mwans infections
against all strains in the field and ma~· introduce new strains and tick toxicosis emerged as significanrcauses of monalicy.
into the em1ronmem. Selection of local strains for immuni- In an endemic situation. imensive dipping to comrol East
zation avoids the latter disadvanrage but necessitates the Coasr fever will disrupt rhe acquisition of immunity to all
characterization and cross-immunity tcsJing of a number of tick-borne parasites and will lead 10 an upsurge of tick-
isolates co establish the mosc effective combination for use. In borne diseases if the dipping programme is interrupted for
the short cenn, immunization of highly susceptible cattle is financial or political re:isons.1 ~ To be lasting!)• effecti\•e.
the method of choice for the control of the disease until an control programmesmusr be aimed at tbe whole complex or
integrated control programme is developed. ricks and tick-borne diseases.

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rm,rblftlgj tn \he .nllv•ry slru,c!s or Hhipic,pltnlr,$nppe11dlcula1t1S 1lclos
Eplwotill>-
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; ll'A.t.n\,'t.~, C:,L., Ut.AO:, $,I,, n.ROWN, \\,C:., C0:0:kAD, 1'.A,. (;()OOtll:l\tS, 8 M.,
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5:l~UlfftA. ~.w.. IAI.OR, V.A.. M
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8 th\lJ)W'lS, r..L, C'U')l)l")Jlljj(JS, B,M.. ,.. MOIU\l$0~. w.1., 198;=-. Bovine he-lptr
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ftl\'Cr ITh,lll'l'fa part'<I) of ea11!•.Jo11mnl of l11f,c1ious Dl.lco.ses.107, Epl:ootk 01,,,.,., q/Ajrlc<1, !3. ~7--1~.
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buffalo {Sy11ccnc,caff,t1 to tn/C<lnon \\ilh Tl:rl/cr/11 1,arr,a IThcllcr. 19(),1). tr HUUIMI, H .. h..'-ds}. />1nctlt:t1I Tf~'tlt' 01/ltlrt AppJ"'11io11s. New Yot~
8rt1is/1 Ve1erinmyJoumn/, 1:12. 379-386. Academic Press,
i3 &\R~ETT. ~f., UkQCK~flY, u.w. & \1.DLEJl. B,O .. l9fil, S1udJes on 24 UKOws, <;,G.D., 1981.AppUcotion ,.,r in ~·11,a 1ec.hm.ques to-vaccinm1on
46-l ,rr.rn ·~ mo: l'rotozoal di,ca,t'l'

olg;!flh: tlleiUerto,-l!-. Im ur•r., , , ..1.~ ~u~ ,,:-.,.1&.ut. ~.ri&l'Ov,v. , .")., lc.~,l. CryvptcSC'f\Jlit,n o(lnk..:1h't: pan1dC!> o! 11,d(t'f-in ]}<ln'1 t,m.·m(l!Jo,:;a1
.-\dnmces In thr• C:QntfQ/ ofTl1t·llor1fJ>;s, llw l logu~. OoMon. I on don: form,aI ufP,unmu Iogr J. 583-3R;.
~14rtiOU,"' ~ 1Jl1off PuhJii.hc,•r,,. , It Ut ,1'\trU,1, J.1' ..... \lOUl.lO'\, H ... l~i.J, l{C",,pOn-.t.-sofchc bu\'1tH.· 1\·mph,ifl(
;:5 tumw~. c.r..1,.. l!JtlJ J1w-,/,!r111 l}(mW. tn~ Jl'1'"' "-· J.U (1.•d.1. In \"1Uo ..,~ ,h•m to Jnfc.oetJun h)' 'l11(ift·ri1J ptm'tt. 1, IH~rnlr)g)' an&J uhra...trut.1ur<' u(
O,J//rv,//tm ofPrmn:.u,111 l'nradtt,, flo<:11 R~mn, Flurid., C!\C f'rc» Int\ I) 111ph nod,~ lo 1.·~pcri1mm:aJJy-tnfott\""t1 c•l\<C~. Jmmml u,1 Ccm11mr1m1·t
:..~ KAO\\' <~r,.n.. 19tJ_; Jmn11.inJ11o1tfon~g.;iin,1 F~t t:o..t..t ft,\~t./11:11:t\'J'i:, , "·~ Pml:f!lll'HoB..l. :?~t-298.
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:u UOklUt>CiE. ~,.,•.,. a.J\HtUl c.u ... 1973. Ourolion uf sc-1-<>l1>¢c.1tre...}k.m"it.: u-.

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pari-n Jnfec1rnn. li,~M,.d1 in tt:mmJr;· \c11•1h 11: 1-l. 270-:.!~l. .;.1 ou torr. P.1. ,c., \'JI 1or, r.tt.. 1p). Oipp1ng .l~ .1 mC'lh<xl oc uadir-...··Hing
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57S-~7i 11.nmum.• rl~pon5e.'tl ro l1wifrrlt1 /No'Tt'fl lf.J.)I c:o.ao,1 r.,.,..cr> durinw
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fnUY1u11ional Jounu,I o/Par&acl~·. 16. 22'3-~0.
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39 (.(,)~M.AU. P,, ttU •),tC>l\or, O.J;'., 11.U.n\\'I'.\, t' ,I., lltlt..-\,. T.r•• (l'C"'..\U.AGII ""'·
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\otl'<(';.A.S,. 198~. Ch.arac1eri"'11ltut of bulfJ.IO..d(!ri\'l'd thcU!.!rial p.a.rol\ite>,
dah)' f.amt'- iii ~1ur'no~·:t r>i:..rric1. Xtn~'ll: ,1t:t0';,•:«!C'tfom11 ~,ud\·.
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S9 (ill.w, (A,>. .. J•1-m1r. U-Jl, • \1CDut\ton, 1.J.. 199'9 I ht• inc1dcncc, calf
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U (.OWUiW L\', • UJ.\1, .\,\\ l9.,U. :.tudihl')ll ~ 1 Co.,~\. kyc.•r, 1.1 h.t Jif'e \ ·i!;uin:1,:.· \lttl11Ci11l!. 39. t;;-;g
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n. 1 .r. KAut .,,•• 0.1. ,, 19;3, 1'1r~Uerl'1' pan'cr. C.omparu.tl\'c lnrC"Cthiry of a anu.s,en ..\.lul«41f<Jr ,md /Ji«iu·,r,(4'fll P11rns1tology. ·lO, 10..l-l l:?.
ground 1,d. -.iobila1r 311~ • das.,ic;il IO·ticl. ,hruf,ngc. 11,,.,.a,ch 111 nm,u:,, Jl.,i...~ . l~b Th,·
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E\aluatin11 "' .i cell culture schlzo!lt ..ntlgen tb~ und ,ror.'<l In taU<ic.-d b}' Tlt~·,tc•nn /h?rl'U . 1 II,~ h•t"•rlnut)' R,wrtl, 126. 255-262
,u,p,11,1011. IIN<fln:h /11 Vtralnar,· SC'ience. 33. 360-.16.'i, 81 uc11n,·1u.1.r,,(i .. 1~11. ltrcllmlnar~rcommunlt'l.ttionon the fixing of
62 oonon.JlJ~. n,M .. )tOURISn~. w.1, t. ruu. A.J .. 1906. Gonomtlon ofbo\in~ compll·tn~nt in horse~kJ..ne$\ and ~1 Coast fC\·tr. Report of1/i,•
~,010.~fc c~II !in..,. ,pccifie for n>ll> lntocmf \\ith the pn>towAn \Mra<!lc Go1..'M'm11to1J1 \.·eu•1i11lll)' &tth.'fiologist. Trannw.ll. 1909-10~
111,•llmn parm •nd rl1'tdc,lon by produci$ of the mnJor s, ,ouxssuR>', ,.,., 190, Tr.u,•mhsion or .vn= Co~s, f,•-.,. ,·'6'fl,11/wrol
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12-13-12-19.
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63 t:R..~:Y. C.C ,1. ~Ottt::JtrSON. W., 190,. Rtport tlJ)CJII T1!.,ll.t Fnorrr,r Rcd11.1tr}t'T iu Cuhl\'tnton fn 1·1tro of bo\'ine.splcen cell lines infected nnd 1run:.rorml!d
F/Jr,x/cs/11 Copt: Town: \rgu$ Prfnrlng & Publishing Compru1y Ltd by Theih·rlo p,Jr,·n Tro,,i~a/ 1111,mnlff'fl/llt and l'rod11ctlo11, 2, 139-145.
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65 JU/LUC.at. L, W1U)l'~ J,l--.U,. fthO\\'' C,C.,0. ~ TU11~Ul. t..., l~b4 ;\lod ..•or r:a.,1,rn. Ccmrol and S0111/1cm rlfticn. :,:a1rob1. i.:enya: lmcm~uonal
mulupllc:nion of 'ntc'tltuu, jn, cuhwes of bcn·mc lymphol.·yue cells. Uws1ock Ri'>e.vch ln$Ulllle.
Sow,.- London.203, 726-730 8-5 'I.\\IE.. '\l,G. t>QL\~. TT., JIJR.\. W,<l.l.\8tt.. Jt. ~· FLO\\"i'R5, "-f,J,, 198~ A
66 WIS,,J.,P,, '\OU~.(;, J.R.. '\;f~f.. \., IJF-.\1, f. \\ll5TLK, f',, 01 f ·!ltOl\01, 0.K. k comp31ill1\'t <1irudy of tht.• dtw~M"'- m canJe c.sused by 71:rll..~rm pan'() or
,1uwx£. ,\.I,. 1990, Ch•l'llclcns,,llon of lhc-gonc ,'l1tudlng • T lau'TtJJtti! II. Hacm1uulu~')', clinical ctwmi$il)'. coaguhJ.,jOn ~u.sdih.
104-l<Jlodalton mlcronemo-rhoptry pro:eln or Tl,e//rno parm. ond compli."mtm Vtterinlll)' P11rasrrQ/r>g>•. 10, 1-19
\folccula, mu/ 8/Qt'ht·m:(o/ Para.<110/og:,. 39, H..f.0.
86 'IRO<'<>,-... 199,i. The lnf~tlon ai,d treatmem m<tl>od or immur\l'llllOn:
&; uw1s, ,\..D•. 1,S;. Chamttt•rtr.i1dor. of 1-p-t.•de:s and ..uujn.., o( Tht•ilf'rln pr-urtkc 3nd prob!~~ lw MO~AKI,\, ... 4, \\1W..'\.\L.."'-l~.... (t"C.h ,). U,·,t
M,wm:es ,,. Pt1rn.,lro/1>gy, 26. 14~19,. mcci11,s/orThollcM• purva. Dcp/Qpnt111 In Er"tcm, C4111tr<1/tnrd
68 IR\1~. ,\.0. .L .MOIUU~O~'. \\.I,. 1987. lmmunop;:uhoJot)•. lmnHm.Oll)g}' n.nd S<>mhcm Ajri<n ~alrob1, Keny:l:. lmcmatlonal Lk·es1ock Research
lmrnunopM1,h>·l:,,:t,; or Tl1t/fnll2 inti!<t!on,., /11: sou,.""· ,.. 1.1.., t~d.\. tosrnuo:.
lmmwt~ Rd.JKlnSd In Pnr"irtJ lt1ftytf.D1:I,; tmmunoJ,,10,, 8; '1C11AtU>\", , .. 1.~89. BuparvJq~1ont. tht newantitJu:UC'ria11 u tc\1twofh~
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Ro1on, Florida: CRC !'11»-<, In<. u11d S01111r,,,n Vrim Nairobi. ~en)'a: lmemn1tonal Labornu,~- for
6g JA1Ut£t"f, \'I,_£ .11 .• r:n.101110..,, (;.\\. &: MA.lll. ll.,.L, ig69. Thtllt•r/JJ 1"ffl-'d; ReseJith on Animal Oheu,.,.
~ine&IC'$of rc;,11t..1!inn. E:tl}l•ri11wnral Par(Ultology.1,,1, 9-2.5. 88 MCllMP\'. , .. H.\IC.H, ..q.11-. 1. uou", ~-T-1976. Chrmmhl•napyof J1rrJl;~rzn

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D~A molerul~ oft. f,art•n h~ scrambled rUru\ ttequenas .md Ot'Jtin 89 !.!CHAAr>~. , •• IIUUSO..~, A.r•• MOR<'•.AS. o,\\·.t .. R..\L ll.C. & OOLI\~. Tr.. t~3.
ffi!dlng fr:,mes for rnilochondrlall) encoded pmreJns. EMBO Jtmrrwl. ActJvl1y oi 10 t1,1phtl1oquh1011es. includlr.g P""-'quon .. (\19:IC'J and
I 3, 8!18-905 mt•noc-ton.,, In cauh: anlficiall)' in!l-ctL'1i •,\1th T1rc.dlttln pNn•o. Ht1S.,'<lt(h
71 ~.\ftfU):J D,P.. YOU SC. A~$.. ,\tOR7.AkJ,\, S,P., LU.\S1 .\.C,. Ml~IS<.i, S.~ •.
iu Vtwrlnn!) Sdw1c..., 3.S, 3-ti--352.
OMWO'tO. 9,, \\'.U-"lll.A. r.L.'1.J. .}lOLTh"UIX, D.H,, 19i).)~ Tlrtllcrfr: pan-a go MCHtunn· ~. t. \\a..n.S.o\.1..s.• 1~5., aupr,rvnqu,one rB\V720CJ. t\ nc,.:
C.ltrh..7.Stma- in nuwmll>" i11.fcctcd .and anitidally in1muni.1ed c~mlc. llnllth..,ilfl13l n:.1phthoquino11c lli ro?e: En ;he therapyund prophytu,:-. or
frop1uil :\mmot J-J,,afrharrtl Pfodu(lto,,. 27. 15-25, lhelltrio,c;l). /u: 1KV111:. A,D.. (cd.J. J,,ummiJ:nr/011 ii&ttitm 11,cili!.ri0Ji$ Jn
7Z K..-\1 E.:-.o~. J,M,, {'.()l)l)ff..lU~. 8..}1 .. ).10:RZAnl!\, !'!,P,. ~l((),'(,f (".C.. $. MIJSOXE.
Afri,t,. ~:tirob,: lr'Ul'rf!.\HOnnl u,bonuory forRese~ueh on An1mr:iJ
OiSC;t..,es
>.,1 19!t0. Idemtflrn11on of~ T1u:ll.eria mmans-~pec-Uk .mnsen ror use tn
•n antfbod)' and :m1lgen drtocnon ~o\. l'nf(0;1to· lmm1111ulogy, I?. 91 MCHA.l\O\' S-,. \\"l:KUA, I .S,. IIUD~O:-;, A..I, &: .RA~:lAU. /1.,\\',. 19,8$,
:J!l->33. Anut.lwll•nal nc1J\ ,w of sw-noc bup•rv;r.quonc.A comp•n~ol\ \\1th
:J x.,n,"'t)f, f,, \fOR'l.c\Rt.\, !', TOVL P•• OOt TO'S, ll S[Sli, \ , ,,;,o'SG(, C. 4 pnn~o1quon~~ Rt':INt.rcfl 111 Ve1c11Ju1Q' SttenC<', 39. ~33.
,iuSOJ.l, \., 1998. ,\11 c111)1nc-hnk~ 1mmunosorUcm as..~y tor dttt."Ction 92 .t>.J. ;\ftJr,(, H.1\.1.,1 , .. ,,~u. LJ.., !-tA.CHU(;>t, !'1;,0 .• A\\T\O, r~.
MC);U"l."1.1':,
o(Tlitllmn 1,ar,v, nnubodtos in c:mle U$lnga rccombmam 1>0lyniarphfc c;onoruu, a,M "" MORftJSQl\". w.,., l99J, Adopm·e tra.ns-(1:1 or ,rumunity 10
tmmunodonun3m n1okeull,. /'tU,7.«:ology R;•S((lrr/1. 84. '108-l !6. 77rell,n.• p.11rn rn 1he CD8, frnclfon of r~pondJng •ffor<nt lymph
r .1 t:QCU. "·· 1tl9&. Rl!ls,rlN,rftlm· OJxtr Rlnd,·r~r. 8ul,Jqm•11pNt Ill Jnrhru und PrtJ,xedlngs 0/1/11• Noao11n/ r.<(1//mwoJS<IM<t US:\. 91 19oS·l9t'l3.
.-Vrf4·a. T~lst>-odttr Stlfflll.·rankhelt, TeA·tU/iel:J..,r, 1mpl•r.;he .\lctlari«. 93 MtHUIUH"li , 11. '"- !"';.lltl-.:. r .• 19a~. The J>iroplusm~: Ufe c~'Clt.!tand .s,·,u~\J
/klr«·or:u'(I.S,-rjiewr. Borlin: J. Springer «:tg,., .-td1'111rrfs m l'nm,itology, 23. 3;- 103.
-n xocu. ~-, lijOJ, tnh...rlm rcpon on Rhod~1-l:tn rt"dw11:tc:1 C)r · ·\!r1c.,n (:..<r:t.)t 94 >U:<OQ\E, T "-., nsmvs. e, .. u.. 1!17<1 1,oent}me varfa1lon In plropl•~m<
'•"~,·. /nurnnl ~I C,,111µarnrh'e l'a1/:o/ogy1md nrrmp,.•ut/C1, l 6, 2111-280. l,olm-d lronl bo,inc blood lnfcc1od with Tl:eilttih n11111tlllM :ind
76 Ll)U!Cll', G~. lCIOJLS. a;., \'A~ ,•_UJt.L-.:nau,;11. a. & :O.L'-.\IMt.fllC.K.\'. ,, •. 1~r~o. r p1m-a. ltt1eo1th ;,, \'ewri,tm.,. •w:1,mct. 2'"' 379-38l,
Scn,ibilil~ du hum., d'Asln aux princtpal.. molnd!,·, b 1>ro10,.o~;,..,. du 95 )rn'1' \ "' ft\\·. ,1. &l.'-.R\JlCIL\LL, 1.. 1936. lumtng-"d:Jie~ n µrotozo.ui
br1•il •u Con8u l>elgt. Es-<ai• ""Jl<'rimen1;1ux , i oli.~notlnn< cltnlquf< enuph.Ji1i, nf ,anl< ,n Uj:(uttlo; ft< reTatlon;hfp \\ilh i:n,i Coa<1 f~<r.
,4111,n/o, d~ h1 S<it/Jte Belg,• d., Mo!dir/11, Tro11/c,1/t, ~O. 189-IS, Paras11olol:). 28. 25-1-?83.
r ~ L\\\~Cf I.,\.. 1992-. H{~tot;· ofbo\i:n(.• thuilerles,is in s:outhc-m Afriru. /n: 96 )UN-\.MJ. l •• \ro<):ri.,·t11t.. PSI. JR\'lS',A..D.,OCA.\lA.J.(i.R,. 0081:.ifl.At'R(. 0.\.t.-a.
SOl\\'AL R.r\.l.. 11£:RI~. ll.t>. & \'OUSG, .\.S.. Nb . Thu F.t1ick'll2tol~· o.f FUJl~AG.\. 1 •• 19&3, Ch.arac1c:,.rl1i-a1lon of Stotko of Tlntlh·rio pa.n'O b"·
n,.i/,•rio.<I., /11 ,',fr/en. 1.nndon, Acadc,nrc Pt~-'>$. rnonnd<">n,d ru1ubndy ptofilr~ Rt'$t'Urtl, In rtrl"rlnm,· Sri,111r-t. jS.
334-340,
78 l.\W.R!NCf., f,1\ .. :.:Oft.VA.I. $\_\.I . .& UlU..-.:Bt'f\G. o... ,933. Rhiplt'(pl:nlus
:.tm11J1,.•;{t11tl~ 3,. -:a ,•t.)(-torofbovlne thcllcriae:-. frQJ>fc:<11 Anfttml n,:alth 9i MOHAN, R.~ .• 1968.. Oi'M~:\t«:Sand p~nisit~ofbulT.alot.:S., P.atl (I(. Por.isltic
111:tt Pt()dt1ctt'n11. 15. 39~. and mtsccllaneou~dl\castl!t \ftftrt'infl'}' Bullt-1/11. 38. 73..:;.....756,
:n unc.u. au. & ,·ou~c, A,S., 1981 nrf'1l.uin in(t..'ClianS- in Rl1111fc1.11l1Q/us 98 MOf..'~ c .. AC-AS, 1. & l,OIUUS(;, /\.. l~S-s. BOVlOC: ccrt'bru1 thcif('.fll»U 111 pun.~
fllJIJtmircutaws uck1 colle<ted in the field. Im un 1s•.\ 0 .. cus:,\~1s-<;H,\,\1, Unmn nnd $ahiwal,cro~~ cnulc tmmuni~ ag..a.tns, &s, Coils\ fever -tnd
\l,P.& >'OUMl, ,.s., tc:dsJ,Advanr~rfl tltaCom,al o/1'11rll<rfo1l,. The kep1 under conunuou, field cholhmge. In. m,,, ••~~.. r,~1.1.
Ha,rut'. Bos1on. l.ondon, M•nlnus ;>;ijhoff PubllShcl'$, Tmm:ml::ntion Agm,:.si Tititih:rWSJs m .l{rica. -Xrurobi: lntcmotio113J
&) U-)'!o;\AD, P, L'l'.PLATl'tNlER. f',, SOR\''\t... fJAt.. lCVNOfR"T, Jr.'., 001....A.~ T.T••
1.:ibur.uo1, for R<:.w.ud1 on ,\Jtlm.,1 o ....~ .
c.R.01.C.. ,, •• W.\Ui.U. J.6.. IK\1.:,.0. 4\.0, fl'. i'(H.ft\'. (\,J)., 1990, (j(•ographk:d 99 ,,ou. o.. LOJ101:-.<;, ~ 11,uoso, 1o.s.• 198-~ f.pi.demioiogrof ,heil"r,osi~ ir,
inform"3uon sJ~tenlS forinsdymg the cpida.miolom·or caule c;U,;c-s,;.es 1he Trons•Mor~ Orn,iun. l:cny-,1. llu,b;uu!ry .md disease h3ckground
466 ».cnus nm: Protozoa! diseases

and prcUmmnry 1m--tsHpliOrb on lhdlerioses in c3lw,·s. Prc<:-cmlt~· u8 .SUTl. w.o.. 19!.i, Theilono~1s. gondtmoscs A.nd c:yt:au:a.oonctse~: A
\'oli'rinary ,\tcdicf11,. z. 801~3?. re\i~w Ond~rsle.JJOOrt Jnumn/ fJ/ \'l'ti-rlmtn• Htrc:turh, ?7 ?r.:,-J30
too ,1ou. <,,. t-,1;JUQlNt,;, "'·~ rouse., A-~·" U'JlCH, s.1,., 1.986, l::p!demlo!ogy or 119 '.t:W~U:",, 1*--M .. om·~, ,.w,, 'U.1U~(i. ,.s UQL-\~, T.'r.• r.:US'.1'"CHA..\ :, M P.il<
theU1mosb, in cal,·es.in 3n endemkeft'a o(~cnyu. v,wn,mry· KA01~,·. 1>.1:., 198.1. Suni,';)J oi l(/1/plctpltt1hu t1ppi'1t/JiJ:ulnws Ucnnnll!
Parn.<1tol<>gy, 19. 2S5-Z~.; lxadlc!aeJ and per.-isc<nc• of 77,ri/t'nll J1(1Tll(J (/IJ>icomplc:u: TheD~riid>el
101 ,,oaar.so~. \v.t.,. utl~ou;,t. (,_
f'!.1hJIY, D l-~ ~(..L'S(t~. ~t ~ ,:uRAAY, \1,. S9SI, in 1he fie!d. /nttm1uio,wl /ourna! ofPma,lro/"1:,1, 14. 483-189.
Th~ klnocfc,; or lttfl-ctlon "'th J11,•Jlerfa p(ln'<I in caul~ ond ch~ rciw.,11,:,, 120 'o'(;,!~U. P !\., 'tOUS(;, A.~., t.,.\}1P,\JU>, D., MTSCS(., $,X.. ;o,.OOSCllJ. :;,G.. U~'.1
co the.• de\'elopm<mt or rmmunit~ Jn· lfl\'IN, A,:,., CV~Nl~GlL\.\I, >t,1•. " ,.c \\'U.UA~,~~. $.M •• I~ , .....0~1. ,\ lo KAAIUKJ, 1,.p•• l,!r.)2. Further
1ouM;, A.s.. (eds). Adivmm 1111/r, Comrol u/Thrilmo,ls, The Hogue. ',"vah.i.ation of the u.se of b~pilf'\';.tquone in the infoc,i,,n and
Boston, Lllndon; ManlnU5 :0-ijhoff Publi,;hcrs. 1rea1mcm method or hnmunlsrngc•nlt ag:,.lnst 1'/t~ikm• por:w
102 MORkt1;0S, \\'.I,, aO~:flr:.R, (.,.MURRA\", "'·· fl..,\1l!IIY, l.).t.. '"'·'KE, Pt.A.. r.oo.:. dorleC<11rom Alric:m bulfulo 1Sy11,t, ,u C<l{fl'rl, Vturinnl)' Paras11oloro·,
11..H, • wr..u.s. P..w .. 19fh. flutfltrl.a µari't1: Kinetics of in fee don in the ·13. 15-24.
l}mphaid ~ tom of caul~ /;xp<'rlmw11af P11rru/rol"I!}·, 52. ?48-260. 1:u ~om·A~ R,A 1.. 19n. iid:. problem) tn rehnion ,o lnnd udJiM1ti0n in
l03 '\!C)RRISO~. ,,M., GOODUJU!. 11-~t, t'I.At.E•.\.J•• ~noo«:.n~. c.,.. .:OtP ",.J. 4& Rhode~"' llhod,,,irm l',rert1ui9• Jounw/, 8, 33-38.
ttACG. IM, 1987. Cytotoxic T cells ..Jkhcd in <~ulo challcni;cdwith 122 , o""'· • A,1.. 19,8. The effec<sofpanfal breakdown ofdfpplng ln
111~/ena pnn'O t~111guga): c,1den<c for re.<1rfc1ion b> das; I MHC A(ritan art'•> 111 Rh<1d c,cl~ Rlt,xk#nn V,i~rlrUII)' Jouma/, 9. ~16.
d~tcnninants tmd pJtbltl' )tr.tin !tfhtCifldt}', Pmosltc Jmmtmolo~·. 9.
12J -.:01w.u.. ,u..1." 1•11nkv. n.o., 1990 The inttoductlon, spread and
~63-578.
<Ub'<''lll<'lll dt~appearnnc;, oflhc b:o,m ""' cick !Rlti11ici,pltalu,
lo.1 MOlUUSO!':. \\' ,I .. MACliumt. s.o. I- I.Al.OR, 1•.A•• 1996.. P4t.hogcnic1rv of npJHntltcrJ/ntu.•) from 1hc SQUlhcn1 low\·t:ld c,f limbnbh'c, £,:,wrfmemnl
T/i,i/trin pnn't1 Is Influenced b,· the hoS1 .,..1111-pe infoctrd b)' d1c 011d App/i~,I AroroltJC)', 9. 103-111.
p3nt5ite. ltt/«#on and Imm unit}·, 64. 5Sf..562.
1:24 -.:ORVAL H.A.I,. Mtnnv. U...0.1 CfJUlt,\0, F4 &> U~Akl), I••• 1991. f:.ost Co~t
105 ~lOR'l.ARIA, s.e .. uw 1:i., A.U.. TAMCU.\, r~• .s1,oo-.:t1t. P..R.., \''(HGT, w.t .. re-.~ a prubfcm nr 1h1• fu ~ur..- for1hc, Horn of ,,.\ (den? Pro,-,mlt•,•
f\lJJN/IG~. T. ~ J.AT~Oe..J,, l98?. ts:nmunii..atiou agailts1 Ei"lS: Qia~1 (l''t'(•r. l'crrrl,wl) ,\lrdirl,w, 10. 10:1-li2
The use or selocttd S1ocks of 1Ji,f/~rit1 ponv, ror immunir.01!0n uf conic
u,; ~OR\'Ar. • .A.A.I .• ,•1:.1mv. 11.11, ~rouse.,\.!-. ,cos.), 1992. 1'ht lifJidt!mi"loff>'Of
exposed to ~eld challcniw, Vereru:of)• Pwnsirology. 23. 23-4 l.
11,,ll~IDsis 111 .4/rfca. l..t>Odon: Ac:tdcmfc Prt.·i.S..
106 MQM7... klA, ... ""-'Ii.,,..
MU,01'!1. A.I, i, " ISHOP, n.. 1~93, ~loclero
126 ...:vni'LL. c;,H.F, & 11t~r)LC. c., 1913,Condiuon:,,, inOucu.1cing the
biot«hnoil}gj,;al mtlhods fo r diogno.,is of bovmc thclterio.-w and fuwre
1ra.tt.8"Jnbjion of F..ast Coaitr fever. ParaJito!agy. 2. 208-:? 10,
trends.. Procg,•d/ttf§' of tar F1\0 Expert Co11s11/ra1icn 0,1 rhe UseoJ
12';' ),u~.s T \\'., WNl>lN. I ,fl., l)ULAN, T.T, l;~'"f,\(;(;, IJ.A.. Lg19. CcH-mcd,ared
.~;//>llCII/JI~ H/01«h110logiwl ,\Wltodsfor Dit1g11o#s ofI laem/Jpflrt<SIM.
fnlmunl[)' to nu•lft1r it1,t.rno:.form~ cell Unc:i.. Nm11rt, L-andon. 231.
Ocro/Jar 1993, Meridb . .We.(lco. Romt: f<>od :md ,\grl<:\llture Org:inisauon
&;8-(i80
of the Unhcd !\~tion~
1::a P-ERR't~ s.o., 100N<:, A~.. 1993, The naming game: 1he changing ronunes
ao; MOJU'AttL-\, 5, "' \\1t.U.\Msos, ~. l~d$)., 1997. Uiic t('tcclnc,-Jor TheJh.•nll
oif4~1 C-0as1f<"'cr nnd Tlrtllfrta pan'ft. 1'i1< l'rr.ri1111f)' Record, 133,
pon-a: t>eplo,.wnem t,; Easttrn. CM1,ala11rl S<Juthf·m ..t}ricn ~wrob1.
Kenya: lme.r(t3tioma1 Uvmod Research !nst1tutt, 613-616

1ot MUti.:Ut.Bt. A.W., r l!ltAY, IJ.r,." i.titUSIVI, R.. 1.99i . Eslimatcd economics of •l9 NllRl . a,n. ~ Y()tJSG, A.~ 1995. rhe pa.st o.nd fu ture roles or epidemiOIC>lO·
lhcllcrio,1, conirol In Eallt.o\frlca. Pn,i.•nrl,,.. V;rerimuy .\/wlfr:/111', 12. '1nd <Conom,c, 111 th• conttol of ud-~omc dise:ucs of Uwsto~k an
73-85. .Africa~ tiw ca~ of d u;dcriosis, Pn.•w11m't Vtflerinary .,n•dic,,h"•• 25.
107-IZO.
:og MUSlSJ. F..L, \.10RC,+\\ D~W.T.&SCH[b, Jl,F',.1935, Tre-~tmt:ltO!tMUtrfosh
&JU Pr..Ml·, 8.U,. KRUS:~,. ,C., U ...."'-)A.1\D, 1'.. SOM\',U R..,\,I, k KOSOlRT, l.,. 1991,
\\ilh p3n·•quone in Zan1blll. T!ic 1,wrfllnry R<,a,rd, I l, 338-339
E.st!moring the dis1n b11don and abundance of Rit/J>iet1•f,r1lus
no MUSO~J!• .\., M O'R.7.MtlA. s.. .SKONC-£. c .. JOX.ES. r1. & NlSC. \"•• 1992. A aµ1,cru/ic11lmus in ,\ fri~. Prercntwr Vtftnt:nl)• NTr£llcint, 11 :?61-Z68.
recombmam sporozoite, ~urfnec :mtfg~o of '/1ttllt•rl.a pan:n induc~
pro<ec<ion In cnttle. P!Wtfflln,,s of1hd .\'m i/J1wl ,·IJ:tuh,mrof!kim« USA. IJI rt>'RY. H.C>,, I.E.~\RO, 11,, ~<.rn,\',\L ..., ••• ti:u:i-.:uurr. ~- & KHU:th:.-\. ti... 1990,

89.SH-518. Ulmatc, ,-cgomlon and the dlstrlbuuon of R/Jlp/cephnlus

n11P•nr1iet1law$inAfrlc,i. Pnrrultolog; Today, 6. 100-104
IU :'otV50KJ;,.,q .. X,\~,.UI.Y-', \.M BU~CHtn.G .. ,\ ll\.'-AJa4 It.A... ()TI)I. a.. sgS:t.
Bovine immuna ,espons• ,o 11rl'il"1'io pnn,a; Neumiliztni; an<ibodws to 132 ,rnDn1 M .. llf\\ rn . tM··• 19-80. Mo:ioclonaJ n.ndbodle-,, dete<:i andg1m;c
sporozoite$. tmmunol~·. -t5., 663-668. divt~tv In 1'1rttlm« I"'""' p:ara,,11~ Jo:murl a/ lmn11mology. 124.
I 000-1 00 I
IU \HJSOXE. ,,\..J,, ~.\:'\'TUI.YA. \ .M,. RUM..'HilR\'tA, f .n. .f. u(J-~JfFR,Ci,, 1~,t.
Evidence for• common protecch'C onligemedctcrml=c on 133 PUfl~blJ.. n.u.• UOAJ\~ c.D,IL £.-..lf;k.(L M-"·· 1971, ·n,eiltritl p()rt'(I:
ftP()rozoltes of.se\·cral T1rellcno panx, nrain>.. lmmmwlogy. 52,231-238. Cornp:a:.11\C inloct!on rot<'> of •dulc and n)'mphal Rhfplctp/10/us
f1Plh'll(liculaws. Uaw.dtt>lt>l:{\1, 62. 3.;9-353,
UJ >IUSOl<E, ;.., N(SI!. \. k\SORL\1U,\, ..... lll'JJ, -~ >f)Oro:eoitc-ba,cd \"Otcinc
(or 11r,l/,rin parm Pmasuqlog)' TOIi")·, 9. 386-388 13J rUIL.~H1., R.L, IAVIN, A..IJ,. Xl\18tR, C:,D., OM\\'O"t·6.. P.L ltl1AV,...'f. R..C.. , 1914
E:lil Co.bt ff'\"er. Funhl.:1 l:\bC-\1,110~· 1n\·cstig;Jtlon$ on thtt u~ of r.tbbh~
11.: ~111soa.., ., .. r At..\ rn.R. c.u .• ,1cnwA1~. ,- t'.. :,-,."t..,-r., ,._ .. ,tO:Ft\'tat. 1,.. 1"6.
3" vcliklcs for inft!Cclng 1Jd;, \\11h ,hollerial pir011lu,ms.. Tropicttl Anithlll
Prni>pccls for subunit \'llcclnes :1g3ins, tick-borne dl;e3<<"1. llrirllh
H,:,1/1/1 nnrl Produrrfo11. 6, 145-lSI
v,,~rittflf)' /o:mttll, 1;;1, 621-639.
135 PURSEU. R.t. & JO't'N•-A. LIJ~, 196&.. Thtt devch,ptncnt 01 Tht/lrrfn ptutvz in
n.s \tt)'TIICl, 1.J.. I.A.>1P,\.RD. D., YOU"c. ,\,$.• .sou:-.:cu. $.(O,. u~·tos1 . .\.. ~HArrlM.
.,.c... Ml~ lNC. S.K.• ;i,,:Gu~u. P.S •. 1.:,\RlU) a. 0 .1·.. \'IIWA..,1so~. s . \1., A\,101. J, J.t, ch~ $:lll\':try gl;nd.S or tht lick. Rhtpk;p/:alu, 1tpf1<'11rliculnrus.
., u:s"-' · ,.,:.. 1990. Recent lmmuni;.itlon trial> otilln.u T ,,,.,.., puny; ParQ.ittofo,gy. 58. 725-73:?
lrlfee1lons in Ktn)'.t. /rr.1-ousc. A..S.. Muruo1, J.J. • ,~urrt,!. A.4=., cdsJ. ~6 ft.A.nu, . tu1.. 1.9;&. Cht rnoprophyf:ictic inHuu.nJ:mltorl dgulrt.~1 East Coast
P.rogms tOWtlf'cl$ 11,~Ct,,:rrol o{EaJ, O>a,r l·'<w m11·ll,rlosl.t1 /11 Krn.,w. req:~r~ 111~ \\1.WL.. 1.x.u.... (t"d.), TfcJ.;.l,ornl' DJ.s~IUP$ and tlzttlr V.:rtotS.
X4irohl, l:~nyo: Ktn)'• Agr1c111t11ral Rl.'S<larch lnstlcutc. Edinburgh: Unl\crstty of P.dinburgh, C:enlrt' for Ttl1ptral Vocerlna!)·
v.. M U SOKli., A.. GOBkfGHT, £. .&: M()K?.i,\fflA, .s .. 1996. Constn,auon of
116 N,£..,"t.,
~lodlclm•.
thosporo,.oite p67 Vllccine e.nugtn ln c:acclc·dcrivc-d ntdi<'rla pan'fl t!7 fl\.Otf.:\. D,£... 1901. lnfoclion ,1r.d U"t:fnmrnt immunis.idon again-st
stocks With d ifferent ero.s.s,lmmunity profilts, Jnfi.-rtinn and lm,tttmity, thrilerfo~is:. In: tR\!lN, A.n .• cur,;:\'1S'm1A\t.. ~1.ar." vnust;••,.5.- (eel:;,)
64. 2056-2061. ·tdt,'iltt(i','. l,, llu1 Cm 1ttt,f ufnicllerio.s/s Tht' H:,tgu~. Bo:1;.ton. t.ondon:

Uj :-.wr~ w.o., t~S3, Aurcomycm In 'ntf!ilt•ri,, JNlfV(I infcc:oon. 1\'mttrt. Manlnus XlfhotrPuhlishors.
London, 171. ;14-3S. 138 RADLEY, n.r.. 1981,. E1u1Co;ist fc,cor ,mmuniiar,on -held trials In
 ~1 Coast fover 467

\lala"i, /11, 1MV1'>,A.U., (td.J. lmmu11i:.ario11 Asa/11.<1 T/11:ller/o!ls Ill lofr/((J. mcan<ofblood. F/f«'l'11111,\mum/ Report QJ1/rr Dirr,i:torofVurrr/11(),y
'1ufrobi: lnrc.rn3tic'mal L.ab<lrntory (or Rt.1~C~rth cm Animal Dht".\'b.. ~~IV:<d, Unio•1 of.'11111111 Afrlw, 15-ll,
l39 AAn1· n·. n.Fi., mtOws. c.c.o... 8HRR1ovr... ~'-'•· cus~,rsr.1-tA~t !\t,11.. ),."fRl\11. l8o TO\"£. P.G.. c.QOO£E.R1S, 8 ..M., JA\tl.,;. K.V.. \IUSO~t.. A,,. k ~101-att.:oiO, \\.I,, 1991.
1...,11, NHt}<.:fiU., R.E. & ,·oosc,A.S., 197,s.. East Coast r~-e:r.1. Charocttri:;utlon of n rolymorphic lmmunod(lminani nll>ll'cule on
Chemoprophylactlc hnmunlt.ition or can le againSt nitll,rin vnn., ,i;porozohes and .sdtizom~ of111t'l/erla pan'tl. J>nra.s!rt /mmunofogy, 13,
(~1ugugal ond n,,,.
thcllo:ibl sir.tin>. Vctui11nry P,ims/11,/~·. I. J:;-41. 49-$2.
1-10 scnr::JN, £. &- vorcT~ w,P_ 1979. Chemotheo.ra11yofbovinetheilenods-with 161 urLL~unac, G.. 19;6. T:ck-bome !l,-estock dl<e.ses o,nd ,heir vector, 1.
hruofuginonc. ,itm Traplcn llas<l, 36. 391-39•1. Splzootfologyof rick•born,• dl,u~..., lt'or/d ,\11/mal H•tiew. Ii. 8-15.
tJt SOft;,,."lJLJ:R, R, & WQN\TSC:f, R,, 1.96.;, \'cnr.urhe 1,ur Difrc.renzlcrung<ier t62 u1uxa6RG, G., 1985. Possible impact of 01her 11cl<·bom.e diseases
Thel!er!enspe,;lcs de, Rinde> durch '"'rologl~cho Umer,.uchungen. foll0\•1ng Eost Coast rc,-ccimmaniY.ntlon.111: """"· A.v.. 1,..i.1.
Lefuc/irij1fi1rT,opem1Jt:d,.:;111 rmd Pn,n.sirolog,e, 16, 17-23. lmm11ni:.1t1ion ngal11Rr '11,i!l/,_•riosi,,in .'1/rica. Nairobi: lnteto:\tiontt1
142 SCHR!Ul>llR, 9.t.C. & UIU.'<lltl\G, G.. tg;S, Studl~s on Th~Uertid~e Ulbora,orr for Research on Animal Dbeascs
(Sporoio•l in ronzan!a, V. Prelirnlnary e,perinwnr.,. or,• new method t63 UILLXBtRG, a .. S.CTR£1Jl)[•R, a.c.e.. MPA!\("~U. c., Sil.AYO. n.s.. TO~DEUR, w ..
for lnietting tick. \\ilh Tl1<//m" 1mn·11.uid Thdlur/<1 n11111>11s. rA'tC'H"E.U. n.J.&S.\.~GA, 11.J.N•. 1978. Inununiutlonngnlns1 Eost Co:l~
Trope111ned/::h1 um/ J>araslrofogfq, 2~. 422-126. fever. In: w1Lor:.. J.):.H., (C-d.}. Tid:-btmm Df:si!h$r:.< anti their \'ttt:tnrl.
,.._, s11,,w, M,K., nt.~llV. LG. & >1uson. ,.,.1991. 1nc 'entry of771e/~rln par,'<l Edinburgh: uo1,·~rs!1; ofJ!dlnburgh. Cen, ,~forTmpicol ve1erinar}'
sporo,oltcs into l)'mphocyte>: e,ideace for ~1HC cl:m I iuvo!vemenL ~-todlcb,~.
Joumnl ofCcr/18/o/ogy, 113, 87-101. 16~ un.1.s.1:U!JtG. G, & zw,orr. v .. 1.979, Skin l\odulos in E.\..,:t Cnatit fever.
l"4 smTA~HJt.. V.M .. NAN"l'UL''·' · \I, \1 .• f',~USO'-:R. A.I,. f.c.\MAS;\ \JY, Jt, l,' o0S<.:tt-tiH, G.. Rt!$cttrch /11 Veti•n,u,,y Sclt1J,1Y, 26.. 2 ;3-215.
1983- Coinplement actlvolion and fibrlnol)'Si< durh1g Infection with 16$ WAGShft. (.,<,;,. Jl:..'\!-1:lT, 0,M,, kJtOW:-.:, :C.Ci,U. 4' MAOIJ 'r, 1),J:., 1975,
Thtfleritl parm (EJisr Coa$t fwtl) In canlo. V.iem,n,y Jmm11111>~•t1ttd o,mh,lshed anubod)' res})<)nSQ 10 rinderpes, Vll<cinatlQn m cnnle
f1tlfllU{)fJj}al/ru/f/gy, 4, 361-3i3, undctgoing cxpcrimrmal F.a.~t Coa~:t i<:ver ~s~n..rrh in Vtwrfnary
145 51JOftT. s.1. & sonV.i\L R..A.L, 1981. Re,gu!tuion of sc."3..~-0n.al occurrence in Seiet1Ci'. 19, 209-211
th~ tick RhiplaJp/111/us «J)pcmlktd1J1u11'eurnann. 19111. Tropkal.~mmaJ 166 WAJ.f(Jlll, .\,fl:1, }H.XE1.I..At,, :>,1)., e:~J 1,. LJ. ~ 81\0\\'°', f"...,(;,I>., J9i1). R3p1d
IIM/tlrtmd NMuc<i(m. 13, 19-26. qu;anmatln~a.s&:~mem of 'nieilerm in(e1:lio11 in ticks. rm,ucalAmmnl
\\'.J .. BUHfUD<';E. M,J. 1"' 8KQWS, C~G.J),. 1973·, Some
146 SP00~-.;1:!:ft. R.1... f'U.."'"IIALC!. 1/Mltlc and Prod11c11011. 11, 21-211.
serum globulin changes !u East Coasi fever. R~eard, /11 V,:,cri11ary
167 \\'ATttX·S•J"fTCHfOftf), 11, (IS.~.• f-1911),;\,J 11/unmted P(lm,,h/010,1
sc~m:e, 16,368-',374. Tick-de,,1rucrio11 nn/1 the Eradl(nr,011 ofE(:Jr CQII.« Fe,-ernml 01/ier
147 61'1Uiuu. r.. 1$14, !:a,tCr.»w. fever inoculation in tbeTronskl.~an S.Africn111)4,•nse,< by f)1pping. Na1al: P Davis 1< Sons.
Terr!torlco. South Mri<::a.J()11111al of Comparoti•.,. P111hologycmd
16U W+\1T, o.. ~J'\.RACA=<01 0,, IJUO\\"N, C.C,t>. & \\'rUJailt, /1.,R,.. 1997, U~t! OfthC
Thempcutks, Z7, 2'Jll-304.
pol)'tncnb,c. thaln rcaciion for fdcmtifk.:uion and qu,uuilk.itlcm ()(
i.8 STAGG, n.n.. 19112. SrudilJS o{Thcl/crill/ Pnrnsl1es o/W1:rerb11d· /Xobu~ Thellerla.panl(l prQtQ1.oa In Rhlpicehalt,sappe11dfculmHs licks.
defossa) and thefr Possl/ik Ro/~ m /11~ E1>1<1em/ology ofF.,141 Coasr Fer,,r. P<1rnsirolog)· //~~ch. 83. 359-363.
Ph.O 1hesls. Uo!vc,slty ofS:ill'ord UK.
t69 \\'EDSTER:, t•.. nonaev.E!.Rt. D.i\.e.'" F1\\\'CF.T'r. o.w.. 1985. ·111e emry of
l.C9 STAGG. O.A,. 1)()1.,.\!-:, T.T.• urrcu. 9.J.. ~ \'OUS-G. A..S., \981,. The initial S-lagei spo10,oltt1S o! 11..1/trla pnrm Into bo,ine lpnph0()1es /11 c,frro.
of infection of tattle Cells \\ilh Tht'i/eria pa.rm sporowltos In uitro. lmmunrn:l~<:uon n,lcro~t'()J)lr ob~cn,uion~ limo1~w Jot1rmtl ofGW1
Para.t/rolo~·. 83, 191-19;. Biology.36. 157-162
150 S'tt(;1'. w., 1928, I i!Stolog,cal nudle$ onEast Coa.t r~v.,-. 771irrrarr1J1 und 170 w1w, .• J,..-..u,. 1966, C-.h:mgC'\ in bovine bone: mtin'ow during the cc,1.u,c o!
T'Ourtrenth Reporcs ofthe Dir11uor ofWrerin«ry Ed11ra1io11 rurd R<1$l'nrc/J. EBSt C:03~1 fe.·er. /ll'J<()tth i11 Vorl'ri11ary St(<'llCI', i, 213-224.
Sourll lofri<a. 243-282.
17J WJU:f!. G,M .. IIROWN. C..G., J:IIWAtt. R.f. THo,u,-, \t,, \\ UJ IA\ISOS, S,.,: .•
151 5VTHF.RST. sc,,,,., & ;\lM'\\.ALO, c...., 1985, A C'o,mµuterited $.}'Stem for llEIJ.-,,L)'I, ... ,. ~ SMM(',Ar{(I, o•• 1998, Chel'(lopr<lphylald$ o!111rilerifl
matching climates in ecology. Agr,cultun:, /:(:f½yStilmJ mid E:m,fro11mem~ n11nula1n 311d n:tileri111x1r1,n inf~tloru. or eal\01 with bupamaquone
13,2111-29~ l'ereri"«f)' Pamsiwwgy. 78, 1-12
1.5!!. TI\Rt\CHA, 1;..t.. ~., C,QODURlb, 1M,!,, ~lORZAtUt\, S,P, &: MOJlfUSO:-:, \\',I,, 1995--
an \\1LSO~. s.c.. 19.u;. Some rac;tor$ affttcdng the incldenrl' ot F..ast Coai.t
1'1\raslte >train specificity of pr~·=r •1•1011>xlc T <ell• In IndMdual (c.....'Cr ln ~•>rthum ?ro,ince. :,;!yri:t'al.and,Jrmrm,t of1/Jr $owh Afr/em:
animil,is correl<ltes \\'ilh cro»·protcction in c.mlo challons,d "'th Ve1tri11a7 .Wetllcnl 1\.tS«i111lon, l 6, ·17--72.
11reflerla f)(lrc•a, ln/1'<11011 n11d lmm1111/ry, 63, 1253-12&2,
\'t.-'(U.>r !)lud!ei. of t'pi-:,oucic [;.J.!)t Cn:l.(t fto\l.:t. I
173 Ytro.,:.,!':, c,11,. 1966. Field
153 TATCltllU, •• ,. ~ EA$l'O!<, ·-1986, Tick (A,:arl: lxodid••l ecologlcal 6tUdk>s A qll.o.mha.th·e reladonstiip bet\\'l.!~n It npp£11:d!culn1us and tho
In Tanunla. B111/eri11 ofEmomolog/aJI Rest<1tcl1, 76, 2:?9-2,6.
opl1.001lcfiyof E<1,1 Con.st fever. H11Jl,1i11 ofF.piuxJtl< Dim,s., ofAfr/m,
15,,t THEILE1t .\ .. 19011. En~t Coast (e\.'er. Tfi.lllSl'(UII Atvftrtllural /OUNMI. 2. 14,5--2:.
,21-138.
174 vioM,\~. <,.H.. 1.96.6. Field vct1or$1Udi<"S of cpt7.ooci~ f."'!,.r CQ:.1!:t t'm·~r II.
l5S rHl'.11..ERt A .. .1.9Q5, Rf/pert ofth4., Go:-emnumt Vcmrina,y &ze1rriolagti1, S1.'asonol otudi~ of /1, ,,,,p,mcl(J;u./nw,on bovlnt: and non•bolline hoi,r;
rran.slf(l,IJ/. 1903--04. in !!ast Co3jt re,er enzootlc. cpfaoqtlc and fr,-e ;uca,. liulll!li11 II{
,55 nnni.m1. , .. 1911. n,c onlfichll 1n111,ntl>0lon ofE.151 Co•st (c~or. Report of Epi::.oorli: IJl=tof;IJ'rleii, 11, I 13-140
rMGo,r<mm,m Vet,r/noryBacreriologis_t, 7ran.••wl, 19011-10, 7-55. 175 ,,:o~t'-~· cu1.. 196<- Freid vceror srnd!e-s o( epl7.ootic ~1 <.:o.,~t flove.r. 111.
157 rnEILlffi, A., 1911. l'rogress rcpon on the pb,;siblll1)' of vaccinating c:anlc Pusture ccolol!)· in rel~LIOII to R npp<Jndfrlilou,s infestation rote~ on
'1g31nst Eas, C<>nst ,.,..• ,. Flm Report ofziie TJfl'<C'torofVe;,r/rr/11')' tau I•. Bullttm o{Epitoorlc DtSerm'J oflo{rlett, I;, 89-113.
R'e.w111t:li, Soutlr Afric11, 47-207. 176 -YOU:-.'O, AS,. LfJTCH. SJ.., SE\\SON, n.,,.,. cuss:-.:<.-'UA.\1,M.P., 1$66..
248 nmLDt. 11."' ou T01T.. P.J •• 1928. "11\\: t.mn.<4imi~ion oftlck·bomt dfsca,e1- Maintenance.of 11udll1ria /Jllfllilpmi'(I mftttlon In un r.ndemk a.nni of
b)' the intmjugulor Injection of !he emulsified intermediary nos1 ii.elf. K<-nya. Pnra.dtofog:,'. 93. ~1- 16.
Th!rlttnlh and fiJurJt'CJllh Ri!ports of1h11 Din.,ctorofVeti!rinary
177 roo:-.c;, A.S., LF.JTCH, -...... & oMwo,u. P.L, 19;9. lndu~tion of infocth-c
Edw:.arfon nru'I Rtstarth Union ojSomhA/f"itll, f>nn l, t 7-i•i. stoge,s ar77:ei~ril1 pnrv11 bye.po.ore of host ticks to hil\h temp,rnture.
159 ruEILEtt, A. &ou rorr. r .. t .. 19:?.9, The tr.tn!miss-ion of East CoaM rc,·cr by Ti:, Vtrcr/1utf)' Rttortl. 105. 531-533.
 30
Corridor disease
Synonyms: Buffalo disease, buffalo-derived Theileria parua infection.
T/1eileriC1 pamc, lawrencei infection. buffelsiekte (Afrik.)

J A LAvVRENCE, BO PERRY ANDS M \.VILLIAMSON

Introduction an identical life cycle (see Chapter 29: East Coast fever:
Figure 29.2). It is mainly transmitted by I?. appendiculacus,
Corridor disease is an acute. usually fatal disease of cattle re- although R. zc1111bezie11$iS replaces R. appendiculatus as the
sembling East Coast fever and is caused by infection "1th btlf· vector in the more arid areas of southern Africa.9 · 11 Epide-
falo-derived 111eileria paro,1 strains transmitted by ricks from miological e,rjdence suggests that in Angola the disease is
African buffaloes (Syncem.s caffer) (Figure 30. I). The disease transmitted by R. dmto11i.6
was firsc recognized in 1934 in Zimbabwe as a form of patho- Buffalo-derived T. pama can be considered to be a group
genic cheilerial infection distinguishable from Ease Coast fever of T. pan/a strains which are adapted t0 Lick transmission
on clinical. pathological, parasirological and epidemiological within the African buffalo population, including the red
grounds.8 Previous occurrences in the region may weU have dwarf buffalo (Syncerus caffer nanu.~) of Angola. 6 It is univer·
been obscured by rhe widespread prevalence of East Coast sally distributed in \\~Id buffalo throughout eastern, central
fever. Twemy years later the disease was recogni7.ed in South and sourhern Africa. except in the Addo Elephant 1'ational
Africa 15 and the causal organism was then identified as a new Park in South Africa.20 11 is usually non-pathogenic in this
species, Thelleria /a111re11cei, by Neitz13 in J955. It was later pro- species although fatal disease can occur follo1,1ng experi-
posed as a subspecies ofT. parva. namely 1: pan,a /awrencei.z,; mental infection.2 The parasite persists indefini1ely in in·
Subsequent investigation has revealed cbac the parasite is fected buffaloes in both schizonc and piroplasm fom1 and
T. parva and is rhe same as that causing East Coast fever and can be recovered consistently by tick transmission or cul-
Zimbabwe tlieileriosis, all.hough che distincrion between these ture of lymphocytes from such animals. 24
disease forms has been retained and remains useful for de· Buffalo-derived T. pt1r11a ls not well adapted to tbe ox and
scrip ti ve purposes. 19 The disease is described by :\'eitz. t4 when transmission from buffalo ro cattle occurs it usually
The disease has been named buffalo disease, for obvious fails to comple1e its development: ma ny cattle dle before
reasons. and Corridor disease. because the first outbreak in sufficient time has elapsed for development of piroplasms,
Sour.h Africa occurred in the corridor between the Hluhluwe and in rhose chat survive. piroplasms are very scanty and
and Umfolozi game reserves In KwaZulu-Natal. It occurs spo- recovered animals are not commonly irlfective for ticks. !n
radicaUy throughout eastern. aemral and southern Africa souchern Africa the disease is self-limiting within a tJOpula-
wherever there is comact between cattle"3nd infected African tion of cattle once it is removed from the source of infection.
buffaloes in the presence of the ticks, Rhipicephalus appen- Onward transmiss ion from cattle has been demonstrated
diculatus, R. :u1mbeziensis ;md R. duuoni (see Chapter I: Vec- experimentally ancl in eastern Africa repeated passage of in·
to rs: Ticks and Chapter 29: Eas1 Coast fever: Figure 29. l). feet ion in cattle has resulted i11 a change of the character of
After foot-and -mouth disease it is rhe most imponam disease the organism on a number of occasions. the parasite behav-
transmitted from the African buffalo ro cattle and is a major iour and the disease that it causes becoming inclisrirlgtliSh·
constraint to tl1e acceptance of the presence of buffaloes in able from classical East Coast fever. 1• 10 This change is
cattle-raising areas for purposes of conservarion or recreation. auributed to a ·1ransforma1ion' of the parasite as ir adapts to
the ox. but an alternative explanation that the phenomenon
represents selectio n of a T. paroa strain from a mLxed popu-
Aetiology and life cycle
lation in the buffalo must also be considered. ·n1ere is no
Buffalo-derived T. parva is morphologically and serologi- B\idence that ·transformation' of buffalo-derived T. parvn
cally Indistinguishable from cattle-derived T. parua and has occurs in the region south of the Zambezi River. where
468
 CorrldQr disease 469

classical East Coast fever is absem.20 Tois may renect the in- Possibilities for infection occurring in southern Africa
frequent contact between buffalo and cattle in the region. are limited as buffaloes do not usually inhabit densely
Funhermore, the seasonal appe·arance of the various tick populated or intensively farmed land. Conracr is most
scages 22 reduces the likelihood of an animal being infected common in the vicinity of game parks or In extensive
by one stage and infection being acquired by the follo\\ing caule-raising areas with a substantial wildlife population.
stage. since piroplasm~ are present only transienti~' and at a Increasing density of human population and the policy of
low level. In conrrnst, in eastern Africa. all rick stages may be prohibiting contact between buffalo and cattle as a foot-
presem on the animal at the same time. 18 and·momh disease control measure are likely lO reduce
Buffalo-derived T. parva has been shown to infecl a Lhe pre\'alence of the disease progressively in the future.
waterbuck (Kobusellipsiprynmus defassa) from which infec, However, in South Africa game ranching in conjunction
tion was transmined co ticks: chis was not achieved using with. or in the absence of. cattle is becoming increasingly
cattle-derived r. parva.23 Buffalo-derived r. parva \\ill also more popular in certain areas. As one of the so-called Big
infect and cause fatal disease In the Asiatic domestic buffalo Five. African game trophy anlmals. the bttffalo is amongst
•
{Bubalus bubalis) .3 the most sought-after game species for introduction co
many game farms and game conservation areas in South
Africa. Buffaloes that are free from Corridor dis.e ase and
Epidemiology
foot-and-mouth disease are in shon supply for stocking
Buffaloes in T. parva endemic areas become infected soon purposes and consequently command an eJi;ceedingly high
after birch and the vasr majority survive to become ex- commercial value.
tremely efficient carriers of T. parva and remain so e,·en in ln eastern Africa, the absence of game fences, the mode of
the absence of reinfection. Almost all buffaloes in endemic life of cattle-keeping pastoralist communities and the ,'!(de-
areas in caslern Africa arc T. parva carriers25 and ticks spread distribution of the buffalo populations, alJ contribute
feeding on carrier buffaloes become heavily infected. Cor· to Corridor disease being a more significant problem.
rfdor disease occurs only in cattle Lhat graze pastUres on
which African buffaloes are or have recently been present.
Clinical signs
Such situations occur where buffaloes and cattle regularly
share the same pasture, \\'here canle are moved into or Corridor disease exhibits the same clinical features as East
through buffalo areas, or where buffaloes stray from their Coast fever except that rhe course is usually shorter, death
natural habitat or game reserves into farming areas. It is sometimes occurring only three to four days after the onset
not always easy to detect the presence of wild buffaloes in a of the first signs. The emaciation. diarrhoea and regression
farming area. They may tra,·el long di.stances and remain of lyniph nodes. which are features of advanced East Coast
hidden in wooded areas. and the presence of a single buf- fcV'cr, ate not commorily seen in Corridor disease. Severe
falo for a relatively short period may be sufficient to cause a pulmonary oedema precedes death. The monality ra1c is
serious outbreak of disease amongst the cattle. about 80 per cent.

Figure 30.1 A;rican buffalo

iSyncerus caffeti (By counesv of
Di V de Vos, Kruger National Park.
SIWk'.na. South Africa!
470 ,,tTios " ' ": Pr0to1.oal diseases

Pathogenesis and pathology analysis·' have shown that isolates of T. p11n1a from buffalo
are antigenically more varied than those from canle. TI1e
The pathogenesis and pathologyofCorridordisea~e are very ·~uguga cod,tail' and other cattle-derived T. f)arl'(1 \'accine~
similar to tho$e of East Coast fever. Pulmonary oedema is a
will thus often fail 10 protect against narnral challenge in
prominent fcarnre bm lymphoid hyperplasia and lymphoid
sirnations where buffalo-derived T. parva infections are
infiluarions are generally not ,·cry advanced at death, and prevalent in cattle (see Chapter 29: East Coast fever: Con-
macroscopical lymphoid aggregarions in the kidneys are
trol).i 1 Experiments indicate that the disease is better con-
m1common. trolled by the infection and treatment method of
immuni1.a1ion using local isolates (see Chapter 29: East
Diagnosis Coast fever: Control).7 • 27• 30 Buffalo-derived T. pan1a is
more difficult to control using convemional oxytetracycline
The diagnosis of Corridor disease depends on the recogni- dosing during immunization and higher reactor rat~ tend
tion of the characce1istlc clinical 5:ign~ and pathological 10 occur. Bupar\laquonc concroli;. the post-immunization re-
changes. and on the microscopic clemonsmuion of actions bener chan Oxytetracycline bur can be over-effective
schizoms in a situation where contact between bulfalo and Jea,·ing cattle partiallror fully suscel>tible. 16 Buffalo-derived
caulc is known or may be presumed co have occurred. In T. prm1t1 transformed by serial passage in c:anle. protected
marked contrast to East Coast fever. the parasites are usually cartle agau,st chal lenge with the original parasite isolated
Yery scanty and a prolonged search of smears of a ,,election from buffalo ln on e experiment. and was mucl1 easier co
of lymph nodes and spleen may be necessary co demon- handle than the original isolate. i 0 such a par:1si1e might pro-
strate schizoms. It is usually more rewarding to examu1e a vide a useful source for vaccine material.
number of different lymph nodes for a short ptniqd than a In eastern Afric.i. wbere susceptible animals are being
single lymph node for a long period as the distribution of kept in simarions of high risk, and separation of buffalo and
parasites in the bod} is very \'ariable. The presence of piro- canle is 1101 an option. strict rick conuol must be practised to
plasms does not provide supportive e\'idence as these are prevent disease outbreaks. Jn indigenous Zebu-type cattle,
more llkcly 10 result from incidental infection 11ith non- endemic s1abiliry may be established whereby cal\les born
pathogenic species such as T. tattrotragi or T. 11wta11s. The to immune dams arc infected ,,·hile young and most be-
immunofluorescent amlbody test (11',\T) and T. parva- come immune. with fe" deaths. 11
spec llic enzyme-linked immunosorbem assay ( ELISA) are In southern Africa ~ontrol of the disease is ba~ed on the
both applicable for detecting antibody after eiqiosure to buf- pre,·entio1l of contact between buffaloes and caule wher-
falo-derived ·r: parr,a (see Chapter 29: East Coast fever: ever possible. Game reserves in canle-raising areas which
Diagnosis). The scarcity of piroplasms in recovered animals contain buffaloes should be securely fenced and the intro-
makes the use of a polymerase chain reaction (PCR) assay an duction of buffaloes imo farming areas for purposes of game
a11rac1il·e option fordetceting persistently infected animals. ranching or safari hunting should be strictly prohibited. un-
·1he$e tests do noi distinguish buffalo-derived from ca11le- less the buffaloes can be certified free from infection. The
derived infections. In the limited number of buffalo-derived growing demand for buffaloes that arc free from Corridor
r. pcm:t1 strains examined, the p67 sporozoite antigen con- disease and foot-and-mouth disease. nnd therefore suitable
tained an extra insert of 43 amino acid residues: this insert is for non-endemic a reas, far exceeds limited supply. In an at·
not presem ln caule-derived strains in which the sporozoile tempt 10 help satisfy the demand for such animals in South
antigen is highly conserved. 12 If this holds true for other Africa. Corridor disease-free buffaloes have been bred suc-
buffalo-derived T. pama parasites. it will l)e the first geno- cessfully from Corridor di$ease -infec1ed parent stock ob-
iypic dilference found between buffalo- and cattle-derived tained from a fooi-and-mouth disease-free area by rearing
T. pan•a and could lead co a method for differemiating be- them in an em~ronmem free from tick vecLors. Although
tween the rwo. transmission of infection by tick vectors to the progen} can
be eliminated in this war, lhe calves must be monitored for
possible transplacemal infection with T. pan'll.
Control
When an outbreak of the disease does occur, the usual
Corridor di~ease responds 10 1rearmem with buparvaquone procedure is to remove the canle to uninfected pasture, to
and halofuginone, but as the course of the disease is usually institute s1rict tick con1rol procedures and to attempt 10 lo-
:ihon it may he difficult 10 insrinne trea1mti111 in time for it to cate the source or contact and tht1s prevent future recur·
be effective. In South Africa. the trcaimcnt of diseased rences. As the disease is usually sclf-limitlng in canle.
animals is not pem1ined by law. prolonged quarantine of infected herds is rarely indicated.
There are no vaccines readily a,·aJlable ttt present to con· but outbreaks should be monitored carefully for several
trol buffalo-derived T. pnrva. Cross-protection experiments months 10 ensure that caule-to-carlle transmission does not
in cattle, 28 ruonoclonal antibody studies~ and genotypic occur.
 Corridor disease 4i I

References
a.\R:,;rn-. s.,·.,ulloc:..:tusv. o.w.. s96ij, The p3ssagt• or·nunh•ria o,-:ilu•tiun Mth, u-., orbupor\'oquone in lh• !ni~l!>r,;ind 1noatmen1
law,11:u/ (Kenra1· thtough c~ute. Bri11Jh Vrtl'ri11n,y /011rn11/, 122, method or lmmunlslng cottl• ngalrm Thdl~rln ptll1/(I dori,·cd from
396-409, African buffalo (S)'ll(truunffer/. \'e/eri11my Pttt/lJUology, 43ll-2). 1;;.2~
2 aROCl..LnlW, n.w. Sc 81\IIXt."rr, S.F~, 1966. l'hc l.b-OlnUon of 'T11Mlt.·tfr: 1; NOR\"Al. lL\.l .. rt\lA%, ll.lJ,, t.A\\'Rl!SC£. J.A. & URO\\·S, \,f., 198$.
fawrencPI tKcnyaJ' from .i wild buff;!.Jo (Synurtd j"/JjJ,:r) :tnd It._ ~e:r1al lipldcm,olo~•y of aie.k-hnmo dl~OS<" of cattle in Zaml>.ibw~, Ill. Tht'/u•r/11
pu~agc lhrough c~p,ivc buifo.Iocs. Brlrlsl1 l'ttrrl1u11)' /ounull, 12?. J)lln'YI group, Tropiml ,\11/ma/ llc/1/1/r ntlll Product/011, 17. 19-28.
l87-39S ,a s-oxvAL n..\.1.. PtllR\', o..o. $a vom:r.. ,1,.,s., 19s,. Tit~ EpitltmlologylJf
3 BVRRJUGU. M,J ~ oov...:v. G,M,. 19i3 77re/lrr/a lm<,•n:nc..•i· fnictt1on in the 11wllerlwl• mAfrl<t1 London: Ac,;odemle Pr~
Indian wuter bu.lfalo. Briba/ll$ bu/wits. E>v,mmemol Pnmst10/ogy. 3-l
19 PfRR\". o.o... YOU:<G. •· , .. 1993. The naming game: the ch•ng,ni fom,r.cs
hl-261
of E:ist Coast fever ano 111t·ilerltr P""'"'· n,e Vtrtrtnnry R«()rt/. 133.
4 cosaw. P.. QU·Mon-c,1, o.~ .. 11...tJ.l)WT~. C..L. l)()UN, T,lH o'rAtL-\.GIU~, 6134;16.
(;.J., ~JA..,u.r.-.crorrw. rc.r....... r.Roon:i..,w,s, 1.a.. STI\GG, o ..,. urror. o..t.. .-.
20 P01G1m..11. r t .. srotn."7.. w.u .. 1:nou:s. £.JI. , Roos. J..,.~ 1988. Corridor
vous4, .\..$.., 1989, Ch.irach?·~\ation ofbuffwo,dcnv,-d. 1he1tcrin.l pnra1,:tu
distt-351! in South Afrle,; r\ reviC\\ of tht.• t..'Urnmt srntu~. Jtmr,wl ofthe
with monodnnol •nllbod,~sand D:'IA probes. Pr1m.,/10/Qgy. Sil. 17))-188.
South ,A{ricfw \ 'tttdutuy1'~<f>Ciu:fon. 59 15.5-160.
S COMtAD. P.A .. Sl MW, Uoi'\., GKOOTD.'ttUIS. f.G., lftVl~. ;\.U,1 Nt.\\~os. , .•
2 1 RAOI.I.Y. o.t .. \'OU:S:G. A..S .. r:woorrSHUtS. ,.G.• CUS'.NIS(atAM, M,P•• DOL.AS,
XJl\.\~UXC.(.1111, R.P-~C.. ~S.Stl'(H, ..... f<i' \'UUNG, A.~.• 11J81 h,l)hUlon of
T,'t. & -'tOft7.ARL..\, s.r,. t.919. Furtht!r jtudles on the immunb:.,uion 01 t:attl~
n,ellerln porusltcs from African buffalo tS.li•<<"1'J <<>Jf•rl ond
chamcterir.G.tfon with aml•schlT...ont monoclonal nntibodlrs. n~lnst 1'heflcri11 fiiwrP11~i b) infection and chemopr();>h)•l:t>.:h.
Parrul10/ogi•, 11-1, •113-424, V~1,ri,u1r;.· l'afiur't()/ugy, s. I J7... 128~
6 QA CAA~ H,.):. "SERRA.~O. i;,:,.s,H .• 1971. C".ontrlhulqt10 p,1rn r> d1udo da 22 SJ1oitr. ~·.1. ~ ~oRVAI 1t...u .. ,s&. The seasonal aetivi1y of Rllipie£fplu1l11J
thcllerlos• s!nccrina mallgna dos bov!no,. em ,\r,g11la. Acrn v.-unnnru~ t1pfl"11tfle11/n11u Neumann. 1901 fJ\carl: lxodido•J in the hil!fwdd or
Vo1Y1 [.l$boa, Z!mb.ibwe Rhodt.lln /1><1rru1t o/!•111,1$/tofoy;,·. 67. :7-84.
; oou;.•, t.?.• 1<135- lmmunlz.ation against thoilerlosls on Ol P~j<t.l R.tnch. 23 ~T.\(!(), tu.. 1992. Saudies of aheliedai pam:16 orwa,etbuck £Kob111
La.lkip¼n. i<t-nyn, /11: tR.\'1~'. .\..o.., (td.). Jmmuni:atlon 1'&,,ains1 r1;,,m,rf01.ts d,j,m«1 nnd their pMsibtc roto :n the epldomiology of E.m <:oo$t fover.
ill ,Vrf<a ~olmbi: lmcmarlonol l.abo"'tol)· for Rc,carch on \nlmal PhD the.ls. Uru,·er,lt',' of Sallord U_<(.
Dl""'ses. ~4 S'l'.\C:(,. n.A., BMO\\'S. C'.C.o.. C:nA\\'f..O)U). J,G .• J.:\..._;ltAJ. (,..)., i,; ~ou:-m. A.~
8 IA\\RlM:fi, O.A.. 193.S. lkport t>ftl:e Dirttrorn/Vnr.rimiry RHtnr<lt., 19;.;. In vitro c.ultiva1lon nf nu•iforia ltw·rtrttf'i•infecttd l}'n'IJ1hob1Mtold
Sourhun Rlrotlcrslh~fet rltt Jt·<1r 1984. coll linr. dtti\;,d from a buffalo (SJ11cmts coffi·rJ.Rtsttur/1111 1ctm,w,y
9 LAW&XO., f.A .. ~ORVAL. H..A,I. t. UILf..."l;JJl,R(,, (;.., 19~3. Rhlpicep/Jnlus
Sd,mc..,. 16, 125-127
:1unlN:1<msl,.~ns ~ vt."C;"tor orbo\inc dwll1.mt1r. 1'topic,,t Animal 1/c:a/11: 2:j STM".G. n.A .• ('OSR,AO. , 1.A. ~ l<OSSJTUII, P.O,, 1987. lsolation or 1111/ltrln
<111d Pr0durtio11, 15. 39-42. port'1 iowrcncc, htfl'CtCd iymphold ctll lines from fr~ rnngingAfrici111
to ~lA1tn1M. A,(.• t'OU·XG, A-<i1, 1,.1.S..\.X, A.C.. ;\QUN:GU. S,G .. ,q,,v<h P ..\ I, ~411,\:.ll, buifulo CS)nc,rra tnffor). Res,•,1ttl1 Iii v,writra,:,· SdM«•. 43. 12.:-126.
r,:,; 1992. 1 r.tn,fotmattor\ of Thi'fl•rfn P<'""
dcaiv<,d from Alr!C"an 26 UILL*'B~RO. r... 19;6. Titk•bome. li\'~tock dl!ii~35eS nnd lhl.."ir \'CClO~ 2.
buffalo Sy11rmu ,:;tjferl by tld: passage in cattle Md It> u<e In Infection Epi>.oo<!oiogr or tltk•bornc d:,e;ue,.. l'lor/lt,Vrlmol llel'i,11•. I; 8-15.
and uca1mcn1 ,mmunSr..1.uon. v,r~rinary Po,a.511oloK)·. .i3. 1-1..;,
r, ,'ffll,\:t.. •,.s... ;g.&>, JmnmnfrAtion of can.le ag.imsa thcHcrlusis 1n the
II >IOU. c;, lOllnJ~C. , . & YOU:<G. A.S., 198~. Ep,dtmfolUJ:)' of lh~Oeno1ts ID
Trnns•.M:1.r,, di,·i~inn of Kt'n~"· At'Omp~.ri"on oftri:t.ls undM' trMlhinn:ll
the Trnns-~l.ira Division. K•ny,>. Husbandry and dlsca,,e bocl:ground ~lnosal m;maicrnonr \\ilh tn'nl~ on• til/lth dt,·clopmttnt ltt: m,~~. , u ..
•nd prelimfnM)' ,nvcstigauon, on thdlerioles ,n c•lv~. pr,,,..,1fi1 ,. l~d.J 1t>m:u11/.:11rlo11.·l;:nlmt n,,11.-,iotu /11 ;,.fr/ca. :-.alrobf: lntcmotioMI
Vr1tr/11<1ry .Wrdlrlne ~. 801-&Z. Libomnl')· ror Rescarrh on ,\ninul Olse:isH.
l2 :-:r:.,t.. v., ,,u~cna., A,J., r.,oBRJmrr, n. 1. ~1ou1AEUA, f.,, ;996. Cnni;crvution of 23 \'OU:\G ,\.~ BRO\\, c.c.o.. c.uxNJSCiHA)-1,.\f.P & R.Al)UiY, 1>.l:.-, 19;8
the ,porozoite p67 v•ccinc anrJg~n in caale·dcrl\1.'<1 Tlwilerio pan,n
,,ocb. ,.,.ith diffCTCnt cro~·fmmunhy profiles... /11fiir1l1,n mul tmmun:ty.
or
F..,,tluat!nn of methods lmmuo!.dng canlo agmn11 n,t'il<'rfll li11ur,11cel.
Im \\IU)t, 1.,.11., (td.l, 1"/cl:-lx,r,,., D,s,yn,,;a,ut 1/i~i, \-.Ctont. F;lh1burgh:
64. 20$-2061.
Unwwlt)· Q(!!d!nburth c,,nrrc (or Ttoplcal Vo1w1ury M,:drc:ln•
13 ~em:. w.n,.15sr; Corridor dlsc3S~: :\ i\no.l lorm ol bovlnu thcilerlo<ls
29 \'()\JXG... ,\,S... OA.00n:.N'uu1s. J.C .. '\IUTUGI, 1,, .. 'lAJUTl~J, A,C., .:.\ftlVKI, D.P &
encountered In ZuluLlnd. 811ll•tf11 ofEp/UJ()t/c V!st•o,,,. ofA[rlrn. 3.
I.AMPARO, 1>., 1990, Th• CorrldQI di~•= problem lltld lmmuni,atlnn
121-1i3.
trials3.i3.ln~t Thtl!vrin pnnw lnuir..mrrtinrtttlon In Ki!ny;t..ln· ,·ousc;.A.<
14 SEIT7.. w.o., 19$?, Th<:lfctfO.Sli. gom1tno~ and cyt.D\J;t,:ZOOOO!es:A "UTuo1. J. 1 • ,.,.,am,,•.,.c. (eds). l'rog,m 101<'tll<b 1llc Comrol ofEa.<t
rC\iow. Q,rdersrepoon Journal oft"tririMfj' B<Set-t,rh, 2.r, "2';~30. Coa$1 Fnw m,-1/,rio,is/ Ill KPll)'t1. Xairob!: !.tnyo.~griculu,.,.1 R..scorch
ll '\'.EJ'T7., w.o.• CANlL\'4. AJt &c J(LUGt. Ii.a... 1955. Conidnr disease:..\ (ah'&J lm1irutc.
form of bovine thelleno>i; encountered In Zulul:lnd /ournal 0/1/r~ 30 \'OUSG. ~&.1J:rrtH. LU... DOI.AS. T,T .• ~ROC.O, :.,,.;,., SUU'lr{OU. S,G• .&. OE
So111/1 Afr/e(t11 l'mrfnt>I')' Mrd,c,,/ .~s,ocw1io11. 26. 79-87. CASTno. 1.r.. 1\!90, E\11lu•11on of mfet:lion and tn:a1mert1 mcthodi. [n
16 sr.U).11 . ... ~ .• \'OUSG. A.$.• u.,u,.um. D.. MIM~G.~.);,•• "\OU~(,U, S.G. U,.\,\~, immunf.5.atlon o! impmvcd ca1dlt again lit th1!llerio~s jn un t!n<lemlc: area
\.C.. \\'tUJAMSOr-.. !l..'•t.• Ll'-'YOSt, A-~ i..UUU>.'l, 0.1•., I~ Funhcr ofl\enya. Vtttrirtary l'tJtt1.drol()~1• 35, 234-25-

,
 31
Zimbabwe theileriosis
S111onyms: January disease. Fonuna disease. n1eileria /)(Irita itol'is ln.fec1ion

J A LA \VR :E:--i CE, B D P ERRY ANDS .\-1 W1LL1 A "-1S 0N

Introduction T. pari•a strains. TI, e life cycle is identical (see Chapter 2.9,
East Coast fever: Figures 29.2 10 29.8) and transmission is
Zimbabwe theileriosis is an acme. frequently fatal di~ease of principally by adults of Rhipiceplw/11s (IJ}J)e11cliculat11s.
cattle caused by Theileria µnri,a infection which occu~ in Transmission of ihe parasite by Rhiplcep/1(///ls zam/Jezie11si.<
Zimbabwe (see Chapter 29: East Coast feve r: Figw:e 29.1). has been demonstrated e~'J)erimentall)'.; Several isola-les of
The disease was firs1 recognized and distinguished from the parasite ha\'e been studied and differences in virulence
classical Ease Coast fever in 1936,5 ahhough outbreaks ma) and in cross-pro1ective immunity ha\'e been demonstrated.
have occurred previously. It has been variously known as Some isolates are relatively benign:1 The parasite remains
January disease, because of i1s seasonal occurrence. and stable in regard to its ,1rulence, even after rapid repeated
Fortuna d isease, after 1he farm on which it was firs1 recog· passage by ticks th rough caule.H Epidemiological and lim·
nized. 11 is known officially in Zimbabwe as ·theilcriosi~'- itedexpcrimen tal evidence indicates thar recovered animals
The disease was described by t,,eit1.1° u nder the name Rho · are frcquencly carricrs.3
desian malignant bo,ine gonderiosis. and attributed 10 a
new species. 711eiltrin (Oo11deria) /Jovi.s, bu t the parasite was
Epidemiology
declared in a footnote 10 that paper to be s~11onymous \\1th
T11eileria lawre11cei. the causal organism oiCorridor dist>ase. Zimbabwe rheilt>riosis occurs sporadicall)' on the highveld
The name T. par11a l>o11is\~a, subsequently proposed for the o(Zimbabwe (see Chapter 29: East Coast fever: Figure 29.1)
parasite by Lawren ce~ 10 differentiate the disease which it during the period December 10 t>.Iay. lne strictly seasonal
causes from Corriclor d isease. The d isease resembles East occurrence of clin ical disease coincide~ with the seasonal
Coast fever b t11 the seasonal oc.c u rrente of theileriosis in dis1ribu1ton of adult R. apJ)l!11diculat11s." The development
Zimbabwe and the fact that infections can be milder than in of antibodies to r. µa1w1 has been demonstrated 111 cattle in
East Coast fever justifie~ l11t continued use of 1he terms the field during the p~riod September to November. Indicat-
Jnnuary disease and Zimbabwe 1heileriosis. However, there ing that transmission by nymphs. which has been demon·
is at p resent no good scientific reason to retain 1he subspe· sLrated experimen tally. may cause subdinit:al infe<"tion.'
cl~ classificacion. T. pnrt,a /Javis. as rhe parasite causing the ~lost cases occur be1ween January and March and rhe
disease is indisti nguishable from cattle-derived 7. par11a disease usually affects cattle over the age of one year: iL is
from elsewhere in eastern and southern Africu. 1~ h is worth rarely seen in calves. Primary outbreaks usually o~u.r 111
noting l11a1 T. pnn1fl isolate~ have been made in eastern herds with a moderate to heavy tick burden and are often
Africa, which show either the lower virulence or parasho· assotiated with new addilions 10 the herd. The disease may
logical fearures associated 1dth some T. pan·a strains caus- appear in the residem catlie csuggesring initiation of in-
ing January d isease.1• 13 The disease causes a significant fection by introduced carriers) or In the introduced cattle
number of deaths each year in Zimbabwe, necessi!aring the (suggesting introducrion of susceptible animals !mo an
1mplemema1Jon of inrensive dJ pping regulations to control endemically stable en,~ronmem). '.\1orbidity in an outbreak
its vector. is usually less than 10 per cell!. The disease is likely ro recur
each year on an infected property unless good 1ick comrol is
achie,·ed. but shows little tendency to spread 10 neighbour-
Aetiology and life cycle
ing properties.
Theileria panm strain$ causing January disease are morpho· Serological evidence suggest~ 1ha1 infection \\ith the
logically and serological!~· indistinguishable from other parasite is widesp read in both rhe highveld and lowveld oi
~72
 i.'.imhabwe theilerlosrs 4 73

Zimbabwe. l1 It is not yet known whether th<! limited occur- immunosorbent assay (E LISA] and polymerase chain reac-
rence of clinical disease. in spit~ or the widespread prel'a· tion (PCR) can all be used {see Chapter 29: East Coast fever:
lence of inrection. is e\idence of the existence of a\intlem Diagnosis).
strains of 1he parasite or of an endemicall~ ~table balance
between \·irulem strains and c,mle. There is no e,idence that
Control
,,ildlife plays auy role in the epidemiology of the disease.
7.imbnb1\'e 1heileriosis responds reasonably well to treat-
ment with tetracyclines ifit is initiawd at the first onset of
Clirtlcal signs
the febrile reaction. Parvaquon<? and buparvaquone are
Zimbabwe theileriosis exhibilS the same clinical features as more effective 1herapeutic agents and it may be assumed
EasL Coas1 fever except that the course is often shorter and that halofuginone would also be effective; th<?re is no expe-
death may occur three to four days after the first onse1 of rience of the use or the latter drug against the disease.
signs. Sometimes one of the preseming, signs ls blindness Immunization by 1he infection and tream1cnt method ls
associated with corneal opacity. Clinical disease is often ac- practised using cattle-derived T. pan,a Boleni as the vac-
companied b} evidence of hea\'y infestation with R. appm- cine. This is a mild siock which has declined rather than
tlic:u/ams. Monal icy in the field may reach 90 per cent in increased in virulence on passage. 1 14 As a result of the
severe ombreaks, but many animals recover from mildness of the stock. the vaccine is used \l~thout the need
experimental infection and serological e1·idence suggests for simllhaneous treatment with oxyte1rac:ycline and with·
that subclinical or mild inl'eccion may he common. out the occurren ce or
react.ions 10 immuni1,ation. whJch
makes the method less cumbersome. cheaper and safer. 9
Vaccina1ion is carried out well before the season of disease
Pathogenesis and pathology
and only \,ith the permission of 1he Director or Veterinary
The pathogenesis and paLhology of Zimbabwe theileriosis Services. 15 Tlieilerfa pnn,a BolenI has also been shown 10
are very $imilar to those of East Coast fever {see Chapter 29: protect cattle against several cattle-derived 7: pan,n stock$
£as1 Coast fever). Emaciation and diarrhoea are not a from Kenya and 1wo buffalo-derived suains. 2 This raises
feature of acute ca'Ses. Renal lesions are uncommon. the possibilhy of using T. pnrim Bolen! more widely as a
vaccine against East Coast fe,•er, although the fact that it
induces the carrier ~1a1e in immuni1.ed animals may make
Diagnosis
some relucram to use it. The degree of protection afforded
The diagnosis of Zimbabwe thelleriosis depends on the by r. pan•a .Boleni 10 buffalo-derived iniections is probably
recognirion of the characteristic clin ical signs. lesions and variable.
epidemiological feamres and on the microscopic demon- The disease is notifiable In Zimbabwe. and infected
5tration of parasites. In contrast 10 the situation in East properties are quarantined for a shorr time and the cattle
Coast fever. schi:wms may be scanty and a prolonged subjected 10 intensive acaricide treatment. Where the dis-
search of lymph node and spleen smears may be necessary ease causes severe or re.curre111 losses. Immunization and
10 confirm a diagnosis. Piroplasms tend to be presem at strategic acaricide use are recommended. It is also recom·
1011 levels if they a.re there at all. The immunonuorescem mended 1ha1 cattle from properties 1,ith a history of
amibody rest ((FAn and T. pan•a-specific enzyme-linked infection should not be introduced onto a ·cJean· farm.

References
AltOC,.,1.tiS8V. u.w. ~ RAU,£\'. t.:.P•. 1968 . .A 1nild form of East('..oas.t ftt\·~r G Lm11r.so. J.A•• 1979. In~ <lirrt.'renunl d1•g11osis<>f the bo,1ne thctlesfas of
ITi1t•Jl4•rln 1mr:t1 htf«tinnl bt.~oml~ ,in.dent on p.1,~ose LhrouG)t c:nnlc. ,oulJ11..·m Africa. Joumhl ofll:t' .South :\fr/Mn V1.•1t.•ri,:a?' Af..\.11rl,uion. 50.
Bridslr. t'etl!r/1Jn')'/ounIC1/. 124. 236,,,,238, 311-.313.
~ JR\'t,. A.D., \1'0St7.J\RL\.-S..P,, .,:U~1'SWA, r.c. 6o ~ORVAi.. R...\.1., 1989. -;" t.\\\1\f!\'O' !-"-·• ,011\'.\l.. SI..A.1.4, UIU:.'\'8EJIC.- c. .. 19-1}$. Rlliplr.;*JJ.ltllltu
lmmuni-.11,011 of c:,,nle "ith • n:,I/,•na.pam1 lwvlsSl<><'< from :ombci~1ul.< .u a ,·cctor or ba,·inc tl1t'IUL1ri,11.• Troplrtd J\'1imo.l Jlt'Vllth
7.Jmbab,,c prote<:tsagnlnstch,1ll~ni• "!th ,1rulcn, r. p. panYland r. ,,. ,mt! Prc,d11c;lo11. 15. ~2.
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J ~9(:11 " r., 1<J90.1\rl)<'Crso/rlr< li(lid,m!olog,,9//an/1ary 1)1stm< <pttlnwn~ O\Ct l'\..·o sMSO!l-i.. Joumal of:/J~Snml: .4{f1(tm \ 'tttrl,:nl')'
'il1.Uena parvu hmi>J III Zimliab,w. PhD 1he11;. Uni,·cr<ity of Uu,-.ln .lledlt11/.-wodati1.J1t. 38. 93-101.
.S l{(><'"Jf, H.1,. ()CAAi,\, J,(UL. \UJX \TW{,\, F.(:., 8YTl('J!>I, H., Ml!!\\' .\t.. R-'i..L,
g ,mTIJ<'it J,, 1999, lmmunfs.ation 3g.;>ln'111 :hctforf()tj_~ in Zhnbntm't.' min~
;SPOO:O.l!H, P.K•• c:o~AAD. ,. \, ... 1k\'JS, .\,1),, 1936. l.sohutun and th\" Uoh'7J:t Wi.c duc.- \\hhOUl ox·y1elr:i~Cline. Jrt: M0n7AHH. ~. 61 \\"IUJ,...\,\!•
charncwr~tlon ufbovine /Jirllwi<> pamshe., ln /Jmbabw<'. l'aurinm; ~~ ~-- .~ds,i. Llr't• \.~tt·mtis/orThtlltri;'I parva.: lkpto,,ncnt (11 Eastern.
Pwt1si1olog:v. 28. 19-32. C.•111ml 11111/ Southem Nrlrn. p,,,,.,.v1111~s ofn toim OAU. FJ\01111d ILJU
:, l>.\\1lE.~CLu....... 1u37.R~/>tlrt ojtltr. Diu.rtorof l'i.!U•rlunry Rt,Jettr,·li, t1.i11J..--1Jrop lr.ri'd-m /LR/ t/mttmmf<>u,1l lii•.-s(i)\"J.: .rlruvurh IIMti:UI,•
S011rhcr11 RltrHleJln,forill,• war 1936. ,\"airo/JI. K~11.1"11. IQ--12 V/tul"/11997, ILRI "lllrobL Rcn)'a.
4 74 s1.Ct10:nwo: Protozoal diseases

,o :<OT:t. \\.0.. 19;;. Tbcller!osls. sonderiosesund cyta\lX%00noses:A 3. Cbemop,ophyl:ittJc: lmmunlsatlon or c:utlc ming <lX)-1Ctracydlne and
revfo'\\'. Ondvrs.tipoonJoumnt o/V~tcrlt1ruyHesearch. 2i. Z75-f30. 11 rombinatiou ohhcll~rial itralns. !.'ettrltUJf\' Pa_riJ$(tofogy, l 51-60..
U SOR\'Al.. 1<..A..I~, fJ\'"21 8,H- L,\\\"l'P~CE. J.A. II 8RO\\~,A.F- l98.S, 1.: Ul~--.;ff{R(,, C;,, l'E.~11 :.-~M., L\\\"Jt~CE., I.A,, Df V(>~. ,q,, PAUNG. R.W it-

Epid,miologi• of tklt-bomc dlscases or cnttlo hi Zimbabwe. Ill. ~,\SIBR. A,A.M,, 198!l. rnusat agents o(bovml1 thCif(!riCJ:,lS ln :j()Uthe:ru
Tl1•1llt•ria 1>1mtt1 group. Tropl(a/ A"lmal HNi/1/J anti Ptaductum. ti', Afnoa. T'ropicol A11/mal Nealrhnnd Prod11ctfon, M, 127-MO.
19-28. 15 U-)Ht:\\'Oll'.UXU•OBA'l'Ol.1J. u.. 1999~ lmmuntsarton 3S;3!ru.1
t.heUcrlosis In Zimbabwe. In: MO!t1.\RIA. s. k WII I IAM80~. s.,
,;; Pf.1\R\'. a.o. "vousc. •· <. 1993. Th• namln3 3:uno: th• (hong,ni fonunes (eds! Uv, l'Ou/,1ts/orl'hc1!criapi11VD: Deploy~m In F.astcm.
of E:mC<last fovl!l 11nd T1i,i/,,,1,. p(lmL Tltr \lewrlntuy ~ro,d. Ill.
C..nrral ond Southern Afrirn Procfflling,of11 Joim 0,\/J.
61.Hl6.
l.l""'"'t
tj Mbt.ti\', o.r.., A-fl()\\'~, C,G,O,. cu...:;'l.c:,.-C,(A."1, "t,V., >!IMIIUl, C.O., MU.,15.1, f,I..,
l).\\""'l.. .R.C.• ..PUft.:'1:1!:U.. k.[,, ST4\"V, D.,\. <&t ,·ouNGA.$,., l9:S.. fu-u.1 Coa~1 fo\'Cf,
~w,
F.AOmtd /UIJ ll'orhltop htld 01 ILR/ (lrtwmar/on/11

Kenya
/n11/rur,/, Nnirnb,, i:,tt)'" 10-12 \fa,d, I99~. ILRI, ~olrobi,
 32
Turning sickness
Synonyms: Cerebral theilcriosis, draaisiekte (Afrik.}

J A LAWRE:-aCE ANDS M WILLIAMSON

introduction Clinical signs

Turning sickness is an aberrant form of rheilerial infection, The condition is usually atebrfle and is characreri r,ed b)' the
cbaracterized by the accumulation of parasitized lympho- sudden onset of a variety of nervous signs. including cj1··
blascs in the cerebral blood vessels leading to thrombosis cling. head pressing. uni- or bllatcral blindness, a1a.xla.
and infarction (Figures 32.1 to 32.4). It was first described in opisthotonus and paralysis. 5 Death usually occurs after 2 to
Uganda b~· Menam and Carmichael 3 in 1936. In East Africa it 21 days. but occasional cases become chronic and may live
is usually caused by Theileria parva. but in South Africa the for up to sLx months, 1he principal signs being blindness and
condition is attributed to Theilerit, rauroiragi. 1 The initial muscular incoordination. In cases in which the spinal cord
hypothesis that It "''lls caused by Thei/cria 111111ans 2 reflects is involved there is posterior paresis or paralysis.
the confusion surrounding the identity of that parasite at
the cime.
Pathology
In the acute to subacule fonn. areas of intense congestion
Aetiology and life cycle
and haemorrhage are present in the meninges and the brain.
The life cycles of T. parva and T. raurorragi are described in and are associated \,ith localized sotienings and yellowish
Chapters 29 a nd 33 respectively. discoloradon of brain tissue. 5 ·111rombosed meningeal vessels
are often very prominent. There may be e:..iensive haemor-
rhage into the ventricles (Figure 32.3). Histological examina-
Epidemiology
tion reveals the obstruction of aneries and arteriole.. in
Turning sickness occurs sporadically, usually In young cattle affected areas with large numbers of parasitized lympho-
Jess than three years old. in regions which are endemic for blasts (Figu res 32.5 and 32.6). The blood vessel.s are fre-
T. pan•a or T. 1a11ro1mgi. A very high incidence has been quemly ,hrombosed and necroric, with perivascular
recorded in Zebu cattle which have been immunized by haemorrhage, and the brain substance which they supply
the infection and treatmem method and introduced into shows ,t spectrum oflesions ranging from ma1acia. character·
an East Coast fever endemic area.' ized by spong:- change. infiltration of numerous Giller cells,
,1nd proliferation of capillaries and glial cells, to haemorrhage
and infarction. Haemosiderin and lipofuscin are widely dis-
Pathogenesis
tributed free or in macrophages through the tissue. In ani·
Proliferation ofschizom-infected lymphoblasts occurs in the mals whk'h survive forany length oftime the necrotic tissue is
blood vessels of Lhe meninges, brain, spinal cord and some- resorbed. leaving cystic spaces wilhin the brain. Lesions
times the spleen,5 causing thrombosis and infarction. There which occur ln tl1e spinal cord are similar. In a few cases
is usually no evidence of parasite invasion of other organs. infarcts may also be present in the spleen (Figure 32.4).
This phenomenon is generally considered 10 result from mas-
sive reinfection or a partially immune animal, bul the mecha-
Diagnosis
nism$ which pi;,m1it the parasite to proliferate in the blood
vessels are not understood. It has been suggested that local- Turning sickness may be suspected when an animal in an
ization of the parasite in the brain may result from intercur- area endemic for 7'. parvn or T. ra11rorragi develops the
rent infection or stress in a chronically infected animal.4 characteristic clinical signs and lesions. The presence of
475
~ 76 sr.cuox l\\o: Protozoa] discoses

Figure 32.1 Throm!Jos,s of meningeal blood vessels Figure 32.4 S~Jemc mlarct

.. . • -
•
•
....-·
"
•
. •
.... • .•
• •
....
~
..•. ..
.. .
• •

..
.. .·... ..,,. ~.,.
_.,
•.
••
...
:, •
•
• ••
..
·. •••••
. ..
.. '
•

..
..
.. •• . .•.
..... ~., .
, ' , .. .... .,.
! ~
....
Figure 32.2 Infarct In the thaJam..is Figure 32.5 Tnrombosisand sludg,ng o' aarasitized lympho~asts tr. an
area oi encephatomalacla

1
";:,.
•. 'I
,~
~

• ~
•• I
•
A ,,
Figure 32.3 Exrensive inuaventnc~'ar haemorrhage Figure 32.6 l'erll'ascular 1nr1lua11on o: Girtercelis and lymphoblasts in an
arira of e~.ceptta,omalac,a
 Turning sickness -J.i7

circulating antibodies specific ro one or other parasite pro- ri11m pyoge11e.s infections) are among the diseases with
,~des supportive evidence. Diagnosis can be confirmed by which it may be confused clinically. Diagnosis is easily con-
the demonstration of schizoms in large nu mbers in smears firmed by necropsy but identification of the species oi para·
or histological sections (Figures 32.5 and 32.6) prepared site responsible is only possible by immunological or
from lesions in the meninges or brain. Schizoms may be dif- molecular methods.
ficult to find in chronic cases.
Control
Differential djagnosis
Cerebral theileriosis caused by T. par11a may respond to
Turning sickness must be differentiated from other condi- treatment with buparvaquono or halofuginone, but re·
tions which presen I clinically with the sudden onset of ner· lapses are common a nd tlie animal may eventually die
vous signs. Cerebral babesiosis. heartwater, rabies, after rwo or three epi~odes of clinical disease.4 There is no
listerlosis, thromboembolic meningoencephalitis. sporadic experience of the use of these drugs against T. taurorrngi,
bovine encephalomyelids, lead poisoning. ' 's1001siek1e'. Prevention should be considered in die context of the con-
caused br ,'vfatricaria nigellifolia poisoning, diplodiosfs, trol of theileriosis in general. Immunization against
thlamine-depende111 cerebrocortical necrosis, and spinal T. par,1(1 by infection and treatment does not appear to
abscessation ('sitsiekte', see Chapter 193: Arca11obacte· provide pro1ec1ion against the condition.

References
T1,~lltflD.? tnuro:ragi: A
DEVOS. A.J,. ,l1~)t\(,E.ft, M," 11..\~•,;n,r., LJ. 1981, ~ \IOU. .....\(;", 1.. £ lt>IIDISC:, .\., 198;. Bo,in~c~n,brnl th~i!Cri<ljll• in PU!~
probable ngcn1 or bovine cetebr.\l U\dllrriosls Ond,m~poor1 /01mu1I of Boran aald ~11,•l-cro<S ca1tle unmuni,ed ag,ilM1 Ea.51 Coa>1 fC\·crnnd
V1•1,r/,u11yRt.<tnrch,,18, 17'"-178. k•pt undor conunuou,. fitld ch31l•ng~. /11· '"''"'· A.O•. r<d.).
2 Fl,..\S.\(h\..',i, 11.0. &LC. IWUX. J, \t,\\"., 19t!i'- B0\1.necrn:br41 thclltr'iOiiS- :!
/m,111111/:.t1/lo11 ;ga111« Iheilvrlosls 111 ,lfrlca, '•arobh ln1crna1h1nal
repon on tn·o cMcrncrurrint: in th~ Union. Ondlftsctpoort/oumal of lAb11ra1oty ror R-rch on Animal Dl•<a.e•.
l'.-:,riluu)· R'9t't1rtli. 27, ,ISJ...461. ; ,·•~ kD.S"1tu~c;. t,ft,J., 19;,;;, Bovine ccrcbr4( 1hclltri1>~>. A r•pon on th'<
:,: ,tITTAM. 1t.w.:,.i. & <".AAMf<'J«An. 1.. l936, Tummgsfc:knrss: t\ proto:r..o.sn ,~~ .,,,h splcnk 1nfao:tlnn fmmml of1/1~ Somli ,lfricnn l'<M'illnl)'
encc1>haU-tis of cattle: in Uganda. 1t:S celatlotl5hip with East Coast fl'ver. Atsodntwn. ,It ll'i'-1,11
Par/l$il()/ogy. 28. 25,t-283,
 Thei!erln uwrorrngiinfcclion 479

Diagnosis and differential diagnosis several immunological and molecular rools exisr which can
distinguish the two parasites, as described fully in Chapter
Thcileria rnurotrcigi infection may be suspected when a mild
29: East Coast fever. For example. one of the ami-schizom
febrile rcaccion. with enlargement of superficial lymph
monoclonal aluibodies in the monoclonal antibody panel
nodes, occurs in cattle exposed to Rhipice[}ha/11s infestation.
used to characterize cell cu Itu res of isolates of Theileria
and schi1.oms are demonstrated in lymph node smears.
reacts exclusively with T. taurorragf. 1
Differentiation of infection with T. wurorragi from Infec-
tion with T. [Jarva has been discussed previous!}' in relation
co East Coast fever (see Chapter 29) and turning sickness Control
(see Chapte r 32). The 1: [Jaroa-specific enzyme-lin ked im-
munosorbent assay (ELISA), unlike the IFA1', does not suffer As 7: taurotragi is usually a benign parasite, there are no
from cross-reactions with amibodies to 1: raurocragf and indications for its treacn1em or control.

References
BISHOP, R.P,. SPOONER, P,R.., 'KASKA.I, c;.s.. J.:IARlE, J,, I.\Tlf. A.A., HQ\'E. r.. 9 L\WJU!XO:. I.A.. ~()liWAL. It.A.I, • UIU'KRtltG. c.• 193!, Rhip,ctphahts
' ""'""'·,;. r. DO"'-~· T.T., 199,. ~Joleculnr cMrnctcrisa!lon of nielltrin :nn:be:tRns,..~as a \1.>('tOr of bovine cbc-iltriae Tropia,t .-'t.n,mat Het,/11:
parasll~ appli03tiOn to 1he epldtmiology or th<,ilmosis in 7Jmbabw,. 01:d Productum. I ;), 39-12.
P(;fa,/fotogy. l~. 573-581. 10 MARTI:<. If & UIIOC'-U.SDY. 11.W•• 196o.A """' parasite or the eland. 77,.
2 81tOCnEStW~ n.w.. 1965. E\1drn~ that Ririp;a..pllalu.f_11ulcht1lru V1•ttritttlf)' Rtcord, 72, 3.31-332..

(Gersr.itker 1873) may be 3 V<ctot of some plroplasms. 811//Mw of 11 sf.IT'/..\\ .o.. ,95,7. TI16ilcrit1sls. gond~rio$e$ 2nd q13ux1..oonose$; A
Epi:writDim,s,40JA{rka, 1:),3;-44, review. Ondtr111poort Journl11 o/Vtrtrlnnr:.· R~a.l'(li. 27. 21-5-490.
3 SURRJDt;K, ~.J., bfH.)W~, c.G.D,, CKA\\~kfi, J.c;., IOMIMlt 1,M•. MOftUAL\, $.P.. t:!. STACG1 D.A.. ~!\"HM, (;,J',. \'OVN"(,, .\.~& 8f\O\\~.C..<i.O,, l9i6. The

PAV'Nf,. tu:.. • SE\VSOS, K,, 1974,

Prclimln::Jrystudics on an acypkal .srrain t~1abbshmem of 111t.•if,:na 1nfet:.ted cell lines from an el.and
0

ofbo\'ino thollerfa isolar.cd In Keni·._ R•..,,,,d, /11 \leterlrwryScle11c,,, 17, cr,wrorraguso,;,r, Lydekker 1906), Hl'S.-nr(h Ill VewrmnryScltn<~. 20.
139-1-14 122-126.
&3 STACC, U,A., \'Ousr..A....,, u . rrt:H, H,1., GMtlOTILSHUll,;, l,C, lit--001.AS', T.T••
•• "VRRJDG1.. M,J.. BROWS, C.G.U, & 1ri:1,tfllik, r..u., 1974.
111dilt>-rla a,,nulma.;
19l13, lnfe<tlon of mamnu1.ll1\n eel!, ,.;,h n,.;1.,111 ,pc-de•, /!wmftal~g.
Cross ~anions bctw,..,n • ocll cultu,e 1ch!,ont ontigcn and antigtns of
86, 243-254.
East Africa,, Spilde, In the Indirect 11<,~rcscem nn,lbody cesL
Exp,:rl,11e111at Pat1Uitolagi1, 3S, 374-380. 14 nu:11u11• •, .• s906, Piropfa$1>lt't mman! tn. spt.•c.) of South African c.tttle.
Jnumnl o/Compnrori,,. PtllhOIQ/:)'nnd nitrnp,ur/c,, I 9, 29'l-300.
s OE 1·os,A.J.. UtsSl\l'I<:~~... • PASTl\'G, c..r.. 1981 11:,1/eriu 11nurotragi: A
15 nnuutt.>... 1!tOi Funhcr notes on Plroplnsmn murt11u. a new ~pcdrs or
prob:ibleag~nl orbo,inc ce1cbrdl chcllorlo>'li. 0 11dcts1i/poor1 /oumal of
V~ttritt.aryReset,r(J,, -181 l.7-li8. piroplasm in Sou1h African cc.mlc./oum,r/ o/Compnr~t(w Patho/og', 1md
11,rrnp,,urlcs, 20, 1-18.
6 o• vos, A.I,• poos, I.A., 1!181 TI1e ,sol•tion or77r,//erln 7ttrunmag/'in t6 UILE.SIIERG, G., PENf-:. S.M., lA\\'Rl~C~, I.A.. tJE \'OS. A.I., P..UJS'(:, IU\.J.
South ,\lric'1. Oudomt1101,1rt /oumal o{l'ti,rinary Hcs,tarc/1, ~8.
~P"'"IFR, ,.,..,,,. t98:i. Causal agemsofbo\1ne theUerio$ls inl'Ou!hem
U!l-!S3.
Arne•. Tropicn/Ammal HN1/1/1 n11d Producrio11. 1.;. 1.27-140.
7 GtlO()TJ ~HUlS, f.(,,. MOIUlJ~OS, \\".I. ~1AU, I..., :,A\'ti". P.O., YOU!':(;.,\.$.,., 17 YOlSS(;,A,S., GR:O()lf~JltJJS. J,G,, KIM8EJI. C,.Q., ~\!\'IW, G.X. &STA(;(i., 0.A..,
~f\lRR.i.Y• ._.,,"' HAW;"" A.O.. 1980, Fatal thi.-Heriosis in t?land (Tuurom,gus 1977. bolauon of• 771<//tr/11 •pede,; ftom eland (Tnuro1mg11s: Ol)'Xl
o,yx): Pa1hol0Jl)' of natural and <Xp<:rimenuil case'- Rt'!<'nrch In infective' fo~ tattle. Trt>JJMmttli.:m und Par11Jftologf1J. 28. ISS..19.;.
\·.rrerfnurySgf<m:~, 29. 219-229,
18 \"(>US(,, A,$ .• CkQOTE,:.-.HUIS, f,G,. ~MrtH, ..... f!t.OWFRS, ~1,I,, 001.AX. T.,,-. $.
a. ui.,11£.~Cli, ,..-. . .,. ~uc 1n ~ 1,11 .. p,x.1,. ,gao, h,ol.:nion oL.l nc>n•pathogcnk •~oc•usav. ow.. 19;8. Scrutture; ""'°tlotcd \\ilh 77i,lwrla parusl1cs In
thellcrla o( cattle uansmitted by Rl1ipfcepha/us nµ,,..ndlculoni,. eland cry1hr~1es A1111a/~of1roplral MtdCctn,and Pamslrologr, 72 .
.liml,abwtr Vertn'nary Jourrml. J J, ?7-35. 443-45~ .
 34
Theileria mutaris infection

J A LA'\'RE'\'CE ANDS '.'vi \\'ILLIAMSON

Introduction size and shape than those of r. pan·n. 1• 23 11 has no1 ye1 been
possible to culrure T. m11tems-infec1ed lymphocytes in vitro.
Theileria mu1a11s was first described as a benign parasite of
the ox by Theiler,111 but its identity was inexrrlcablyconfu$ed
Epidemiology
with thai ofother benign species of 17reileria for mann•ears.
It was assumed to be the only benign bo\ine 111eilerfa in Af. Theileria mwans is 111despread throughout Lhe range of its
rica until 1977, when il was demonstrated that Theileria w11- tick vectors in sub-Saharan Africa and on some Canobean
ro1mgiwas also capable of causing a mild clinical reaction in islands. It has been sho\\'n that in an endemic area 99 per
caHie. 2b Elsewhere in Ihe world the benign Theileria of cattle cent of calves have become infected by the time they reach
is considered 10 be r. oricmralis.21 Theileria 111r1U111sis trans- sb: months of age. 12 Amb~'Ommn cohaere11s. which is com-
mitted by Am/Jlyomma ticks and is now known 10 be con- monly found on African buffalo (Sy11cem$ Cfl/ff!r) and devel·
fined to eastern, western and sou1hem Africa and to the ops high infection rates with T. muui11s, will frequently feed
Caribbean Islands when! it was introduced in cattle from Af· on cattle and is likely to promote high levels of T. mum11s
rica. The parasite also infects the African buffalo CSy11cerus transmission where buffalo and cattle come Into comact. 1
cafferJ. in which ii was first described under tl1e name Fatal infections due lO T. m11ca11s have been described,;
Theill!rin /Jnmeui. 1 Its only praclical significance in south· often in situations where an lmals have been exposed to
ern Africa is the confusion that it causes in the differential T. mur,ws infection after immuni7.ation against T. pnn•a
diagnosis of Theileria par.·a. In eastern Africa. pathogenic with a reduction in uck control. Iii, 18 IL is probable that stress
strainsofthe parasite occur, which maycauseseYereclinical caused by poor nutrlcion, intercurrent disease or other fac-
illness and death. tors predisposes cattle ro anaemia when infected with
T. mmnns. Field and e.xperimemal evidence suggests chat
strains or T. m11ra11s derived from buffalo may be more
Aetiology and life cycle
pathogenic to ca.ltle than thOSI.' derived from catdt.>.6· 12 15
Tlieileria mu1a11s has a t)-pical theiJerial life cycle (see Chap ·
ter 29: tast Coast fever: Figure 29.2) but prollferatton in the
Clinical signs
ox occurs mainly at the stage of the piroplasm rather lhan at
that of the schizont. lt is transmitted by several species of Infection with T. mu1a11s following tick transmfsston usually
Amblyomma: A. uariegatum. 2~ A. cohtUJrens, 25 .-1. gemma, 15 results in a mild febrile reaction and slight sweUing oflpnph
A. hebraer1m 4 • 11 and,\. asrrio11.?0 Amblyom111n hebmeum is nodes commencing after an incubation period of 20 to 30
the principal vector south of the central watershed of Zim- days from the time of attachment of the ticks, and lC1Scing for
babwe, and A. variegnrum is the principal vectorto the north two to five days. Pa1hogenic strains in eastern Africa cauSe
(see Chapter I: Vectors: Ticks). lnfectlon occurs trans;tadi- severe anaemia and icterus and soml.'timcs deatb. 16 • 18 The
,ally from laiva to nymph. or nymph to adult. Recovered ,l.ni- parasi1e may also cause a marked anaemia and stunting in
rnals remain carriers indefinitely. The piroplasm is readily young calves. 13 It is unlikely that the clinical disease de-
transmitted by blood inoculation. 1~ scribed in 1937 at Tzaneen in tile Limpopo Province of
The piroplasms of T. m111a11s are larger than those or South Africa 2 was, in fact, caused by 1: m11ui11s. There is no
r. pan,a and dividing forms may be seen durini; the acute e,idence that 7: 111wa11s causes cerebral theileriosis and re·
phase of infection. The schizonts are also larger and the port, of cerebral theileriosis in South Africa a11ributed to
chromatin granules are more numerous and less uniform in T. 1111111111s should be ascribed to r. uwrotragi.3
480
 rh~iforin 111111tms i11fcetion ~81

Pathogenesis and pathology specifiC' piroplasm antigen 9 11 or hy indirect lmmunofluor-

escence 11>ing piroµlasm antigen. in. -These tests provide n
The pathogenic cfft'cts of r. muums result from the i.11,11sion specific rctrospectl\'e diagnosis and are also useful in epide-
and prolifcratlon of piroplasm, in circulating erythrocytes. miological studies.
Pathogenic strains in eastern Africa arc presumed 10 cause
intra\>ascular haemolysls associated with a hea~ piroplasm
par-<1Sitaemia resulting in anat·mia and ic1erus.: It has been Ojfferential diagnosis
sugge~1ed that this reaction may be associated with an amo-
Differentiation of inf,•ct ion \\'lth T muram; from mfection
immune re,ponse. as with T. 11111111/ara.8 The degree of
with T. pan•a ha~ been discussed in relaiion Lo East C:oa!:.t
anaemia observed in an animal correlates closely \,ith the
fever (see Chapter 29: f.as1 Coas t fever). The pathogenic
le\'el of pirnpla,m pamsitaemia.
form of°/: 1111111111s infection seen In eastern Africa must be
differentiated from other haemoparasiric infections. n0t-
Diagnosis ably babesio~is. a11apla~mosi~ and 1rypanosomosf~. by
demonstration of the piroplasms in blood smears.
Theiler/a mwans in fection may be suspected when a mild
febrile reaction occurs in cattle exposed to Am/Jlyomma spp.
and when theilerial piroplas111s are demonstrated in blood
Control
smears. Schizoms, although characteristic, are , e11 scanty
and are present for only a shon period. TI1e)' are therefore 1\s T. m11mns is usually n
benign parasne. there are seldom
rarel> detected. The presence of large number:. of piro- indications for it~ 1re.itmem or control under normal
plasms in animals showing anaem ia and lcterus may rouse comlitions. The , irulenc~ of field infections with patho-
suspicion ofinfeciion with a parhogenic strain of T. m11U111J. genic strains m:11· be reduced by inoculation of blood
if no other cause c:an be dcmon,trated. Confirmation ofdi- ,(mtoining pir<>pla,m, of the local 5trnin. prior to ei.'PO·
agnosi~ b posslbic using \'ariou,; method~ (see Chapter 29: sure. Para,Jtaemia is reduced aiter t!xposure and 1he re-
Eas t Coast fever), the polymer.1se chain reaction (PCRJ sulting anaemia is les!> severe.z3 is The piroµlasms are
being best suited for testing blood smnpll'l> from the field destroyed by treatmen t with a number 01 8-amiuoquino-
Detection of anrihodl~ b possible using an enzyme-linked linc compounds Onclucling primaquinl. parvaquotie5 and
immunosorbem assay (E11SA1 based on u T. m11m11s- buparvaquonc.

References
'&ROCl.:1 ~~'B\-. 0.\\ 196-!i, \ 0£.'\\ t~h.~llt"rfaJ patMit(I' or lh(' ·\rrkiln buffalo lmmw1• n.w10nfil"J i11 Puuultr 111/tV"llruu. lmmllliOlfll:)'.
5y,rf"·ms wffi•r>. Hr,ll~h, o/fpu:QOtlC' T>s..tt·tt~1·t<,fAfm:a 13, 325..330, lmmw10JA'ffi1t1l11~•mul lmmu,mpmphyfa:(1s. \'al, Ill. Pmto..:0'1. 8oc.t
2 u~ ft.or.,. c. v.-., 11u.1m1 ,, c.r. ou ron. P \\·,n.. t9.li, Bovinr:
to· ,u1'l..
R.1111n. fl•>ridn CR(" Pr<"->. Inc
the-Uerta11i, in $ou1h Afnr-.i ',\11h ,J.'h.;t·fa) td.:-1t!nc~ tu n1ril,~Jt1 muraus 9 )..\ft \bt. J,\t (.t.tfHH-1 Hl"t. fl \f,, ,mHf,\IU.\ ,.v. SJ.O\O[. t-G. N "IU'-())a.
Ondf'ffitl!poort Jo11nt1:1/ of Vt·u·n·,,,,,,,. .S<'iim«, 8, ,-12;,, q .. 1 ~ Jdc•n1ific.1uon nf .:a th~tlrriu ,m,tatrs•'-1)1."Cdk antigen for u,i.• in
) Ol: \U\, ,\.I,, IIJ ..M•",lf;t1c, f', '° lH'.\'.J1~4, 1.1., 19ftl. n1,•J/uia? tJlUllltraJ!i~- \ iD111ntlbod) and Mittgen dt."t~Uon fli'·\. Pnra.'l"l:..· lnwu11m(o~. l?.
pmb.ahl,! ~gtnt l'Jf bovJn,· ccrcbr,'1 th,"'n,.,na~i .. <J,ult•1#t1/}f)(lft Jaumal of 119 ..;:u
l',•1,,ri,:my /IMYlrrlt• .:fl. I i7-l'i11. ID )r,J".1111:.N.. C.U ~· ,·ou,, •.
.\..'-,, tljj7. S~.m.dOJZll:.:!l "iUdi~ on o,:tr.iin~ Qi n1~/kr1t1
.i. or. vo:.. ..\.J.,. rtoo~. J.A... 1981. Ubwrt:mons.on uw trnnsndu1on of muuu,:'" isol~tc.id in t-:a~, \fr~t',, u~ln:g 1heo mdh&:-c;t Ouonttnont amlbody
Tl:l"ilnia m11um,.1n South;\frka. OndirJ/1'J)O(Jrt}m,r11,,l oft rt.-rlnnry r,"t'hniitm• 1\1111nl.11,f Tm1•f.ul ,\krflf1111•,1,ul Pilf<t.'\ithf&'i}'.. 71. t-lO
R1•~1td1. ·18. I -<, U ;.\\'!;Ill 'f"l . lw\,. \l,\(lr,;1..~7..tt. 1•.>i..l. 4 '(JJl\"c\L IL LI,. 19$). f'°"hU{on of
llw/U'r/11 t1mrm, en lJmlMh\,'1.· /1111bahuy \ ~·wr/nn~· /tmnwl. 12. 2; -30.
:; Oc)r,....._,. t.1 1981, l1t~rn~ ln tltl! chcmotht.trnpy of 1hl11c,r!u~t.s. /11: mn,
,.n.. c11!\::-,:l!\·c.1L,M. ).1.1• ,.. ,c,v,n..\,!o., '1.-ds.) .~h-'tlt1(1•:.' tfJ 1;11• Co.•urol oj l:t )101.1... «;.. U)HUiS(, .,. & "I (')USU, \ .S.. 19th. Epuii:mu:1togy n( d11:1ll'rfOSiS. in
r,,.,fl,•rlo.rlJ. The llaguo. 011,1on. lJ>ndon; '-lartlnu, :1:IJboff Put,11,hers. 11w rr.1n,-~1.u.1 Divi,thm. iu-1\Ft, J h1-thandt1· ,mJ disca:~c b.'lekgroUnd
and prt1lll1liUGf) ill\'(•.,ifgutlmt, nn thdtc.•rio.,P.i<i In c~l\•c~. Prt•tre>JJffl.'i·
6 \,1\00t'L,11UJS,. 1.0, ti! '\OUM,.'-"·· Ht1'1, Tht' lmoh·enH:nl Oh\1.11.ilifo in
Vt•1,,11•J1WJ"\ Ml'tll<mt•, ~ 80 l--8~2.
'Jili..'ill!11d infoc1lo1\~ of domc;,tit fflUl\OJ'- u1 t::a~t Afric.-:,. Tn: tK\ 1,. A.n ..
C..:U.\:~l~'(;ll,\.\l, \f. I"~'- ~OUSG. :\.~ :l'<ls., \dtrtJUl.'t'.t h, tlir.• Cmllml o[ l3 '1UJ t. <,.. I 01m1,,•.., )'OU~(;, J\.~. ~ t rrtOI. R.1~, !ftBti, Epid1.1n1iolo~ o(
n1~kt!c,.fi,: J>tM('l."t/inR,( af1m 1,m'fl:miom1/C.,011/('rtmre Jli!ld at ILR.W.. nr
thellt•rlth>ili tn c,1h es In un cnd.('mlc arN1 Kt'h\ .,. 11(•rluar, v,
\:,tr<>h,, 9-1.!I h•im.Wt)' l!JBJ rh\.• HaJ,_,1u.•· Ma.rcinu.., .'.'i'IJhcul P&.1hJl:,:.ht·r:,... 1'11fl1.<;trH.r1f!J,.'. 19, .2!",S..:?'i'-1.
pp. 71-i'.t, J.1 MOft;l,\WI.\ .... , ,1 ~l~ v .. ,mw~, l ,\ ,I • tu,li(,)11," 199:s, Modl'rn
'; lk\'.,L~. ,\.0.. OAC1W~. c.n.n.. nunnn,c;r. ,1,r. I u:--.NfN<.,IIA,\I, ,,.P.. \fUSO~ h1otL-chnotot:1,c.al m-:-1hods ror dli1_ijnn~1~01 bo\·me 1heileno!-is und ,,uurc
,\,J .. PU RU \L\.. f 1 Uk.,1u. lU ... 1UULH. 1.u.. 197~..\ p::uho~ntc trend., p;°'~..-rtltngs ,,f1hr l·A<J t.'X/Nfl ron11ul•t1t(<m ()JI th<"·'"'''}
thcllt1cfal 'l nd!''1m,· nr r111th.· ln chn "'Jr-o.k D1.suic1 ofKi:~-J I. n(1111fr,11'!,• hwu•i:11 nUTOJ:iC"dl ,,w1J10.,J.J fm di1~'1(Ufr, of110emoJ)(Jto.sit~
r~nu..<.slo1~ ~tud.it'-tl. 1'rrJJ'!(11t ·lninwt 1-lc-ultlt cmd Pr~/uctlo11 -1. 0(tobt-r lm. ~h:ndn. Ml'xico
22C,...2l9 1s ''"' 1s,; ,c,w ,ekoo11 ,m1LS. J.(., ~ \UU'.'-'.t,, ,.., .. ,,~:. hoJarion of T11cd1·nn
a tfC:\·ss ..,.u . .:..· ~~c)Jousos. \+. .I,.. l98'7 lmmunop:uhology, 1mmunol~· 4nd muuuu from ~cn~·-,n buffatu. ilnd u;an"-m~son by llnrbl.wmmtt gc1tnm(;,
immun1,praphytaxi.s or n1rtl,•ria 1nf<.'Clit:1ns. J,r, _'4>Ul.!'BY. E.l,t ...ed.J. VdrmWI')' P{1ht.,iu,1tt~·. 8. J1-.s7
482 ,t,:;110~ 1wq: Prt110203l diseases

16 )\08-~0~. J•• Pt;.Dt.JtiL~. \' 0DttL r..,L kAMVA, t.r. ~HKO\•,"\:'. C.(;.t).. lC};:". ~ tmr.,nr Rf,.<;•• M.011.:.0~. f. & Pt.n1,ut,. ,·.• 1974. S6mc('xperim«n11son Lht1
En.t Coo<i re,·cr irnmuniu1io11 trim lo Ug:mda: Fidd ,xpc,,,.,., of 7..,bu lr.u1~ml~<ion of 11tclltri() mutnn;. 'Tht-ilt:r 190G: :ind 11tdforia vunia
c.attl~ fmmur'l~c."tl \,hh three isol.:stc.S of J'hc.~UtJrla ptVL-'d, 1 ropi~lll ·i.nimu/ 'Thdlcr 190·1 b)· th~ tkk~ •.\mblyomma 1virlt1ga1Um tFBbrfdUj. l i9..; .ind
u,111t1ia11d fJIOllurua,1. 9.219-2ll. Rh/pi,,·p/11111,; «p/1<"1d1,•u/1rtµJ x,mn~nn 1901. In Uganda. /.,·,:•.-J1rifi fllr
17 ~111~ou:n.. R, &, WQJ.:.\l"Sf"II R.. 19~~ , ..eJ~urbe 1.Ur DtfiereaZ:itmm; dc.,r Tro1~11m1edi:J1t wul Pfllll~itnlt;gf41, 25. 20i-2J6.
Thdlcri•n,pc1J05 dei; Rlnc!es d1uch $<:r<Jlojpl'rhe Uniersurhung,,n 23 UIU ~.. uu,c, G, .. 11,\\f),. k:,t.., \ltt,\~(i~L\,C f\),~l)I,UR, \\',. T\1t:Hll.L 1\..1, I-
r.,•wclirlf1 /llr I 10f}llmnrdr.:u1 u11d Par1u:1oloSlt. 16. I~-~3. u.1.x,. 1977. Studict on lbellttHd.;e ;Sporowo.. 1n r~ni.1.nfo. X
S.-\..'li:C.A.,

18 f-XOl)(';'fV\f-~. D.R 111.fr\, A,I sow,n. w..,\,, 8[RG.\L\,."', f.tl., t)-'fl, I &-'\\','\} L \ l.,ri;i,·«·"1c fl.rd aria! un imrnuni,auon •~nlal,t enuto 1hl'ilorl<>~J..
,1r ,. 191a. E+1:s1 C.,.l>I Cen•c f"Jtld c.·h;1.ll1:0Kt! oftunle immuni7..ed ei&a,ms1 ·1101w11mi'tli:.lll :nul t'flmtllnfQglr. 28•.;'l9--.506.
1'11rJlniu /Xff4"-' fMu~'Ug.at. 1'rupi(al •.\mmal JJralth and Pn>ilurricm, 4, 2.1 ,ou~o..\.~•• ~9-8,..,.. Jiumuniz1>iion of i:3Hlt.: agains1 1tu:ilcrio,ct..,tn tht
112-151 Trnru,M.uu dhislon of i.:011) • A componsou of trlols under uatlltlonru
1y no!ltJ'll. A.. 190&. l'lrop/asmn 11111w11., rn. ,pc..:.) ofSouah ,\!nr:an r.iulc. ~1a.n.1t mano:~1.1Uh!nt ,,1th trt1l!'I on a mnch t.11.l'v~lopine-n,. /Jr.1n,·J'.".; ..\.-0,.
/011111"/ o{C:01ml(1"1li1-r Pmllolog:/<111d fh~rnp;,util:e. 19. 292-300. led. lmm1111L:mla1111.gnu~, n,,,if.urJoiis fn.A{rir,1. ~::ilrobl: 1111.,matfon:ll
20 111&.t:'l.tWtt(,., <;•• lU83, AcquMiJom.. 111.lm'C.'UC'~ d.ub la runn.ahnntc du r6l1,-•
l.Jlx-Jn*'r):' ro, R1. '«.'ilfCh on .".ntmal Oh,c-a.i,e5.
1

vcc1tur de 1lquc.., du genre .\m/i(1'111tm1111lxodldae), Rt1·11<d'l,l,~V1~,·,i 11, ~ \OUM.t, \.:,.., lUJRR.11)(;[, M.J. & P.\'(X"l R:.C.. 19"."7. Transm1.«ion of a
Mtdt.'t'inr Vt!ttri,tnir.• dtJ Pay,, Tru11itm,x.a6, 6 l--66. nwU.·rln ,pccic,. tfl mttfl1 h> th4.! bndld ,r,~.
,\Jub(l:vn:mtt<Cira,,,,,,,s
21 UIUJ~tsrrnG. c;. 1~11RII' ~."1 ,. 5-VA'.\Jlft. ·'- U,I. A pjlt.,\.~SSr'\, r..r.J .. :.183.
l>onil.G 1909,, l't<tpi{ttl Auimlll lh:,Jltlt mul Pmrluaio,r, 9, 37-t5
Tit'1llc1la<Jrk.nralls• .:\ coMnopoUta.n blood pa.msitc of cattle: ~0 \'OU~&. \.S. r.MOOIF-,JIUf..._ p., ~\t.lJtJl C..D.. )...\MlAJ. C...K. &. ST~\G-0. D.A..
Dtmons.um-Jon or u,,.. ,chi7.ont ~ta Re. Rf'St"l1~h fn \'t•r,,nna1:1Srlrm·r. l8.. i9-:-;. J!i,,QJrstion ot A'nwilt•ria~~tw~ horn eln.n.d traurotrtt_JJm(lf\·.t
3S2-3~1 mf"'''h."" for cnu!t- Trt;111•11mi.•tfl:J,r mut PnmsiU>/(Jgf~, 28. 185-1~4.
 35
Theileria velifera infection

JA LA\.YRENCE AN OS M \VILL IAM SON

Introduction

Tlleilerin velifara was first described by Uilenberg2 in 196.J in

:vladagascar, under the name Hnemawxenus ,,e/iferus. as a 0
benign parasite of cattle charao1eri7.ed by 1he presence of a
veil-like strucrure associated \\1th intra-erythrocytic piro-
plasms (Figure 35. I). The parasite is widespread in eastern
and southern Africa and also occurs in the C..aribbean. le has 0
no prnctical significance oilier than its role in confu ·ing the
diagnosis of Theileiia pan•n. The parasite also infects the
African buffalo (Syncerus cajfer).1

Aetiology and life cycle

Theilerla 1•elifem is presumed to have a typical theilerial life
cycle (see Chapter 29: East Coas1 fever: Figure 29.2) but no
detailed studies have yet been undertaken. The parasi1e is
uansmi1ted br ticks or the genus Amblyonmu,. namely
,I. variegamm, ,I, hebraeum. A. lepid11111 and A as1ria11.3
4

The de\·elopmem of T, l'elifern in the gtn and haemolymph

of , 1. 11ariegarum has been shown 10 be typical of the
Theileria genus.1' The piroplasms are similar to lhose of
Theileria 111111m1s except mat some are associated with Lhe
presence of a veil-like structure in the C)'lOplasm of !he
erythrocyte (l'ig11re 35.1), This structure is believed 10 be
composed or a crystalline derl\·ative of haemoglobin.5 The
schizonts have not been described. The piroplasms are
readily 1ransmined by inoculation ofinfecced blood. 2 0

Epidemiology
Little is known oftheepidemiologyofT. 1,e/ifcm. It is assumed
tO be widespread 1hroughou1 the range of its tick vectors.

ClinicaJ signs ThSilfll1a sefliJrata Theile1ia wlifera

No clinical signs have been recorded in intact infected

animals. Anaemia occurs in infected splenectomi1.ed Figure 35.1 lme dra,·.~ng of p1r0l)lasms ol Tllell:maspp
calves.4 whrch de11elop ~e1ls

483
48-1 ,lo10~ n,o: Pro1ozoal diseases

Pathogenesis and pathology SlTUCtures are found in el)'lhrot)ies. Howe\'er, as yet, nose-
rological test or spedes-speclfic monoclonal antibodies are
~o studies have been undertaken on I.he pathogenesis or
pathology of T. 11elifera infection. available.

Diagnosis and differential diagnosis Conu-ol

Theileria w lifera infection can be differentiated from other As T. 11efifem is a benign parasite, 1here are no indications for
benign lheilerioses if piroplasms associated with veil-like it$ trea1ment or co111rol.

References
SOUll:;UOI R. u.u.t oiu....,ut-J<C., G, .. fO\!Ut-Uh. \\ ,, JS~ Smdih On -1 oru ...:1.+1~u o. '"'~(,011 uou. 1u..~. 11;t;6. Stt:dle,$ on TheUeriidnt-
The!Lerlldae \Spuroroa ,n T:u1,anla. \'Ill. fu<porlmont> \\ilh Afritru1 iSpotozoa tn fanLant.3. 1 Tid.tr.:insmU.iion of Harmatoumu :~li[tnu
buffalo iSJ·ncti.tuJ ,:nOu~. 1°'op..•nuwdl:l,t um/ Ptu·wtwlogu, 28, Th)pl•,,m,'111:.in wul Parasltt,lf>RI,·. 2;- 106,.l l l.
36i-3,I. ,3 \'\~ \'OR.)o"T(SUOSCH. C.l..,Ul.\",, UII,.L'-:t'IER(';. C..&\".\(I.: OJI._, f..E., t~78
2. UJU:-;BQtc,c;.• 016.i, f!t11•mmo.u.•nu..<,>lllftr11.1, nig., n sp,, paruit~ f11a_•,u,e f.t')'thttw:)1fe forms of T11~llcrrt(l tl!lifi•,ft. R~Mrc·h ;,, Vttetina.,- SdPm:,,
stt/U.du s:ing de txnin.s a )tada~scar. Rt11 1uc.•,rEk:.Y1glt!I tic .\fl!ifldn4! 2-t 2tt-221
\ '/tlrinttln•des Pa)''J Trapitflu.'C. l; 655-662.
u \\ \Jl:'\U~ ,, .• ~ltlr-., {. \'OtGT. \\,P... UUL,'IJliR(,1 G,., 19i'9• On the
3 u1U$ftU<C, r.., 1983. Acqu,;111on; 11011,cll<:> dnns l~ conn3u.....u1c., du ruk Lifcciclc of T,1r,f1•rit11·1•/ifn11 IUilenbeccg. 196-1) in 1ho gut and the
\'etu•ur de uqul"s du 1,:.t-nre ..~mbl_\'Ommn (lxodidaeJ Reru~ d 1tfct'tl.gi' r1 dr hacmotrmph of 1t1e ctck \'fcwr .-tmhl)YJmuw Jl'tlfr,gmnm Fabnc,u.s.
M~der.tn~ \tt1/n11111r.rtl-.tS Pl1!*1 'frOJJICtrUX, 3b, 61-tl6 17~1 Tro11Mmffll:.i111't1r(IS.11olog1~.30.318--322
 36
Theileria separata infection

J A LAWR ENCE , A J DE V O S AND,\ D I RV I N

introduction Epidemiology
Theileriasepnmta is a common benign parasite of sheep in J..inle is known of the epidemiology ofT. sepamra. It is assumed
the eastern half of southern Africa. It is of no practical sig- lo be "idesp read throughout the range ofits tick vector.
nificance but pi roplasms are encoumercd occasionally
during e:~amination of blood smears. The parasite does not
Clinical signs
infect goats.
Infection \\~th T. separata results in a mild febrlle reaction,
which persisls for 4 10 16 days, ar1er an incubation period of
Aetiology and life cycle
9 10 13 days. 1 There is moderate swelling of the superficial
Theileria separata shows the characteristic morphological lymph nodes. A mild anaemia may be detee1ed.
and biological fea tures of a Theiler/a and is rransmicred by
Rhipicephalus everrsi evertsi transstadially from larva
Path ogenesis and p athology
10 nymph, or nym ph to adult. 1• 5
The piroplasms are characterii.ed by the presence of a Enlargement of lymph nodes and rnlld anaem ia a re the only
clearly defined veil. often situated outside the erythrocyte lesions to be expecied.
membrane but lying close co it. opposite a corresponding
gap in the erythrocyte cytoplasm (see Chapter 35: Tl1eileri11
Diagnosis and differential diagnosis
velifera infection: Figure 35. 1). There mal' also be a mall
granule. possibly a reduced bar structure of parasitic Tllei/eria separata infection may be diagnosed when piro-
origin. within the erythrocyte cytoplasm. Veils can be seen plasms or schizoms are found in sheep exposed 10 R. et'erui
associated with a proportion of piroplasms in Giemsa- evertsl. The characteristic features of the p iroplasms permit
srained smears but mar not be obvious. They are more differentiaiion from other Theileria spp. in sheep.
easily demonstrated in wet preparations l;>y phase-contrast
microscopy.6 The schizoms resemble lhose of T/1eileria
Control
pama.2
The piroplasms are readily transmitted by blood inocula- As 7: separata is a benign parasite. there a re no indications
tion.4 for its control.

References
JA...-..:~.1:.-... H.C.., Sfitf7., \\ .o.. 1956, rhc r~pe-rfmenial rransmisslQn of ($porowa, Theller11d;icJ. a ne-., blood p•ra;,ile of doml'S1k sheep In
m~lltrit1 <Wff by Rlup/c.p/111/1,s ,wrrsl. 011dtr1wworrJo11rrutl of Tanx.unl:z. Rl•t.1111· tl'ilttllJ.gc? ti drt ,\1«1,~lnr \ Whiuaire ,!es P,1>1,
v;urintTl)1 Ra.,arch. 27. 3-6 rroplt/111;(, !i ·15!1-165.
'2 ~£IT?. w.o .. 1957, l'hefl€!rlo~i.s.. gonde.rioses nnd cytmL,..,,oonoses: A 5 Ulll881!J<C, (.. ~ SCJUUiUDER, 8.t.c., 19;6. Funhc-r studit.s: on
revie,,.v. Omtcr,.1epoon Journal of\ l1wrinory Rc:tt·nrch. 27. 21-5-430. lfn,n:nio.r,•nu, W/Jt11ntt1' ISporo>ll:l, Th•ll•rilda,1 or •heep ln Tarueanta.
J u11.ia.,••.11c:. c... 1981 TiieUcrilll$J>eci~ ofdornesuc lh~stoe~ he R"1!11ed·t!tr1'0l/n1 dt M"tlrt111, Wt,Wna,,~ dr, Pll),~ 1Wipinmx, ~.
lft\'1:0., ..u,., (lJSS(SGJ-fAM, M.P &, '\'OU!'.G,A.S.. {eds). Adwmccs rn tl:e- l 19-126
Conrro/ o/T1mil,'ffosls. The H•gu•. Boston, London: Mautlnus :-:ijho!T 6 ,,ou_:.r:c, ,.s~ a. MCJUXtA. s.1•• 1971, Ol>!lm':ltto~ on H1u,mmoxa1us
Publl~her>. ~fJ{lr.nus Uilenbofll & Andiuscn 197~ In :he e11'throcyt.esofKenyan
.t U11.E~IIFRG, r..&ASURl'ASf. ..:, M .P., 1914. Jlflt.WUllDX~lltlS'!tpaTalUSSp. n. sheep. R,~JiUltrh in \'"l!ti..•ri11nry Sr:fonct. 23. 387-388.

485
 37
Theileria annulata theileriosis
Synon}'mS: Tropical theilcriosis, Medjterrancan coast fever.
Egyptian fever. theileriosls annulata

E PIPANO AND V SHKAP

introduction schizonts are not as common in T. m11111/a1a infections as

they are in those of Theileria parPa. This is true both in in-
Tropical lhcilcriosis is nn infectious disease of cattle caused fected cartlo and in culwre.s. Schizonts de\'elop and grow in
by the protoz.oan 1'heileria amwlmn. which is rransmincd sii:e as a r~ult of binary fission of their nuclei. The intra-
by ticks of the genus Hya/omma. Invasion b)' the parasite or cellular locarion of rhe schizoms stimulates multiplicarion
cells of the macrophage :ind lymphoid series resulls in their of the hosr cells in a manner that resulls in the disl!ibu11on
replication o r destruction, and invasion or erythrocytes of schizont material to each of rhe newl>· fonned cells (Fig·
results in anaemia. Peracu1e, acute, subacute. ntlld and ure 37. l).8 ~ Some of the schizont~ develop into merozoite,.
chronic forms of the diseases arc recognized. Animals that which penetrate crythrocytcs.u;
recover rem(lin latently infected. 110
Theiler/a ammlma was first described by Dschunkowsky
and Luhs in Transcaucasian caule in 1904:33 1her named it
Piropla.sma c111111ilawm. During the following two decades
fatal theileriosis cau:;ed by T. m11111/Mt1 in the i\ledilerranean
area and India was often mistakenly attributed 10 Theileria
muums infections.124
Tropical theileriosis occurs in northern Africa, including
the sub-Saharan territories. ;.i Sudan and Eriuea. southern
Europe. the Kear and Middle East, Central Asia. India and
northern China. The extent of its distribution in the Far East,
where it overlaps with Tlleileria sergenri infection. SG is not
yel clearly delimited." 1 The disease does not occur in south-
em Afiica.

Aetiology and life cycle

Three or 1he developmental slages of ·r. a111111/ar(I are infec-
live for cal tie: the sporozoi1es, which develop in the sa]jl'ary
glands of ticks and, in the bo,ine host, the schizoms in mac- Figure 37.1 The1/eria annulata,infec,ed call Ill cu!IU!e. The sch,zont ,s
rophages (monocy'tes} and lhe merozoites in erythrocytes. stiart?d ce~:.ee~ ·re new form ng cells /\c!1d·ne orange s,ainitQ "2 000
However. in nature, the disease is lransmiLted only by
spo.rozoites. lmra-ery1hrocy1ic merozoites (also referred to as 'plro-
~lature sporozoilcs are oval-shaped. measuring about pJasms') are generally spherical, oval- o.r comma-shaped,
1 µm in length. Schizonrs are multinucleated, round. vary- bm anaplasrna-like fom1s measuring 0.5 µm may also be
ing in diameter from l to IS µm (in some cases up co27 µmJ, encoumered: each is bounded b~• a single-layered cell
wilh an average of aboui 8 µm. Each nueleus measures membrane and posscssc, an oval m1cleus.U7 Spherical
abou l 1.5 µmin diameter.67 70 In culmres. schizoms contain forms \!ary in diameter from 0,5 10 1.5 µm. and 0\111 forms
an average of 12 nuclei. but some with 80 or more may be are 0.6" 2.() µm and comma-shaped forms 0,5" 1.6 µm in
found.89 Schizoms containing mlcromero7,oites (micro- si;>;e. 70 During muhiplication of rneroz.oites. nuclear division
48G
 Tlwilerin an111,lam thuilcriosis -18i

precedes q1oplasmic dl\1sion. in a process characteristic of stadially. bu1 infected male ticks that accidemal!y become de-
schizogony.i 1 tached from one host may rcauach 10 another and. thus.
The life cycle of T. n111111ltlw is similar 10 that of T pan,ar;; 1rnnsmit the di~ase m more than one animal. 110 Not all H_v·
(see Chapter 29: E ast Coast fever: Figure 29.2) and only (l/omma spp. 1hai transmit T. mmulam under experimental
some characterbtic.> specific 10 T. <1111111/ata are discussed conditions are ,·ec1ors in namre. since the pre-imaginal
here. stages of some species have a preference for hosts 01her than
T11eileri<1 a11n11/11w is transmiued transsiadially by two- caule. 11 · Rn 111ei!erir,. mmullllfl is maimained in nature by a
and thre.e-ho~1 ricks ofthe genu$ Hyalomma. After the pre- cattle-Lick-cattle cycle. The water buffalo (811/Ja/us lmbalis) is
imaginal stages ingest blood comaining intra-erythro~1ic considered to be th<' natural host of this parasite and co be the
merozoites. spindle-shaped bodies rconsi.dered to be mi- one in which II evol\'ed. 120 Theilerla mmulma usually cau:ses
crogamonts. which give rise by fission lo microgametesJ. no apparent infecLion in water buffalo, but lhe lruter mar act
and spherical forms (which are macrogmne1es) develop in as n reservoir for infection of ticl,s. The yak (Bos gr1mniens) is
che lumen of the gut.Ii; The zygote-like stages which lacer de- highly susceptible 10 r. a111111fc1u, theileriosis and suffers se-
velop in thegul wall transform imo club-shaped kinctes that vere clinical signs and mortality when infected.~
migrate to the salivary glands of the adu lt ticks afcer the The disease has a seasonal occurrence. which is related
moulc. After che moulted ticks have begun 10 feed. the ki- 10 the ecology and biology of its vectors. In most endemic
netes become polymorphic and undergo repeated nuclear areas. the majority of cases occur becween June and Sep-
dh,;sions until ~porozoiles are formed. Generally. infective tember.11 35 IIO. no. 11 i although sporadic cases may be
sporozoi1es appe:ir in the sali\'ary glands after che 1ick has found throughout the year.
been feeding for 48 to i2 hour,. 116 However. ·activation· of f/1i-ilerit1 m11111/ara may complete i1s life cycle in ticks on
ticks b>• close proximity to canlc or by being warmed to pasture lands or in bams. Engorged nymphs of H. derrir11m.
mammalian body temperatures6b· 104 may also trigger I.he one of the main vecmrs of T. m11111lata, hide in cracks and
rransiormation, so that ~ome 1icks may be infective before crevices of clay or s1011e barn wall~ and. aftermoulcing. adults
feeding on their host. Tickl. can only become infected by in- infoc1 ca11le introduced into these bams. 110 Susceptible cattle
gesting imra-ery1hrocytic merozoites. 105 on zero grazing in tick-free premises are. in Lheory, pro1ec1cd
,\fter inoculation imo c-aule. sporozoites pene1ra1c from iufecllon, bm infected ticks can be introduced on ha}' or
mononuclear cells and de\'elop into schi1.onts. These are de- by stray ca11le 1hat have grazed on 1ick-lnfested pastures.80
1ectable in smears made from superficial lymph nodes and Engorged adult Hy<1lomma ticks produce large numbers
the li\'er i to 28 days after infection. 110 Merozoites are de· of sporozoi1es: c. 50 000 have been es1imated to be present
rected in erythrocytes one to three days after the first ap- in a single cell of Lhe type 111 acinus of the tick salivary
pearance of schizoms. and may persist for years in infected glands. Since rnany such cells are generally infected. a bo-
caulc. Schizonts also readilr Infect caulc by needle inocula- vi1ie host mar receh·o tm enormous inoculum from a single
tion . Following their successi\·e passages i.n cattle or culture. cick feeding to reple1ion. 6; This high lnfec1iviry has been
schizonts may lollc their ab11lry to produce mero;i;oices. al· demonsir:Hed by infecting cattle experimentally ,,ith a
though they retain their infeccivicy forcaule.S:.. 86· 110 l: I00 dilmion of a single macerated rick.83
Isolate~ differing in virulence and cross-protection ability In comrast 10 1hc siniation in some other tick-borne infec-
have been reported from various geographical areas.?. 92. ~9• tions. such a~ babesiosis. where many Licks :m:.- needed to in-
IO'J Fifty-three Nort.h African isola1es orr. mmu/ac(l ha,•e been fect a single animal. 118 in the case ofT. 0111111/ma. a single tick
characterized by antiparasiric monoclonal amibod~ reactiv- is capable of transmitting a fatal infection.88 Examination of
ity. isoenz>11w elecirophore,is and S0u1hem blotting. l\o 1he salivaryglandsof Hyalomma ticks showed thai a high per-
identical isolates were detected by these methods and the cemage of tick.~ harvested in barns or in the field were in-
majority of isolates contained more than one pal'\1Site popu- foc1ed ,,~th 1: t1111111/arn. 36• s; 102· 123 Thus. a small number of
laiion.9 II appear:;, however. that lhe reported diversit}' in infected ticks ma~· cause extensive outbreaks in cactle herds.
field T. ammlaw populations is not necessarily related to im- Calves are considered by some authors to be more resiscam
munological strain differences and. therefore, It has a limited than adult catcle, but others consider them equally suscep·
impac1 on immunization against tropical theilerios!s. tible/1>- 112 In our experience young cal\ es are highly suscep-
tible to tropical theileriosis. while exacerbacio.n of neonatal
disease~ may further complicate the course of the infection.
Epidemiology
The epidemiology of 7: t11111ulma thei!eriosis differs in
The occurrence of tropical 1hellerlosis is dependent on 1he different grographicnJ regions. In the Mediterranean area.
presence of infected i.~odld tlcks. Fifteen Hynlomma spp. where the two-host 1ick H. decrimm plays the major role in
ha\'e been incriminated as vectors. 101 although some of lhese the transmission of T. a111111lara, the disease in young calves
may be s~11on}mS for identical species. Among che most im- is relatively rare. During the first tick season tha1 calve~ are
pormm are H. detrit11m. H. anmolicum, H. exca,mnun and e~1)osed to, 1he level or infestation wirh adult ticks is low
H. dromedarii. 70 TI1e infection is generally transmitted crans- compared 10 that in older cattle. and they are therefore
exposed IO a limited natural chnllenge. 11· J; In corurast. in leads 10 an allogcncic graft rejec1ion response. rcsulLing in
Indio \\'here the main vector is H. (1111110/lcum. calf morbidity relea~e of th.: ,chizom~. /11 t'it'o. the recipiclll ho~t cell, for
and mortalhr represent most of 1hc theill'ri01.is cases \\'hich T. mwulma are phagoc)ies, whereas 1'. p<1rt'iJ L-. fo11nd
occur. since the infestation rate of calve, \\ith nymphs is largely in T ct·II~. II was, 1herefore. assumecl 11· Lhar efficiem
high: and nymph-s ~ well as adults of /I. 1111wolic11111 can transfer or ~chizonts is a\:SQcl;ned \\'ith their capm:ily to pen-
tran~mlt 1'. u111wlma infection.tt· ;z etraw into new host cells. and that phagocytic t'\'lls actively
f\nrhhd lerial antibodies have bet>n detected in colns- englllf '/'. ,1111111/ma :;chizoms. The prolifera1ing r. m11111/mtt-
m1m of infecced dam~.-a buttherc is no ,wide1K~ that colos- infec1e<l cells multiply and me1astasize in the bod~ ,3!' ~o that
tral Immunity can engender significam protection agah,st the ~chizon1-infected cells become disseminated rapidly
sporozoite infection in young calves. EndPmic (enzoo tic) through 1he lvmphnid tissues. from the lymph node drain
stability de\'elop~ in herds that show n ~erocon\'ersion rate ing the site of inoculation to more distant lymph nodes.
of 100 per cent i1, cah·es in the first season that the~ are ex· spleen and thymus. f>ara.sirized cells also ~pread to non-
posetl 10 theileriosis.!' Older cartle become more hea,·ily in- lymphoid organs. such as the li\·er. kidne}'s. lungs. abo·
fested \,;th ticks35 and If they ~cape infection in their first masum, adrenal gland~. and later 10 the brain and
theileriosi\ season they are more likely than young calves to myocardium."~ Memzoiles released from schiioms infect
be infected by T. t111n11/t11a. 11 • 37 red blood cells. and in severe infections more l11ru1 95 per
cent of red blood cellb are parasflized, each comaininit one
or more mcrozoile.70
Pathogenesis
The severity of infection flethal or subh.'thal) follo\,ing
Sporoz<>ites introduced into s\1sceptible caule through the tick tran:.missfon depends upon the number of ~porozoites
,;ali\'a or infected ticks infect cells either at 1he site of inocu- inocula1ed. Jhe proporcion,- of ,chizont-inf~c-1cd cell, and
lation or migrate to the lymph noclc which dnun~ the infec- parasitized erythr<>C)1cs. the duration and ~evcrity of
tion site. There is no evidence that they pass through Lhe pyrexia. and the size of l!nlarged lymph nodes are directly
lymph node into the efferenr lymph. ·1 he rate or l}1nph !lo\\' related 10 the ~izc of the inoculum. while the Lime, to orbet
and total cell outpur in the lymph node lncre:i•e ~ 10 10 fold of fc\'cr and to the appt:!arance of schizon1s nntl <irythroc)'tic
from day 6 onwards.73 rnerozoites arc invol'liely proportional w the number of
·n,e sporo1.ohcs invading ho~t cell:. transform 10 tropho· sporozofw~ inoculated.9 "· tuJ
zoitcs, which develop into schi1.on1s. 11wilerit, t1111mlat(I Significant reductions in erythrocyte count, haemoglo·
predominately infect~ CD3- and CD 11 l>+ pha~ocytic cells bin concentration 1111d packed cell volume. with C\ idence of
imacrophages/monocyte:.) that expres~ major histocom- severe apla~tic anaemia.Ii:! occur during the cour:-e of the
patibilhy complex (.\IHCJ clas~ II anligens. 311 Rece1uly. it has dlsea~e. Anaemia \'arics In dt!gree. and ICll'!rus dr\'(1lops as a
been show11 18 that primary infl"ction 1,ith T t1111111/ma may result of erythrocyte de~t-ruction and liver damal(e. II ap-
induce an ab.:rtant T-cell activation. which result~ in failure pear,; l11a1 nm only merozoite~ are involved in ery1hroC)1e
of immune responses and which ma}•subsequemly rt-suit in desrruccion: caule infected by nt>ctlle i11ocula1ion with
an acute lethal infection. Schizc>nt-infected cells non-speci- strains of T. a111111la1a schlzoms that do not produce ernh·
tic:ally activate naive T cells: within 48 hours 101101,ing infec- rocytic meroznites may also suffer from a marked anaemia.
tion. large nu rnbers of macrophage,; del'elop in the medulla This suggesN rhat schizoms alone can trigger erythrocyte
of the lymph node thac drains the site of infection. Within dcsm1ction. II has been st1ggested that amo-lmmune
four day» of infection. these macrophage, \\Ctivate inappro- mechanisms pl~y a role in the parhogcnt"Sls of the
priate!) large number, of CD4 T cells \/4a MHC cla~s fl ami- anaemia. :H s:i Lcukopcmia occur,; in lethal and \Ublcthal in-
gens In 1hc medulla. but 1101 in the parar<>rtcx <>fthe lymph fecrions because of:i decline in circulating lymphocytes and
node. These events lead to diversion of,m effectiveinunune neutrophil~. fl coincides with the onset of fever. which tl- ac-
response. a, T cells are not primed in the nom1al anatomical companied by initial evidence or schizoms In lymph node.~.
site and .intigen-drin,n T-ccll stirnulacion undergoing An initial brief period of monocyt0penia i, followed hy
clonal c1<pansion does not take place.20 marked monocyto:,is in bmh fatal cases and rcco, cnng ani-
In culture. ·1: t1m111lma sporozoites in\'ade and transform mals. lhe monoC}1c1sts peaks becween three and four weeks
macrophages. and also B cells characterized by the CO2 after infe<'tion.'';, ion
lymphoid antigen.311• ~9 - 119 lnocularion of schizont-fnfccted In general. the pnthogencsis of the lesion• \\hich occur
cells int<> cau le results in the transfer <)f schizoms co cells or in organ, such as the lh·er. spleen. lungs and kidneys is
recipient animals. This was demonstrated by the iact that poorly understood. Ir has been suggested thac pulmonary
when ~chizom-infected cells from a cow \1C're inoculated congesrlon and oedema arc related to lymphpcytoiysb of
imo a male calf. the schizontS were lacer found ,,ithin the infected cells within the lungs ..J Toxins ha\'e been
lymphocytes of the recipient 121 The mechanism by 11hich suspected as causing haemorrhages and lesions in the
parasites arc transferred from infected cells 10 host cells is lii·er. 112 • ·n but the occurrence of such toxins has not yet
not known. lnoculatlon with T. t111m1lm11-infocted cell~ been demonstrated.
 Thel/Pria n111111lnra thelleriosis .J89

The tumour necrosis factor fD.7Fl-o. is com,idcrcd 10 play toxic cells an: kno\,11 to lyseschizonc-infected cells. but mar
a major role in the pathogene~is of 1ropici1l thcilcriosis. 1"· .1~ also act direetly on 1rophozoite-infec1ed cells by producing
The level of TNF-a produced spontanc:ously by mncroph- IH,-y. The ccll•Hne vnccim:isK1 induce resolution oflnfection
ages from cattle with either primary or challenge infection becau$c they generate consistent and prolonged macro-
increases upon exposure to Interferon CIKF)·"(.'1" Abnor- phage acti\"ll) against s<'hi1.ont-inrec1ed cells. when only
mally high level~ of INF-y1hat occur in lethal T. r1111111/am in- transient cytowxic T-cell responses arc detectable.M· 9 '
fections may induce excessive production of T:'\F-a, 1' 1~ The mechanisms 1ha1 control im.ra-el)1hrocytic mero-
and thu$ conrribute to the clinical sign~ and tis~ue damage wites are unknown, bu1 since nitric o~lde stops inva~ion of
in itlfected can le.'· 1"· 39 Exce;,,ive production of 'f'\rf .c,, may peripheral blood macrophages by sporo7.oites. 122 II may
be related to leukopenia. dissemina1ed haemorrhages and also pre"em invasion of ery1hrocyte, by meroioltes.sa Anti-
ulceratl\'e lesions.3 '1 T/1ei/eria a111,11/aw-infec1ed mo111l- hodies 10 parasite antigen!>. which de1·elop during primary
nut'lear ceUs produce !F:'\-y,9h whith may ha\'e two anLago- infection. 11 may inhibi1 in l'itro infection of cells br $poro-
nistic effects on the host immune response: zoi[es.1b but are 1.hought not to play a role m protection. be-
• to ,;tlmulate production of T"\F-a a1{d nitric oxide that cause they have not been shnwn to interact with 1hesurfuce,
protect cattle against infection: and of eitlwr schiiom-infected cells or of mero?.oite-infected
• to suppress proliferation and dampen the generation of cryt:hroc-yres.''8 Recovery from J. rm,111/ma primal)· i11fccuon
T-ceU responses. The suppressll'e acthitiC!> or the T-cell result~ in the de1elopmem ol a persiMent carrier s1a1e in
responses might over.vhelm its beneficial propcrties:i9 cattle. This carrier state is of cardinal importance in main-
taining immunity. and 1he accurate idemitication of carrier
Increases of calcium. chloride and sodium ions nnd of rotal animab could be useful in as,es.sing their immunes1a1m,. 30
protein in the com ems of the abomasum and duodenum dur-
ing the acuce stage ofT anmtlmainfection have been rcponed.
Clinical signs
It has been suggested that the los:, of thel..1! ions and proteins
from the interstitial fluid into the lumen of the digcsch·e 1rac1 fn natural infection~ th11 incubmion period varies from 9 to
may explail\ l.hc rapid debilitatio11of d.isensc<l cattle.7 -1 25 days. \\ith an ,l\'erage of 15 day~. I lowcver. the prepa1e111
Latent infeccions with various agents are oflen activated period may be ru. shon as ~even days in cattle inocula11:d
during the acute stage of tropical theileriosis. It i~ possible with a ~uspcnsion of mac:~·ra1cd tick,. nw prt'patenl period
that the massive destruction oi lymphocytes by the schiz- following inoculation of Infected blood varies from 12 10 30
onis !eads to a deficient immune respon~e. 113 dai•s, \\ith an average nf 17 days: on rare occasions it ma)'
Cell-mediated immunity appears 10 play a major role in extend to as long as IO month~.110
the protective immunity against T. 0111111/aw infections.2 l lnitinlly. ihere is fever, swelling of the superficial lymph
The l'arious parasite stages (t;ophozoite.5, schizonts. merb· nodes and incrcases in thc pulse and respiratory ratt.'1>.
7.0ites and erythrocytic mero1.oitesl and the cells they itl- Anorexia Md decreased rumen movements arc followed by
habit are affected by different mechanisn1s.1:1. ~a Tfleilerin dt:hy<lm1ion and constipation. Diarrhoea ·often occurs in
an,111/ara infection stimulat,'s innate (T-cell independent) young animals. possibly because of exacerba1ion of latent
and adaptive or a<:quired (T-cell dep,mdent) immune re- inft•ct((m. Emaciation is rapid. l'he erellds ma)' be swollen
sponses. The protective immunity against the infection tlC!· anti hypcra,:,mic, ;ind lachryma1lon and nasal discharge
pends upon 1he co-operation between the innate and occur. \nacmia, which becomes progressively more severe
adaptive immune S}'l:>tems.98 as ,een in infections \\ith other during the course of the disease, causes mucous mem-
parasites of macrophages.J·• brane~ 10 become paie. lcterus may bl' present. Pctcchiae on
The innare immune responses. stimulated by inJection the visible mucosae usually indicate an unfavourable prog-
\\ith sub!ethal dose~ of T. mwu/aw sporozoite~ or by hnmu· nosis.
uizalion with ~chizont-infected ceUs, include ac1h-a11on of rhe urine i~ dark brO\\~l as a result of the presenec of bi-
'cytostatic' macrophages, which suppress the proliforaiion of lirubin. In dairy cO\\'S there is a sudden and marked drop in
schizom-infecced cells.~; These macrophages spontaneously milk produc:1ion. .\bonion of1en occurs. Rare!~ , ner.·ous
produce TNF·et,Jt; and nitric oxidc. 1:t:? and. in co-operation signs ma~ appear in acute 1: (11m11/a1a infections one to
\\ith natural kiiler cells. lrsc schizom-infected cells.'"·"' The three days before demh.r'° Peracute. acute. subacuce.
nitric oxide, implicated as an effective mediator of pro1ec:tive c:hronic and mild forms of the disease ha,·e hcen reported. ·o
immw,ity, t:a has also been sho\\'n to inhibit the proliferation rhe peracute form,~ most like!~ to occur in highl~ suscep-
of schi2om-infected cells. thus causing schizoms to disap- tible canle breeds. after a massive mfection. ·r11c clinical pe·
peor and host cells 10 become apoptopic. 100 riod lash for bel\,·een three and five days and end~ 1,i1h the
The adaptive immune responses include production of death of the animal. Typical gross pathological changes gen-
cytotoxic T cells for schizom-infecwd cells.3 :u. "1· 98 anti- erally do not ha\'e time to develop. but numerous schizonts
gen-sensitized lymphocyres93 and CD.J · I cells. 1" The t')'tO· may be present.
490 ~ti.(.lto:,.. mu: Protoz.onl dist"tlses:

The acule form is Lhe type mosl frequemly seen in sus- disseminated haemorrhages. :\ccording to Sergent and col·
ceptible breeds. Clinical :,igns are well defined and the dis- laborators, 110 the lesions in 7: mm11l11ta infection are similar
ease lasts for one or rwo \\'eeks. ReCO\'ery is «low. especially to those obser\'Cd in hncmorrhagic sept!caernias. being
in aduh caule. and morcalily is high. characterized by haemorrhages and congestion.
The subacutc form resembles the acute form, but clinical In typkal case:~. the carcass is emaciated. anaemic and
signs are lcss incense. Most animals recover if complication~ icteric. \1ith yellow gclntinous connecllvl' tissues. !':umer-
from secondary infec1ions. such as bacterial enteritis. par- ous petethiae or larger haemorrhages are seen in the subcu-
ticularly in calve~. do not occur. taneous tissues and in the mucous and serous membranes.
The chron ic fom1 is rarel) encountered. ,\lfected canle All superficial and internal lymph nodes are swollen and
are recumben L. emacia1ed. and show incermittent fever. ln often haemorrhagic. The spleen is enlarged . mainly as a re-
most cases death oixurs af1er two to three week~. rarely after sult of hyperplasia of lymphoid tis,ue. The liver is swollen,
four weeks. brownish or orange-yellow in colour. and friable because of
The mild form is encoumered in naturally resistam parenchymatous degeneration. Subcapsular haemorrhages
cattle, in cattle protect~><l by passi've immunity, in vacci- an.' onen observed. The gall bladder is distended and
nated cattle with only partial immunity. or when the infect- contains dark-green. thick bile.
ing strain is of lo\\' ,1rulence. Cli nical signs are generally Epi- and endotardlal petechiae and ccch~'lUOses are
unnoticed and cattle recO\·er spontaneously. found. The myocardium is onen degenerated and shows ex-
In protracted cases. skin lesions consis1ing of lenricular tensh-e haemorrhagic areas (Figure37.2J. Excesshe amounts
haemorrhagic spots or skin nodules. which are generated in of serous fluids may occur in the pericardia!, thoracic and
the pei;i\'~c-ular lymphoid tissues of the dermal anerioles. other body c3,ities. The lungs are frequemly oedematous.
are sometimes observed. Ulceration <>f the nodules may ·n,e kidneys are congested and may show haemorrhages or
occur and Giemsa-stained touch preparations reveaJ small (I to 3 mmJ. whitish-red foci oflymphoid hyperplasia.
numerous sch izom-infected cells.&.· 110 Deprndlng upon the durntion of the infeC'tion before deach.
Rarely, infection may occur, even ,vith an otherwise viru- red patche, or ulcers may be found in the mucotc, mem-
lent strain of T. amwlma. in which there are no clinical signs branes of rhe abomasum and small and large intestines. The
and often no detectable parasiles, because the animal is urinary bladder contains urine pigmented dark brown
strongli• resistant to infection. This condition has been re- because oft he presence of bile pigments.
ferred to as an immediate latent lnrect ion (infection lme111e
d'emblee).10A
In practice It is not always feasible to cla,;sif) a field case
Into one of the above categories when it is fir$t diagnosed.
:-tonalicy rate1, of ~O to 90 per i;ent ha\'e been reported
from various geographical area$: 0 This range is apparently
due to variations in susceptibility of cat lie and In the viru-
lence of 1he parasites in\'olved. ,\dult animals which have
apparently recovered may die two 10 three week$ after the
disappearance of schizonts as a result of complications.
such as li\'er dysfunction. 63• 11' Growth may be impaired in
young calves that recover from the ciisease.

Pathology
Significam increases in the levels o r sc!rum gllaamic oxalo-
acetic transaminase, sorbitol dehydrogenase, isocitric
dehydrogenase, 1r; alanine aminotransferase. alkal ine
phosphate. creatlnine kinase and y.glutamyluansferase. as
tvell as increases in the concentrarions of uric acid. blood
urea nitrogen and bilirubin IOG have been found in cattle
suffering from tropical theileriosis. Haematological
change5 consist of se,,ere anaemia, anisoc~-cosis, and
basophilic stippling.
In animals which ha\'e died. the severicy oflesions is pri·
marily dependcm upon the course of the disease. In fulml-
nant cases. lasting only a few days. lesions are limited Figure 37.2 Ecchvmoses and petecttiae ,11 ma myocardium
mainly to hyperplasia of the lymphoid tissues and to Expe•imenta 1nfeclion w,thwulen; sch•ronts
 111ei!uria t1111111/t1w thcileriosis ,191

Exceptionally. p apular and puswlar skin lesions (mainly antibodies to other Tliei/eria spp. may cross-reac1 \,ith the
in the non-pigmented areas} which may be ulcerated or ne· 1'. m11111/ma amigen. Th~ IFAT b u,,.eful for identifying
erotic, may be observed. These lesions are infiltrated by infected herds containing carrier animals bur is not suffi-
neutrophils and lymphocy,es. as well as cells comaining ciemly accurate to decen all iniected individuals. /\either
schixoms.liS the ~chizom nor meror.oite IFAT detects antibodie~ in every
The histopathology of tropical theileriosis has been anlmol "ilh cry'throcytic merozoites in its peripheral
blood. "·"°
2
poorly studied.
In cattle that have died after ha,·ing shown cerebral dis- rhe enr.rrne•link.ed immunosorbent a~say [El.IS.'\! ha!:
turbances. clogging of meningeal and cerebral blood vessels been successf-ully adap1cd for detection ofanci -T. a1m11/ata
by large numbers of lymphoc)'les has been observed.60 antibodies. 15 Jn con irasl to che U:AT. schi2onL antigen used
in ELIS/\ Is ]!;!s~ sensitive and specific than the a ntigen pre-
pared from erythrocytic mcro%ol,es. since the latter detects
Diagnosis and differenliaJ diagnosis
ant ibodies for a longer perfod. 5A, ·'9 Serological tes ts are
In areas endemic for tropical theileriosfs, the non-specific ma inly used for epidemiological screening for T. ammlara
clinical signs of fever. anaemia, icterus and swelling of pe- infecrion and as~essmenl of 1he results or lmmuni7.atlon
ripheral lymph nodes justify a presumpth•e diagnosis of wilh lh·e ,·accines. 11 appears lhat Im munity ro
acute T. annulata infection. T. a111111/am theileriosis i~ mainly cell-mediated. but anti-
For laboratory confirmation. thin smeah ~hould be pre- bodies can be used a;. an indication for establlshment and
pared from biops}' material of lymph nodes or the liver. multiplication of Theilcrin paraslres which induce a
Blood films prepared during the initial stage of the disease protec1i,e immunity.
show rare or no erythroCytic; merotoi tcs, a picture also ob· ONA techniques based on ~equence analysis of genes en-
se!'\·ed in recovered comer animals. On rhe other hand . rhe coding for panicular proteins ha,·e enabled the develop-
presence of schizonts in lymph nodes and the liver is often ment of molecular methods for the diagnosis of T. ammlata
sufficient 10s11ppor1 a diagnosis or acute thei leriosis. Smears infection. Specific polymerase chain reaction (PCR) prim-
made from liver tissue are considered by Sergent et al. 111 to ers. dilrived from a gene encoding for a 30 k.Da merozoite an-
be the mos1 suitable means for the detection of schizonts rigen that is used to test samples of whole blood, detects a
and for the evaluation of the ~c\'erlty of 1he lnfec1ion. parnsitaemia as low as 0.000048 per cem.3t and can also be
fn T. wmulmt1 infection. schizonts are rart? in the periph- used LO detect the p.irasite in l Tyalomma iicks.30
eral blood, and 1heir presence in blood smears indicates a

• .'~ ...
poor prognosis. T he numb£'r or parasitized erythrocytesfa- •
creases progressively during 1he course of the disease. and ;
l' ~
'
-
as many as 95 per cent of the red blood cells may be infected
.. •
(Figure 37.3), but 1hc severity of the disease does noc neces- '-·.,. :
,,
~- ...,
~
: '
sarily correspond to the percentage of parasirlzed erythro- • •
•• , •4
cytes. The Giemsa staining method is considered 10 be the •• )

most suilable one for demonstrating parasites in smears.

•·¼" •• ¥ ., .-. : :).

"'
To detect schizonts and merozoites in fat.al cases. impres-
sion smears of the liver. spleen and several l~1nph nodes ' , .., ...,
'•
..• .
~ • I
.. .. -
.,: ,
:,: •
showd be examined. Both intracellular and free-lying
sch!zonts may be detected. the latter having been released
"i
•
' ..
•

from parnsiti7.cd cells during preparation of the smears. Most v

••
of the merozoitl!S in 1heacme s1age or the infection are spheri-
cal or o,·al-shaped and are smaller ( I to 2 µmJ than the mero-
)
- t
-•
wites observed in the chronic carrier condicion. /1 lerozoites Figure 37.3 El'/tl1rOC'/1•C meroioites of Thellerie emw1a1a to11ow1ng
uek·,nduted infet110n. " 1000
persi$t for moml1s or even yeors In recovered tattle, but they
are 1101 always detet;tahle by direct microscopic exam inarion
of smears. On the other hand. the presence of schizoms is aJ. Endemic regions of ·r a11111ilatt1 and T. par11a infections
ways associated with die acute stage of infection. do no1 O\'erlap except. perhaps. in East Africa in t11e
Ttie most widely used serological test for detecting latent proximity of the fourth to eighth parallels north. Schizoms
T. a111111/am infection is the indirect fl uorescent antibody ofT. n11n11/atn are very s imilar to those of T. ptu-11a. but those
te$t (! FAT}. 80th schizonts and erythrocyiic merozoiles are of the former arc us11all)' scarce 111 peripheral blood.
used as antigens.~6• s9. 8-1 The schizom antigen appears to be Mic:roschi1.onts are more often observed In Eas1 Coast fe\'er
significantly better than the merozoite antigen for detecting rhan in tropical rheileriosis and. in addition. rhe majority of
animals c-drrying 'f. <1111wlt11a infection.26 Th,e lFAT is spe· the ery1hrocytic mero;-;01tes in the fom1er disease are
cific only in areas where 7'. a,11111/am occurs alone, since comma- or rod-shaped.
 Theiler/a a111111/n1n thcilcriosfa 493

Growth and serial passages In culture result 111 attenua- immunlt~· against field-transmitted infection in indigenous
tion of Lhe virulence of Lhe scllizonts.2' 91 At first. cultured su.sceptible cattle as well as in a high-grade exotic stock
schizoncs or originally virulent strains cause severe clinical breed introduced into theileriosis-endemic regions..s 1 How-
disease and evcnrually death when inoculated into suscept- ever. field observations ha\'ll shown cha1 some vaccinated
ible cattle. After several weeks or momhs of cultivation, with cattle suffer se\·ere theileriosis four or more years after vac-
passages being performed about mice a week. milder clini- cination.911 Jt seems., therefore, thac the immunity acquired
cal reactions are observed when schizonts are Injected imo after a single inoculation of culture-derived schii;om vac-
cattle. ,\licroscopically, only a few schizonrs are found in cine is not life-long, and revaccinarion might be required for
internal organs and rare erythrocytlc merozoites occur in canle in herds with low tick-lnfcstation ra1es. The response
peripheral blood. Complete arrenuarion has been achieved to primary immunization with ami-theilerial vaccine con-
when calves inoculated \\1th cultured parasites do not de- taining schizom-infected cells is barely influenced by the
velop clinical signs, and when schizoms or ery1hrocytic histocompatibility of the schizont-infected cell and the
merozoites cannot be demonstrated in smears made from recipiem animals. I lowever, together \\'ith the immune re-
their blood and tissues. 7 7· 80 • 9'.;\"arious field isolates of T. a11- spons.e 10 the 711eileria scltizoms, an allogenic response to
1111!a1a may require three montl,s to three years of continu- tile cells carrying the parasites occurs:51 • 5i Uponrevaccioa-
ous cultivation in order to become completely anem1ated, lion the allogenic antibody can block the parasite transfer to
and quality cont m l of the culture-derived schizom vaccine the recipient animal and prevent rhe enhancement of im·
comprises testing for safety, s1eriliry, potency and efficacy.es munity at the time of a second vaccination with the same
The cell culture vaccine is safe for all breeds and ages of culture-derived schlzom-infected cell line.71 On the other
caule, regardless of their physiological sracus. such as preg- hand. heterologousschizont-infected cells from low-culture
nancy and lactation. 86 No erythrocytic merozoites are de· passages conslderabl~· boosted the immunity ngainst rick-
1ec1ed in vaccinated cattle and they are. therefore, not a transmitted infection.117
source of Infection 10 ticks. rn~ Vaccine prepared from fresh As with other tick-borne diseases. tropical theileriosis can
suspensions of cultured schizont~ has a limited shelf life or be prevented br contcol or eradication of the vectors. The
only a few days a1 4 •c, but storage in a concentrated fonu in sensitiVit)· ro aeariddes of $011le 1iyalommaspp. has been re-
liquid nitrogen extends the effeccive life of the vaccine to at cently evaluated,"· 5 1. 127 but adequate rreatmenr regimes co
least two years.81 control or eradicate ll1ese ticks have not yet been critically
Vaccine contain ing completely anenuared sch i1.onrs in- evaluated. Since a single infected rick can cause fatal theile-
duces a marked production of specific, but not necessaril)' riosis,88 tick control must be highly efficil!nt if ll is to be effec-
protectl\·e, antibodies that can be detected serologically. 10 tive. The time schedule for acaricide treatment should take
There arc indications that some mulliplication oi such at- i1110 account ti1e effecrJvc residual artivity of the acaricide and
tenuated schizoms occurs in inoculated animals. since the ve1yshon period required for mamration of 1: amutlara
equh·alent numbers of killed schizonts inoculated withom sporozoites in attad,ed ticks. 116 Under some environmemal
an adjuvant do not induce the formation of significant circumstances ticks may carry mature sporozoites even be-
serum antibody levels. fore they commence feeding,Gll and such ricks may transmit
Labonnory challenge of schizom-vaccinated cattle \\ith the infection "itbin a few hours after attachment. In tick-
tlck-deril•ed sporozoitcs results in moderate or no clinical in fcstcd barns, periodic dusting of the floor w[th acarlcide
manifestation.42· 75 and immunity laS1s for at leas1 rhree- after removal of litter and manure is recommended. in addi-
and-a-half years.1 26 The schizont vaccine induces effective tion w acaricide 1ream1ent of the cau le.

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1nreC1ed lymph6id cen hne.. Pt1rmirolllgy R/!5,wn:11. 79. 1,s-182. 'i'AKUf', .!t.. 19,1.. 'R~pon.~cs in unim:il,., c1cc1nntlod \\'Ith 1hc 7'/1(1/lt'ntr
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lt.P,, CNUUXGlON1 )l.D.. ~rt.\Z.:l'.\L. n. ,>..,SPOON.CFC, r.R .. 1%3 J>l'fl(l11cr/o11. 29. 109S--1 l:IS.
494 ,;uni), 1wn: P(01or.oal diseases

9 JU!:'\ \UUiJl, I... UHl11AC,1, t..,,. Hf::M:'\'.\f{nl, (;,, '11ilM()Sf., 1.JII,, UAK(i()llflt, \\., c::1lvts: ., comparl1Jon bt·{\..ttn hup.trvnquon(\- ;tnd Ions ;'!t'ttng
U.0\1,\MOUR, ., •• t;J~ ~\JJ\0, , .. ~llll.J.S, n.~
urr, A, & HflCI\\~. C:.\o,U... 1994, 0).)-1actac:)"C"Jlnc. Ret~1re11 itl 1/ci.Wiunry Sroln,·o, 49, 110-112.
C:.,nomic a11d ph~Ol}l)IC dh"•r<lt) ofTunisi•n '111ef!trln 11111111/au; 90 I>£ ,n,. 1.. u·ou>1•uu. c. & JO:-<GUAS. F. 19,3, IJoiectlon of protow:i.n
bol•1c,. Pflf/t5ttolog;•. 108. ~l-<50. p.,ruirt!' 111eili1rin ttmwftJCfl in liJ'lllommn tickS by polymtraS\.' t.hain
to e-1suor, I\.P.. .-\1.t-\.OPP, !I••\.., Sll()Qsrn, P.tt.., snuA:,w,,i., 1u:,, )tOk~-\klA. s.1•. 4 renctlon. E.xp~rimcmtJ.J ,ind AppJJrd Ar111·olog\·. 11. 839-8-i6.
coaRlrJIT, ,u.. 1995. l'lldleria· !mpro,·ed specl;; dlsc:r!minatlon u,;lllJI 31 T>OlA'X, T,T,. 1989, TI1C'ilcnasis: a comprchcms1n:: n•vit"""· Rf!,•ur
0llgonuclcu1klcs·dtn""<I f,o,n lnrgc ,ubunh ribo-.om•I RNA scqucnees. sdi11111jilquu1 1'«:hniq,,e. Offie< tm,mnr/om,l rl~ Ep/UJ011cs. 6. I l..36.
Expmm,1110/ i'tlm.1i10/ogy. 80, 107-115,
32 o 'oU\'1!Ik.\, G. \'AN"Of..R WUtlf. M .• H.ABEL."4 \1.A., J,\Q)Ul f;T. P. « 10,m.JA~.
ll OOtJ ,nouR,.A•• O,\RGHOUTII, \l-i\., & BE.~ MllJ..1), t.... tS96. C...ttlc... infbC.lrion ..... is-ss. 0~1cctlon ofT11ullt·rln nmmlotn in bJo.o d$mp1Qof c.srncr
b)· ilJwlommo tlcL<and pr,,-..lonce of Thtileritt in H th:r/111111 •~cio<,n <11<tle by l'CR.Jo,miq/ ofCl/11/cal Microbiology, 33, 2(;w-2669.
Tunl,,I~. V<1n111n,y .ot,ro,11n/t,gy, GS. 23:1-2-1-,;,
33 D~f"HU:".K(.)\\ -,I:;'\', t,. R UJH~. , ... I~.:~ Of(' pfrupfa.)me)'t.(111 <lcr Rlndcr.
lZ 80ULT01t, X.R., <it...\$$. t.. ,_,,c;m, P.. 8tJ.L·S~Jm. L. lJk()Wl<. C.G.D. < 7.m1rnlbla11ftlr /Jakteriologlt Parasitqn~wula lnfek1ionkrtmkJ1eilm. 35.
N.u1.. 11.. 1995. 1'11ei'ltrib unnulma ~porot01te antigen fus.t..*d oo hepami!t 8 486-l92.
core antigen used rn n \'9:t'C'irlatlon tri.a.1. \laccln'1, l.13. 1lS.2-1100.
J<: fl!..\ttC)'.\ . D,T.-. l..C)Q.Spt\, A,M, 1996, The inMTIJt:th,-e N>lc afinn~te
13 sou1.TER, N.R. ,:..it.Au.• a. 1999, Tmmunl:y and ,,accinedcvelopmcnt rn the immurut}' in the acqulr(!d immunt re,,.pon)e. ScJi,11t•c. 272. so-.;.;.
bovine thencri~is. Adt'aucrs in Pa.rtmwfr/tJ•. ,S-4. 4 l-9i.
33 ,.,_,c::J.1. E.J. "' mllrEIJ...t. u •. 199'3. The c11idcmI0!t,gy oftrc,pirnl rh-r!foriosl-s
u k"OO\\"S, c.o.o .. ,983,. Theileru1. In; ,c.~s,:..~. J,O .. (ed.1. Jn \ 1tru C:u/ti1'tUiOt1 l1'/1eilrrio aJ1nulora lnfocllon in conleJ fn on c,idcmlc on,a of ~!omccn
ofPro1oz,x111 Para,11fs. Boca Ra1on. Florld,r. CRCPrcss. lno. Vl!Wl'f111.lry Parasitolpgy. ~.J. 31-65.
15 BT«)\\-:-O', c.o.o. 1990. Control oftropkul 1he.Ucrio,"h. rn,ulleril1nm1ulmu 36 FL\Ctt. E.(., OUUllLl. IL, WAt>DINGiOS', O, &lit. Hi\$XA.OU1, ,1.. 1S93.
lnfccrion) of cnt1i<'. f>m<1SS11,µogln.32, 23-,31. PrC\11lencc ofTI1rilt.1rf11 hl the tlck Ht<tlomma. dc1n'wm detrfrum In :ht>
16 OftOW~. D.. C'.Ar.£P¥£LL, J., avs:-1;;u.. (;,. 1101>):IN"S, ,. liGL,\S$.1: . 19.9S, ,, ce:H f>ould<ala region, Mort>C\:o. •'1etlil:tll a111t \'l!t,rlttflf)' E,uomo/ui:)·, 7.
acth..stfon hy Thtl{rria nmmlnm nmccophilJ;<!S corrtL11cs wfth C)'Wkinc 3,;3-350.
pioductl<>n, C/111icnla11d Ex11ertm,111a//mmu,,a/1>gy. 102. 507-SU. 37 FLKti. tc:.1 .. OUt-ltU.J, H., \\'APm~Cl n~. fJ.. OUIJIC.H, M, k.SPOO:-.:CA ,u.
17 J,, ORQW~. Q., ~!t:.H,\"\'J,A,, lfOWU!, $,, "''00;\"f,A.', ft.. nuss. r.,
CA,.\0181ill. 199.5. Fa<lO/$ mflucnclngth• transmi$Sioo and locidcncc ottrap,cat
i99;-. A non-pr<itcc!l11, T h~l~r l r~ponse ui;a!n,1 the intra· 1heUcnOS"i!. \ T1hrllt•rlr, a,mutam irtff.cllon of caufc:l in Morocco.
macrophage prqtozo:m Th,llt•rin nnmdau,. Ol11icn/ amt £.tptUittu.lJ11t1/ VtttrlnoryP,,ru.,tltOlogy, 59, l ii-188.
lmm1mol(1g)', 10$. ~63-l;o, JU' fQP$\"XlL i... rACl:.SQ:O:. L. \\~·rr G.• ~A-"\"DERSON. "·· snO\\~!\ 0, f"
18: CU1PREU.. J.1 H0\\11£1 S., QDUX<i, t. &· (':I....\SS, J:. 199j, nwt((•r,(l illlltli/(Wl PftESTOS. P.. l99i', Bovine cclb, 1n(<.'C1ed m z•iro wnh Tlrt•it~ra ,m11u/atn

tnduOM Jbcrmnt T crll acm,nlon ill 1•/rro and ;,r """'· C//11/cn/ Md express CO I lb, the C3b1 complement r,:ccp1or Ve,,•rinary Fks,nrc/J
Exp.mmetil(t/ Jmmm1alo[Q•. 99. 203-210. C..'ommuni<-<1tit:ms, 21, 249-263

l9 C\.\U•UUL J,~ NtCH..\.\:"l, A., 8!,1:0\\'~. t), , lCU\\'JE.. S., SPOO'l:£Jt, R 4tGU.;,.-,, L, 39 fO~)'Ttl . L )l,G,1 \-UNN!l:·,·1-',C:•• f.JK\'t\Jt. f,,;\IH"\JW~. n.b., H,\U,. .-.,-..

1997. PamsJtL..,m0<.liatt1d swp~ in 1mmuna respon.s.t-.s !allure during \1CQ1u.s.1, t-.. 11:RQW:r-:, c...c:;,n. ~ PM£5,lD"!\, r,M .• 1999- ·r1s..1.ue d.tmij,£e 1n cattle:
primal')' 11:tllerill m11111/nro-lnf•<1£0n. frtJplc,1/ A11lmnl f/<"<lltl/ nnll infec,ed '"tl'
17,cller,n n111111/mn accompMled by nic1as111S» of
P~ual,m, 29, 13.'IS-l~S. C)'lokine-produc1ns ~h,1.ont-infocu..-d mononudear phagm.:}1Cto.
/01m1nl oJC.omparnl/lH! Pnthol~·. 120, 39-57,
'20 U.MP8l!Ll. J. & SPOO.S£1t. R. 1999. ).13CTOphagc bch-avtn,t badl)": it'l!C'!Ct(•d
ctlls. and sub,·~r.;io,~ or imrnune rcpon.s~ 10 TltPiltri,; i11}t1t1fnf(I. 4U r,i,'-~~. M.. rn•1iso, f.. ft R0$1.i.XRFJtC, .\., 19;-,. TI1cdlagno.sls of Tltl!/ltria
/'ttmsirotogy Todl,y. 15. 10-16. mmulttta infcc:tfon h)· the lndln.'<1 nuon.i..cent .1ntlbOd) met.hod u>lng
$mall qunn1mr,s of blood dried on ob>0rbent paper. Ref,t(lh \',tarlm>rirh.
:?l CO~KAD, l'.A•• Xt:LLY, a.c •• .8AO\\~. (:.(U)... l9ij$, lntr'nt.".l')"lhrocytie
?8, ';$,,,,82.
M:hlzogony of 11ttif~rlutmnu!tlla. Pornsftol~"Y~ 91. 67-82.
-H (.JU ~ ., S&A.~~-\t, O.C:.. 31tAT"tACH \M\1H..U. \' .. VAUJ.\, 0, be ~J~OH, A. 1980
Z2- 00~'1,[I, (,,, C'.AMPttt.u •• , ... S((lf,,s,. , .• t-J,.\$S. F~.Sf'OONfR, fl. f.c hHMll'>, 1.. lnununQlogJal 1cla1lo11shlJ> beu,.,.,n >train., of /1i,l/,rfn a11m1/mo
1,q98. E,idence forsLroin spectr,city inc)'\01oxicT-lymphocyt"· ·o.chunkowslly •nd Lull> !!104) R,""11Yh In l'~1erlmUJ• !klmrr. 2!!,
m<!dia1cd, major his1ocompa1ibUicycomplcx cl3's l·dcpcndom killiugof 93..9;,
Tlnuleriu ,m,,ukt1t1,infcctcd cclls. Pm'usiUJl()gy Rt.'"$tard,, a.;. 593-595.
a:! c;uJ... 8.$.. rn-u.rrAcL\R\1n.u. ,,•. N\Ufl.. o." :.lSCH. 1\0 1g;t;. \'acrln:ufon
23 c:o..x• .,.•E..c.. 1982. C:<'Jl-medl:ttt.:d 1mmuni,y in tlte:Uen9~~ Nm11re. 295. 14 _ig.i1rh1 bo,1rw ttO-p!cal thtllcri?."ob Th1•il,...rl1.1 amrnlatn)• ."VflWI\'. 264.
2.; t>AJWHOUTH. M.r...,\., Ot.'-·~IIUD, 1... flOUATIOUH, , .. Mfl)«)}a:, T, • .H1'C>W.:,_,, 35S-3'5G,
c.G.o. ~ Kll.Al<I, M.. 1996. A prellminarrs1udy on the 011enuation of i1:3 GlU.. 8 ..).., Dl«lTM~lfi\RYULU. Y ~AUS. D, ¥. ~l;XOH. A,. J978.
runi.sfan sch l~<>m ·in!ectcc! ce.U~·lincs. o( T1,eif1.·ru1 un,1111,aa. Chc:mt>i'Hophyf:ixii whh w1rnc:y-dinc-drug~ in 1he immuo:1.1:uh,n of ennil
Pffmsimlos;y R.t•,,;eorclr, 82. 647-..655. agaLn.,t Tiu!lh!rfll t1t1n11/a1a ln!ectlon. /111mu11/rmt1/ /ottrnnl oj
2,3 b.\,RCIIOU11t, '\t,l-A., tU\tl.\tTt)Uk, A,..Jll:.X·MIU1U, t...J.:U.AXf, '\1, 4,- IJk{)\\ :,;:, />nrnsi10/ogy. 8. ·16,-169.
c."'1>,, 1996, Eptdemioloi;y of tropical 1hellerlosls (//tffprin nnm,larn 4.1 GM\'. ~1.A. & oRowN. c.C.t1•• 1981. In 1•irro ncumtli>'JUfon ofthcilt!rial
in(ecuon or cnnlel in an e11dem,c ieg,011 o!Tutiijia; charoeu,ruation on ,poro.to!t~ inff.'Cti\ity with lm.tnune Siin.tt:J. /1J; t9\'tl\'. A,.O .. CUN'\t~·GH.1.\M.
endeuucily •'1ntC$. \'fterinory Ptlrasi1of11gy. 65. 19.!1--211 , >LP.~ rouxG, ,_,.. Ced$). Ad,wnres /1111"• D)mml o/T11e1/rrions. 111~
.:>:6 Ot\R<a-lOUTII, \l.f_ \., &0-.AT'fc;)UR, /1., ftf.N•Mll,t.D, Lat SA.~~ I , iggtt Hague: ~!anlnu, Nijholf l'ubllshc~.
Omg:nm.11, nr 11:t>ilermimnultJ(tJ mrecuon uf cartJe in Tu,nl.!)13.: 4S t~RA'h .\LA.. t uoaxs. A.G., f<At.. ..... & tUt('J\\' S. C.C.1>.1 tg.80, E\'alU3t1on or an
comparison ofserology-:md btood smears. Viierim.1()' Hrs~nrch. 27, cntyoic im.munoassny for serod.J,!ftnosis or iufcctions wh.h Tllt'llcritt
613-t;l?J. J)(l.ntJ and T, t1.rmulm,1.. lleMv1rth in 1'derina') ScUmtt, 29. 360-366.
2:- OIJA'R, ~•• \\>\I i101a.,, 0.\ .• rorn~HA:'\., c. N~UTA)I. 0,f'., l9f\i ,ab nAu... ,=,n., 19~8.Antigensand immuniiy tn 1'heiltrlonnuu!ata.
Chemohnmunoprophylr<;<is "i•h b11parvaquo11e ~gau,st thellerlolNS ,n Pnra:.,uo~·7ixlny. ,,. 257-26!.
c-alves. lfelerinory Rucrmf. 120, 3;~.
,r. HMH~H-1-1.Stf\TO,:f, R, ttsHAD~DEL. f ... 1974, ·1111: therupeu1k,alut> o(
;:.8 r.m..ut. ~.. ~MUH;>Ttv., r;>., .. BHUl:ittA,, c:. «<,i,AUTAM. u.,.. isaa. Trcauncn1 o( o,;wiracydinc h)'drocl>loride lterram)'t'inl ln cottlc ,nfocted
r.x1wrimcnwlly Induced 111,i/eria a11nulnra infccllqn in cro,<s·hred c-xpe.nmemnlly wich Tl,e;Wrlu ummlar11. TrtJpiml Animc,I Ht.Yilrh uml
eah·e.s,'11h hup3l'\'1lquone. Vcierinnl)· Pcuns;u,Jogy. 21. 26;'"-2:S. Pro(/11(1/011. $, 1111-1 ZI,
~ Oli.A.R, f-., ~b\UIC') t"KA. U.\ • OHUNL\:,_,, c. ., c.~u·rAM, (l.J,. 1990. ..a HOO~lL\IA,o•llAO, , ... 1976. nie p:uhogeneslsofanaemlaln l1wll~rl11
Chemoimmunoprophylaxls agi,Jns1 l>o,in~ tropical lhollerlo,ls In young a111111/ar11tnfectlon. Re,corc/1 /11 \ er<rlttnr;· Sdenm 20, 32-1-3:?9
 Thrilerin 0111111/nrn thdleno,i~ 495

,:g HOW.\l\n, c.. ·~(WC•..... rKt:,'"tn~. P ... f1\Ct,i"~C),,. I.. f./ tlMll\\'S+ u ... l~J r,8 " ' LRO"•ti. f .H .. ftRUW'I:. I:.(, u. 1. ..11 ,R,1 \. ft,.U .. 19Uo. Glut OSI!. r,hospltnlC
Pheno"vl< anal)"* of b0,1ne leukuc:.ie !111,.., lnfo<l<'<l \\ilh m,11,,;., l>nm•ia•• boc11zm1,• pa11crns In bo,inr lrmphabl ..<1t1id ,:,,II tin,.,.
or.nulnt«. \''"'U•rlumylmmunoJor> tuid /1111m1,m1KJ.1ltlXogy. 39 275-:?8?.. infttch.-d \\ilh ntrilnM mnmlatt, ,md I pnh·o, wilh .,n fmptQ\\\d
crw.~ m~ YJ .. u,tfri..1fion method u:-JO& meltfula Diec lkwnrc-h mt,m·nnal}
so ,.1t. , .• -..mu"'· 11 k ""ow,, <..r,.n., 1998.
tUJ..,:\. t., \,nu.,~1~0~. , .. "'-'K'
11,~V(u,a amwlma; cnmcr'\tute and Jmmunt1y. Annal.An/ l'nrt .,·,~w :x·umrr. 29.2~30.l,
69 ,,uSl)'L Lt .. ~IC (;.(UfH. \'., IJHO\\ ~- (.t;,,l).," '1,~(Utl'ARl-\.s.,1 • 1981 Prt.limimu:y
.~uufi.'myo/Xumcc~. 8J9. t()9-12S.
in\·t~1i&a1101h. on l'-Ol·nz)nte \':Jtf..1nt~ of l}"rnphobla~totd cdl lhu.~
jJ t!\--Sb,. £,,\., ~m.L.,R. p ,, BROW:'\ c.r...D, Y 5r00~[R. lU- l~ l'hc fnfe,.'tod \\'Uh 1'i1t'l!1.·l'll1 ~p<.-eie,., R,..i,•m(h in \ ',;Mrltttt"' Srrf,11t,1. 3Q. 38-:3
d.,vdopmcnt and «.pc:cUi.cny of ~'toro~u; (:elf'- in c;iule ,mmnn,1,ed \,11h
-o ~1 J 11.. w.o,. 19Sj. lhl>ilcrin"'i~. gundl'f10.,,1....,. und C)'timXL.0()1'10~~~ ;1
autoio-~Qu, or '1.llngl·n\"iC n,Nlr"na-in<ecwt.l lymphobl,l.),t(11c.l ,·cll lin\.'"
Ptu'flsi1rtmmu11olog;•, JJ. S7-66.
rt\'!~\ Ondtr$-1c:1,uun Jour,Jal ofl'1.1:enna,,· 'Rt"St'-4'<'1. 2:". 21..,..430

32: l~~f-; •••• C.mttfol It, H,. OlT\'TR. A.\,, '.lil)S,-~0:'\, '.,,.,P., HMO\\' '• C..<t 1), •
- 1 ~1c1u.,t. .\. RRo,,•,., .,•.n. ;<.-!\t•oo:,,,.'T.R. n.1-, 1997, m,\V.JC'cln:ttlnn \.\ith the
~J•OOX-lfR. R,L l98g. ·n,~ eiTl.'Cl of ~li IC:i·C1mp.atib1H1}' hct\\'et."!1 ~nme nw1Mia 1l111111/111n illr,...~ecJ c(~f lln• may not 1>,, fca.Jblc ru1
blio"lng lmmunh,· ~l!'lin<1 rrorfcal 1h<.'ll<-rio <s. TrbJ1:ml ,\ntmnl H~aitfJ
0
par;;site-;nfec1ed cell line and re<:tp1rn1 1n lmmllnliauon 3g3!1m
tropical U'\cilcr:osl, Pllrmrt,: lmmum>ft'Jf}·, t 1. J7-SO. nml l'l't'Hi11ctio11. ~. t US l 18~

53 1R\1'.'.', .,.n. #.
,10Mm-:,ox. w.1 .. t9&:- lmmum.1pmhol~)~\ immunqlngy 1Snd ;.:. M\:lf,\~I .\., ,;.\~UJ.\. "" );te.Alfll,,.... <-tM,,!'>., IU \:l\\'J\l. ..... c,u•rA. <,. t.:lht\R,
unmunoprophylaxis ot n,-llffw 111fC1CL1Un5. In: sout.Sur. l.l, t 1ed. k ., ,1t.,n ....... R. 1998 f.plcl•mlololl) •nd ,onuol ol 1topicnl 1hullcrla~I~ ln
Irr.mun.? R{!Sponses in Pa,ct$it(c Jnfrttions· lmnumofOK)·. ~t:Jtth 1nd1n. Ptoc,wlf11gsn/1h,•Sl1i J,u,·mntfon«l Vrtt'Tl11nrJ· t,mmmr,fo~
lmmuuopi1tltofo~n1zd Jmmu>:oJ)rcpi:.~·ffttis. \ ul. 111. Boe.a Rotun. ~ymp(J..~iu,11 Pmlj(ll,,lgr1rr1/fUfO/ U111fw~Jl)", /.~1dhim1n, /utlft1
Flond:i: CNC Pro<.<. lllc. n ~t,11\:\1 .\."' ntonp. n..11 .. nnm\•,, c.r. o., c.A'\IPBLU. 1.0.~J., bkows. o.f.
G,4 JI\CQUU:"f. t•., (..OLAS,J ... ca11m,a1, U.. Tifl"t. I', t t.,•, 8,lh. 199.: Mt I nm "· N "Pf,Hl:'11:UC, u,1.. , 19~. /111 ,fr11 dl•\·t.1fopm,,m1 or Tlu•,lurm
Epidemiologie descriptl\.•c de fa 1heiltril):,<?' bo\'inc ,, fheil,m1, a mm tarn 1mn11/11m ma:Jorchnngesin cfforcm lvrnph 1ol10\'1ng 1nr~1£nn \\ilh
t:'n M~urimnlc. AfMquo de l'Que:n ~t:l>-.sahnrlcnnc. Rt!11urd'f:fct·11i<1 ct di! .sf,0101.olU!:i ot .tltoienc:ic s.c:hi1..t)nl•infcc1t•c1 n'lor,onuc:l.tnr cells.
,.\ f'!did,;.t ,lttttri11,llredcs l'tl.)'$ trupft,m..t . ,17, 147- ISS. P11.t11sfu,/o~·- 118. 32f...,Ul.
5S fAGDIS,U, ~.. SJSGJt, r>..Kn ('".__\\ITA'.\I, 0 ft_ 6i OJUFI:• .,,,, 19;9, C:ht.•muproph)"faCII, ": 1 OUll~.1 ,. 11 .. O,\JU..\.~..... "OUAl.;.oASI, H .• 1987 77:l!J!orw nnnulnu,
imnlu.nlS1l1ion ag:unst bovint: tropical ,he,h.,nost~ V~w,inmy R/Jcor(I, Dintrenredu pmerui'-•1 rmnsmurnl 1tt L1bom~um e.t tll1 duodt.mtam t:t
lo.l, 1~0-142 prmwuhilltt• dt· t~r rtHtt(lWU"t 1'.'hn: t~· btn1o info·-~•~
..;6 ,,,..~xu:,,.;, 1,... & \\"L'liMi o.v, 1._ l99f- (..;Jtllt- 1hc1l<.-no.s~ in (.111na.. Trapiml "J>t'nmenialemcm. faµ,,r/111;11111/ Pm·111i10/"lf:t". 63, 2!",J-259 .
Ammo/ Uoo/1/111111/ J>rc>d11rtio11, t9. ~;';-n. 15 n11,,oc \I. 1, ,.,,. \ \'O, 1... 1981 l'rf..111\ wflh ;1 ccll ruhun: \.'11ct·lnc :tl,t4.1Uhl
s; 10:i.'Gf.r,.,·. , ... MQH4.\JUA, s•• MU<tr.u-,.o.r.u.,. 1.Anr, .,.. 198:,. fnftct,on thcJlt:rlo~i'I In r urkt!, Iii: IM\ 1s, A., tU"'-"''<.,H.\'1, !o.U• ~ voo:.:v. A.. !'> 1
T.:llf-> of 11:Nl,:r;o urruulau, in 1hc~.h1,'Qr".1,r l,tln.nd\ or the rick f l_r()Jommn t,"t!,) .,1,t,.111,·.., 1111l1<<:t>111tQ/ufrlt,llerio,h n,e HJjlue: M~iunu,
r,u1rgu:rmmr mfipi."S, htu..'r11rory Pnm~uoln~·. L'J, 121-126. Xijhoff PublNlwN.
S8 J;.\Clf.\,'\t, M , M.AClf, f.,f, \\1Ul.\:'.TSQS, .S,, OUHTlLI, U .• tl II \~X\OUI. ~:. ~ i6 r. 1!4;1. tmrnunl1a1lun ofcal\'~" whh t,m.•ni.1:t1tl(I wild <;.ttuins o·t
Jtlr.\NU,
woo:-:f.a, Jil,t , 1996. Tht'. u"i{· 01 ~n en•·} mc·1lnk.,•d ln1munoso1{)(."nt ass.a) Hteitcna nnm,lara. In: (.Omp1t"$•rentl<1:i du Mt11licolloqt1t• l:trro1,,-vn de
for 1ropic.1l l.h1:iler1os1, N?5ta.cch in ~tomcco. l·'rt1'1'11frrt• \ 'elrrmut') f'llrtl)f(U/ogL· Renne~. t HU I N:'11t .. 19n. Fr~mtt'.
,\J,.·tl:cmt. 2&. 329-339.
':'7 1•1ttAxc,1. ,. 19;..;. lmmunolQgicnl n'Spect~of nk'llmnm1m,fRt(lin(..~1frm.
S9 EACil\Xf,~L .,."l'OOS-CR, R. R:.\f P, 8ft ·~Ur,Yl, 1 ~1JR0\\ ""1 0 .. 1992. l:Jullr11n ,:.,, J'()f!ic4' lm,•rrmtlo,11,J dr,f fl,1J:00Iit•1, 81 , lJ~.. t SCJ.
Sttsg,~-!1,ptafic n:.,pt.J1UC5
.. foUo~nng m(i.icuon wllh 'f11,•1/Ntn tumlf;'fllu u~
:a ..w""o. •··· l97:' B;t)h:' principle&- or Tit<il,•rt,, 11111111/ma comrol. In' "' :\.·
C\11Jua,rd uqng ru~!\. Pora.i1rr;/01:,• R?:wt1rrlt. 78, 13-47. ft;c;h.,J. '111,·lftno$.I,\ Ou"wn. Canada.;
"4,')X. J.B. 4 r.\llt>BtU. 'J
60 J,r.UA~S\\, o. ,1 ._,1A1mt.~ ,1.u., mt.AR, ~- ~ c,u, ,.,1, o.l'.. 1g80. Pn1ho~i:m..,i,- tmcmo1uun;aJ Oc:n•lupment R~·~c-~rch C.i'n.L•r. pp. 55,..6:;,
o! tesjons in buvme cerl!b:al ,hetlconjh. /w {o,u-r,M, n.r., "tltAt4'.\M, n.r,.-,,
i9 NPh~x,. 1 ... ~980. irnroum.uulnn ugaJnst inft.aceUuJo!r blood protozoa of
t>H.\Jc, s eds l{r,~nnp101,no,"ln Di..G.'fjnmoft)Ouu,')Ut:.1lnlnmf.,
caulc. Tn:. \Hl.k\111~ -, .• m fITMA,. ,., ,,;uNm ttr.... '-:..,. & ,ous ... \., Ced$ .\'('u·
O~arxmcnt o!\'ctcrinal')~\1~dicin", Harya.n-1 ..\g,icul;uraJ uru,·,.:nity.
04'h'lopmcms with lf1mf<m mu/\ ·,'1i"rin1Jtj· Vnamt"S. :Xe,, York: ,\b.n, R
lii•~r IUO(M. lndin.
Us.-s-. Inc
61 ,au;Jl,.~ft. K,\... CJL\HfM. ,,.a...., ~~:..,,,.M,·. ,. . 1992.. Comp313u, r ttlk~~· of Ht> l'lr\\O, 1 .. 1981l Bo\1n.t!' 1hcilcri1.1\l, m hntd. Rr1,1,l• ~wntiflq,rrtl
scirm1 ucaritidC'§ .t)rJ.Jn~t Hynlmnma ,pp. tick., in ,,11ro.J1mma/ of
Tttlmf11ur.•dc- I 'OlJir,· lmt•r11ationnl tlN Evi:tH,til"$.. 8. 711--87.
l',r.·r:11or)' P(1m;11oloi:r tlndia 6. 7-10.
6Z L\tBt.J~ c... 1~;8. Cllnicnl lnq~ng;uion~mc;ntlr expcnmeJm:Ul\·infeccc-d 81 PtP\!\O. t., 1ss9. \ actan,uion against nu...llt'fla (lllfmlaro 1.hcilcriosb. /n-
wJttr.1tT. t .(1. rudJ ', 'i•1ttitUU)1 Pt'f)tu::ollft tmd J(.-111opum:.lt1.• V(J1Yim•i.
'-"lth Thrllt•ria tmuulmo. I. H~cl'lUtOlogir.tl m,·t.~lig1;10on,. &·rlltr.:t und
.\ffm~lt~nrr Twtar:1.litfle \\·Oflt~ns-c•hrift 91. 1g..su.
Boe-., R:uon. florfda: r:Rc Pn:".,.., lnc
8.:t. NJt,\...'-'O. L 1-qq.: Pro~pi.'<11:f ror \ Jc<'ln:;1Jon .iirMn~t Tiu:iC.:tM nmmlour
63 1...,nms. c... J:\1\\'~V. :\:.f.·,1uLr.ut. ~1~19-:&.0inir.:il mve!iligauort.incaufr
c.penmeni.alli· inrcc:tcd "i1h 11,~/Nia 111:11utnw. l t:limco-chcmlr41 th~ltt•tlo,-!\.. /n: ..,,oo,fM, u '"t.A\WfULL , .. ft-d~ · F.uropt."".1n Umo11 Third
1n,·esugauons. /krlfncr und .\111r.r/JMtr Ttm,r:tUrlu• WIKl:,·r.,rl,n/1. 91 Coordtn:11,on \leel!ng on tropical 11:r//(.""iOJu. Antal)':!, rurkey. Roslln
25-27. ln<1i111te. l:dmburf:h UK.

64 LIJ. ,,.~ ~ ns, H .. 199~. tinpon'1Jlee 4nd cpld<'"miolo~ of thcilcr1~1~ In H.s PU"A''O· • i~u-\';icdn~, tlJ:ain~r hcmop:irui;lc dhcasc .. In hr:id ,•.1th
Chlna /JtOCit'll.ittR,S ofrh~ 11urd f.urop~·cm (1111011 <A<Jrtfl,uuum ·~t&-.rmg "P'-1!t1l! J1t-l,•rt."nCl" to qu,t,Ht)' runtrul. {IIJtJiUil .'"in,mal /Jtt,tltltmid
Oil Tropt'c,,1 Tlic/1,•rlo#.1., ·9 0Ctobor.A111alrn. furl<<'\. Pmd<1r1torl, 29, ffb,,_90,,_

6s ,r \>:1c14,M, "'· mc:,tt, '!I., ~1NGt1, "-· r, & J.Ji.,ROr ~. , 1.u •• 198.1, l·Us1opa1h0Jorr F11torttsccm ,u~tlbody ft~r for the
H 1 Pl~\~O. L "'t,;.\11.\s.,, ~, •• '!.'~·
of t-ut.a.ntou> lesmi\$ an ·nw,len,, ,mnuJma infect um of cnh'e.. hldicm $Crodl.igno~15. of l'hC'ilfrt4 nnm,lma. /rmrual ofP<muttology. ;;s, 7\i!,
f/11uri11ary•/011mnl, 61, 13-1:».
R5 1'1P.\M,). I· • t~ttt\t'I '\ , 19';'1. Ab.."""~nci.'" c,f cry;hmq1, (Otm"I t;( r/t,-til•nr,
6ti ,t.\ZUP>t~ 1... t~Gg. Tr.:mqn~,;..,10n of n,e1/,...,;n f1nmtlntn by the ~rushed nmwftllu m cah ,.... moculatt:d \\11h s.chaonti trom a: ,indent hetd -;1r.1ln
in(ec1ed unfed Hy<,tommfldromednrii. Pmasirofr,JO', 59. 59i-600. grown ITT ttS"loue cuhurc. l<nmwl o) Prt,10:.~;Jo~·. 18, Svppt 37
o, MfJIUIOl<>C. II.• S<:lllil~, U., 1984 The piroplo,,n-., hre C)ciO artd '"'-''•l 06 Nft'A,o, 1 ftitoPH 11, u. 6'('0111 N, 1t., l!t';'l, lno~uJa1iort uf caull~w1th
:Stage$, ln:i,\U::n, J,R,&:O.IULLl"R, R•• •,ed,,')i,,4drlf.WCCS iu l'nraslmJ~-. Vol hovlnC' lymphoid cell lin(?i Jn(t.l'ctt...d \:\'Jlh 11wl1':r:o nnuulo(a. htH"(trch in
23. 1.ondon: ,lc4domlc Pt'-'>$. \l,•11rimU)• Sti,·11t1... lS. ~l88<~).
496 >T<110~ mo: Pro11>1.oal di~cases

87 PJP-\NO. f:,. KRlt.H, Y•• tJ:JBO\'lY'/. B,. '-HKAJ'. V., 1-'M "-I;, \\. t., H~ft. l.., l~i- u:,5 ~,,u'!<H, \1 .• >-RtlUt. 'r., 1;:a.,~t.. ~.• n1,. c "'rJJtAN0,1:..• 198.s.. 'fnto
Thrlftfi(t mmultrtn; rt:lnfQrt·t"mi.•nt ttf lmtnunicy h)· l\VCJ•/\lep rr.m"lmfi....,fonAl J1wlh,1ritin,1nulma to canlt by llj(llom11u, t•.rem1mum
immunb...i.don, Proc.·1tdi11.gs cf rht J•irst lutdl1n1/on{l/ \~ttrl1111ry lnt•e1NI , ...!h n:ul/erlfl Ctllti,attd ,,, j,ftr(/ R.-[1111'1 Vemiunrlt/1, 41 ,
l'nnirl<'> n,ttl l)lfl..l{nO.<lirs CD1,fer611ce, July 27·3!, ~11,dl;on, \\'i<eon,ln. lt2~.
USA. 106 "-\SOHO. G,!\ .Gnt.W>'1...\,S... !,J~OH •.\... ).:()"-OU.. u:•• SJNGH. 1,At m1..,~. n~~...
88 PIP,\'()· I!.,, MMlSU. ~1. J,;. ~'Rt.Gt.:..\'., t'98'2. RclBli\'C iofl"ctivit)' of 'Jhdll!llil 199S. rlaomo:ologjcal ond b1ochcm{t'31 <tudlL"' un "'1"'<imen1al
n,11111/arn ll)schunkOW$ky and lull>. 190-jl s1nbU,11esderl\·ed from n,,,Jlcrm <uzttulmu inft."Cticm ht ctth8bted c:tlVt.~ Veteruutf'.\' R~nrd,
remJ!t.' arid mall• UyuJommr,,•xrotv,tum t Koch, Jltt rldi;\, , "t1t•rlm1tj• (;()lwnuni<mlon,. .22, J4i'-35,.;.
/'Jlr,i,/zoloJO, 10, 21-27, 107 sc:nns. t- & vn1t':T. W.P.. 1979. Chemothcmpy of bovine 1hciltriosis \\ith
89 F'IP"o. •·" ' II"-'"·,. 1990, Obsomnlons on 1hc lt1 l'/tro multiplh;inlon halofuf!luonc. ,c111 Tmp/a,, 36. 391-394
nf btninc l>1mphofd ce1l, 1n(eeo.•d with 11,.•ih•tU'I m,,wtrua 1-chiJ.unl.\, 108 ~lA:Gt..,-r. I!- 1963. totem infection and..pr~Jnunilion. Some dt?iinirfons 01
Rc.•J.IW! d'Eltrwgq ur de ,\/rdlrinr Vt1crin1tirt tlt1s Pny, lroplco.u~, 43,
~85-188.
microbiology and immunology. /11; G,1t,"•"·
1•.,_c., !'1u1c1 1,£; & nom.
L, ed\l. I Immunity to l>tbrn:011. Oxford. Bl3ck'\1.•ell Sc1ltnlifir
90 VJP,\:'\'('), ~£ .s,o.·:Al1, \ '.. 199-5. ~lmran V~trrln:uyt11q_i1uu:, Bet O;i;gan, l'ublicatlon<,
l~racl. Unpublished ob!M?r<.!tlons. U)9. :r,.t.K(m~'T. L, l>0~.\1ll "· i\.., r~,KROf. L S.tL~'()QUAR.0. f .. l9'35.1helk~riQ~

g1 r1PA.'"\O. e.. ~ rsun. 1.. l~. E.-q,tnmtntal in\munb'~11ion .ig:sin\t Thvlkr/11 bovtrw.s di! t Afriqu1! du ~ord ..:l du Proe:he Orieru.1lrchhY.t.1-dt' J'lnsri:m
am1ula10 wirh .t 1&.-w~ cullufc \·arcinc.1..ahor.uor,.· trJal" R11f11nh /Jllir-t'Utd'A{gtrit, 13• .;72.-188,
~-,,i,rlnar/th. 23, Ulli-la-1. 110 .Sl;tJ(it,~,. a:•• l.>('IS.A1 IITN, A.. ltMCIIO r. 1- .& t£S'£OQU,\R0, 1 .. 1945. Tb:ri1cno.se ;a

92 ,1rl\.'=O, e.. wv~SMAN,,. • B£.~.\00. i\., 19;4. "rhe vtrule-nce.o! iou.r to<"al l'iwll,rln d/f/J(/r.111: /!J.udessur /~JPirop:1>,<m<r1os />0vincs. lruri,u1
-\tmlnsor7'hl'ifl!rlnnmwfma. R,tfualt l11•li!n'n1,rlth, 31. -59-6:..l. P..t.."ttur d'Afgc.ril\ Alger. 243-SGQ.
1.11 Sf...ftGl:.':,,,'T, l,, PO~i\Tif,."i', ,\.. PMtRQT, 1.. .o,. USTOQlJAJltl, F,. J~,$. TC'Chniqut"S
93 PlttS?ON, P.lt.. *' nRQw;\', c,cu, .. 198:;.
lnhlbltfon orl\'mph<.1<1."t<-in\'u1tfon
b;· ,pc,rorhl<c~ ond die •r•n~formalfon ,if ,ro;,ho,;~,c lnf,,;,,od In: Fwd~,$ 5ur IM l'iwpla~moSIOt l,o:,mes. lnsdnu Pasfeur d .Alser~t. Alger_
tymphoq.11:..'= in virm by wrum from n1e/JMi11 mmulmu Immune calt1v. iD--118.
l',,rnsite /11111111110/ogy. 7 aol-314. u2. :-.JtAK~t.A, n.n., 198o. Same epidemJologh:·al obs~JV7\tions on 1.ropkal

94 PKL>TO~, 1•,,. •11011% c.G,o.. 1988. Mncrophugc-mcdlatcd C}i0$1usls and thciltrfo;.--b in lndf:i. /11: to.\UT.\M, IJ..... ,.tt'\'AM ,. ic.u. s rutAJi h .. !c-d,i.
lymphocyte q~otoxldty In cilttl< lmmuni,cd with 'n,vi/,rin,m11uln1n Huvmop,nro:otw f>1S1,r,t1t•>·1,f Dumt·$t:i: A.mmal~ Ptau.-cdin~ of !\eminar
sporozoues ormacroM:hlt.Qnt-lnk..-1ed ce.ll llnes.. Part1J1/1~ ,,,111111110/ogi•. on Hacmoproc()1,mm J)lsta.se:- Oep.i.r1111tn1 of\'eterinary Mtd1cine-.
10. 631--647.
lfaryon• Agncuhural UnivNsn)· Hh<ru 1ioo04. !Hol')'lln:i India

95 ntf.Sf(lS# v.,,.. 1.Ut()\\""· C..(;,f),. JlfU•"'-1'):VI, L. tunl.\l\Dk)'\. \\ ,.

1\3 !'>H \M.N R.T.... u:"l.'(ol "''· \ ,M. 1983, l11e dT~ of '111t?J/c~/(1(UIIJlllarn
1:ifei;1ion oo lhe imrn..wh! re$ponit o! mote to fooMmd-mouth disease.
~--o:otstsox, A.. 1992-, f rop!ca! lhellerio~i~ ln lk>$ llWr:tS and SuJ 111urus
Htiriiil httNmal)· /011mal, l:i9. 37"8-38S1
cro~i&s l,u//rztSca:Jve?i! respunSi! to infection wi1h grudf'd d~t?S-of
.lip(>rt>1.(11lt!!'i ur n,i:lf,•rio a11m,tmt1, Res,,urdr In \f(>fL.,-fn«Q' ~dt',,.,.:. 53. t.S$8
114 !tHA'i\S-, P, IUEIGV.Ni\, R.. :,OIJlt~. t .. Ui;1'DFS. J, fc.-\1fMH), I ,
?30-243. De-11!'trion and dlfk~emforton o! T11,d1 ria cm,wln.111 .smJ T. pan~, wm~
1

m.icroxhltom·dt:ti\'c:d O~.\ probe,. ,.\tmn/,,- ofrllt Nt•tt• Yali: ..Wdlm_\·of

9G PMl:~lt)I\, 1•,'.\1., KftOW~. (.'.t',.n .• J.~11Ur-A -.:, (... SUCHAftlh.O,. \\', tir IWIO, ft,,
)<'/fn(\•s.1149, se-g5.
1993. Symbesh or Ulntnr nuerosis ful.tll>r•ulpha nlld lnierfcn:uh b)
mononudcar ctlfs from n:,·Hma n111111latJ1-!11!cc.ed cank. P(llr,$/i, 11.i "'1Dt.0:mrne•., .. uoms, t... r1P".\..\'O. e. 4.: ru""Gtlff, 1.• 1973. Surum \.'lll':)mc
lmm1mo/1,gi·, 15, 525-534. k-.-cl, in cxpctlme111al thcflerlosi, In"'''"'"- R.ifunl, H'/trl11artz/1, 30,
24-2:-
97" l'f\f$n):-.-. P,M .. UROW:,O, c.c::.o. 11,,M"(K)'\tR, "·'- 198~.. (:elJ ..mcdint("<i
C}10<oxicity in TlrnlrJ/n nnn11/atn infcclionwi1h c11denc~ for SoL\ 116 ~l~(.H, I>.~ .. J•tU""· ~.. ,.AU"tA\\, 1),1', & Ol{AR, ~ .. 1979. ln(eclMty Ill
r~ttfctiun. ClinfMI ,mt! E.1:pnmJe11wl lmm,1110/ogJ·. 53,. 88-100. groun~-up UCk$upema1e, p.~p:ired Crom /1rafl,rln 111111ulara lnil!Crt<l
sa 11 Rl:....<iTO:-.-. 11,M .. HAIL t.t,t., ()~'), b,. C'.AMJilU!.U.. r.o. , t .. 0<\ftGH()Ul11, ,1.A..
flj'illommtt ouarolftum tul(ttollrum.. Trrrpiatl 1\mm,1/ Hn1lrJa muJ
Pta//u<Zfon, 11,87-90,
wooxtm. R.L.. fON'GQAN, F.-"' naow·s. c.q;,o.•
;\HMED, r.~... SJla:t~. D.R.,
l999. fnn.ite-nnd udup1.11.\: ,mmuru." rt."11pun~es co•opcr:tw- to _p ro;('(l 11; "'of.'-(,lf, U ...., 1 IHklJlt \I,, Q,\C,Jt.\\.' '-'-it~."' VA~11!'.t'l'• R.C.0 199),

caltle ag.a.in~1 11ieilcrin nmwl<uo. Pur,n;w!,>gJ' T0<iay, I5. 268--274. Chomer1h<rnpcu1ie trial> wilh rourd.,u~, in cr05'lbrt•d c,IIVl/s
oXprnmomall, ln(rcted whh Tlr,•1/er/<111111111/mn Remm:I, f,: \ <'lerinur;·
99 RAJ.l'I, ·"· MA(>UA.,11, c.. • HOO:;.tL,1A~lrRAD, P•• 1905. Sur !a ,irute-nce de
':-i(/t•nt\' 54. 68-il.
Tlrcilerinammlnm (Psch1.rnkow~kyt'1 l.uhs, 190·0 ct ta p~munfnoo
cooue la 1helletio,c bo,ine e11 Iran /J111/cli11 ,It l'OJ]ic,: ,,,,..,,,011onal 1fe$ 118 ,,11n1.11-1> .. •'lfls.&1b<'$/o w11i" tompmoriunlllntiQn ortho rti~<ionshlp
Eplwofit$, 64, ot31-&4G. b~rwton Ute tick ·,1.-ctor, pnmsh• ond ~int• huM, F.x11mm•ntt1/
Pnras:t1of11gy. 56, 27---10.
100 MCflAR(?_:,()X, 1.. f()R...~l'TU. L.. 8ROW~. D. f< PRFSTO~. P,, f9'93.
~itric oxfde
c--au~s 1he macsos.chiconb of 11:~,l1Yia mw,1.lara 10 dl,:;~1);)t;1r :md ho'i', 119 ~VQO'.\.'HR, M,, •~x• . ;-, ··- GI.AS',, I· "'"HM(l\V(, u .• 19$9. Tl11tifL•rltHm1111!11ra o.uul

crUi 10 bl"COmc 4tPOPlOllt\ Vrurmmy Reseqr<,'11 0)1mmmictlliu,,, '.!2, Tl:~il,n// J)(l<1.i lnfoc1 ~nd 1rnn,rorm dliftr<nt bovine mononucleur
31-45, coll., lmm:mology. f>l\. 28-1-288.
101 80RI.S'4i<>~. P,M,1 1962. 'f1rctlnia amrnl.tlla :md It.~ minsmiss!on: r<•\iC'\v, 120 uIULwtrtC.. o .. 1981. fhcllcri:11 spuci<-soi dom~tle lh'l'stOck...hr. ut\·1,-, >.~D ..
n,._;\'.ISOU+\.'-1. ~~.-P- ~ \'0V1'<'f. A,S., IC~). ,•\tlt'ttlf('(rJ in llle-Comrot of
'/rot//C(I/ ,!11imn./ I lmlz/1 n111t (>r0tl11c1/011. M. $.-12.
Tl:r!il,•rlt>Ji<. Thr liagu<!: :.!arum,, \:ijhnlT 11ubli,hcr,,, pp, ,1•37.
101 ~,m)Q, P.Ji, b M~M. ').(';+, t!f9l. ~:ltural nlfC>Clion nn~ of nu·lltrlr:
tmnulmu in J/y11lrmmm ana:uhtwn cmnu,/i('mn m 8hubmu,:<wur. /n,111111 t:tl \';\,\ OF, i:~nf<. ,,. A F.nl.1!\'fU it, E•• 19r1- c:uhurn d, lig,11£"~s l~1nphoq,-aires
l'cr,,nnmy Journal. i2. 68r-689. bo'-'111~ ittftttt.'-es 1>:ir T11t1Jl,•rto Dmwlt>Ul,,\rchit<t'.'5 d1.1 l'l11-nlwr-Paucurdi!
Tu,11$. .JS-5,1.
103 S..."'\M.A:0,.7AMY. s.~.. fJlL\TI,,OL\Jnvw. v.&t1u.1.• e..s.. 1980. tmmuniu1ion of
12:t ,·tSS.1!11•.,. AftRAIIA\1. A.. ALIL•"''K\·t. L, Uk.<)\\'~·. 0, ~ J\1tt-'Std-X. I••• 1995.
calves again:-t bc)vu'>c 1rop1c.i1 tiu:llttn(~lS cTlwil11rl(I mmufnrt1 \,ith
Ni1ric n>..idc· inhihils e~mbfo;.hm~nt of rnattosehiJ.ont intccn~ cclb lines.
4

g;rodec.1 d0$esor sporoioh~ and, frn,dio1ed spow,.oncs. lmi."rttatwna.l

/011n1t1I/or P11rns/lology. 10. 3S5-3S8. '111d Is produced b)' moctophaiw~o! cal,-e,; ur.dcti,oln!! bovine tropical
thc.itcriostsar F.U!-t <:oo.'ft fo\'er. Ptumlre lmmwwlogy. 17. 91-102.
m.; ,M,11~0. M.. 19n. tnf~o,-.: 1·n.,,t,:tf(lm11ml.o.1<1 m 1he tkk wahout .a blood
123 w.u):t::H, ., .. 1..,1 If••, •• ),.t<JR1..AH.U. s-. g; JO.\G.LJA''· 1: .. 1~3. x:uunal infL-ctton
meal st.Jmulus. Naum.•, 270, 51-52..
r.ucs:of H.w,Imnnm rmmolit:um tmo.wlicum \\ ith '11u:ilt1rl,1 mSuda.n..
Rt-j.Mfl"h b1 Vt•tt>rlluuy S1.•fe1r0!. 35. 87-90.
 T/11•ilerir; mm11/nu, 1lwi!eriosis 49,

12-1 wv,rn.,. C.).l., 1926. rn,11,rlJJ lo c:,mlc. /n: l't<,tb:.Wl"l,,'V, Vol. II Lon,!on: or
126 1.Abu.n~1m. \' r .. 19133. Dw.1lio11 posl\·J.cetn.il lmmunlt} in bovinl'
Bailllr.r.• Tlndnll 11nd C:nx. dWUt,rla:<1!-.... Ttttdf \'°.4•.-l'O)'U.WOH" /1u1w11a ~Jli'rlmtnrnf,10/ \ft·Nrmnril.
~7. ~!.-4il
U,5 WILXU C.:o., .• ISRU\~. (:,(.,,0., lo:IR\'.\a, t , 1HO\I/\>, \\,, \\11.U.\'1\0~. °' \I,,
D'7 .ti\, ,1 , l!\79llle r,~cp1lbilhy 10 Hnd.inr. of field ..1roim, of Hy,,fomm<t
aDJ... ~Ak1\1. t.J. A :,.,Alt.\GAS01 u .. 19Y8, Chtmo1)rophylaxh of
r,,-,i/41ria n11nulmn 1mc1 7lltilt•rU> 11(lr1·n in!cction~of c1.th,\''I. \\1th ueks llx<x!oi.tk·3. l~>tlldnc: in h:r.u.1J. Rt'jut1h \'ettrmantlt1 30, I M-1 It-.
hup•t\-aquon~. Ve11•r11inry P~r1JS:tolu;0·. 78. t-1:?. uu lmmuniution 3R/lin~, nwill'rin
l\U\, (1\:ltt,lt"'l:O:O.:oJU-"ft:J. 1•• 19,lfl
nmu,fM,r. ·Hi:br\'Tw-f"..nll,h\h :,.ummary), Hrfunlr r,•torlrmritll, 5, 69.
 38
Theileriosis of sheep and goats
Synonym: ;'\'1alignam Lheileriosis of sheep and goais

J A LAWRENCE

Introduction by phase-comras1 microscopy. 11 The parasite is transmiued

by Rl1ipicepl111/t1s e,,ertsi 1ll'errsi and R. e. mimeiicus.11 Thl'
The taxonomy of r.hc speeics of 111eileria 1.ha1 infect small plroplasms are readily transmirted by blood inocula1ion.~
ruminants is very imperfectly undersmod. T/1ei111ria le.stQ· fnfecth~ty for goals i:, very low.
quardi (syn. T. hire,) is generally accepted as being 1he only The T. 011is complex infects both she.ep and goars. and its
species of economic significance. 1l1e other species curre1111~, vcciors have not been identified \,ith cenaimy. Rhiµiceph·
recogni7.ed are T. 01,is (syn. r. rcco11dita) and T. se11<1ra1a.8 Ir is a/us bursa is known to transmit one non-pathogenic o, ine
probable 1.ha11he parasite generally idenr.ified as T. ovis may Tlreileria. and,\mblynmmn spp. are suspected EO be 1he vec-
be a complex of several species. judging from the number of wrs of another. 7 while Haemnphysalis puncrara has been
genera of licks tha I has been implicated as vectors.7 shown to transmit a 1hird.8

Aetiology and life cycle Epide,niology

Theilerin /esro(Juardi is 1he causal organism of malignant Malignant theiferiosis or sheep and goais occurs in the
theileriosis of small ruminants and has a life cycle and mor- Medi1crrani:!an basin. Sudan. wes1em and central Asia and
phology typical of the genus Theilerio. It is transmincd by India. and has also been reported from China, although 1he
1-lyalomma m1atolic11m m1arolicum,~ and po:.sibly by olher idemitv of Lhe parasite im·olved there has not been con-
species of Hyalomma. 1 Transmission from nymph co adulr. firmed, as a species of Haemaphysalis is thought to be the
has been dernons1rated. The parasite in fens both sheep and Yenor. The disease is "1despread in some important sheep-
goats. An1igenically it is closely related 10 T. a111111/ma of breeding areas and causes heavy losse~ in both il1digenous
canle. 6 and exotic breeds and their crosses. There is disagreement
n1eileria lestoquardi schizoncs can be grown readily in in 1he literature as 10 whether lambs are more or less resis·
suspension cullure in lymphoid cells. in a manner similar 10 1am than adults, and 10 what extem indigenous breeds are
those of T. pnrtlG and 1: amwlatr, (see Chapter 29: Eas1 resistnnL The dcvelopmem of endemic stability with re·
Coast Fever and Chapter 37: Tlteileria amwlcl ta theilerio- spec1 co theitcriosis may be hampered by the praccice of
sis).4 Cultured schizoncs are of value in immunizalion 5 and 1ranshumance in ~heep-raising areas, and reponed differ·
as amigcn ror indlrec1 immunofluorescc:nce rcsting.b ences in susceptlbility of various classes or stock probably
Theileriaseparam, origi nally described as Haemaroxe1111s reflec1 differences in the epidemiological stams in differem
sepnrarus.9 can be distinguished morphologically from the situations. Goal;. arc seldom affec1ed cliuically. The disease
01her species of r1tei/11ria in small ruminanLS b~· the associa- usuall> occurs over a period of two month& at the end of
tion with the piroptasms or a clearly defined veil. often si1u- spring. when ticks areacrive. 1
atcd outside the erythrocyte membrane bu1 l}ing close to it. Little is known oflhe epidemiology of the benign species
..opposite a corresponding gap in 1he erythrocyte cytoplasm of TIU!ileria. They are \,1clespread throughout Africa. Asia
(see Chapter 35: Theileria ue/ifera infection: Figure 35.J ). and Europe.
There ma}' also be a small granule. possibly a reduced bar
s1ruc1ure of parasitic origin. within the erythrocyte cyto-
Clinical signs
plasm. Veils can be seen as.~odatcd wilh a proportion of
plroplasms in Giemsa-stained smears bm may not be obvl· Malignant theileri<.isis of sheep ai1.d goa1s may be acute,
ous. TI1ey are more easily demons1ra1ed in we1 preparations subacur.c or chrc>nic. In the acme form fever, anaemia and
498
 Thcileliosis of sheep and goms 499

swelling of supt·rficial lymph nodes are prominent rcatures. chronic cases. anaemia. emaciation and generalized oedema
Death follows withjn rour or five days, and the monality rate may be prominent.
may read1 100 per ceru. Recovery. if it occurs. is ~low. Ocher Enlargement of lymph nodes and mild anaemia arc the
signs descnbed arc listlessness. malaise. nasal discharge. onl)' lesions to be expected in benign theileriosis.
hyperaemia of the conjunctivae, 1achycardia, dy~pnoea.
oedema of the throat. iccerus and transitory haemoglobin-
Diagnosis and differential diagnosis
uria. Climcal signs Jn subacute and chronic cases are similar
but less marked. and anaemia and emaciation ma) be 111e diagnosb of malignant thelleriosis of sheep and goatS is
pronounced. based on the chnr(lcterlstic clinical signs and lesions and ma)
The be111gn T/ieileria spp. cause only a mild febrile reac- bt.' confirmed by the demonsmnion of schizoms in lymph
tion. moderate !>Welling of the superficial l)'lnph nodes and node and liver smears: piropla~rns may be present In blood
possibly mild anaemia. smears. but may be infrequent or even absent in the acute
disease.' Specific diagnosis can be made recrospectively hr
means of the indirect immunofluorcscent antibodi•1es1.u
Pathogenesis and pathology
Benign theileriosis may be suspected when piroplasms
The parhogenes.is and pathology of mallgnanr thetleriosis are alone are found in sheep or goars in the absence of severe
similar to those of the pathogenic Theileria spp. in cattle (see disease. but it is necessa ry to exclude the presence of
Chapter 29: East Coast Fever and Chapter 37: Thei/eria 1: les1oquardi In a late111 or subcliulcal form by serological
ammlam theileriosis). Both schizonrs and piroplasms play examination. The characteristic features of the piroptasms
a role, the former by inducing lymphoid h)1ierplasla and de- of T. se1,am1a permit differentiation from Other Tllt>ileria
generation. the lauer by causing haemolysis, anaem fa and spp. in sheep.
icterus. Salient features at posc-monem examination fn acu te
cases include subcutaneous and inrranmscular oedema.
Control
especially in the ventral pans of the head. neck and abdomen;
enlargement of the spleen and lymph nodes: enlargemem. Bupar\'aquonc may be used in the treatment of malignant
pallor and subcapsular haemorrhage of the liver; pulmonary theileriosis. 1. 2 An anenuated schizont vaccine prepared in
oedema: red or white nodules oflymphoid hyperplasia in Lhe lymphoid cell culture has been de\·eloped and used success-
kidney conex (so-called infarcts} and perircnal oedema: fully in Jran.5 Tick control In small ruminants is generally
haemorrhage.~ on :;erosal surfaces and beneath the epicar- impractical as a measure to comrol the disease.
dlum and endocardium; haemorrhagic lesions and necrosis There are no indica tions for the control of the benign
of the mucosa of the abomasum and large intestine. In species of 711eileria.

References
•M<>u11<1», , ., .. 1997. riek·born~dltta~ of$h<:ep 3nd goalScau.,cJ t,y - u1.LE.~8ERG.. c; .. 1~1. Tiu?dcdal ~P'-'cit.~oi dom~nc iiv~!iotOCk. /rr. lk\ 1s. A-»..
&tbc'!S/11. 11:rlltria or J\m111lasma spp. Pan1J.#U1lugt,,, 39. 99-109 cu:,;:,;I\GH ,,1. M 1• . ",·CJu"·c. /l..<-.. {eds), Adt.'tmed" 1,: ,he r.<>nrrol uf
2 1tAS1tf!\1t•fES1L\1t._1, n..i99;. 'Iid:•bome dist:as.t.'- or sheep and ~<'trus 1.1nd n"•//;rl,,;!1 lh,· ll•1,t11c, 80>1on, 1.oodon: ~tartinu, Nljluiff Publl<hcr,.
thc-lr rcluu:·d \ICC\orit in tran P-tucwi10Jogin. 39, 11!,-l ti. pp•.1-3,

J 1t(IOSt1.l-1AXt'>•R.o\.O, r. I> lli\W,\, s.1•• 19;5. Tr:msmiS-$iOn t>f Theilqrin itl,c:t i.n 8 u1uM•ERG. G.1~9;. Gcnc1al rc·vlewol c!ck·bonlCdJ,c~,e,. vf ,hc.:11 •nd
sheep b> /(mlon,m,1 nrwolicum 111:a,o/ft·um. T1uvlcal.~11/11wl 1/rnlrll goa~worh.1-wlde P<1ra,;,ro/ogf(I. 39 161-16$.
and Pf<Hiu(t/011. 7. 103-109. 9 UIU:\IU:kC. C. "'·"Oft.MSE..~. M.1',, 19;-. Hfl:l.,•t1ut:O:tl,·1wss.:pam111~ '.',.p, n.
J HO()StlM.\..~1)•R.o\O; P, & UA\\ .\, :\,J.. 19'> Culm:,:ulon or11wllrr10 irtrd In !SporD1.o;. Tb~l!erildael. • new blOCJd parasite o! domestkshecp 1n
,hc~p lrmpho,d <'<'tis. Trop;cn/ ."11111,n/ H,11/1/1 mu/ P1ud11rt1m,.: Tam.ant., Rrt.·ut d'F.let·agt t!I tlr .\!tdt,:h,.t \ ·11.trinnir,.d,,s P111,
121-112... Tropf,·ou., :r ~~9--:6~
3 L\WJlli.,,c1. l,A•• 1997. Conv~nuonal vMclt1~ for ~tc:k-twn,.: ,o UIUXKfl&C., '"" .. ~HfU.VOLR. ILJ;,t;.,, 19:76. Funher "'1Ud1L", on
hocmoparasllcdb<,o,~ ofshocp ond go.1t>. Parn.<slto/Qgffl. J9, ,,f
HM11wroum11 s.•11t1ra1u.i tSPQro1.oa. Thellorlld~eJ •hcer• in T4l1Unl,1-
I l!>-121. Rer•:i, 1t'Ele1~, ,,, dr 11,YIN:irw l'lt/Jrmozr~d.., Pa>n roplrau.T, 29.
tlll-1~6.
l; ULM-\S~. I,. ll00$T1MA!~D-RAD. v. 61 UGGU. , •• 199;. Th<' indire,·1
Ouorv1<:1•n1 antibody 11'$1 ba>cd on >Chl;ront ftntlg•n for stud)' u vouNG, A...<.• ,,cn1,~. (;.J .• 19;;. Obs.er\•adon~ on flm•maw.~,ws
of lho <hcup pnru.,llc Tl11•1/1rria lell()(fUhrdl. \11>1Qfi11nry l•aro.1)10/oir,•, 69, $<1f)(lrmu, Udl.'nbcrg li.-Andrca,wn 1~74 in ,he tl')1hrt1<.1·1cs or Kcny-,n
9-18. sheep. Rl't-M«I: m \ l'teritwry Seieutt, 23, 387-388.
 39
Theileria buffeli/orientalis infection

J A LA"VRENCE

Introduction morphological and biological characteristics of Lhe group

are similar to those of Theiterin pamn (see Chapter 29:
The 1'11eiler/a buff'elilorienralis group of parasites is widely East Coas1 fever: Figure 29.2). The macroschizoms are un-
distribmed in tattle around the world. The taxonomy of the usually large. i:. TJ1e piroplasms are often longer than chose
group is unclear; gene sequencing has revealed close simi- of other species. and bars are usuallr found in the erythro-
larities and also differences berween isolates from Asiatic cytei: associaced wilh che piroplasms; veils ,mi also
buffalo (Bubf4lus b11baiisJand cau le in various locations-.4· 5 commonly found 1,1~th cenain isolate:;, hut not wirh
but the Interpretation of these findings has still to be others.6 · ~. 15
agreed. The name I'. bujfeli was given to a parasite recov- The main ve.ctors of the group are Haemapl1ymlis pu11,·-
ered from Asiatic buffalo in Indochina by :'seveu-Lemaire 1ara in Europe, t.he Medltermnean basin artd western Asia.
in 1912. and that of T. oriemalis ro a parasite recovered H. /ongicomis. H. jap(mica and H. conci1111a in eastern
from canle in Siberia b) Yakimoff and Soudatschenkoff in Asia.Hand JJ. lmmerosa and 11. bancrofti in Australia. 12 The
I 93 L8 The: name T/leilerin serg1m1i is frequently used to de- vectors in :"<orth America and $Uh-Saharan Africa have not
scribe a pathogenic member of the T. bujfelilorienw/fs been identified and prel>umebly belong to another genus of
group that occurs in eastern Asia, but this name is invalid ticks. ~o Haemnphysalis ticks occur on cartle in North
as it had beell used previously to describe a parasite of America, while in Africa the only Haemapliysc1/is that o.;:curs
sheep.8 The parasites have been desoribed from Asia. Aus- on caule as both immatures and aduhs Is H. aciculifer.' 3
tralia, the Mediterranean basin. western Europe and North which is 1101 thought to be sufficiently common to act as a
America, although they have often been identified mistak- vector. 2 The parasite is readily transmissible between cattle
enly as Theilerin mutans. Under rhe name T. oriemalis Lhe by blood inoculation.15
parasites were first identified in sub-Saharan Africa in Ethi- The Theiler/a b11/)'elilorie111nlis group is assumed to be
opia in 19832 and they have since been described in centra l \,1despread throughout the range of its tick vectors and
Africa6 and eastern Africa, where chey were called T/reileria often occurs in mixed infections 1vith other species of
sp. (Marula)Y Theilerin sp. [Marnia) has been shown to be Theilerin. It is rarely a..,;sociated with clinical disease. except
indistinguishable b) gene sequencing from stocks of the in cattle of e.xo1ic breeds in eastern Asia.
T. b11jfelilorie111aiis group from Aus1ralia. Britain. Japan
and Russia.5 The parasires have also been identified by
Clinical signs
polymerase chain reaction 1es1ing in African buffalo CSy11-
cems ca.Ifer) in Sourh Africa. 1 l n mos1 parts of the world the Clini<;al illness usually occurs in animals s ubjected to stress.
parasi ces are usually benign, or cause only mild clil1ical such as imercurrent infection \,1th other parasites or
signs. but in eastern Asia they may pose an economically vimses? The main signs described are fever. inappetance.
significant 1hreat to cattle production. anaemia and reduced milk production; enlargement of su-
perficial lymph nodes mar be noted. The condi!lon is occa-
sionally faml.
Aetiology, life cycle and epidemiology
The parasites In the 7'. b11jfelilorie11ralis group normally in-
Pathogenesis an d pathology
fect caule. Asiatic domescic buffalo and African buffalo.
Persistent subclinical infection of sheep after transmission The parhog.enesis and pathology of T. /}l(ffeliloriemalis
by blood Inoculation ha$ been demonstrated. 1s The infection have not been described in detail.
500
 T11(•f/(•ria l.mfleli I oriemnll, infection :SO I

Diagnosis and differential diagnosis pathogenic characteristics, as In eastern ,\sia. treatmem

with the amim3.larials primaquine and pamaquine is effec-
T/leileria buffe/iloriemalis infertion ma} be sL.cs pected when tive when they are adminis1ered inuamuscularl) at 1.0 to
a febrile reaction and anaemia occurs in cattle exposed to 2,5 mg/kg of the phosphate? lmidocarb has also been used
Naemaph.i•sa/is in festaiion, and elongated piroplasms asso- with success ai a dose of I H> 3 mg/kg.3 Buparvoquune and
cimed with imra-e rythr0cytic rods. and possibly \\;th \'eils, haJoii.tginone are li!:ely to be effective. Combined t reatmenr
are demonstrated in blood smears. Large schizoms mav with primaquine phosphate (2 mg/kg/day for three days.
occasionally be demonstrated in l~111ph s mears. possibly repeated afler one week) and halofuginonc lactate
Differentiation ofinfec,io11 with T lmffelilorie11wlis from at I mg 1kg or buparvat1uone at 2,5 mg/kg on the fi~r and
infection with other species of 711eileri" can be made by use third days ls frequently effective in eliminntlng inl'ection
ot the indirect fluorescent amibody test. using a range of from carriercanle. io. 11
Theileria antigens, as there is Huie crO$S-react1on.q Where necessary, infccHon can be prevented byappropri-
aw methods to limit expos1ire oi susceptible caulc ro infected
ticks. II Is common practice in eastern Asia lO stimulate im-
Control
munity to the parasites by deliberately exposing cattle to rick
As r. buffelitorienmlis is usunlly a benign parasite there ar~ infestation and treating them prophylactically.J
no indications for its treatment or control. Where it assumes

References
,\U,:,()J,,,, ,,.r.r~s•.. THUi()~ , .• C:OET1Ut.. ,~.l-, UIJ~',l'~Jt\'IW. M,1,', CJ NC'ar~:1. P.,S., U\...\'\:, .\.f.., \\IUJAM).()S, S. ,~.,.\\\lCH, J.J\,. ,\JOR1.,,\Rl.4., S..P.
\11.",,Ql'S". lt..\., 1999~ ·£he occurrcnct• n:" 111.t•Jlrrln and O,wdrla p:tra..itc.. Jn n()tA.'I;, 't.t ,u," ,,,._ •., vo11s1>. \ 1'1,, 1994, tsolut!on rutd pn-'liminm:

.\fni:an butf3Ju ~S;rnern,.( t(lffer· anc.l tht=it JS.-SO<'hucd Aml,tymnmu charom·rl<ation uf • prl'\1ou,t, unldcnti.1"1! /11rilrtin pora,ltc ,,r,,..~~
ln,llftUHrtr ;lc;"1. Oml1trsu111>0n Jrmmal of t;,,1.•rlnnry ReJCnrch 66, in ~n)'a. R,•-ftrJ«lt In l'l"lri'lnn,,· .~i11mY• .i7. 1-9
;!.;.;..2-4•1. ,o strw\.81. 'l;.P... o• ,,o~."" 1 g,._ruu.s. 1•• 1990 Elimin~tlou or f'1wllcrl.J.1
:J_ bt 0.10(/\, ,. ,r.. LGGL~ J1ur(/(·rfn orJrnra/i,i: In
\ •.A.,S ,. DL JtOtl\. M-( .• 1983,. buffi~U tnfl~t.it,ru. uom cottlt• byconc:urr~m trci.ttm,mt with primaqutn~
cattlL• irt l:.thlopfa. llt."'Si!Off'h m \ 'c.•1c.1n,w.ry Sdt 11n,.34. 362-364
1 phosph•t~ ond h.th>f11¢nnnc tor.101~, Tmpkt1f..l11/11:al lltY1l1l, n11d
3 Rft0\\'!'1.', C.GJ>.. \9YO. ThdlCOOSi'\, Irr. $t'WLU~ M.,\t,U « 8f'OQ;lf~flt, 0.\\,. 1•rod1101/011. 22, !0~115.
od<J, /lt111d/)<)(1k 0111111/mnl Osfl'as,·! 111 ,,,.. Trr,pit.•. 1.011do<1: Baill1ero u snw.uu., 1, .. In \·o;., ,\..J.. MCliAIU)\'. s.a. :;r.,~or.~L s.r..• ,~o.
Tin®U,pp.183-199. Fllmin:niou 01 Thrlltt'ltt b11Jfrli inrel!ttOns. Crom t'.1111~ h)' c:onrurretu
f'JH<,;;\,JiU, ~•• l':\W.\1,1,). :-.. t,.\'.\HO, T. Tl·-RAJ>.\. \,, TUll5AK11 .... r111u1111, )l, f.:
an,~unont "1th bup;u,,~quon~ and prim~quiec pho1pha1r. 7'ropl<nJ
~1Jm1,,J Hl'tlrtlt and Pr,xlm.•tfon. 22. 116-122.
<i.\fl<\t.~rH ,,, , .• 1gg9_ ~1otec;uJar ph~~ogenc1lc nudi~ on ntt~1li•rin
p.i1ra....1w., b.$$1.-d on ~oil subunn r1b<1~1m.a.l k~.\ gl'n~ seqm.'flC'e,j •~ snrw.uT. x.r.• u111~1U.HC: r. . .-01:. \()i<..,\.r .• 1996. Rt.wje\, of;\usu·alinn
\".,,m,u,()· J'aro.oirology.1!3. 99-105, SJ)(l'Cfl'S of n,~Ilttia, wtthspccl~I rLofor..-.nc:c to n,c,ltrla lmffi,u oi cattle:,.
S ~,\\\'.VU.>~ J:.\ \ 110, rj. u,:uu..,. T 'fl R.\O\ ....... ,m!IMUl'U, C. (i.· fUJl}>Jrr.l. "" Tropical ."111imol Jfrolrh uml Ptodu~:ivn ::08. 81-SO.
1999. Phylogenetic rdodon,h1p, ol lhe bcnll,Jl Tl,c/wrl11 ,p..:ic, ,n <,tit!~ 13 THFIU R. c.... i96J. n:.· ,~"doid~11 Pllm#tt,s,of \ ·,m.~brmvs in ,\fri,,,Srmtl; Qf
:ind A~nn huffalo b~d on the m:iior p1ropf3qn ... urfo.ce prott'ms rlu, Stil111r1< ·1:1'1/opio11 fl<Ri<mJ. Proje<:15.9938. Repon 10 ah~ Director or
p331p..ll) Kenc sequeotc..':). lml"rmmon((/ /oum"I of/>amsiro!fll!J,·, 29. Vl1 t~rim'U"\' $('n'1(.'C5,, OndCt'SIC!llOOn.
613-618.
1.: u1usat.RG. '"· ,,'8,. Thrile:na.1 species of dom?"iottc Ji\.'esmck. In; mv-1~ . .\.V••
6 ii:IL17-, K.IL, Ul&.r..U.t.tta, ..... ,.R,,~sr,. f 1',f.-., rheilt:l'Jn
f'.CRlf.. ~.M, 198ft. cuss1.,,0,1tA\1, \i.P.,;. l'OU~G. 1',S, lcds· "\11:,'(lttm in lh<' Commt uf
tJrlitm,>lo ocrur~in Centtal:\fnca. R~1rth in \'t!t,•rlr:a,:.·Sdtt11u. ~u. /'lwiltriods: fltoc:t"td£11g,~ ufon Jmt•r,,orfo,uil c,o,1f~tvm.1· ht·ltl ,,1th,·
J~j..20(t,
lmcrnminnnl Lal10rt1tdry•for ltt.fc."tlrc/1 on .~nin:11/ Oi$~tuc-s m sa;mbt.
:- '-!.\TS"U8.\., T .. X.-\W.~~H. \',. I\\ \t, u. .1. O~U~SA, M .. 1992~ G~numtt .tnnl}si~ 9-131/1 frlsrut1')', 1981. Tho lfaguee ~lmtnu, :,;;Jholf PubJl,hct<.
nl T/:rllrrln A'rgmll ,mck< in Jnpon with DX·\ fir<>b<>, l·.i,•rit1fl')' pp. -1-3-.
f>t1,tff.ll()fo8)'. '11, 35-13. JS UIU!~8ERC. c;•• J>$Rlf, !'1,,)1,. Sl·A~JEJt. A.>..:)1.,. FR\sssrx. r ..... J., 198.5.
8 ,10ttlt .. ,,.r_" lULGSBER<,, o.• 1981. Sur la nom.t.1ncfature do qutJqu<!S tl1«1it1.•ria or!emolts. ~ co:c:mor,uht~\11 blood pa.rass,e or c4u1e:
Tlll'ih'rfu (Sporo1.()11. Bnb.!sfofrlt't11 dt.°'\ rumln.mu: domi,stiqut.~ Rc•n,r dtmon~'tOllon of lhc s<'hlYot11 M3Jt~. Rf'1Mrth fn VNrriudf)' Srn•nrti. 38.
d1fkt'(Jl',at de Ml<llrdl!c \'r1,•rl1111ilcue-, />11w Troplqunux. 3-1 139-1-13 352-35:.
SECTION THREE
Riclcettsial and
Chlamydial Diseases

•
 RICKETTSIAL AND CHIAMYDIAL DISEASES

Dise<1ses caused by bacteria in the orders Rickertsiales and Coxie/la bumetii, the causative agent of Q fever. is the
Chlarnydiales are considered in this sec1ion. Many of these subject of Chapter 43. It is noted t.here that the pre"ious
bacteria have recently undergone name changes which may cla~sitication in the order Ricketlsiales, famUy Rickettsi-
make tl1em unfamiliar to some readers. aceae. has been brought into question as a result of srR~A
Traditional bacterial classifkation was alway~ difficult, r.e- studies. and that reclassification in the order Legionellnles
!ylng hea\ily on propertie~ where differences were small and has heen propo~ed. Thi~has been confirmed brf'urther phy·
hard to quantify. Characteristics i.uch as stain affinity, cellular logenetic work based on the rpoB gene·1 and the currently
morphology. metabolic capabilities. and (in tht> case of intra· accepted classification of Coxiella burnerii places it in the
cellular species) hoSt cell type, could often not effectively dis· gamma subdivision (lf the Proieobacteria (f igure 2).
ringuish between species which had convergemly eve>lved imo
a common niche, and it had long been realized that a geneti- Phylum: Prol.l!Obatleria
Class G3mmaproteobacterla
callr based system would be preferable. From the mid· 19,0s
Order: Legionellales
the usefulness of ribosomal Rl\!A sequences to determine phy· Family 1.egionellaceae
logenetic relationships was recognized, and techniques for Family. Coxiellaceae
comparing small-subunlt ribo~omal RNA [srR'.'1:\ or 16S) gene Genus: Coxie/la
sequences began 10 be applied to bacterial phylogeny.' For the Co.viel{a bumetli
first time this allowed the application of statistical ph~lo·
Figure 2 Pama! ohvlogen'{ of 1tie gall'.'lla subdiV1s1on of !be phylum
genetic techniques 10 infer rhe likely paths of baeterial evolu- Proteobac,eri.
tionary dtvergence. The introduction of the polymerase chain
reaction (PCR) technique in 19887 allowed the specific amplifi-
cation of bacterial srRNA genes amid a huge excess of Chapte rs 44 and 45 deal with Eperyr//ro;:0011 a.id /iae-
host D!\A and che numbers of bacterial 16S sequences avail- 111obt1rtonella spp. respectively. and there is now good evi-
able began 10 increase ex'Ponentially. tn 1992 the Ribo- dence that thest' organisms are in fact 1'v/ycoplt1Sma spp.5 · 6
somal Database Project (RDP) was initiated. and the fir~t So fur onl) Epery1hrozoo11 suis, Epe1y1hrozoon we11yo11i.
public rl!'lease contained •171 I 6S rRNA sequences. The currcn r ilt1emobm·ro11t!lll1 muris and J/am11obar1011ella felis have
release of the RDP (RDP II version 8. Il :s held at ~ lichigan been reclassi'ied as ,I tl'coplasma spp., their ne" names
State University: it contains 16 280 bacterial 16S sequences1 being ,"'1ycopia,nw haemQs11is. MytQ/J/asma we11_1'011i. ,\!/yr.:o-
and a phylogeny based on these data. 111e ph}1ogenetic plasma Jiaemo,miris and Mycop/asmn /u,emofelis respec-
identities which follo.w are taken from !.he RDP II. !.he URL is tively. It is likely that the remaining EJ)eryzhrozoon and
lmp·llrrlp.rmP.m.<11.1>rl11/ l·lnPmnlwrumi>lln <pp. \\ill hP. ~imilal"ly rnc:lassified in due.
The reclassification of the order Chlamydiales is de- course. The currently accepted phylogeny of these organ-
scribed in Chapter 42. The phylogeny is sho,\11 In Figure l : isms is shown in Figure 3:

Phylum. Chlamydiae Phylum' Firm1i:vtes

Class: Chlamydiae Class: Mollicutes
Order: Chlamydiales Order. Mycoolasmatales
Family- Chlamydtaceae Family My;:oplasmataceae
Genus: Chlamyrfia Genus: Mycoplasm.a
Chlamydia tracoomaris Myooplasma haemomuris
Chlamydia SIiis {was Haemobartonella murls)
Chlamydia murldarom Mycop/a$ffla haemofelis
Genus; Chlamydophl/a {was H8emobsrtonell11 fell$
Chlamydophile psmaci Mycoplasm;i haemosuis
Chfamydoplii!a aoorrus (was Epet•/thtozocn suis)
Chlamytfeph/18 felis Mycop!asma wenyoni
Chlamydophi/11 cmae (was Eperythrozoon ll<lln;,oml
Chlamydcphila pecorom
Figure 3 Par!U!I pny1ogen~ of the genus Mycopla,ma show,r,g ~pgcies
figure 1 Pan,al phylogeny of the phylum Chlamyd;ae previous Iv ciass,fie!I in Ille gene•a fperythrozoon and Haemobar.one/Ja
505
506 ,,~.,o., '""st. Rick1t11$lal and chlamydia! di ..-as~

Phylum Proteobactecia Chapters 40, 41 and 46 deal with species in the order
Class: Alphaproteobacteria Rickensiales. which has been exlen~il·ely reorganized as a
Order: Ricl:eusiates
Family AnapJasmatateae result ofsrR:-:A phylogenetic anal~•sis.~Thc reorgani2ation is
Genus: Anapfasma described in Chapter 40. "here the agent ofheam,•ater. pre-
Anaplasma bovis (1•,as Ehrlicnie bovis} viously kno\\11 as Cowdria mmi1w111i11111. i~ now Ehrlicliia
Anap!esmaequi(was ETulichia equ~ mmi11t111ri11m. ,\11 the B1rlfcT1ill spp. described fn Chapters
Anaplssma marginate 41 a11d 46 have been renamed, with the exception of Eilr/i·
Anaptasrna phagocyropnilum(was cilia ,,mis. ,\ phylogtmcti~ tree giving the old familiar names
Ehrlichia {lllagrJCyrophitol
Anaplasma pfatyS (was Ehrlichia pfal)•s} is sho\\11 in Figure 40.1 of Chapter 40: Heartwater. and the
Genus. Ehrlichia currently accepted phylogeny of all these organisms is
Ehrftchia canis (unchanged) shown here, in Figure 4. Ofintcresr in particu;ar is that the
F:t1r/icf11a chaffeensis (unchanged) agen1 of Potomac horse fever. previously Elu/iclria ris1icii. is
Ehrlichra ewingi {unchanged] now renamed as o\"eorickeusin riscicii. An important distinc-
Ehrliehia maris(unchanged)
tion b;,,twecn the familie, Anaplasmataceae and Hicken-
Ehrlfchia ovina (ullcilanged)
Ehrlichta rurninanrium [was Cowdris siaceai.> should also be noced. 111c Anaplasmataceae
ruminanrium) comprises the genera ;\11r1plos111r1, £hrlichic1. .\"eorid:eusia
Genus: NIJOfickeusia and IVo/l)(lchia. which are obligate intra,ellular bacteria
Neorickeusia helminrhoei:a [unchanged) that replicate while enclosed in a vacuole derived from I.he
Neoricketme t1s1,ci1 !was Ehrliehia
cukaryo1ic hos1 cell membrane. The Rickeusiaceae com-
risticii)
Neoricketlsia senne/$u (was Ehrlichia prises the genera Rid:e11.~ia and Orienrin, which are obligate
sen11ets;h lmracellular bacteria that grow freely in the cytoplasm of
Genus: Wolbach1a 1heir eukarymic host cells.
Wolbschia p1pienris (unchanged)
Family: Rickeusiaceae
Genus; Ricketrsia
R1cke1tSJa prowaleHi(unchanged)
Ricl!etrsia nckeusiiluncilangedJ
Rickerrsia rh)hi(uncilanged)
Genus: Orfentia
Or,em12 1svrSl1gamusirl(was Rickerma
1wtsugamJJsh1)
Figure 4 Par. ·al phylogijnV of the alpha sub!livis,on 01 ~a J)hvlum
!'rc:eo:,aaar,a showing the order Ricker.siales

References
cnu. J,M,. l ""'· tt. \\ARSH. t.t.. PARR1!-. R.I \\',\.'W. ') .. JmL\.\t, ~ ..\ -1 :.1ourr c:. oft.,:-ri;rouftr, ,1, 1, fiA01111 u .. t9i}8~ Outt-rmint'ltion of CflXi••/111
at.\."<:ttJt.\ ~ \fCfV.RJU'~ t>~~, .. ~rw..uOl' t ,t.,. c.uoun 1.,.\1 .,._ J 1t.11n 1 ,, 1•111111.•tu rpoB scqucmce and h'l Ust" for ph)'lOJ,tcntttc a.n.o:I~~~ Grnt1• :?07.
,003. The Rlbo,omal Dat~bo.so PtOJL"Cl ROI' lit pra,1.,,,ni: a nc" 97-103.
aotu.d!gm:1 lh.n u11u"' r1'gufa, upcluui, ar.d :hr nl'\l ptok3.r}'UI c ~ '\ll'l,,f'\~,. It fOH.,'\"">'.'-0', )..tIH1i.llllM, " · · IUll'I J.c; ::uOt. Prcapo,.LI UI
111.,onon\\. •\'µe/v/e Arid< Ru$Mrcl:, 31. -1-12-111. tr ..u1..f\"t ,um1:m\.'snh\•r,.. oi t.ht: gcnt•rn 1/niJmol"trto,-;~lla ~nd
: 0-V'\fLE$t. J.j.,. BAftBCT. ,.r.. ft(;~l,.'f._fl,,. C. ()Mt:11, C....\.. P,\l'.'\.ffJl. C:. 11 tt,w. ,.("'.... fiJ"11'')·IIJro;orm co lhe grmi, ,\fyrop!asma ,,1th <fesmptfqn:> of
ofGent:a in the
f41h1Hll'~. , ..• RVR,\_'\:(;ntWA, F,tc•• :wo1 R('()fgam7lltiOn 'tlm;thdmtu .\JJ1.·11pkmnu iU1rmt>fi1lls' '('.tmtlldntu.'5 .\f.\'('()J)!aMtUt
FanuHes; Rick~tL~tacciw :.111d Anapla~m~t~~ an the OrdN Ricken).{.llC): Jm.·mtm:u,,l. ~omdirl0111A 1\l_\l'oplasnm ltncmosms wu:: ·Ctmdfdutm
Undk:nfon ofSomespe.caes or F..hrll(ilm ,,·1th .-t11apfrum1,. Co11.vir:11 \,i1h ,\1.rro,,1,'Qma "111J.".()t1I. Jm~·mnuQ11nl JcmmnJ cfSysr,·n:r,rtc and
Eilrlicl:ia. and Ellrlfthfn ,,:1th S00rfrt·~u$lt1. o-,q:rlption,.. of fhe ,.:,, F.t>c1turtonmy \ficwi1it>/ug.· :;1 851)-899
Spao.- Combination>: und D<rslgnotloJ• of /ihr/lchia oquJ ond 'HE 6 ,·toltrA-", tt ... tOJ-1.\..~"'<>'· );..f.. Hn:1111~\. T. ~ TULL, .1.c;.. :002. R\':'\Ulon or
a~~ru· as-SubJl~tive- Synnn)'l'n~ of El1tUch«t piU1t,,~l, !OJJ/rilt1. h•wmo:taphu.~ \,!_,~>pt,~nm ~p,t-dc~ nnm1.•'- lm,•nu11io1ull /tmmnl of
ln1t.mmlonnT /oumnt oj $)'tll'11tntlr muf !it-ollm.LJr:t,,:-· ,\1h'10M11lox,1, 5 l. syn,·murirmu.l l:'i·t1lurI,mr,ry· .Wftm1'lul1.>ty. 5:t ua.,
214;,-2165. 7 ~Alt.I. k,~ (a,u-,,o.,u, .. ,101r•1.~.-bt:fl.\HF.S 1. H1c.tro11.1L.llOK:--.,~.T..
3 ro:-.. c.$t:•. t•1.(;'l,.,,,u..~. K.J. & \\'Ott-t. t.A.• 19;;, Comp~~uve tara!o1t1,1ing of )JULLIS, K,B. &- utw<·n. 11••A.. 1988, r,11ner,direcu.~ cm~~ mauc
J6S nbrt~m:al rlbonuctelr acid. mofccular ;ippio.id1 10 ptoko\l')Ollc: .1mplltk;uion of OX\ wfth u th~nnoi:.rnhlc f>Olyn1~rast.1• 5':t.•nu. :?39.
, S)~tem:ttics. lllh'ftlt1tion11I Jm1r,u1t r,fSysunu,rJc &tc1c.•rfolt>rtj·. 2i I.S-57 .;87-491
 40
Heartwater
Synonyms: Cowdriosis. la cowdriose (Fr.). harrwater (Afrik.}

BA ALLSOPP, J D BEZU!OENHOUT AND L PROZESKY

Introduction 1ing the organism are present (see Chapter 1: Vectors:

T icks). The endemic area encompasses most of sub·
Heamva1er (cowdriosisl is a rick-borne disease of canle. Saharan Africa, including the islands of Madagascar, Sao
sheep. goats and some wild ruminants :hat is caused by a Tome, Reunion . .\<Jauritius. and :r.anzibar., 5 • 102· io3, 152 The
rickensia, previously known as Cowdrla rumiuqmium but disease is absent from the Kalahari Desert and dry coastal
recently reclassified as Elirlichia rwninanrium.92 Typically. areas of Namibia and South Africa. Heanwater also occurs
the disease is characterized by high fever. nervous signs. by- mi the French Antillean islands of Guadeloupe and Antigua
dropericardium, hydrothorax and oedema of the lungs and in the Caribbean Sea,.;..: to which infecred Amblyomm<I 11ar-
brain, and death. It is one of the major causes of swck losses legarum ricks were introduced. probably \\~th cattle from
in sub-Saharan Africa. Senegal. during the eighteenth cenlur}•.22• 1~ 5
The first reference 10 what may have been hearcwater The occurrence of heartwatcr is frequently taken for
was made by the Voortrekker pioneer Louis Trichardt in granted in the endemic areas and definitive diagnoses are
1838.16.1 \\1,ile trekking through what is roday the Limpopo not often performed. This leads 10 the pre,·alence rates of
PrO\'ince of South Africa many of his sheep succumbed 10 a the disease being under-reported.ss In rhe endemic area in
disease known locally as 'nintas· three weeks after they had South 1\frica mortalities from heartwater are three time$
suffered massh·e tick infestation. According to e,idence greater than i:,ose from babesiosis and anaplasmosis com-
given by a farmer. John Webb, co rhe Ca,rle and Sheep Dis- bined.16';· 2:1-, From n li\•estock census performed by the Di·
ease Commission of 1876 in Grahamstown. heamvater was rectorate of Animal Health. South Africa, in 1996, ir was
observed in 11!58 in South Africa in the northern part of the estimated that 17,5 million head of livestock were ar risk in
Eastern Cape Province. Because of its confusion with other the endemic area ofthet--ountry. Goa1s are especlally threat-
local diseases of unknown aetiology that were pre,"alent at ened. and in rnme pans oi the rural farming sector ii is be-
thar 11 ml'. ~nmc> of rhc> m,rliP.r infllrm:trinn rpgar<ling thP nr- li('Vt'rl 1hn1 up m ~O pc>r t:Pnr of gnats hf.rnmP infPnc><I ,drh
currence ofhearrwater is unreliable. 108 heart water annually. It is not known how many of 1hese die
The first imponam eiqlerimental findings came in 1898 from the infection, but a ,mbs{antiaJ proportion must do so.
when both Dixon70 and Edington93 showed tliat hearcwater The economic impact of he.inwa1er is difficull to quan-
disease could be induced by blood passage from infected to tify, botl1 because of the under-reporting noted abo,·e and
susceptible animals. No organisms could be demonstrated because the actual occurrence of the disease may be par-
in the blood or other tissues of diseased animals but it was tially suppressed by a range offactors. These include 1he use
concluded that heamvater was caused b~, a U,ing micro- of acaricides {ill 1996 acaricides costing USS 13 million were
organism 112 at that time believed to be a virus.:!!$ Ill 1900 purchased in Soutl1 Africa), antibiotic prophylaxis,180 i.m·
Lounsbury published his confirmation of the long-standing munization by infection and 1rea1mem.247 the resistance of
suspicion that the bont tick (J\mblyomma hebraevml was certain animal breeds to the discase,511 and endemic stabil-
the vector of heam\'ater in South Africa, 135 but another ity.~23 An e,nima.te of the impact ofheartwatcrwas made by
quarter of a century elapsed before C-0wdry demonstrated the Deputy Director of Veterinary Services or the Eastern
that the infectious agent in the. tissues of infected animals Cape Province ofSoulhAfTica during 1998. 1' His assessment
and ticks was a ricketrsia which he named Rickerr:;ia rwni- was that 10 per ccm of all stock losses, costing up to USS 30
11m1/iwn.61 112 The name was later changed to Cowdria ru- million annually. were due to heanwater, and this was de-
mi11anriwnr;s and eventua lly to Ehrlichia rumi,umtium.92 spite an annual expenditure of between USS I and 5 million
Heartwater occurs wherever ticks ~pable of transmit· for prophylaxis and vaccination. There are estimates from
507
508 •! cr1<>., 1tt•~E: ltickeuslal and chlamydia! di~eases

Zimbabwe of annual losses O\·er a 10-)•earperiod amounting as ro whether there should be more than one species of
to USS 5.6 million per annum. which inclui:Jes the cost of Ehrlicilia tex-Cowdria) is addressed below.
acaricides. milk losses and 1rearment. 1:;, The type specimen of£ r11111inanrh1111 comb. no\ . Is the
Heartwater is a major obstacle to the imrnduction of \Velgevonden isolate. 79 This was obtained irom an
high-pr(iducing animals into sub-Saharan Africa ro upgrade Am/Jlyomma hebmeum rick collecteo in the same geographi-
local srock1:!3· = and is of parricµJar lmporrance when sus- <.-al area as 1he original R. rumi11a11riwn which orfginared
ceptible animals are moved from heanwarer-free to heart- dose 10 the Onderstepoon \lererinary lnstin1te north of Pre-
water-infected area:;. 1ll6, 212 Whatever the actual cosrs may toria. In Gauteng Pro,ince, South Mrica.61 The pathology.
he it is certain that the economic imponance of heamvater morphology'g.; and \~rulence70 in cattle. sheep and goats ap-
as a disease of domestic ruminants in Africa is comparable pears to be the same as tharofthe original R. rumi11a111ium.o2
10 that of East Coast fever, trypanosomosls, rinderpest, and The 16S gene sequence' 1 is the same as thar oi the Crys1al
dermamphilosis.21 · 192 Springs Isolate,"° which is deposited in GenBank under
The possibility of heamvater spreading from the Amil- accession number X61659. The Welgevonden genorype is
lean islands to the American mainland. where a suitable rick common in the field9 and fro-1.en stock~ o( the isolate, held a1
vector is present. is an ever-presenr threat ro tbe livestock rhe Onderstepoort Veterinary lnstitute, date back to 1985.
indusrry there_..a. HZ. 23~· 236 The disease is also a potential Ehrlich in rumi1u111r/11nz was very easlyshown 10 be heat-
danger to countries where the vec,01'$ mar be introduced labile and to lose It,; viabilil) within J 2 to 38 hours at room
and become established.2J· zoo Ir will be of major importance temperature.? 132 The first long-term cryopreser\'ation of
until an effective and safe vaccine becomes available. infectivestabila1es in liquid nitrogen was in media contain-
ing dimethyl sulphoxide (DMS.OJ. u s. 203 More recemly a su-
crose-potassium phosphate-glutamate medium (SPG: 0,218
Aetiology
M sucrose. 3.8 mM KH,P04 , 7.l ml\1! K1 HPO,, ~.9 mM
<::0111dria ruminamlum was traditionally classified as the Csl-luNO, l<))' has been found to preserve the infect.i\icy of
sole species of the genus Cowrlria. rribe Ehrlichiea·e. fam!Iy the organism more effeclively.43 The infecti\'e half-life of
Rickensiaceae. order Rickeusiales.205 h had long been stabilate material fro7.en in SPG. and then thawed and kept
realized, however. that 1he organism had a close antigenic on ice. is only 20 to 30 minures.43 Large numbers of srabi-
relationship with certain Ellrlichia spp .. and in 1992 the lates have been presen·ed from almost all the areas where
first molecular phylogeny of the organism. based on the the disease occurs and these are referred to as 'stocks' or
16$ ribosomal RN,\ gene sequence of the Zimbabwean 'isolates' according ro t.he terminology used for rrypano-
Crystal Springs isolate, revealed that it was related to some populatlons. ' 3 The names and origins of important
Anap/asma margina/e. 05 This was closely followed by an stocks which are in current research use around the world
analysis of the (slightly different) l6S sequence of the Sene- are given in Table 40.1.
gal isolate, which showed a closer relationship to Ellrlichia At one tirne it was thought that E. r11111inm11i11m was a
canis and Ehrlich/a chaffeensis than to A. margirwle.= relative!} homogeneous emity, but within the last decade
As the 16S genes of more organisms of the familr Rickeu- the advem of genetic $equence data has revolutionized this
siaceae were sequenced it became e,'ident that Lbe geous norlon. I, is now known thai there is fur more genetic vari ·
Ellrlichia did not constitute a monophyletic group. and r.hat ability among £. rumi11an1i111n organisms than had ever
organisms classified as A11r1plasma. Co11Jtfria. and Neo- been suspected.
rickeusia were members of three separate cladcs, each of Currently eight different 16$ ribosomal fu'X/\ genotypes
which included various organisms classified as Ehrlich/a are k.110\\11 (Figure 40.2), all classified at prese:n as f. rumi-
spp. 1si. ~:; These clades became known as genogroups I. 11a11ti11m. Howe,·cr. 16$ sequence data cannot be used as
II and Ill Ehr/ichi<1 (Figure 40.1 ) with C. mmlnmirium the sole determinant of what constitutes a species: it is also
being included In genogroup lll. t0gerher with £. cmlis. lmponant t0 be able to grow a stock in tissue culnue in order
E. chajfeensis, E. muris and E. rn11i11gi. 6 A:. the 16S genes .o f to accurately cletermine its disease-causing status, and this
more isolates of Cowdrla became available it was found has 1101 been done for several of these eight genol)'PC'S. Some
that there were several distinct Cowdria 16S genotypes, all of them are undoubtedly justifiably classified as£. rumina11-
of which fell into a tight cluster wilhin the genogroup ill ti11m. but orhers may need to be reclassified in the future in
Ehrlichin.9 the light of their infeclivil)' and pathogenicil)' in ruminant
, The most recent revision of the order Rickeu5iales"2 ho,;is.
abolishes the genus C<m•rlrla and the tribe Ehrlichieae. All It has long been stated that E. rumi11anriwn srock!. dis-
three Ehrlich/a genogroups are moved to the familr Ana- p lay differing degrees of pathogenicity in different hosts.
plasmaraceae. genogroup I as.Veorickeusir1. genogroup II as Unfortunately much of the information ln the earl~ litera-
Anaplasma. genogroup mas Ehrlichia (including Cowdria). ture which relates to parhogenicicywas obtained in the days
Ehrlichin rumi11a11ti11m is therefore now classified in the before a tissue culture method had been established, before
family Anaplasmataceae. order Rickettsiales. The que.srion reliable methods for quantifying the infective dose were
 Heur1wa1cr 509

~
- EM·cli16 sµ Germis1urt1
- f can1s Flw«la sua,n
- E "1slieet1S1S
[ muns
4
E ~Nir.91
Co,•/;ft'a fl)mmenr,um Clvstal Sp«n;s
- - - E sp. '10m Odqco//eus w:g,n,enw
-E sa. Huma~ granuiocv,oLrC!J e
,-- -
....., - [ equl
-£ p/18goc1•1ophiLn.rain FG GENOGAOUP 11
- £ sp Omatjenne
-£. p,arys
- - - f bov,s
'-----Aoaplasma margina/e
_ - -E senm;;sustrain 11908
--1 -E. tis:,c.1i GENOGROU!' I
_ _ - E -SD lrom Sreflamcluismus fa Ce,LJs
-Neor,cke.t,sa ltelmmthaec;i
-P.,cterrs1a ,:'Ci:e:rsu
,-
_-R l'fpn,
--R aro.~~teK't
- - - - - - - Esclurich,s c1J11

Figure 40.1 Max,mum l1i:ehhood tree based on TES rillosoma: RNA gene sequences 1n !he oroer R,t~etts•ales belore reclassihcatron. 1V1th &:nencr.,a
coli as ourgroup. Note Sl!eC•es classified as EhrlicNa occurring in 1hree separate paraphylei,c clal!es lgenog,oupsJ Cowd11a 1uminanm1m is located ln
genogroup 111

--Kiswani
Pre1ona Nonh
-----Maia 87/7
E rummanz,um
- - - ~tal Spongs 1sn?21es:
Ball 3

---0/l'li(j!!l ~

Ehrlichie cJnis florl!fa

~J ---£~rlicl1ia chaffeensis
Ehrlichtu sp, Gerrn,sh1;ys

- - - - - ihr/ichia mur,s
Et.r/icNa ew,ngi

- - - - - - - - - - - Noo,;rkeus,a helmrnthoeca

Figure 40.2 Maximum likelihood tree ba58ll o~ 16S rioosomal RNA gene se:iue= or some spec,es m jte f2m1!y Anaplasma,aceaa. will flic;e;;sia
0towezekil 1iam1ly llickerts1aceaeJ as outgroup. Nnte ;ne e;vni isolates of El11/tch1a rummanr,um wn1ch are mo!c: closely related to eacn cthe; 111an the·1
are to otlle; Enrlichia spec,es
 Henm, mer 511

Table 40.2 Cross-,mmunity CHALLENGE STOCK

pmtei:11on eng:ndere:l in Sheep SiOCKT0 1NHICH - - - - - - - - - - - - - - - - - - - - - - - - -
be!Vl!!en vanous Ehrlict11a IMMUNE Wi:lGEVONOEN BAll 3 GP.ROEL MAAA KWANYANGA BLA<\UWKRANS
r11rnman1wm isolates 87/7

Welgelillnden • •
' ••
Bal 3 • •I- ·'-
-·-.
Garcel •I- + •I-
Ma;a Sin -I-
Kwanyanga +I- ·/- +
Blaau,\ira•,s •!- ,,... -
~ comp!et~ ucss·p•otectlon
no c1BSs-pro1ection
+/- pan,al cross-n:otect,or.

equally infecnve by both the mtravenous and the mtrapen- before It can be said whether they enuse ctass1ca1 heam\'awr or
1oneal rou,es, whereas other isolates were mud1 more infec. c1·en whether they p05e any di~ease threat in mminam~.
tivc by the intravenous route than by rhe intraperitoneal Ehrlirllia r11mi1u111riw11 can be propagated In ti~sue
route, 136 and it appeared to be non-pathogenic ror caule.11(1 culrures. In the first succe.o;s-iul initiation of E ruminnnrium
The stock also resisted all anempts 10 establish it in tissue cultures a tick-deril·<.'tl srabilate. prepared from :\mblyomrm:r
culture over a period of 15 years. Only recemly ha,; it been /1e//raeum nymphae a$ the ,ourcc ofinfectious organi,m~. was
both established in culture a nd characrerl1,ed geneticall~. used:~; In subsequem impro1·emems 10 the technique freshlv
and the stock is now known to contain mo different geno - drawn hcparinized infective blood and suspensions or li\·er
types, Kumm I and KUmm 2. each having a distinct behav- and spleen prepared from infected mice were employed to ini-
iour in cul mre.269 tiate cultures.Z'l .\ technique using plusma from heparinized
One stock, isolated before any genetic characterl1.ation blood was deYised which showed that infective £. mminan·
data were available. seems to lie 1.he mos1. likely candidate for tiwn ls presem in 1he ccll·fr~>e plasma ofinlected animals.' "
reclassification. The Germishuys stock was isolated from a Several procedures have been described to facilila[e mi-
naturally infected sheep92 and I.he record~ suggeSI. that it tial Inrection or the endothelial cell monolayer. suC'h as irra-
caused disease which was thought to be hcartwater. ·nw stock diation and l'yclohcximide treu11nen1 of the endothelial
was therefore assumed to he Ehrlil'hia (Cowtiria) mminan- cells'" or the use or a s low-rocking platform. 29 Alchough
rium mnil l6S gene sequence daia showed it to be a 1>reviously rocking or the cullure flasks is still widely employed it has
uncharactcrizedEilrlichia more closely related 10 H. can is than been demonstrated that these trea t menu. are unnec~ssary
to £ mminnmiwn. 11 This organism has been provisionally for the initiation or propagation of E. rwninam/11111.z~r,. 2fl-1 Jt
named Ehrlicllia sp. Germishuys (Figure '1.0.2) and no 1ypic-dl ha,. also been ,-!)own that continuous propagation of£. ru-
1:.~ mmint111ri11m I6S sequences have ever been recovered from mi11t1111i11m in DH82 (canine macrophage-monocytcl cells
it. However. because it has never been grown in tissue culture was only achievable when cycloheximide was added. w7
its ability 10 cause a disease 1.hat resembles heanwau.'r has not Eilrlichia rumi11a,1rium culture$ nre normally incubated at
yet been properly evaluated. around 37 •c and. depending upon the pH of the buffer sys -
TI1ere are at least rwc, other stoC'ks. currencly considered 1.0 tem used in the medium. either in air or in a carbon dioXJde-
be£. mmi11ami11m but not yer grown in tissue culmre, whkh enrichcd atmosphere.
may need 10 be classified as new Ehrlic/1ia spp. if they are even- The multiplication of £. m111i11a111i11m within the endo-
tually found not 1.0 cause heanwarnr in ruminants. These are thelial cells of infected animals was originally described by
the Omarjenne and ?remria North genotypes. Both arc phylo- Cowdry in 1926.63 ~o it 1,. not surprisi ng that the first success-
genetically more closely related 10 £. n1111i11m1tiw11 at the I6S ful in 11/tro propagation of r:. rumi1111111i11m was achieved
le\'el than to any od1er currently kn0\\11 £/,r/ichia spp. (Figure using bOl~ne umbilical cord endothelial cells as host cells..JS
,10.2) so the reclassification question is more difficult than in Almost all sub~cquem propagation studies of r:. rn111i11a11-
the ca.~e of Ehrlichin sp Genni~huys. The Omatjenne geno- 1i11111 used e11dothelial cell~ from variou~ species and from
type was originally derived from a Hyt,lomma tkk and it docs various anawrnicaJ sites, Mich as bovineaonaand pulmonary
not appear to caw,e disease in goatst I orin ~heep.~ me Preto· artery. 18 01~ne aona:'9 shee1> br;iin," 1 bo1ine hrain. 119 bol'ine
ria ~ orth genotype was origi11ally detected in a dog by DNA hy- brain microl'a~cularurt>ii; and caprlne jugularvein .:!2!1
bridi7.ation7 and it is not rurrcntly available either as on The propai:ation of £. rumi11cmri11m in non-nurunan(
infecri\'e stock or in culture. Both of these unusual organisms endothelial cells:?Z" wa.., fir,t carried out using human
must be gro"'11 in tissue culrure and wsccd in 1livo in ruminantS u mbilical and human microvascular endothelial cells.
512 , 'rir" n ,., · Rkke1tS1al and chlam;ulal di,=es

Subsequently propagation was achie..-ed in end,ithelial cell

lines from a range of" ild African ~pecies. including those or
a bushpig (J>o1amo1•/zoeru.~ porcus).1 •
Shon-duration cultures. the cell~ of which are un,uitablc
forcominuous i111•i1ro propagation oft.he organism, ha~'(' hecn
achieved in caprine nemrophils. 1"" in mou!IC and mminam
macrophages.-, and In monocy1e-macrophage cell !Inc" trom
mice and dogs. 1" ,\part f,om endothelial cell and 01182 cell~~;
lines. conunuous propagation of E. rumim1111i11111 has alS<1
been achieved in a rick cell line from l.rode,:sa1pu/(lri., (IDl'.8).2:,
Various media. including commercially a\·ailable S~'l1-
thetic culrure media. have been used for 1he in l'i1ro cultiva-
tion of E. nm1i11a111i11m. Example; of the la11er arc the
Glasgow modification of Eagle'; minimum essc;mial
medium (GlVTEMJ.29 Leibovitz L-15.~11 Dulbecco's minimal
essential medium (DMEM) 1'w and Dulbecco·s modified
Eagle·s medium nutrient mixmre F-12 Ham (Dl\lE/F-l2).i 7o
Commonly used supplements for a complete culture l
medium are foetal bovine ::.erum (FBS). 1ww-born calf
serum and bovine serum at \'arious concentrations. usunlly
JO per ccm (vi\·). although 3 per cent Fl:lS has been used in
one ins1ance. 2b·l Another supplement frequently used is
uyptose phosphate broth. The media mentioned above also
comain L•gluta111111e and a ntibiotics. ru1d glucose (0.45 per
cem) has heen a<lded 10 an L· 15 ba,ed me<llum'18 which
olherwise only conmil,s 0-galactose.
Work 10 develop a chemit.illy defined culture medium
for E. rttmimmrium passed its first mileste>ne when a ~erum·
free culture system was achieved. using a modifit•d I IL· I
• •

medium.:a~n In ,.ubsequent experiments it was round thar

the serum could be eliminated when l).'.\fE. ' F-12 medium
supplemented with bo\'ine lipoprmein~ and bo\~ne trans-
Figures 40.3 !top) and 40.4 (bottom) light m,croscop1t :ncrphology of
ferrin was used. 268 The final step was replacement ofbovine Ehrl~hliJ r111wnanilum r cao,uarv endothelial cells ,n bra,n S!Tlears
lipoproteins by chemically defined lipid,, and bo\·ine trans- stained wit G,ernsa
ferrin b} an inorganic source of iron; this rewlted in the fir~t
formularion of a chemically defined, protcm-free medium
for the continuous culture of Ii. rumi11,1111illm. 2"" pink background of tick tis~ue. This ,taining me thod is u5e-
Ehrlichia mmirum1i1m1 ti> a pleomorphic rickcttsia, ant.I ful for idemlfving /;/ir/ir:hia-infected ticks and is superior ro
colonies containing from one Ort\,·o to sc,·eral tl1ousand in- the \-ariations of \lo.llory'5 ~,ain which render the colours of
di\idual organisms are found in the C)'loplasm of endothe- colonies and uck tissue) various shades of blue?";
lial cells (Figures 40.3 and 40.4J." 3 167 In ge,,ernl the nw ultrastruetural morphology of E. rnmillanri.um rFig-
colonies consist predominant!}· of small (0.4 µm ). me<lium ures 4-0.5 to <10.8) i:. ~lmilar In endothelial cells of the choroid
(0,,6 µm l. large (1.04 µm) or very large (> l,O·l µm ) organ- plexus of ,hccp, 187 cndolhclial cells in the lungs of mice. 19' en-
isms,18~ bura number of smallcrorgani~msare also found in dothelial cell cultures.19~· 19'' and leukocytes.1l-l and in the cells
colonies of larger organisms. and 1,ice rersa. \.Jost organisms oftic~ya. ii - l!S
are coccoid. except for colonies comaining ve~ large organ- Specie~ in the family Anaplasmataceae. including Ji. rn-
isms in which plcomorphic form• !horseshoe-. ring- and bn- 111i11mui11111 grow in an imravacuolar compartment.
clllary-shaped) may be scc11. 1u7 bounded by a lipid bilayer membrane. within the cyto-
, Ehrlichia rw11innnliwn is a Grrun-neg,nive hacrnrium. pl~~m of lnfPrtNl hMt <'Plls.~2 Thi; di<.1ing11i~hr~ 1hem
o.nd stains purplish-blue wilh Giemsa.v:1 Several Olher stain- from species in the farnUy Rickcusiaceac which grow freely
ing methods have been used or developed to demonstrate the in the cytoplasm or their cukaryotic host celb Individual
organi,rn in its vertebrate and hwericbraw hosts:'"· 187 iu,i. l(>; I:. rum/T1a111ium cells, in common with other Gram-nega-
In paraffin-sections prepared from tick midgut and stained tive bacteria.I cells.26 are bounded by rwo membranes. an
with Mallory's phloxlne-methylene blue stain, F.hrlichia colo- inner (plasma ) membrane and an outer membrane. Inter-
nies and host cell nuclei s{ain dark blue against a uniformly nally the cell contain~ electron-<len$e and electron-pall'
 1-kanwater 513

areas. with electron-dense material occup)ing the greater readily fit into one or the other category are referred to ns
part of the inner strucLUre of small and intermediate-sized intcrmed iate bodies. 122· 1117 191
cells. whi le elec tron-pale areas dominate the inner struc- f:hr/ichia nm1inrmri1m1 replicates 1nainly by binruy fission.
ture of large and very large organism~ (Figures 40.5 and and possibly by cndo,pont.lation. 167 lt appears that the reticu-
40.8). The latter are referred tc> as reticulated bodies. and h1tcd bodies are predominantly proliferntivc1i;. •~. i,i,,_ 197
the smaller organisms arc described ns elcmcnmry (clcc- while the clcmcmmy bodies roprcscm the infocti,·e stage. 122
tron·dense) bodies (Figure 40.7). Organisms \\hich do not There is some C\~denc-e that £. rumina11ri1rm undergoes

Figure 40.5 Colony o' Ehrlich,a rvmmanc,um in a~ endotnalla, cell Figure 40.7 E'e..,enta11 ,e,ec,ron-Gensel bod,es

Figure 40.6 'Mixed' colonvof Ehrltchio fl/mm;;nr,,;m Figure 40.8 Retculated b<ld,es
51-l ~crro, m-. t: llickettsial ru1d chlamydia! diseases

a sequential de,·elopmem in both the ,·ertebrate78 and the omma spp. capable of transmitting the organism occur in
invertebrate hoM. ii, Transmission electron microscopic Africa. The major vecrors arei\. 11nriegawm and.el hebmew11,
studies of in uirro culwres of E. rwni11m11i11111 in endothelial the latter being the main ve<.tor of heartwater ii: southern Af·
cells have revealed the presence of intracellular rericu la te rica. Amblyo111111a 11ariegarw11 has the widest distribution
bodies rwo 10 four days post-infection. and intermed iate in Africa and is the only originally African Amblyomma spe-
bodies four to five days post-infection. l !!Z Large numbers of cies that h~ established itself succc~-sfully outside the conli·
elementary bodies are seen afrer rupture of rhe endothelial nent 011 two Island, in the French Antilles. Ho The species A
cells five to six days after culture initiation. l:!i sparsum. A. t/,ol/oni and A. 11mrmore11m were at one rime
Heam,'llter is transmitted by ticks of the genus Ambly· considered 10 be ·accidental' vectors which do not nonnally
omma (see Epidemiology, below, and Chapter I: Vectors: reed on domeslic stocl:, 166 but more rccenr evidence suggests
Tlcks). :\lost Amblyommtl spp. are three-hose ticks. Larvae that this ,s not tnie for A. marmoreum at least. and that rhis
and nymphs become Infected when lhey feed on domest.ic species can be an effective vector or beamvater.183 Ambf.r·
and wild ruminants and possibl~ also on certain game birds omma sp~rsum tick:. collected in 1999 from leopard tortoises
and reptiles31 at a time when E mmlnantillm is circulating in (GeorhelDne ,>ttrdalis) imported imo the USA from Zambia
the blood of these hosts. 11\e immature stages of the tick com- were also found to be carrying£ n1111inn111iu111.'' although it
monly feed on smaller species of domestic and wild rumi- was not demon~1ra1ed that the genotype detected wm, infec-
nams and game birds, while the adults prefer catlle and the uve t0 ruminnnl!,,
larger game animals, such as African buffalo (Syncerus cajfcr) Vectors of lesser irnponance include A pompo.mm
and giraffe (Gimffa ca111elopard11S), as hosts. 1s,; Xymphs or which is pre,•alem in Angola: A. lepid11111 wh!ch occurs in
adult ticks transmit £. ruminamium 10 susceptible hosts East Africa nonh oflatitude 8 •s: A asrrion. whid1 primarily
wilhoul losing the infection. lntrastadial transmission has parasitizes African buffalo in Central Africa; and A. cohaer-
been demonstrated. 12 and rransov-dl'ial transmission was ens and A. gemnw, which occu r in East Africa.ZS(,
once demonstrated in very heavily Infected ticks under labo- ,'\mong the .-l>11blyommt1 spp. native 10 the US,\. A. ameri·
ratory conditiom,34 but it is unlikely that it occurs in Lile field. camon and: \. cajo1111e11se are only marginally susceptible to in·
The developmental cycle of£. rumi11anrium in lhe tick, and fectian and apparently do not cransmlt lhe parasite.Hl while
the infectiviiy of successive stages of the tick, are poorly under- A. mat£1lat11111, on the other hand, has long been known to be
stood. It is lhoughl that after an infected blood meal. initial rep· capable or transmitting the disease23:l and has a vector effi-
lication of organisms takes place in the epithelium ,,r
the cienC)' in sheep which is similar to tlmt exhibited by A. V(lri,,ga-
mtesrine of the tick and that the sali\13!y glands eventually mm.142

become parasitized.127 Colonie,. of £. ruminmuiurn in tkk The impo11anceof.-\m/J/yom111aspp. as vectors ofheamva-

haemolymph~9 may be imennediate fonn~ in transit from in. ter depends not only on their vector cfficlem:y but also on their
1estinal to salivary gland cells. Transmission oflhe parasites to distribution ai1d adaprnlio11 to domestic stock23" ai1d on their
tl1e vertebrate host probably takes place either by regurgitation acti\ity and abwidance, the latter being profoundly innuenced
or through lhe saliva of the tick while feeding. The minimum by temperature and humidity. 1116 An increased pre1'l\lence of
period required for F.. rumh1c1111i11m to be cransmined after hearrwarer usually occurs when peak numbers of ticks arc
ticks ha\'e anachl'd to susceptible animals ls between 2; and 38 present. and good rains are often followed by a transient in-
hours in n)mphae and ber1,,een Sl and iS hours in adults.3 1 crease in the occurrence or the disease. Its occurrence is not.
howe\'er,strictlv seasonal. a fact which is panicu.larlv true in re-
gions b.l<e the C:aribbean50 and in parts oi Africa which have a
Epidemiology
temperate climate and n poorly defined rain) season.
The epldemiology of hearrwater depends upon factors Am/J(romma llebrao11m ricks fcerling 011 E. mmi11m11iwn-
relating to the tick vector. the causative organism. and the ver- infected sheep have been shown to become infected during a
tebrate hosts. Important considerations relating 10 the tick period from rwo day!. after tbe commencemem of the
vecror are infecrion rates In 1he ticks. seasonal changes lnnu. temperawre reaction 10 two days after tl1e animals have been
encing tick abundance and acth~ty. and the intensity of tlck treated fer heartwater.3 1 Tbe transmis.-aon of E. ruminanrium
comrol. Significant factor-s concerning the parasite are dilTer- b)' A. 1J(/riega111111 feeding on Crrole goats in Guadeloupe! b ap-
ences be1ween genotypes which may affect virulence or the parently somewhat dilferenr. with the organism beinf:! trans-
stimulation of immunot()gical cross-protection in ruminants. mitted tothetic~afteradelayofLwo to three days fornfmphs
l)S far as the venebrate hosts are concerned. the ava.11abiliry or and four dar,. for adults.53 and the disease iS apparently more
,,1ld animal reservoirs and the age and genetic resis1ance of oflen fa1al when transmicted by tlcks than by needle transmis-
domestic ruminant populations are ofimponance.234 Some of sion. Ostensibly health)' ruminant hosts have been shown to
these issues are summarized here. while lmmunologkal re· remain infective 10 ticks tor long periods, at least 361 days in
sponses10 heanwater are dealt with further on in the chapter. canle t?- and t I months in Creole gonis:'~1 In [he ,aner case the
Heartwater occurs only where its ,·ectors are present (sec carrier status could not be detected pennaneml)' during the
Chapte r .I: Vectors : Ticks and Figure 40.9). and LO Ambly- I !-month period. demonstrating the danger which could be
 Heanw;ncr 515

D A. hebraeum

D Apomposum

D A. YSriegotum

D A. vaneg,mun elld
A pomposum
D A. heb:aeum aridA. var,egarum

[:] A tep1dum

Figure 40.9 Distrlbu!ion of toe rr.aior

Ambf)'omma spp. vectors of Ehrtithia
111minan1ium in Af11ca

posed by the movemenc of apparently negative carrier ani- African isolatl'l: than co a western African and a Caribbean
mals to areas free from the disease. isolate. 140 On the other hand, asimilar level of susceptibility
As little as 2,7 co .5.5 ml of blood collected from a host of A hebme11m to illfection has been demonstrated in all the
during the febrile stage car1 infect Am/1/yommn larvae_'.n isolates tested. which may explain why hcartwater is gener-
Ticks retain their infec1ivity for life.ss. 113• 166 so a small ally a more serious problem in those areas where A he·
number of infertrd rirks 1·n111<1 prC'~umahly m:iinrnin 1h!' hra1111m i~ rhP prinripal vertnr
infec1ion in a particular herd or area. The infection rates of When a pathogenic genorype of£. rumi11amium infects a
licks vary according to che season and locality in which susceptible vertebrate host, either inapparem or oven
they are collected and may be surprisingly low. Survey~ in disease may deYelop depending on the pathogenicit)· of the
South Africa have found that 1 co 7 per cent of A. hebraeum organism and on the species, breed, age. degree of nacural
in some parts of the endemic area were infected at any one resistance. and immune status of the hos1. Young cakes.
time. to. 79• ~ While somewhat higher rates have been deter- lambs and goat kids possess a reverse age resistance which ls
mined in Zimbabwe 169· 165 and Senegai, 1°3 the infection independent cf the i.mmw1e status of the dam. 3· Iii. 167· ll0. l63
rates are in general low. When infected sheep are used ex- This resistance usually fasts for only the first four weeks of life
perimentally w feed A. 11(1riegar11m during their larval or in cah·es and the first week i11 lambs and kids,167 although it
nymphal stages, they show infection rates of 100 per cent at may persist for sLx to eight months in cal"es. 89 This age resis-
the follo"~ng ins\ar. 10.l This suggests that the likely reason tance is not absolute as infection of some cal\'~ less than
for low infection rates is that many ticks feed during the 1hree weeks of age and of some iambs and kids less than one
larval or nymphal stages on non-susceptible or non- week old may result in fatal disea~e. 167 • 23Q
infected hos:ts. The susceptibility of differen1 breeds of cattle "aries, Bo,
Amblyommn 1,ariegarmn 1icks also differ in their infec- i11dic11s (Zebw breeds being in general more resistant than
tion rates "~Lh different genotypes of c. rumi11ami11m. and are European (Bos rauru.s) breeds.u. The resisrance oflocal
have been shown to be less susceptible to two southern Zebu breeds, such as the ;'\guni and $anga, is probably due
 llcanwmer 51 i

regurgitated gut comem.30• 127· 129 Initial replication of the heanwater generate lymphocyte responses characterized by a
organisms seems to take place in reticulo·endothelial cells mixnrre of CD4". cos· and yo T cells. and t.he expression of
and macrophages in the regional lymph nodes. after which T helper type I ci·iokines. 14"· 158• 228 \,\lhen peripheral blood
they are disseminated ,~a the bloodstream and invade en- mononuclear cells (PBMCJ from heartwater·immm1e canle
dothelial cells of blood vessels in various organs and tissues, were exposed 111 11ilro. to either autologous-infecrod enclo·
wl,ere runJ,er multiplkation occurs.n In domestic 1Utni· thelial cell~ or Infected monoci'!es, PBMC proliferation was
oants £. ruminantium most read ii, infect~ endothelial cells observed. 1sa Under tJ1e same conditions. howe\"er, ~ nimi-
of the brain, and thi, coincides with the onset of the febrile 11ami11111 elementary bodies did not induce PBMC proiifera-
reactiort.73· 75 Gene1ic variation within both parasite and rion.1511 The endothelial cells required pre.treatment with
host populations plays a role in the degree of endothelial cell 't·cell growth factors to induce class U~I 1-iC expression prior to
colonizati<ln in the brain and other organs.~ infection and lheir subsequent use a.~ slimulators of PB!'-IC. 1~
Increased vascular permeability with transudation is re· Cells Crom immune cattle have also been shown ro respond
sponsible for effusion into body ciwities ~nd tissue oedema, specifically tO recombinam antigens of E. rumimmti11m, CD4'
and this is panicularly noticeable ir\ the lltngs. pericardia! T cells responding to major outer m1lmbrru1c protein (MAP} I.
sac and pleural ca,1ty. Oedema of the brain is responsible and y& T cells responding ro MAP2. Borh proteins induced a
for the nervous signs. hydropericardium contributes to car- T-helper type I immune response. 159
diac dysfunction during the terminal stages of the disease. Peripheral blood mononuclear cells from animals made
a:nd progressive pulmonary oedema and hydrothorax result resistanr 10 challenge by vaccination with inacti\'ated organ·
in eventual aspbyxiation. 178 The effusion oiOuid into tissues isms (see below) contain £. n1111i11m1ti11111-specific. :>11-iC
and body ca,~ties also results in a drastic reduction ln blood class II-restricted, iF::S:·",l·producing. CD4- T l~mphO·
volume ..;& cy1es.22.q CD4' T-cell lines generated by the use of E. r11mi-
The pathogenesis of the vascular permeability remains 11<1111i11m lysa1es responded strongly 10 whole lysate.s bur not
speculative, and in mice there is no apparem correlation be- 10 recombinant :vtAPt or '.\·IAP2 proteins.:u' When these cell
tween the number ofparasitized endothelial cells in the pul- lines were Slimulated "~th soluble E. rumi1u1111/111n proreins
monary blood \'essels and the severity of the pulmonary purified by fast perrormance liquid chromatography
lesions.197 It has been suggested that cndotoxin241 and in· crPLC). a ~Ingle pC1al:. whkh included proteins between 20
creased cerebrospinal fluid pressure~ 5 play a role in the de· and 32 kD::1, induced proliferation. 220 Tn the same study only
velopment of lung oedema. ·(6 T-cell lines responded specifically to recombinant ~LAP!
It is known that the parasite causes down regulation of and M1\P2. Whiie flow cytometric analysis of PBMC has not
imcrferon (lf!\'J·"! production and inhibition of major histo- shown an)' significam change in the immune cell popula-
compatibility complex {MttC) e.xpression.2.'lll This would be tion after vaccination with inactivated organisms signifi·
expe.cted co slow the development ofimmunocompetent cells cant changes do occur when the animals are challenged.
and. although no £. n1m/11a111lwn specific L)'!Otoxic cos· Initially there is.i depiction ofC0,1·, CDS" and y5T·cellsub-
T cells have been demonstrated, It is known that cos• T cells sets, and a rise in numbers and activation ofmono~ies. and
pro!iferatein animals vaccinated with inactivated£. mminan· only later when the animal is fighting the infection. is there
ci11m and chen challenged with vinilem organisms. 146 It Is an increase In cos· T lrmphocy1es.1•1~
possible. therefore, that if T-cell proliforation is slowed until Immune cattle produce if;,- ·Ci when s11bjected to a lethal
man)' endothelial cells arc infected the subsequent specific E. mmi11a11tillm challenge. while non-Immune animal~.
killing of these cells could lead 10 blood vessel pem,eability. similarly infected. that died did not produce this cytO·
Against this hypothesis. however. is the observation that bot.h kine.:u6 Even so IFX·« alone could not complete!)' Inhibit
infected and uninfected endothelial cells appear 10 be severely the in 11irro growth of£. rw11ir1n111illm, suggesting that other
damaged in ruminants which die from the disease. 1117 facmrsare required for complete inhibition. The type2 cyto·
The earliest work on the hnrmme response 10 £. mmina11· kine IFN·yhas been shown to be a very potent inhibitor of£.
tlum were attempts 10 1ransfer immunity from heartwa1er rumi11a11ti11m gro\\ih in endothelial cells. but the mecha-
imnume canle to naive animals by transferringsemm.2. i2. ; 3 nism of this inhibition is unknown.22'1 Another important
These auempts failed, indicating that antibodies do not con- role of IF:-S-·1 is the upregulation of MliC class l and II ex-
trol the course of che disease. aJtJ,ough antibodies may stili pression on various cell t)11es. 60 This effect on monocytes
plai• a role in opsoni1.a1ion. complement·mediated killing or will lead to enhanced presemation of E. ruminamium anti·
;mtibody dependent cell mediated cytotoXicit)·. As £ rumi gens to the immune system. Ehrlichia rum/1w111irm1 has
11amiiu11 Is an imracellular parasite it would be e>.1)ecced chat also been shown to elicit de 1Wt'O synthesis of Interleukin
both cos- cytotox.tc T cells and CD4" helper T cells would be (II.· 1p, 1L-6and IL·8 mR:S..'\ in bovine brain endothdial cells
important in the development of protective immunity. in 11irro, an effect which wa~ potemiated by l:'ff··r 36 Since
TI1e first confirmation that cos· T cells play a major role in IL· I and 1L·6 can ac1 a~ CO·Stimulatory signals for T· and B-
protection came from adoptive transfer studies in a mouse eel! acuvation231 it is possible that by secreting these cyto·
modeJ.78. 81 • so It has now been shown chat cattle immune to kines. brain endothelial cells may contribute 10 protective
518 ,u:no.'< Tilll[l! Hlckensial and chlamydia! diseases

irnmune responses. Antigens of E. mmi11a111i11m capable of d1a1 a control strategy oased on a vaccine would be lhe most
inducing the production or these C}1.okines b>• immuno- cost effective way of combatti ng the di~easc. 1•i
competem cells should be invesrigaced as potential vaccine The only commercially availahle procedure for hcan-
antigens. water immunization is an infeccion and treatment
It has been noted 1ha1 CD4• T-cell lines.from animals im- technique, the use of which is described more full~· below
munized ";th inacrivaied /J~ ruminant/um proliferate in the (see Control). It was developed more than ;;o years ago at
presence of a fraction of E. mmi11a11riw11 proteins between the Onderstepoon Veterinary lnstitllte. South Africa,;:, but
20 and 32 kOa.22'1 in addition£. rumi11amium proteins of 11. it has a number of drawbacks. The procedure is panicularly
12. 14 10 17 and 1910 23 kDa specifically induced prolifera- inappropriate for use in rural areas in Africa or the Carib-
tion ofPBMC from ca1rle immunized either by infection and bean b.ecause of the need for continual frozen storage
1rearmen1, on,1th inactiva1ed organisms.2411 These proteins facilities.247 and both production of the infective material
should have potemial as vaccine candidates. but none of used as antigen and the methods required for its applicarion
them has ~·et been purified 10 homogeneity for amino acid are expensi\'e. Despite being so ou1dated nothing better is
sequencing. so the ge1ws coding for them have not yet been currently available. so there has been a great deal of research
identified. In fact. relatively few antigens ha,·e been charac- over the last decade aimed at developing an improYed
terized 10 the ~1.em of having their genes sequenced. vaecine. These research developments are deal t wilh here.
The antigen of E. rumi11a111ium which is currently the The Senegal Isolate of£. rumina111ium has been attenu-
best ~haracterized is the immunodominant tvl.APl. a protein ated in cul cure, and animals immunized with the anenuated
of about 32 kDa:is 1 which va ries in size from isolate to iso- organisms are immune to homologous challenge. ' 1' Un for·
late. 19 The protein appears as a prominent band on Western tunately cross-protect1on between different isolates is far
blots of total E. rumi11a11ti11m proteins 2oi; and is located on from complete 101 and it would be necessary to attenuate a
the surface of the organism. 120 Theantigen is rt>cognJ.zed by number of genotypes to cover the entire amlgenic repertoire
immune antisera and by CD4 • T lymphocytes from immune in any panicular area. The fact, however, that some stocks
animals. 159 has been used 10 de\'elop serological assays have so far proved to be resistant 10 auenuation by i11 1rirro
(see Diagnosis below), and has also been rested as a poten- passage11 ' limits the possible usefulness of this method of
tial vaccine (see below). vaccination.
The mapl gene appears t0 be present as a single copy Inactivated organisms have been successfull~· used for
in the genome.4 1 lt is polymorphic ber.veen differem experimemal vaccinat ion against homologous chal-
E. rumina11tium isolates and was at one time advocated as a lenge.':ia. 148 This type of vaccine consists of inacti\-ated
marker for isolates from different geographical areas. 20 ' As non-viable protein from lissue culture material which can
more data have become available. however. it is now known be stored at ambient temperature. Two doses are needed.
that the ,'3.Jiants are not geographically constrained. 8 Elirli- and large amounts of tissue culture material have 10 be pre-
chia ca11is and £. chajfee11sis. rwo close relatives of E. rumi- pared. which makes production expensive. if production
11m11iw11 , have orthoiogous families of immunodom inant costs can be reduced b}' industrial-scale culture techniques
major surface antigens in the 28 to 30 kOa size range, 176· 21>2 such a 11accine may be practicable. At present the most im-
blll it is not kno,\,1 whether these genes are polymorphic be- portam problem to overcome is that, in a field situation
tween different isolates. where genotypes ha,~ng differing immunogenicities are in
A second immunodominant major outer membrane circulation. the protection levels have been disappointing. 143
protein gene Cmap2l has been identifiedt39 and the protein In one trial the administration of an inactivated vaccilie re·
evaluated for use in serological diagnostic assays. 38 The groES duced monality le\'els in nah·e sheep and goats from 78 to 54
and groELgenes have been sequenced.130 as well as numbers per cent and in naive cattle from 67 to 19 per cem. Results
of clones from genomic libraries which contain open reading such as these suggest chat die present fonnulaaons for inac-
frames (ORFs). some ha,~ng homologues in the public data· tivated vaccines are not \iable as a commercial proposition.
bases. and others currently being unique.20. 4 1• : i TI1e 1576 PotentiaUy. it should be feasible to include in an inacti·
kb genome of E. rumintmrium has been mappedll6 and whole vated vaccine as many imnnmotypes as would be required
genome sequencing is in progress.• Once complete this will for effec1i\·eness in a parlicular area. It would be necessary
be an invaluable resource for further gene discovery. to isolate organisms from any natural£. r11111i11anti1m1 infec-
An economic model for Lhe prevalence ofheartwater and tion appeari11g in animals immunized w1d1 an exisl.ing vac-
its impact under various far111ins system~ has been devel- cine 'cocktail'. The new isolares would then need to bi,
oped as a result of a 10-)•earstudy in Zimbabwe. 157 The esti· cultwed and added to the cocktail. prior 10 re•testing in the
mated ann ual losses in that counuy amounted 10 USS 5,6 field situation. This would be costly, but it could be a useful
million, with economic losses per animal in the commercial 'stopgap' measure u mil something better is developed.
production sector being 25 times greater than the losses in The fact that immun!zmion with Jnacth-ated E. rumi11a11-
the communal and Lraditional farming sectors, largely as a rht111 organisms is possible implies that the development of
result of the cost of acaricides. The conclusion was reached a subunit vaccine is feasible. In principle all that is required
 lleanw::ner 519

is 10 identify£. rnmi11ami11m genes which code for antigens "qth tl1e protein boost. but it is disappointing to note that the
which stimulate a protective T-cell response. Unfortunately 88 per cent protection level ob111ined using only the gene in a
there .ire-no known directed strategies for doing thi1-. DNA vaccinc Vl'CtOr in earlier experimems172 was not auaint)cl
A naked DXA vaccine expressing the :-,1,\.PI antigen of E. even with boosting. This casts doubt on the usefulness of the
rumi11/'111tium has been shown co proce<:r mice against a le- lll(lpl gene as a ,·accine candidate and underlines the non-
thal homologous challenge. 172 Sple11ocy1es from the inunu· n:pea1abill1~ which characterizes these results.
nized mice. but not from control mice, were sho"~1 to A cUs1urbing obser\'atlon common to all the vaccination
proli[erare in 11irro when treated \\1th recombinam MAP! experiments carried out in mice has been the unpredictable
protein or a protein e..xctact of whole E. rwninanriwn. The variability of t11e results. In addition, some e~-perimental
proliferating cells also secreted enhanced amounts of IFN-y immunii.ation results obtained in ruminantS have not been
and IL-2 as compared ro cells from control mice. There " 'as repeatable when the mouse model has been used. One ex-
one serious concern, however, in that the level of protection ample of the Ianer, as discussed above, was when ,-accinauon
was not reproducible, varying from 23 to 88 per cent in dif- \\1th inacti\'ated organisms produced good protectfon against
ferent experiments. homologous challenge in cattle. sheep and goats. but attempts
In an attempt to overcome the lack of a directed strategy 10 reproduce the nndings In mice were unsuccessful.i;.i Sim-
for identifying E. ncmi11a11ri11m gene" that code for protec- ilarly. encouraging protection IC'Vcls up to 88 per cent were ob-
tion-stimulating proteins. a strategy was developed 10 screen tained \,~th the fll(lpl gene in a O:S:A vaccine vector in miceHz
thewhole parasite genome, using a S<llmo11ellc1 vaccine deliv- whereas irnmwtlzation with denatured ;\lt\Pl protein appears
erysystem to idemifyprotectlvegenes.'12 A cloned segment of not to protect goats and sheep at all, despite very high antibody
E. mminanrium D!'-A. which contained five ORFs, protected titres l?eing present before challenge.z.m These observations
1-1 per cent of out bred mice against a leLhal homologous chal. suggest that the murine model may not be suitable fore\'aluat·
lenge. ln subsequent e.,-perimems. using the same deli\'ery ing putati,·e protective proteins or recombinant vaccine$
and challenge systems. each of the indi\idual ORFs was which are 10 be used against heanwacer in n1minanrs.
tesrcd in Balb/ C mice, \\'here thera were no survivors. and
C57BLl6J mice, where the results were not reproducible:'n
Clinical signs
In a comprehensive series of experiments two E. ru111i-
11amiwn genomic expression libraries have been isolated lnfccrcd domestic ruminants may manifest a wide range of
from a set of3,I unique clones which reacted with sera from clinicaI signs. The incubation period, course, severi~· and
hcamvater· immune sheep.20 These clones were sequenced, outcome of artificially induced disease arc Influenced b)· the
giving a total of abour 73 kbp or sequence, and were found 10 species, breed and age of animal affected, the roureofinfec-
include 33 complete and 23 panial ORFs. Selected ORF$ tion, chi! virulence of tile stock or T!. rumi11ami11m i1wolved.
were cloned inco an expression vector 10 g1mera1e rccpmbi- and the amoum and source of lnfec1ive material adminis-
nant proteins. which were evuluated for their ability to react tered. a. 166 2.14 Peracute. acme. subacute. and clinicallyinap-
\\;th heart\Vatcr-immunc amisera. Bacterial lysatcs con- parent forms of rhe disease occur.
caining these recombinam proteins were rested for their In the naturally acquired disease in a herd or r.ock. the
ability to stimulate proliferation of PBMC from cattle inunu- morbidity a11d mortality rares are inOuenced by the species.
nized by the infection and treatment mel11od. Many clones breed and age of the animals affected, the virulence of the£.
were recognized. although the response?, between different rr1111inmuiwn s1ock, the effecti\'eness of immunization, tick
immune animals differed even for the same recombinant. control and specific chemotherapy programmes applied on
Recombinants which had botb reacted whh heartwoter- the fann, and. in soma instances, the season.2 - · 55• 106• 231
immune antiscra and had stimulated proliferation of P13MC Death usually follows in animals which show clinical signs if
from immunized cattle were selecred 10 immunize mice, they are not specifically treated for heanwarer.
which were chen challenged. Disappoincingi~'. significant
protecrion (58 to 89 per cent survival) was only obtained for Cattle
cwo pools of recombinants, but not for any individual re- The incubation period in nacurall)' infected caule ranges from
combinant, e\'en those from the protecrive pools. 9 10 29 days. with an average of 18 days.~ Cows of B. 1aum$
Recently, a successful prime-boost strategy has been de- breeds. such as the Jersey. South Devon. Charolais, limousin
vised for inducing protective levels of CDS' I cells agninst the and Friesland, especially when in tl1e advanced srages of
liver 6tagC of the malaria parusitc.208 1\ similar $lTOtcgy has pregnancy. arc pnrdcularlr pro no to develop pcrncute heart·
been used in mice with the nu1pl gene and tl,e ;\L-\Pl protein warer. a fonn of the disease \\'hich mar sometime, also occur
of£ nm1i11anri11m.H 1 Priming was carried out with the gene in 6- to 18-month-old animals of different breeds. 10!<· 241; Per-
cloned in a DNA \''llCcine vector and then some of t,he animals acucelyaffec1ed animals die\\1thin a few hours after the initial
were boosted \,ilh the protein. and others were not, prior to dc\'elopmen1 of fe\·cr, either without any clinical signs having
virulent challenge. Protection levels were lncrea~d from J3 10 been manifested, or ha,ing shown tem11nal. paroxysmal
27 per cent without tile protein boost. and 53 10 6i per c:ent convulsions and marked respiracory discress.2· 166
520 ,ic 110, nmi,, Rickcttsial .ind :hlami'dial diseases

Acute heamvater. the most common form of lhe dis- acute fom1 of the di~ease. ~lost animals collapse suddenly
ease. mainly afl'ect~ cattle between tl1e ages of 3 and 18 and die after a few paroxys.mal convulsions without ha,ing
months. 11 is characterized by a fever of .:o •c or higher, been obsel'\"ed 10 be 111.234 Some, however, are anorectic. ap-
which \lsuaUy persists for three to six. da)'S. showing only pear dull. exhibit nys1agmus. bleat. and twitch their tails
small nurtuations before the bod)' temperature falls 10 continuously. The~· may show forced respiration. frequent
subnormal shonlybefore deach.2 ·2.l' While some observers urlnatlon and defaecatlon. chewing movemems and. when
consider that a mild mucold diarrhoea is a regular occur- recumbent. leg·pedalling. 1-H 218
renee.2 64 • 231 others regard it as an infrequent sign.z45 Cer- .~ is the case in can le, ncute heartwater is the most com-
tain breeds such as the Friesland. Jersev and Simmentnler, mon fonn of the di:,ease in sheep and goats. The majority of
and four- 10 eight-month,old animals of all breeds. appar- animals manifest nervous signs. bu1 these are generally less
ently de,•elop diarrh()ea most common!)'. A profuse. often pronounced th,m In cattle.:! lnirially. affected animals show
haemorrhagic. diarrhoea may be the most prominent clin- fever. a progrcssh·e unsteady gai1 and listlessness, and often
ical sign in some cases of h.earrwater, particularly in stand with their legs wide apart with the head lowered and
'2·• • ears drooping. They eventually become prostrate. assume a
Ierseys.•· ··•
During the later stages of acute heanwater. nervous position of larcral recumbency and show intermittent leg-
signs occur which range from mild incoordination to pro- pedalling. chewing movements. opistho1onus, licking of the
nounce-cl con\'ulsions. 4 The animals are hypersensitive lips. and nystagmus/
when handled C)r exposed 10 sudden noise or hright light. Black-headed Persian sheep possess fiome natural resis·
Slight tapping ,,~th a finger on the forehead of the anjmal tance 10 heanwater, and lambs and kids of all breeds under
often evokes an e.xaggerated blinking reflex. They fre· one week of age have a degree of innate resistance. Such
quently show a peculiar high step?lng gait that is usually animals. as well as those which are partiallr immune. may
more pronounced in the front limbs. Calves may wander develop 1he mild fonn of heartwa1er and. apart from fever.
around aimlessly and walk Into fences. and some, pre,1- no clinical ~igns are generally manifested.2 • 234
ously unaccustomed to handling by humans. may be
approached with eaM.>:. Animals ma)' stand with their heads \.Yild unguJates
held low. make constant chewing movement,, and pu~h CU11ical sign, of heanwater in susceptibl1.1 wild ungulates
against objects. in the later stages they often fall down sud- [fable 40.2) ha\'e not been well studied. but are generally
denly, assume a position of lateral recumbenC)'· and show similar Lo those reported in domestic ruminants affected
opis1J1otonus and either have frequent bouts of leg- 1·.i1h the acute form of the disease.9 7• ,~. 261
pedalling movements or the legs may be extended and stiff.
in most cases the animals weaken rapidly and death usu - Laboratory mice
ally follows soon after the commencement of a convulsive The incuba1Jon period in mice infected lmravenousl)' \\ilh
arrack. the \\'elg~vonden stock of E. r11111i11t1111ium ranges from 10 to
The subacute fonn of heanwater is characterized by a 14 day,. 197 ~o febrile reaction has been no1ed ln mice ex·
fever which may remain high for !Odays orlonger. The clin- perimentally i nfl!cted with any of the stocks LO which ther
ical signs are simllar 10 those described for the acute fonn of are su~ceptible. 1::l<i Twenty-four hours prior 10 death af-
heanwater. but less pronounced. During the course of the fected arumafa are lethargic, and develop tach)'pnoea and a
illnp,;,s :mimal~ mny di<' sndrlenly. llr gn\duillly retover ruffled coat lncoordinatio11 i~ nrr"~ion:tll)' ~een. 1"
within a few days. and dcmh frequently results from compli-
c:nions such as h}11osuuic pncumonia. 2
P ath ology
The preval1311ce or Lhe clinically inapparem form of heart.
\\'lller is difficult to determine under field conditions. Apart A slight reduction in haemoglobin and haematocrlt \'alues is
from fen:r. apatl1y and slight tachypnoea, the animals appear observed in sheep. goats and ca.Ives. The reductions coincide
10 be normal and most recoven,~thin a few days.55 Calves less with. or s1an shortly afLer. 1.he commencement of fe1·er reac·
than three weeks of nge.2..11 cattle infected ,dth a stock of low tion and continue throughout the course of the disease. but
,~rulcnce, 166 and infected but partially immune animals2 are neither ,aluc reaches critically low le,·els. t .e. 1' 8· w The
particularly prone to develop 1his mild fonn of the disease. anaemia, which is not clinkally discernible. is usuall)' nor-
moq-tic and normochrornic.242 although a microtytic, hypo-
Sheep and goats chromic anaemia h~, also been reponed in goers. 1 The
The incubarion period in sheep and goats ill()Culated intra- pathogenesis of the anaemia is 1101 known.
venously wilh 10 ml or infected blood ,·aries from 5 to 35 Du ring the cour,e of the disease lhe colour of tl1e blood
days (a,·erage 9 to 10 days). and that of naturally infected plasma of calves. sheep and goats may change progressively
animals from i 10 35 days (average 14 days).?, 23'1 10 a dark orange colour.a 100 asa result of increased le,·els of
Exotic goat breeds, such as the Angora. and two- to six- bilirubin and/or its conjugates.15-1 In cah·es the bilirubin-
month•old Boer goats are commonly affected br the per- aemia appears 10 be associated wilh anorexia ralher than
 lfearrwater 52 l

with haemolysis or liver damage during 1he initial scages of increase while the haematocrit dropped progressively.2~~
1he disease.242 Lesions in cattle, sheep and goats an: very similar and
Mild leukopenia, mainly resulting from a decrease in 1he differ only in their severiry and frequency. 2• 195· 2 18• 331 Gross
number ofneurrophils, develops ln calves and goats prior to lesions may be inconspicuous or even virtually absent in
1he onset of fever and persists throughout the course of the fatal cases. 195
acute form or the disease. 111 An eoslnopenla develops be- Severe hydropericardium (Figure 40.10) and hydrotho-
fore or during 1he onset of the febrile reaction, and is accom- rax (Figure 40.11), and in some cases a degree of ascites. are
panied by a lymphocytosis which commences shortly after striking changes in most fatal cases of the disease. However.
the onset of the febrile reaction. 1 53· i.a, 242 h}'dropericardium is usually more pronounced in sheep and
The total serum protein content is reduced in calves. The goats than in carlle. 108 The transud,He is a transparent w
commencemem of this reduction coincides with the start of slighlly turbid. light yellow Ou id which may coagulate on ex-
the fever reaction and persists throughout the acute s tage of posure co air. Several litres of ir may be present i11the thorax
thedisease.242 High concentrations of albumin and globulin in cattle, while in sheep up w 500 ml, and in goats rarely
occur in the oedematous fluid which accumulates in body more than 20 ml. may be presem. 218
cavities. 24 ~ A moderate to severe oedema of the lungs occurs in most
Blood urea nitrogen58 • SR and creatimne elevations have anjmaJs that die of the disease (Figure 40.12). bm it is par-
been detected in the blood shortly after the onset of the fe. ticularly severe in animals which have suffered (rom the per-
brile reaction in sheep, whereas in calves these are not acute or acute form.2''5 Frothy oedematous fluid oozes from
higher than the upper normal levels. but they do suggest the cut surface of the lungs. The trachea and bronchi are
some interference in glomerular function durjng the acute often filled \\1th a frothy serous foam occasionally ;accompa-
scage of the disease. 242 nied by a fibrinous coagulum. a nd their mucous mem-
Blood glucose levels rise terminally in the disease and branes are often congested and contain petechiae and
may cause glucosuria in sheep and goats.58· sa. 1 t,, ecchymoses. The media~1inum and bronchial lymph nodes
Terminal increases in blood pyn1,·a1e and lactate levels, mny also be oedematous. A slight io moderatesplenornegaly
and a simultaneous drop in bicarbonate. have been re- :resulting from congestion and lymphoid hyperplasia is
ported in sheep and goatsf.S, it, During the acute stage of found in mos1 anlmals. but in sheep and goats it is often not
the disease in calves respirarocy alkalosisand hypoxia occur, as severe as in cattle.
as evidenced by a lowered oxygen tension (pO:J.2..1z Sheep Oedema of the brain commonly occurs in animals suffer,
infected with the Ball 3 srockshow retention of carbon diox- ing from the peracute and acure forms ofheamvater. 188· ?,ts
ide (increased pCO,.) resulting in respiramry acidosis on the Occasionally the entire brain, but particularly the grri of rhe
rwelf1h day pos1-lnfecrion. 1n addition. there is a reduced cerebrum. is prominently swollen, and this results in a par-
oxygen diffusion which gives rise to hypoxia (101,·ered p02 ) rial herniation of the cerebellum through the foramen mag-
that persisis uncll death. 178 In other experimenrs in sheep num. Most fa1al cases show varied degrees of co:-igestion
changes in blood gas included a decline in arterial p02 com- and oedema of the meninges. TI1e choroid plexus is swollen
bined with a respiratory alka losis. Although the sheep be- and dull greyish in appearance. In some animals, petechiae.
came hypoxaemic, blood-gas changes associated with ecchymoses and sometimes suggilations are evident in rhe
respiratory fallure were not observed.249 brain substance. panicularl)• of the midbrain. brain stem
Increases in both the respiratory rate and tidal volume a11d cerebellum 188 Microscopic lesions in the brain ;ue
result In increased minme volume and a slmuhaneous In- characterized b)' changes compatible with oedema. such as
crease in \'entilatory equivalent in sheep.178 The tendency widened perivascular spaces wmch in tissue secuons some-
towards Increases in both physiological and alveolar dead times contain material representative of oedematous fluid
space may be associated 1~ith lung oedema, and could ac- or prolein droplets, swollen and often necrotic astrocytes,
count for che moderate reduction in pO~ which occurs dur- swoUen axons, and microcavitarions, pa.nicularly in the
ing the acute stage of the disease. 242 white matter_Occasionally small haemorrhages in the neu-
In sheep, changes in blood-cloning beha\iour, blood ropil , scant cellular perlvascular accumulation of mainly
calcium and protein levels. haemacocril. and white-cell macrophages and lymphocytes. fibrinoid vasculiris. fibrin-
counis occur. A marked decline in thrombocyte count was ous choroiditis and multifocal glial nodules (particularly in
recorded during the acute stage of the disease, associated 1he white matter around small blood vessels) are eucoun-
with incceases in both prothrombin time and actlvated par- tercd.188 \ Vide~prnad status spongiosus .o f varying-severity,
tial thromboplastin time. Fibrinogen increased, wl1lle there 1hat main!)' affectS the larger tracts of white matter in d1e
was no detectable increase in fibrlnogen degradation prod- brain, may be found in animals that have been recumbent
ucts and wtalserurn protein, albumin and globulin dropped for days. and particularly in those tl1at have been specifically
very sharply. Total caldum showed a progressive drop bur but unsuccessfully treated a rew days before death. 193
ionized calcium rose initially and was then followed by a ter- Nephritis of varying degree, sometimes accompanied by
minal decline. The total leukocyte coumshowed a terminal pen.renal oedema and petechiae in the renal cortex. occurs
522 ~,,10, rnru: Rickcusial and :hlami·dlru disease$

Figure 40.10 Hydroptlricardium Figure 40.12 Lu~g oedema

Figure 40.11 Hydrothorax Figure 40.13 Seve,e cecema of abomasal folds

in most fatal cases ofheartwater. 195 Renal tubular epithelial ence of a diffuse catarrhal enteritis of \'8T)~ng intensity. In
cens are swollen and occasional tubules may contain hya- olhers. particularly Jersey cattle breeds, severe diffuse
line cast.S. Severe nephritis is particularly obvious in Angora emerorrhagla, together with intense congestion of the
goats that have been unsuccessfully treated for 1:.: no11irum- mucosa of particularly the small intestine, may be the most
1i11m after the first dar of lhe febrile reaclion and killed in promineni of all the lesions.
exiremis four to seven days after the onsel of the iebrile re- Many lymph nodes are moderarely swollen. and cut sur-
action, or which have rued three to £our da)'S after the finn faces are oedematous and congested, and maycomain ~mall
e,idence of fever. 1!•~ haemorrhages. 2 Petechiae or even larger haemorrhages mai•
Congestion and/or oedema of the abomnsal folds (Fig· be present in serous and/or mucous membranes of tissues or
ure 40. t3) are regular findings in cattle but are not a$ com- organs such as the epi- ond cndocardium, urinal')' bladder,
mon in sheep and goats. Petechiae may also be present in vagina and conjunct ivas.
the mucosa of the abomasum. Cattle which have sho"~, Variable numbers of £. rwni11a111i11m colonies are
some clinical evidence of diarrhoea often reveal the pres· discernible in the cytoplasm of endmhelial cells. and
 Heartwater 523

par1icularly those or the brain /Figures 40.3 and 40.4). lungs pletc. Flxed or unfixed smears are still suitable for staining
and kidneys but, as a general rule, they are difficuh to find in and diagnostic purposes for at least one month after they
haemato:-.·ylin and eosin stained tissue sections. have boen prepared. ~3'1
The lesions in game animals that die oi hearrwa1er are Colonies of E. r11mi11anriwn are generally easy m find in
very similar to those reported in domestic ruminants. 1.:i. 1~5 smears of untreated cases of heanwater, but their numbers
In mice Infected with the Welgevonden stock the lesions may val"}"\lidely from one animal 10 another. Or1.<anisms can
closely resemble those in catcle, sheep and goats that have often still be demonstrated in brain smears prepared from
died from heanwater. 197 1n tissue sections endothelial cells or animals in an ad\'llnced s1are of putrefaction. The examina-
the lungs contain the highest concentration of organisms. fol· 1ion ofbrnin biopsies in live animals for the confirmation of
lowed by those of the myocardium. Organisms are only rarely a diagnosis ofheanw,Her is useful in experimental animals.
detected in capillaries in tissue sections oithe brain of mice. but is not practical under field condilions." 2• 219 • 239
Transmission electron microscopical studies of the lung For the histopatholog!cal diagnosis ofheanwnter in ru-
lesions in sheep, goats and mice reveal the presence of minants by the examination of [issue sections. the kidneys
minor cytopathic changes in endothelial cells. :\pan from and brain are the preferred organ,., E. rumi11anti11111 particu-
mild swelling of mitochondria and endoplasmic reticulum. larly being $Ought in endothelial cells or the renal glomeruH
no other changes occur in most parasitized alveolar or capillaries uf the grey mauer of the cerebral conex.63 Be·
endothelial cells. Non-parasilized endothelial cells are cause of the low numbers of organisms in these tissue sec-
sometimes swollen, or even necrotic. and are separated tfons. ii Is a time-consuming method of confirming a
from their basement membranes. Oedema or blood vessel diagnosis. and may even yield an inconclusive result. Vari-
walls is infrequently seen. 197, I$& ous staining methods may be applied to demonstrate F.. ru·
111i11anti11111 in tissue sections.46 but those Incorporating
toluidine blue or Gicrnsa are preferred. 193
Diagnosis
The confirmation of a diagnosis of heanwater is often
The traditional method of making a post-mortem diagnc1$is difficult in animals 1ha1 ha\·e been treated. A small number
ofheartwa,er is the demonstration by light microscopy of E. or treated animals. panicularly those that have received in·
ruminamium in the cytoplasm of endothelial ceUs of blood adequale chemotherapy. do not recover full). remain re-
vessels in stained smears of brain tissue.200 Organisms may cumbent forda~-s. and eventually have. for humane reasons,
also be found in tissue .sectious of the brain, or other organs to be destroyed. Although most of these cases reveal a status
such as the kidneys. spongiosus ofvruying degrees histologically in the brains. 1~3
Brain smears are prepared in such a way that segments of the typical macroscopical lesions or hcamva,er are usually
capillaries remain more or less intact and can be examined absent at necropsy. and identification of E. n11111'111111ri11m in
after staining.A small piece (less than 5 >< 5 x S mm) of hippo· brain smears 48 to 60 hours after an animal has been treated
campus or cerebral grey maneris obtained either by opening is often difficult. ln these cases brain smears should be
the sb.-ull or by scooping some brain material through the fo. stained for 30 minutes in a 10 per cent aqueous solution of
ramen magnum.= Smears a(e made by crushing the sample Giemsa. The electron-dense hodies arc more severely af.
between two microscope glass slides umil the tissue has a fec.1ed b)• chemotherapy than are the reticulated bodies, and
soft. pasty consistency. The material is then collected at the the organisms are poorly delineated and appear to fuse.
end of onr of 1h<' slidr~, whirh i~ hrld firmly in a hori10111:;I making h riiflk11h 1n di~1ing11ish 1hem from phagn~omrs
position. The other slide, angled at abom 45 •, is used 10 make and chromatin in endothelial cells, and from groups of
the smear by drawing the tissue along the horizontal slide. blood platelet, and mas1 cell granules. 193
While making the smear it i.s preferable 10 lift the angled slide The first serological test for sun•eys for 6. mmi11a11ti11111
slightly, about every 10 mm, so !hat the smear has alternating was the indirect fluorescent antibody test (IFATJ.76 The tar·
th.ick and thin areas. This procedure stretches the capillaries gel antigens were peritoneal macrophages from mice in·
linearly and iacililates their microscopical detection. Smears fected with the Kumm stock. 87 or goat neutrophils or cells
should be air-dried before staining. zoo. :!J.t from endothelial cell cultures. 150 r\Jt enzyme-Jinked immu-
Various stains may be used to demonstrate heamvater nosorbent assay (ELISA} was also developed}"° In both of
organisms, such as Giemsa or CAM"s Quick Stain (C.A. these 1es1s cross-reactions with antibodies agains1 related
Milsch (Pt)·) Ltd.J.:!'.l-1 but Giemsa is the method of choice u11- lilirlichl<1 and A11aplas111a spp. occur, resulting In the com-
less large numben; of organisms are ptesent. ror G.i emsa mon occurrence of false poslllve resuhs.:;o· 9o. 109 · ' "
siaining, smears are fixed for about one minute in absolute A com)X"lilive El.IS,\ using a monoclonal anti-MAP I an-
me1hanol or ethanol. and a re then stained for eithcr30 min- tibodr was the next de\'elopment 120 bu1 this too gave false
utes in a lO per c.ent aqueous solution of Giemsa or IO min- positive reaetions 18 with related Ehrlich in spp. fl is now
utes in a SO per cem solution. CA1'l's Quick Stain ~·ields known that the reason for this problem is the existence of
acceptable results where a h igh concenuation of organisms homologous families of immunodom inam outer mem-
are present and requires only two 10 three minutes to com- brane proteins in Ehrliclzia and J\11aplasma spp} 76 · 1; 9 . 26l!
524 serTI(lx nmn: Rickcusial and chlamydi:ll diseases

In an attempt to overcome the _problem the compe1ifive to be Ti. ruminan,ium. There a re two oddities among them.
ELISA cest has been modified by the use of a fragment of however. the Oma1jenne genotype. which does 1101 appear
1'1AP1. designated ivlAP 113, in an indirect ELISA fonna1. 2S.2 10 be pathogenic co goats,J I and the Pretoria :S:onh geno-
This has been shown to have a higher specificiry for E. rumi- rype, which has been found in dog.s.; The infectivity for
mwriwn than any other serological test. and it d,oes 1101 ntminams of this la11er organism is not yet known. All eight
cross-reacc \\ith amlboctlcs against lhree orher orgarusms In of these genotypes give a positive reaction with the pCS20
the order Rickeusiales which commonly infect ruminants. probe and. because of the uncertaincy which chis may in-
namely A11aplasml/ bovis. Atltlplasma ovi11a and Anaplasma duce. It Is routine to use the !6S ribosomal R;'IA probes as
phagocyrop/zi la.= Note 1ha1 these three organisms were well as the pCS20 probe on animals which are being exam-
classified as EhrUchia spp. before the recem reclassifica- ined to obtain permits for imponation into :ion-endemic
rion .92 The MAP 1B ELISA test does, however. detect ami- heartwater areas.
bod1es to other Elzrlichia spp.. notabl)' £. cal/is, E. The 111<1pl gene has also been used asa 1arge1 Tor molecu-
chaf/eensis, an unidentified Ehrlicllia sp. infecting white- lar genetic diagnosis of E. rwnitiamiwn. 9 Because of the
tailed deer (Odocoileus virgi11ia1111~) In the south-eastern extensive polymorphism of this gone it has been used co
USA,125 and the organism currently known as £. r111ni,u111- characterize different genorypes of the parasi1e.8 There is no
tium (Omatjenne). 11 evidence of any immunogeneric or geographic clustering
Apart from this caveat 1he MAP I B ELISA 1es1 works well among the variants of the gene, and other gene targets need
in sheep and goats.li1', 152 i n canle. however, antibody levels to be soughrin orderto better characterize stocks ofE. rumi-
against £ rumimmri1m1 can be very low in heanwatet en- 11mui11111 which are to be incorporated imo vaccines for field
demic are.as. even in caule that have been vaccinated or are use.
under conrinuous natural challenge by infected ticks.67 · iw
/\ntibody levels decline in cattle to such an extent that they Differential diagnosis
become seronegative 14 to 33 weeks after initial exposure
owing 10 a down regularion ofMAPl-spccific antibody re- Nervous signs occur in most animals suffering from heart-
sponses posi-recovery.210 Similar results from cattle have water and they must be distinguished from a wide range of
been reporccd using the IFA test00 and have been confirmed infectious and non-infectious conditions 1ha1 manifest
°
by immw1oblo1ting. 67• 21 Care must therefore be taken similar signs.
when using any serological test in cattle, especially if the In can le nervous signs may be caused by other infections
animals are being tested in order 10 decide whether ii is safe such as rabies, the nervous form of malignant catarrhal
10 move !hem co a non-endemic area, since it is known that fever, ce:ebral babesiosis, cerebral thcileriosis, chlamydio-
tl1ey may be tick. infective subclinical carrier.s of l1eart- sis. meningitis and encephalitis caused by various bacteria,
water.12 especially Strepcococcus spp .. Pasteurella spp .. Arcr111obac1e-
There has been a molecular genetic revolution in diag- rium 11yogelles. and flaemophilus spp. ln sheep and goats
nostic techniques over the last 10 years and it has become meningi,is and encephalitis are caused by a wide range of
possible to detect the presence of minute amoums of para- bacteria, and abscessation of the hypophysis occurs. par-
site genetic material amid a huge excess of hos1 ONA. The ticularly in goats.
critical factors which al.low this are the specificity of Oli!/\- In southern Africa neivous signs in cattle may be th.e result
nN I\ hybridi7a~ion probes. 1he sensiti\~ry which follows on of poisoning \\~lh pl:inrs suc:h as Alhizin 11e~irr1lnr. Al/Jizia
target amplification by the pol~'lnerase chain reaction (PCR) 1anganyice11sis. Cynemclmm spp., Euphorbia mauritanica.
and the ready availability of public darabanks of generic in- Sarcosremmtl viminale and Cynodon dacry/011, and fungi,
formation. For E. rumina111i11m diagnosis three families of especially Aspergillris clavatl/$ and Clauiceps paspaii. 126
probes are in use, targeting the pCS20 gene. the small sub- Other poisons which may also induce nervous signs are
unit ribosomal ~Ni\ (16S RNA) gene, and the mapl gene. pesticides (e.g. chlorinated hydrocarbons and organic
The pCS20 gene is an open reading frame of unknown phosphates) and heavy metals (such as iead and mer-
function254 which appears co be specific for E. rumi11an- cury).59· 220, 221 In sheep and goats plant poisoning (espe-
limn. giving no cross-reactions with other Ehrlic/zia spp. 9 Tt cially by Cyna11chw11 spp., Euphorbia mauri1anica,
is the most sensitive or 1he pr<>bes for E. ruminanrium detec- Sarcostemma viminale and Cynodon dact_v/011). and hea\'y
tion and has been widely used to detect E. ruminanrium in metal and pesticide poisening. induce nervous signs sim-
domestic animals, \\'ild game, and ticks, 10, HJ lH 18t 182.184 ilar to 1hose seen in caule.
The l 6S RNA gene has been ,·el)· \\'idely used as a caxo- l.ung oedemR. hydropericardium and hydrothora~ are
oomic and phylogenetic 1001 for classifying bacieria. 177 common necropsy findings in cattle, sheep and goars that
There are eight differem 16S ribosomal R,'\iA genotypes cur- have died of heam,•ater but ti1ey are also regular findings in
reml.y classified as E. ruminanriwn (Figure 40.2) and six of 1he case of gousiekte ('quick dise3$e') caused l>y 1he inges-
these are known 10 cause virulent heartwacer in ruminants. tion of the rubiaceous plan rs Pachys1igma pygmae11m, Pach-
which suggests that they are what is classicaily understood ystigma 1han111us. Pachystignut /a1ifollus, Fadogia homblei,
 Heamvarn, 525

Pal'eua har/Jorii and Paireua scl111mo11nia11a. 1lb Lung cated for marginal areas where Amblyomma spp. were only
oedema is also found in sheep that have succumbed 10 fbund occasionally. The l(>gist:ics and expense of ~uch opera-
pulpy kidney disease or bluetongue. and in cattle suffering tions are formidable if they are applied over large ar<'as. and
from Corridor disease or East Coast fever acaricide resistance is a widespread problem.235 The main
The diagnosis in cattle, particularly Jersey~. which have dis-advantage. howei:er, is that animals may lose all immu-
died from the form of the disease in which intestinal nity to Lick-borne diseases because of the lack of a natural
changes are the most prominent sho1.lld be differentiated challenge. Any breakdo\\1, of the intensive control regimen
from poisonings by. for example, heavy metals (arsenic, d1en results in heavy losses from heartwater and other tick·
mercury), plants (HOml'ria, Moraea, Urgi11ea, Omithoglos· borne diseases.
sum and Omirliogalum spp.) and organophosphates. Strategic t.ick control implies the control of lick numbers
Chlamydophila peccrwn and E. rumi1uu11i11m may be so that naiural infection of livestock occurs and high levels
difficult to dilferemiate morphologically in brain smears or ofimmuni!) are maintained. The aim is to achieve an epide·
tissue sections made from animals whic/1 have died of the miologically stable situation with respect 10 heartwater by
respecti,·e diseases. Both organisms arc pleomorphic, and the regulation of the numbers of ticks present so as 10 pre-
elementary, intermediate and reticulated bodies are found vent the debilitating effects of ~evere tick infestations. It is
in both of their multiplication cycles. As a rule. the usually recommended that animals should be dipped only if
organisms wi tbin a colony of B. rumimmriwn 3re of the they are carrying, on average, more than JO adult Amb/y.
same morphological form - either all elem£intary bodies omma ticks each. The animals should be monitored weekly
or all reticulated bodies. 187 A colon}' of Ch/11111.l'dophiln, in summer and autumn immediately after or during periods
however, may contain e lementary bodies {0,24 to 0,4 µmin of good rains. and every two to three weeks in winter. h has
diameter) and reticulated bodies (0.6 to 1.5 µm in dia- been sho\\11 that thi::. approach can lead to endemic {or en·
merer).16 The his1opa1hological lesions in the brain of the wotic) stability, even when the strategy is somewhat errati·
two diseases usually differ markedly. Whereas those caJly applied.223 Economic studies have demonstrated that
caused by C. pecorwn are generally cnaracterized by a strategic tick control is both a more economical and a more
moderate 10 se\·ere muhifocal or diffuse lymphocytic men- practical option for limiting losses from heanwat~r and
lngoencephalitis. vasculitis and. in some inscances. also b)• other tick-borne diseases.1~3 In fact, tl1e counter-intuitive
rhromhosis llnd smal l fol'i elf ne"rn,i~ of th!' neurnr,11, ohsNwHion h~s hPen mRde rh~t direct lick-borne disea~<'
oedema of varying severity is usuallr the only lesion that is losses increa~e with increasing use of acaricldes. 5 '
encountered in lilt' brain of animals that have died of
heanwater. Antibiotic treatment
While the characteristic colonies of E. ruminantit<m arc Several drugs have been used 10 treat animah suffering from
caspo detect. ifpresem in sufficient numbers. it is not pos- heanwarer, but the ietracyclines, especially oxytetracycline,
sible 10 distinguish between different species of Elrrlichia by are the most wide!~· used.240 During the early 1950s the
light microscopy. If one suspects the presence of a specie.~ minimum recommende.d therapeutic dose was 5 mg/kg live
closely related 10 E. ruminamium for 01her reasons. such as weight. given eilher as a single dose or in a divided dose with
the clinical plciure or geographical origin of the case, then a 24-hour i.mervat.107· 259 Some years later it became appar·
the 01,ly way to characterize the parasite is to use molecular ent that rhis dosage rate was no longer effective and a r:ue of
genetic methods. IO 10 20 mg/kg was recommcndi:-.d. Despite this chc:c is ap·
parendy no resistance of E. r11111i11ar11i1111110 tetracyclines.
Short-acting formulations of Oll.ytetracycline are most
Control
common ly used at a dosage rate of IO. to 20 mg/kg body
Four methods are available for the control of he:mwater. weight, either administered intramuscularly as a single
tick control. antibiotic treatment of clinical cases, prophy- dose, or half the calculated dose is given intra,·enou$ly and
lactic use of antibiotics. and immunizacion. the other half mtramuscularly. This treatment is usually re-
peated 24 hours later. A long-acting oi.')'letracyclinc prepa·
Tick control ration has been shown 10 be equally effective.z:i.;
Tick control has been advocaced as a means of controlling Do:-.ycycline has been used successfully at a dose rate of
heanwater from very earl~· days.2 • 7 1. 222 Even after the in· 2 mg/kg body weight in the treatment of experimentall}' in·
fecreci' blood-based vaccine was developed. tick control duced heartwatcr in sheep.llij Th.is forrnulatlon is more
was still advocated as a supplementary or alternative lipid-soluble and has a longer serum half-life dmn has oxy-
means of control.' 08 • 170· 217 tetracycline, and $mailer therapeutic doses with longer in-
Tick conrrol can be either intensive or strategic (see tervals between treatments can thus be used.
Chapter I: Vectors: Ticks) 33 but intensive t:ick control has When treatment with tetracycline is instimted during the
largely fallen into disuse. The objective was 10 control all incubation period of heamvater in cattle (before 2pproi.i-
stages of ticks throughout the year and it was a system advo- 111ately eight days after infection) the course of the disease is
526 »1:no~ nuu.c: Rjcken.sial and chlamydia! diseases

usually altered: animals usually develop no fe,·er or other advocated. in the form of a tabler imptamed behind the ear.
clinical signs. except that, in a few. a low-grade transient fe. Jn practice this is most often used as pan of an immuniza.
brile response will result. In these animals no. or at best only tion-and-ucatment regimen. 131 The dosage is critical so the
a partial, lmmunitydcvelops.ea. 1~0 Trea1mem of sheep and body weight or 1he treated animals must be kno1,11. since an
soats during the incubation period may also give rise ro a inadequate amou111 ofnmibiotic will not prevent fatal hean-
delayed febrile reaction ( it may be as long a~ 20 w 25 days water.
after infection compared to the more usual 9 10 10 days Jn
untreated animals). and the reaction may be so severe that Immunization
additional treatment is required. The economic need for a control system based on vaccina-
A wide variety or sul phonomides has been used in the tion. and the steps being taken 10 develop a suitable ,·accine,
treatmenr of heanwater;3 · 106, 163 in facr. 1he creaunem of are described above (see Pathogenesis).
hearrwater \\ith uleron by :'ieitz in 19~0 ,va.s the first use or The only '\'accine· curre111Jy commercially available Is a
any sulphonomide against a ricke?tt$ial infection. Sulphadi- cryopresen·ed preparation of blood from a «heep Infected
midine gives good results in severe reactions to heam,:ater with virulem £ n11nJna11rium organisms of the Ball 3 iso-
that persist in spite or repeated ox11e1r..icycline treat· late.32 The blood is injecwd imravenously in animals to be
mcnts248 bm because tetracyclines are normally so effe1;clve immunized, the rectal temperature is monitored dally. and
sulphonornides are not common!>· used. RifamyC'in has also amibimic treatment is administered at the proper time. The
been used to rreat E. mmi11anri11111 infections in mice and infective blood must be preserved on dry ice or in liquid ni-
sheep at a dosage rate of0.2 mgtkg.1• 8 trogen and thawed shortly before inoculation. and the
Supportive therapy in clinical cases ofheanwater is often whole procedure must be supervised by crained s1aff. The
inefficient because of the poor unders1andingof the p11tho- duration of Immunity is uncertain, and because li\'e organ-
genesls of the disease. Various an1i-infiamma1ory agents isms are involved the procedure cannot be used in non-
have been used bur the onl)' published informiltion con· endemic areas. The procedure is. however. successfully
cerns the use of dimethylsulphoxide39 and prednisolone. 201 used to protect susceptible animals against rhe disease.
Calves develop halitosis and dark yellowish-brown urine as especially when they are first introduced Imo endemic
a consequence of the use of dlme1hylsulphoxide but no real areas. or if they are particularly valuable.
improvement in the recovery rates occurs. 10• Betame1ha- The Rall 3 stock was originally isolated in the Limpopo
sone can be given at a dose of O. l mg/kg bod~• weight and Province of South Africa 105 and was cl1osen as the vaccine
treaunem c:an be repeated after 12 hours if necessary. stock because it produces an early temperature rise several
In addition tQ specific therapy. diuretics have been ad- days before any other $erious clinical sign~ appear. This
ministered to counteract oedema. and In the past furo· makes it relatively easy 10 decide when ro treat. Some stocks.
semide was used for this purpose. 156 Howe,·er. 1.he drastic such as the highly ,irulenr Welgevonden isolate. can cause
reduction in blood volume which occurs as a consequence death very shortly ar1er a rapid temperature rbt' and are
of heartwa1er ls exacerbated by the use oi diuretics. and the therefore not usable for the 111rec1Jon and trea1me111 method
adminis1ra1lon of furosemide as a supponive 1reatmem is of immu11iza1ion. Unfortunately the Ball 3 \'accine does not
no longer recommended. 211 protect against all the isolates which circutaw in the field. ll
confers only ilmited protection against challenge \\~th the
Prophylactic use of antibiotics Wclgcvondcn91 and Senegal 121 stocks. l 1. is 1101ed above thot
Routine oxytetracycline injections may be u~ed to protect the Welgevonden isolate offers a wider specm1m of cross-
susceptible animals against heanwarer when they are intro- protection than other stocks, but its virulence makes it diffi-
duced imo a n endemic area. 100· 201 In goa1.s it is ad\'oCated cult ro control and therefore unsui1ablc for an infection and
lhat shon-acting OX)'letracyclines be administered at a dos- treatment immunization procedure.
age rare of3 mg/kg body weight on days 10.20. 30. 45 and 60 lmmuniz.atlon of pregnam cows, especially if their off-
after their imroducdon, and that the animals should not be spring are valuable. is not ad,~sable as it ma) result in the
dipped until day 60. coo Injections or a long-acting o>.')'letra- peracme form of the di;,ea,e. orabonion.2{" Uncomplicated
cycline in ca11le are sufficient to protect them from conuacl- immunization of ewes that were two to four momhs preg-
ing heartw,uer. while -at the same time allowing them to nant has been reponed.J 2 bu1 in such instances II is ,·et")' im-
develop a natural immuniry.201 • 202 The dosage is 10 10 20 poriam lo monitor the rectal temperatures c~ref1.1lly and to
mg/kg body weight given on days 7. 14 and 21. or days 7, 12 treat the animals al tho tlrst rise In tempcramre.
and 17, or even on days7 and 14. Tiuianimalssbould be kepl Two 10 five per cem or more of animals. especially cattle
under close scrutiny and given appropriate 1reatmem if they that suppo~edl) should have lost their age-related resis-
do develop oven disease. The succes, of this regimen de- 1ance, fail 10 reacl after immunization. 82 This could be due
83
pends upon the animals becoming natural!)' infected during LO persiswnce of the innate age resistance in some animals
the time that they are protected by the drug. or 10 high conglutinin levels. Reinununization of non-reac·
Slow-release treatment \\~th doxycycline has also been tors sometimes produces the desired effect. bm lhe risk or
 Heartwater 52i

anaphylactic shock Increases with repeated inoculation or determined O\"l?r a period or about six days for both •,mall
blood or tick suspensions.ii. 2 3 1 stock and cattle. On the first da)rof the temperature reaction
In the absence of periodic stimulation or the immune the rectal temperature will usually be I •c or more above the
system. resultlng fro111 the bites of infected ticks. the clura- average.Thi.s can be expected 9 to 14 days after inoculation
rion ofimmunity after immunization \'aries great!) between irl the case of small stock and 14- to 18 dars in cartle. ln ex-
different domestic animal species, and also perhaps be- ceptional case. it may be a$ early as seven days i:i small
tween individuals \\ithin species. In sheep the Immunity stock and as long a~ 25 to 30 days in cattle. A rise in the early
may w,me after six momhs, 77• 1~ bm in some cases it may morning rectal temperature 10 abo\,e 39.5 •c in caule and
remain sufficient 10 protect animals against a fatal outcome goats, and 40 •c in sheep. is usually regarded as an ir:imuni-
for at lea.~t four years. 1M ution reaction.217 The general habitus of all animal~
The duration ofimmunlcy in goats following immuniza- should. in addition 10 the temperamre reaction. be dosely
tion is poorly documented. In Angora goats the degree of monitored after vaccination.
immunity seems to depend largely on the. time at which the It is unnecessary to treat as earl}' a$ che first day of the
animals are rreated therapeutically during the reaction reaction. except in rare cases where, after o sudden rise of
subsequent to the immunization. In one experiment, 1,5 •c or more. a temperarnre or 41,5 •c is reached. Treat·
treatment shortly before the commencement of the mcm should be given only ff the temperature on the sec-
febrile reaction resulted in the development of a poor im- ond day of the reac!ion equals or surpasses that on 1he first
munity. while animal~ treated on the second and third days day. Should the 1emperature on the second day be lower
of the reaction were immune 10 challenge 107 to 205 days than tl1e pre\1ious day the animal is either resistant or par-
later.a; tially immune. in which case the temperature on the third
The duration ofimmunity in cattle after immuni;,.a1ion in day will again be lower than on rhe first day and !reatment
the absence of challenge appears to be approximately two would be unnecessary. Occasionally the tempera1ure on
years.82 It has been suggested that some animals may be the third day is much higher than on the second day and
susceptible again as soon as cwo months after immuni7.a- even higher than on the firs[ day. If this occurs the animals
tlon15 but this observation could have been the result of in- should be treated immediately.
fection \\1th an immunot~'Pe against which Ball 3 does not Treatment eonsis1s of an intramuscular injection of tet-
offer pro1ectlon. rac1•cl ine a, LO mg/kg body weight. and short- or long-acting
formulations can be used. The record ing of temperatures
Jmmunizalioo of ve ry young a.i1jmals Ad\'amage of the must be continued afterire111mentand if the temperarnre48
age-related innate rl!sistance of young animals 10 heam'll!· hours after treatment equals or surpasses that on the day of
ter is taken In this procedure.as. "" Immunization of calves trcntment, a second t reatmcnt should be given. A high tem-
under the age of one month and oflambs and kids younger perature \\~thin 24 hours after treauuent can be ignored. un-
than seven days does 1101 generally result in clinical disease. less the animal show~ other clinical signs such as listlessness
but the animals develop immunity. While il is advisable to and lack of appetite. in which case it should be treated a sec-
monitor their rectal temperatures twice daily for a period ond time without delay.
after immunization, a, is advised in the case of older ani- Relapses of vaccine-induced heartwatcr 1113\' occur after
mals. this is still the med1od with Lhe lowcsuiskoflosses due treatment. It is !herefore advisable to monitor the rectal
10 fo,al heanwater reiictions nr1er inoculnrion."3 tcmipcroturcs. cspcciolly of valuable nnimols. for an addi-
tional period oirwo weeks after the last treaunem. 14 A small
Practical aspects of the infection and treatment method number or animals may react severely despite the Jack of a
,\ft~r the \'accine Is administered to the animal a definite fever. and other diseases. such as babeslosis. may compli-
fever usuall~ develops and if this is not ueated it may be cate matters. 16• Blood smears tand pfl~:,ibly other examina-
fatal. Inoculated animals are rreated early in the course of tions) should be pC'rfom1ed when animals do not respond
the disease, but if treatment is effected too early no immu, sa1israc1o rily to specific hearrwarer therapy.
niry will result. There is no inditatlon that tetracyclines in·
terfere with the development of immunity 10 hcartwater. 213 The block method ofimmunization The block method of
unless ther are given 100 early in the incul>ation period.88 immunlzarion 88• 11 ~- 189· 190 also known as ·systematic treat-
It is ad,isable to immuni7.e individuals or small groups of ment'. 1·1• 234 is \,idelr prnc1ised in South Africa. especially
animals because large groups presenr serious m,inagemem when immunizing large numbtm, ofkicb and lambs. !11 tMs
difficulties. method the \'accine is administered and this is followed by
Starting on the day following vaccination. the rectal tem- trenunent as described above on a predetermined da)'
peraiure of each animal is recorded daily. preferably early in without recording daily rectal temperatures. Treat:nem is
the mo ming and before the intake of food. In order to iden- recommended on the follO\\~ng days, assuming that \·accl·
tify the first day ofche temperature rcactlon the average nor- nation has taken place on day 0: exotic Bos raurus canle
mal temperature preceding the reaction must also be breeds and their crosses, day 1~; indigenous Bos indic1u
528 w·no" n••fll: Rickeu~ial and chlamydlal diseases

cattle breeds and thei r crosses, dar 16; sheep and Angora breed and age of the herd, and record 1heir daily re.era! lem·
goats. day 11: Boer a nd crossbreed goats, day 12.88 peratures in the usual way. A~ soon as lhe temperature of
This procedure makes immuni1.ation on a large scale five or more of rhem is above the average for rwo conse<:u·
possible and it is panicularly suitable for the immuniza1ion live days, the whole herd is trea1ed.
oflarge groups of animals being moved from a heamvatcr·
free area into an endem ic area. The animals can he 1mmu- The do:q•cycllne implant m ethod In this merhod animals
nized on the farm of origin or immediately after their are inoculated and at 1he same time dox·ycycline table1s are
arrival in the heanwater area. Care must be taken. how- implanted under the skin. usually behind lhe ear, using a
ever. as the procedure has some disadvantages. In older special applicator. Slow release of the acti\'e ingredient
animals durable immunity onl}' follows after lhe animal means lhat 1he animal does not become clinically ill. but at
has developed a febrile reaction.2 · 38 Treatmem performed tl1e same rime develops an immunity. 131 Since the animal is
100 early in the incubation period results in failure to de· handled only once and daily 1c111peramres are not recorded,
velop immunlry88 and if 1reatmen\ is given 100 late heavy this method is suitable for the immuni1.alion of large num·
mortalities can result. 8s ln exceptional cases some animals bers of animals at one time. The dosage is critical. so the
may bemoresusceprible 1han 1he rest of the herd and may bodi• weight of the treated auimals musr be known, since an
show clinical signs before they are created. inadequate amoutll ofantibioticwi ll not prcvem fatal hean-
A prudem modificarion which may be applied to lhe water, while an excess wiU prevent the subsequem develop-
block method is 10 selcc1 10 animals. rcp,esentative of the ment of immunity.

References
ABD21 kAtll.\l, A,I, & .SltUM~UIZ... A,M .. 1~;8. Hu.cma\ologic-aJ .>tUdlC-$ in i 1 ,,xu~,1,.mu$. i.~.c \!cihod,, o(immuni7..ltfcm og.ims, Qm·tlrit1
it0•1:> •"J>'nmenrally lnrec,ed with l1ick,.rts,o ruminnnrl11m. 8111/e1in of nm1lnm11ium. hr. Mo. Tfck$and Trrk-8<Jm'1 ms.. asesC<:mtrQI. ,'1. l'mcti<nl
Ammal Htflltl, (1/Jd Ptoductron ,n .;Jnu1.. ~. 232-23$ F"-id ,1ra11110L·Ti<'k·8Qrr<t D~nwc:011trQ/. Romo. FIO. pp. ~-s:s.
:: ALm.t,"muN. fl.A.. J9l1. Jlt.•tutw,m.•r. ·nw prttent u.1w a( c,ur k.nuwleds~ of f.5 \fl~OLI>. It.)!, N .\~(1..ffl<N(i'.'t, M,, 1981. Tid-hornc dtiC-Mt'!"-
the dhe:i-. 171/, //tporr of1hr Dire/CW o/\'t1m11ary S<!rt•lcc, a11d /\11im11I lmmuniztnlon of catd< Imported Into Mo,,mblqur. l'.'~rldAnimal
lndusm·. U11iorrofS0111l1A(rl«1, 17.i1!-1$0. R~irfm,· 40.. 23-:?9.
3 \1,f.,X.\Sl)}R .M...A. ~rtTI,. \\'.O. i. AOF:l,AAR. T.F .. 1cu8. H«l.ff\\1UUr 1-'nrming ill 16 "''""'-"\,x., A. t- ro1'0\', \'.L, t98,f Ric.kcm.,tja.«:eaeandCh'nmyd'i;tren,e:
So11Jl, ,Vrlru Z I. 548-55.2. compm,ith~ Clccuon mieroiCopic ,tudlcs. A.,:to \'iroloti"""· 28. I59-1 ;J.
..i \U..S('WP, Onde.nHcpc>on Vctcnnary Tn~titurr, Prh·;.·ue Hag XS.
11,. A.. :tt)(>t. ,, ,xs.LL G .. 199& D•i>ut).· or«><:tor ofVetcrinruy Se"1c~ Prl\',11e llag
Onderstepoort 0110. S<>u1h Afnrn Unpublished results X12999, C~ntr•hil 6006. Sou1h ,\frit:o. l'tMnal eommun!c4uon.
~ ~u11 r. li.A.,Atu-QPP, M,t-., l)U \'lJ '-l,I_!- l,li. lr\'1S!!itR. J,,;'I,. 19.96, l8 MROL"'T ,\., TtBUJ:. '·~ "'f'-1lf. '-·• Pl'T[lt. T,. \\'.A$SINJ..· . LI. ~t,\RA~. '-, 1993-

Uncha.r.lcctrlxcd F.hrlichia spp. may contribute to cUn!c4l hcarn\nter. SorolosJr.tl dl'11lnos!, o!heanwatcr Jn YJmbabwe-problem< and
,lnnnlso/11:,· New l'ork,w>r/cmyo/S<'leu,e.,. 791 17-23 ptrs.pCCU\~. Rtt•uctf"tJ.t•J,.YJ,g~ «'t di! ,\l&l«J1tf! V~tiri,tt1/lt' dr1 P(t\'~
l'ropkm« 4ij, 12 L
t. AW.UPI', !.\.t., ~ooo. Ondcrs1cpnon Vct~"flna..-ry tru-s.uu1c, Pri\'lU: Bag XS.
19 U.'\Slart, .U-.. SE\\U • .:. M. c mCi\GVRP. S',, l<i.H.L\', , •.,. lO~QfJ,\:\, ,.,..~~\H.AX,
Ondc1>1epoort 0110, Sou•h Mrfc., Persooal cornmunlta11on,
s.~t... 199-: Size \~Jriation of the m;sjor tmmunodomimu11 prou~m of
; AU.SOPP, lt..T. & Al.I "-OPP, ll.A .. 2on1. A novt·l Ehrllci11a gN1orypc detected
Cl>i4",lrlt, rwmunmlum. ClilliCDI nnd Dl«RI/OStir Lau<Jfl'>tOr,·
In do;~ ln St,uth Afrk.-.. /011mnl nfCl,meal M/r11>l>loln8) • .l~. ~2o.l-4207. lmmutt.of~·. I. ;44-;~6.
8 AU.SOPP, .M. t ... DOPtfUSG. C,M .. )tAII.L.Mtn. J.O,. BSS'M.10. A.. HA\1)0S, D. t .. :,n R-4RRM, t. t- WIUT\Utlr W ,, t,,11,. 11..)t. r.,,"T,, kR,,
\:'\Ml') K, ' " ' ..
\'A!\: HlEIWL,.., u. &.,u.sorP. U,.\.. 2001. F.Jutt<hf<t n,m1,wnrmm m-ijor ).f01U.1A.1't0. A ..L MWA.St;I. D,M.. .MCGUOU:, 'I .C. 4r ,uUA..'-, '\-.:\,1., :mm.:\
nntigt'nic prottln gene (mll/JJ J'\':lrinnci ,lte not ~t'OMnlph1'-'31ly wbse.i.o! Cm,'drla rwnimmrium gum.·> important tor Im mun~
con$truinl-d o.nd shttw no (Widencc nfh11\IU1gei:o.l\"t.d under poshiw recogni<lon ond protctdoll, Ut;ne, 275, 28i-t98.
<eltttion pr~»un,,/ocmwl ofCl/11/<ol M/crobiolDgy. 39. 4200-4203.
~) -OAf\SAlt>, " '·C l9SS, Hettrt\\1llc:r fmrnur\.tro1!on under field conditions in
g AU..~J,p, r.. JlATTI:-..~ou, c· M ,, vooei.. !,W, &AU.SOPP, 8 , •• 199:9.
~~. !>\<i121land. 8uU,rfo oj' F.pi::ootl< Dls.-1>1:s t>fAfrl<n. I. 300-313
E,'llluauon or 16S, mnpl and pCS20 probe,. (or dtt,>clloo of Cowdrro and
~ 3IJlAL. " · o,\JlRI"-, G. ~ c,,)ius, t .. 1.9~5 Prop.,g:auon of 1hc- nci.
EJrrlid1io ,pt!Clb. Epidmuol"l{l·nr>tl lrt/,Yl?OII. 12?. 3"..3-s.128. Am(,{yomma 1'<ln,~orrm1 in the Curibht"an. R1:, 1ue ScicmriJicJu,•11;
10 AU..SOPP, ·M,_T.• THE(lO,S, J.. con,a ~u... DUSSTETt\1.u:£, M ,T, &. .\USO~P. T,'Cllllt</ltr, I l, 6tl-85S.
D...A•• 1999., The occurrence of 1111Ul#rltl and Cowdrm paro:,itC.$ m African 2.~ U~'\ft.Rf., ~ .. Ul&.L.'\'Df:J\G. (i,. \fOkll.1.. p.c. ~c.-\MUl\, I., 198;. Oangc.rc,f
buffalo l.Syt1tl'ru1 ct1ff~n 3nd their a,soci.!lcd Arnblyomma lt<•l>rutum inuoJudng ht•anwatt.-ronto Uw AmrrfC"JO in.ilnlanct. pm~ntia1 role of
licks. Ond,•mcpoor:Journnl o/Vi'r,•rl,wf)· R,Jtart"Jt. 66. 2.S.5--249. irutig~~OU> and cxo:rc Amt,11,mm11, Ucks Orulm:apoo•t /ou,1111/ •I
11 .-.u.snrf'. ll,T•• \1$SfR., M .. nu PLE!tSl\. J.L., \'OG[L. ~.\\' l:t "L.1,b,OPf'. t'l'lerlnnr,• Rt-s~rrh. s,;, ,;OS-H 7'.
a.., .. 199;. Diff~runt org.1nisrn,- :h~Ociratl~ with ht.•.arn,11u.·, 3.s stlo"in b;- u Nil.L.4"3..\k"\"L L. l.:o,n. L.U..., DOGBn. ,, ~ ,uM.Pno'". Lr.. tQqf,. Hei.rtwa.ter
analy<i> of16S nbos<un,il RNA1,enescq111·nce, Vererlrro,:,• P(lra,<irolog)', Jn Gban3· lrnpllc~tio:>s for eomrof 1>f cic~s. Trop/1111 .~111mnl IIMltll nnd
- , . 283-30/J. Prodiutit>II, 28. S9S-64S: dl~uSS!o.tJ 74s-86S.
1.2 A~o1u;w, H.tt.,. ~on\'AL. L\., ,,sg. Thi.• c.arrlcr ~uuu" or .:.hccp. caalc and n.s RtU•~1. l. P,\.,'TO'.'i., l.-'., )IUNl)fAIOH. U.(i, StS-U\IP'TTO~ , J; ,l- 2000.
Atrkan buffalo recovered from hoam,·nt~r. 1·,r,1111111)• />llt'/>$1/0log,, 3•1, Lro\,'\h o(C,ou"Jr,a rw,mumrium, 1hi: cous3tlvto ;tK~nl o(lwam,-;~tcr ln u
261-266. tiri. cc I hnc-. Juunu,I o/CJmu;{l/ .\1/crolnofog:l, 38. l2.J8...J.?40
13 A;\"O~.,·Mou.s, 1rt Propt,~als !or the nomtncl:uur~ of qll.quinn ~ Rb\'EIIIIX,f .1 • .., O.\\'lE>, , .... 19$3, Ccllutor rnpo= or Bac/1111.,sw,/ll/s
trypnno,omc, and fur 1ht m>lntonunc• of rcitrenc.,collt-ction.<. 8111/Nin and F.«lldrrt1tln coli 10 lhe Grarn >1al11.Jo1m:a/ of8aet,'riolo$)·, 156..
oft/:, Wor/11 NM/rli Org111ti:ur1/011, 56, •lb".-180. 1146-856
 l!carm,mir 529

2; llCZUJOExuoUT. 1.0.. 1982,. Ol<1.?aseS or tht: ccntruJ Jh:n.·l~U!lo. <),~l'rn of ..16 u,u1u'>1~. ,t.L. t962. St•lt,cu,e ,u,1ntn~ or Ri,:Avttsia rumfnam,um in ;i,,ul"
tittle u.nd~r tropical conditions. Proet\'ti/ngsofti:, 'Jtli l\ 't>rl,l Co,,g,,,s.., "CC.1.i(ln> Tiu.•\ ·,11,•.ri,Ultj Rt"\"Ottl. ;' ;. 1371- u;a.
o,,Dis<VlkJ ofC.1111£,. Arnm•nl,,m: The ll:c1herl.ind,. pp. 991-Y!i:I Ii IUJKtcll>nt . \1 , ... ~t\t\10'.\~ t \, ,1,1b1. ll H., Ml-rR, TX. &.lttuu, ~.M • :iooo.
28 8EZulD£'.\1tour.1.:>., 198.t O~monstrauon or Co:tY1rw r:,mi,wruwm in E\'ld&.1nu nf Cln1dri11 mmuumttum tnl1.'<'ticm <h<'1lrt\\4lCC. In
\mb{n'>mnu, l:tbttl<':tm by tluott"S(('m ~ntibod)· h'tt:hntql1t:-. lf~1 and Ambf1,·,m111111 $J.'IQ~Um Uck.s found on 1urtolM:S impmtl-d inro Jilorida.
dn:uon mirrosco~ . Ondc•r.flt!/NJt>rt Joumnl oft'tttrfnrtry R,rk.Vlrtlt, S1. /m1r1111I OJ l'ttrosi:o!c,,:.~ 66" 113~·1136
2:3-21.S. ~8 fi'l1t0'1 n, -~ n,,a;1 "· c.t t ~ lmpro,cd rul1urc condUioni for CQ:ntrfa
29 1:1(zu1otxtti'litn, 1.:1 .• 196;. The pr:uscm1 ~ta(e of Co:1·,1r:a mmmtmmuu nmwimmum 1R1c~tt'-t~lt .. l. 1ht'..r~n1 etfh"·Url\,·oue, d1wa~ of
cul(,v;auon in cell lines. (J11dtf1ltJJ(Xm Jaur,u,/ o/i ,•rt•rmnry Rl>5ff1rrl:. ~ , domr'!-t11; nJmmant~~t:)·101,•rimolQKl·. -1. :t8S,..:t90.
20.S-?IO. ltt H\11()\1, H.. \l~,u• l .. 00'.\'.0\',\'.\, l'.l~ .. '-'IITll. ~ J_. 1,t:tt, /,j 1·1 tn,
so at.zu1or,ttotn. 1.u., 19~;. X~tur-JI lr:uhmi.»\rJn ofh1.art\,·.ucr i'l.cthtunn u( O.m-tlri11 mn,l,mntum, fmn1 pla>nt•L n( inf<.-ucd Nmlmmts,
Ond,·tilqx>art /oumal of\ tflermary R,:~,mrch. ;i 1 3-t9-3Sl. \ 11trri,1my .\l1rrol1to(og•. 26. 26J-2ti8

31 nuu1ol-. ~Ho1n. J,u. 19ft8. C.cmtin aspccn of ih~ uan~mt,.,ion or so U\\\I'- t .. 198,. Con1rtbu1lon ~r~1ude ~pld~mlol<>glqw, do la co-,drlu..,
hnrtwate:. the- occurnmt'4.' of th~ org;.,ni,m in 1ici;.. .1nd 1n :·/rm cutture ret,1t1/rla trwm1amtu11:i t•n Gu.idcloupe. o..:sr-. thMt,. Unt\"t':.;116 dc-
DX Sc. thtsa. Ur,i\t~I~· of Pte1oriu.

32. 8£2U1Uf.XHOln. f,U., 1989-- Cht\J)JN 2·

.
CouYiri,, \"dC'Cine;,->. J,r. \\!'Jt:GUT. I,(~.•
P11ri ...~Sud.
;1 C\).IU~. L. 199:t. I.~ pt.•rtagc .s,1 mptnm.i:titLUe de bo,·in) c: chl'Vrt~ Ctt.'Oll·
ii,m•rb de 1,1 cowdrio"c l"n GuadL1loupt•. H~l'tli.' d'E.lr1-as:1 t'I ,I(, .\lttb,•tf11~
(td V1'1~inary1 Proto;l;Km ""'' ll1.•mo1,amsiu,• Vtt«int"..<. Boe~ R~ton,
\f,•n)ti1t11lt1:1(('.<P11p TrlJPfU111.'t. .t~. }33-l3S.
f1Qr1da, CRC Preos. Inc. pp. 31-12.
:,J n.czu1oc..,1u,ut. 1.0,"' n1r.Alli.t. n.u .. i9JJ;o. ThC" ('Ont.tot of hc~rt.....Hcr h!- 52 C-A~u,, ..... ., :• ouu .. ~ .19a;. DiuJWQ..'iJ.) of hc..artw;H('r an I.ht? li,·c amm.U;
menn~of iJdo co'n1rol. 011tlctS1ep11tm Jm,r,ml of\ ·rh~rl,m') kf'f,•m-tlt, !w, C.);ptricm:c~ wuh ,:ou.h Jn Ciu~dcluupc:. 011d,·~r.,,Kl(Jtf Jaurual of
52J;-528. \ «umumy llts{(lrch.;...; ~l ~·l~H
34 uuum~-.:uOuT, 1.0. ., J.u::no!..7,. C.J .. 19R6. VtofJf uf mrn,cw.arfaJ tJ 1.:AMu;._ 11. , , 1u1ou, ~. 1992. -0,t1' rQlr- or ,\mhl,i'Ommn l'flrf1'1t'<lrtnn In ch..t
t.ran.&,n,ibion of Cowdrla r,1mmcmtium by .'1..mb!yommu l14!btW!tun. trnn>ml,-1.ion of hcauwtUl'l with ~fH..'Cinl 1-..:Cttuni.·1.· U) Gu;td.doui;w
Onderlttrpoon /otmtal ofV1.•1t•.rlm1ry Rt-.51N1rth 31-3•1. ». ,\111111/J t1f1h,• ,\ ,•w l\,rk .\(t1//m1rofSd,·11<es, 65:\. '.<J~I 1
5-l t.A.\Ht'I-. L~ ~ uAnRL. ~ .. 1~.;. ,·eemr~irn1uton or uck•bornr, dIW":t~ m th!!'
35 h&.IJIOfSltOUl". J.D,. J>,\1(n$()!\\ C.I.. &, M.R~,\kb, ll.J., 1!)83, {tl 1'/(IQ

C'U1ti,·-~u:Jon of Co:t'tlrin rum(11aml,w1. Ond1.•1$W/>tJOrr J<mrmtl t,f C:arihh~n hl.-nd, \ ttc•1ft1nry· Pm'l1.«111f,'8>'• :>7. 1f,i-t 76
'Jo1,1n'ntu')' Rl!'St!t.1ft'h. 52. I Jl-120. r,:; c~\\J1J~. L"' EWt1U.- , ;!)81. l..a «J\\Urio.sc thNn1w;itt1r,. Rc,·ue i::en~r~t!<.-
36 80UltPQUU.IIJ). $"., 8liXS\H), j\,, )11\fl"l"INI"'/.. u .. .,.11111,:urnmou, r: TAA.r, , ..
dt> connnt);\dr.C\.l-5. Publ~hetl br 1,u11tuc ,n;k'-"'t(!at d~· i\tM,-rmt.•
VtfUirmnm t/1.'.tPilJi T"OJ)fMIU fO(i/N1fl1•JJ11•111 do C/IW)). C:IHAD,LM\T
::),fKO~UJ•:JU.i, .\ I), .. WUU,-\VO, r.o.. U)4J;',., Jn(eciion orb<)\ Int" br;\iO
BP 503:; 340'.J? ~l<m1pc,lller Cud,,. I rr,111<·,·. 147pp.
mfcro-vL!ssel endo1.ht"hal ('cll:-wHh o,wtlrl<, rum1,,ami11m clic:tL~ ll·1
b~1'3.. ·6. and ·8 mR.'XA produ.cuon and rxpr~lon ,1r on unu~ml !\\> C".\,'\IIJS, J.,, \\ \11.L\HIJ. J,C.., HUI ... t; J.'t 1°1~, I., X,\Vt,.:r,.0 \l, ~ 'IA111l MO~.~,
~UIC ct;µ> n OQ Cllph::s 1r.tn._«dpt~ Journnl tJ[lmmmwlo,"ti· 1; 1 199i:. <;,,nt•fh' r,~,~J;)nn~ n: \r,-o!f' non," 1n t'n\\·,ino-~1< In riunrt,>.looJl,~
-!032-1033. St,11u~ 111 t9gs, .-4mmls uf rlrr S1•m )(Irk ·\cculrmy fl/~H·11c~s.. ~l.
16·SJ.
r BO\ ,R!\K\... ,1.N.. MU I Ett, J.C.~ ::,.;r,;YTJFJ\, 1,r.. 1q30. the intluen<"t•of cemun
salti>, amino acld:;., suga~ a.nd piOt<'iM un the .,,o:,Un-y nl rit~n.,iai:' 57 1:1tM1h0KO. T., ~l0~1iUtL ,\,\\ .. Ci\U.\MIAI\, 1;.J,, l'fTtllt. f f,, );S.IJ~M, ~
ftJuro:nl uf&,(1molOR1·, .59. SO<J-S:!l. ,;1 uu 't. r...1 :\1Ut.\, ..... ,, ~ 111 MM\' H.u, 1 ~ . Tht! euntral ol'h(':&n\rnttr
on largc·.>Otlu rnnundcl.il :1nd ~m;sllholdcr fom1, m /tmb.u>wc:.
38 OQ\\1E ,1.\\ RlOD\', (Ht. $f:\tu. '1-,M .. \l,\H.,\!\', ~ '1, 4 Bo\,.!SfT \ f. Jqc;g,
P"•i·tnm •" t',,unmrry .\!Pcltont 39. 191-210.
PotontiaJ \aJue ot m.tjor:.ntl~enlc pmtcm 2 for ',,CTO[tJgftal dlaprtO)L\ o(
ht,am,mor ~rid related chrlldtlal u1rc~tron, Cl/mm/ ,<.c Di"871ostl< .;3 u.,.•. ft.. iu~:
Tht patholoi;kal pht,!ololl)' ofh,anwat.r 1Co11•/rin
1.nb<,rotory Jmmuno!ogy. 6• .?OS-?15. ,Rid,•usln n1Ml1i,m1fom. Cowdry. 19261. Jtmrtwl oJ'tl:t' \outl: J.fr:a111
V,•1,,l,:m; ,\t,,1ltct1I Ar:ttk'inunn. 33-. 18)-191
39 BR.\\,.O~. c.r.. 1986. OinMhyl ,ulf<lxflfo IO~tSO:· .\ rC\icw Ctmwll
~<1lfri1U1rlnn ;..,, 6HiO. ar,. )9 GL\lll:l l t, t:' '- r.1.u1:u:,t l9':'0, Gmm·,·~· \",,~trrinnt)·1"t.,ri(ot,,g,.
\t

London: S:1U11tzt rind.lll & c..i.~~tll .

.40 bAA\'TO\:, X.A. 1$.~· Collq;t of\'rtcrinru')" Mcdfcin~ \\ ashingtO!l ~hHC
Unl,-e~ity. Pul.tm.1.n US..\. Pt<n..onal rommunfcu1:,.m 60 (;0Ult'ft0. G.( DIJltJ:f:U•TI\.\U1~1A'-'-\ o .• !>-, ltf)'-OUt(';. A--t't,, C:OUk\.UD, r.o..
l!1,1 ( Jlt."Ctwh.minc,. ,1hn\1hU~ 1h~ If.-; g:muno rnductd "lM~ 1t MHC
..;1 81l\no~. F,A fl,'H~t.s-. 1. nE\'IWE-RS, E..l"•• ,·.\,-.; ncu. \I A ,\U.~M· 8.A.
~~pr"-~1tm on OO,"lne bro.tn cainllory endo1heUal cell,;, J()tuiwl flf
Hl9i' Coruuuction nnd iniu.tl :.u1;d~~is of n ((l]lrc.scmt.:..tf\·t.· lumbdn lA!111
lmmtmolog>·. M:- ZS:!S.-2329
l!Xprcssion IR>ra~· af1he imrucellulo.r R.1cl'Ptuia u,:,•,trw n11uir.mtrwm:
c!onmgof mA.pl ,md thn.~ 0:th<'r Ccwdrin j:.t'nes.. l t'tt.·rm.,r:· l'nr,t•rlOlog\'. tit c:owuur.1.,·,.19:i:1. 'S10dfc,. on thc1..•uolob"}·c>fht•;inw:m:r II. Ri,~n.iin
;2. 185,-199. rummr1111Jum n. sp m thi: u~uL-:::,, ot 1,cl-. t.ran,m1mng lh~ db:cn,e
foumul ofF..1p<rini~·mnl 1\rrdtcfn.:. ..JZ. ~:;.~z;.;
.;.2 B.lL\\iQ,, >. A- \'a.., :U It.. \\'-:\1 t'. M•• \'QGEl.• .;,w , \I.I.SOPP. II \ .,.. l998,. \
panJ~· ptOtet'U\ e clone ftom C:uwdrifl rumim1mirmr 1denut1t.od b~ 6.2' cowmn , .,· 1~2~,. ~iuUl1.·~ on the cutth)l!)' ofh\'3rt\\at.Cr I. Ob\t'I'\ ,1110:1
us-in~ a SalmoncU.a ,11.ceim:.dtli\'el)' sysuim ,.,nuoU ofth\' \4•:•• \'ork of ,1 rid.e1t:St.,. ;u, ~w,111 ru,mm,mlum fn. ~,u.
1n chc ti...,u'l:, oflnfc.'ctl-d
·\C.lulcmyof&Jrnces, 849. 2-r· -2.S2 .. rtnima.1-.. Jtn,rm1l ofE.iJNrfmtnuH .\fedlcfth", 12 231-:!...~.
..al DM'liO:,.. K-\- cow;x$. ~.~.. ,·.\..~ ~-nuw. f . $< W -"iOl•P. n,.\ ... ?003. 6J f()\\Ul-l\, t.\· 19.:Z:b ..;mtJ:c,.. tlll tht r1Jnl~y<>! IW;trtw;ur.r. Jll. ftw
Prt1parmion ofEIJrlichia rumlnamiwu chaJJettt;c mni<-ru.,1 tnr mulupllc~1ion of Rf,-J..,.,u;r, 1tan1111m1irm1 ,, ilhtn tlw cndo1hdu.l tt-l!,01
quantili3blt- and reprndudble ch:lllcngc in mice an~ ~ht'Cfl. I ,•i.•rirrnrr lnl<'<l<'<I animal< and thrlr d!<elia~e in,o liw cir<ufalion. /<>11Nr,1/ of
P,1,a;/10/"I()', 11? 63-73. f:xpi.•rlm••t1tt1/ !.ft•dir,11'(9 ..:.;, 803-81.1

44 aul'T• .:... 'BEZUltll.'...'-!lto\JT, f,n. '"- oe W<\.\l, o. t .. l9~ Th,.-

e:,ta.bhshmrm of t..a nA <;11..,c..\, 11.,t., 190.:- some asp«"t'- of ht-:,rtw:11cr ht ,\11,;ol..t IJuT:l'tm.
""o,ineccrcbral cndo1hellal cell line {SBEl89, lhat •upp<>n• 1he gnmih Offetl' lnf1•runtto1tnl ,I,•.( FJ,i:nmit,:. Ci2, 9b3·9G9.
orQ:m'tfna n,mmmm1m1. /ounz(I/ o/1hq South /\Jri,·,1,i \'t:l•rmal} 6S PA\1L. J.n.. \I.\UI\S, s:. ,,. ~ \'O\\U 1 , ,,. 19g-.,, Phylogcnctli;
rdauon~ip o:
AsstJCtal10Jf, 63. 87. <...<Jwtlr/l, rumtnnmtum. np,rnr ot h~:s:rt\\'lHN. toAr,apl(Umil mt:'l,(fno/t
43 B-Su)'.,~. c.c u :r-1."0\\hO~n ••q .. rgcJO. H~tolo~fc .11~0 lrnmunocht.•tnitai und miwr in ember.,;. of the· 01<Jer Rfrl,;e-1~1ntc~ d~rcnmn('d on It~ bas-:~of
IOS rR~A S&..~uenc.e t,,r,;,:a,lwml /<mtnul c,fSy111:mutlc Bac,rrwlo~.·. 42.
1-tudr of the p~thogco1.°'i"' nf h1t:-inwo1u.•r c<:onrlrfo r11ml11amlum
tn(cctlon) tngoo.ts.ond mice-. ;Unerlcm, lrmmol ()j\ tJterinary HcSt'(lr(lt, 2ro-:n.1
51. 14,6-1480. 66 Ul1 VIU Un,~ l.t1, .. HRAYTO~, L.\... /Wl\'(.O\Hl'll, l,...P. f;' ,\U..'l':OPP, 8.,.\ 2000.
530 •u:no"nnll'I': Ric:kensfahnd chlamydia! diseases

GenomeslZedclCrmin.ufon. rrnd ph)'iSical ~ndgcm~tic rr.ap of Cou,vlrla 87 au PLESSlS. J.L & ., ,.,uAX, L, 1987.. The appllanton of Lheindirect
rum!mmrlum..Mlcroblolt,gy, 146.1627-~1. nuort!Secnt anribod)t lt!lt in r~1.~rc:h on hcart\\it.ter. 0,2dersu1poon
67 or. w,\AL o.T., ,.1..,·rrunn. o.,,. 10-s<;llJA.~. F.. 2000. E,~tui:ttion of lhc mapl b Journal afl',i,•rl1rary Rcstorr:h, &4, 31!).325.
ELIS,\ for the diagnoSl, or hcanwo,er in South ..Vria. •.:,1111nls of1/r, N~,,, 88 DU Ptl'sSls., J.L tc ,t1u1.~. L, 1987. TI1e block niethod of1·ao:lnftllon
York Acadtm;• ofStle11res. g 16. 622-627. ogalnst he.i:n\\'11wr. 011dt1Nrcpaurt Ja,11nnl n/Vet~inan• RtSt."Q.rth. 54.
68 DYF.\l. s,t•• 1998, Areview of he,,mvmer and the threat oflmroductlon of 493-495
Coufdrin ruminamlwn andAmbl)'Omma:spp. rick.-. to 1he Ameriam 89 ou PI..ES'JiS.. J,L. & )J.AJ,A,v, L.. 1987. The non·spL-cific r~i~anc:e o( ~ule w
mnln!and. Journal of?.Mm1d ll'lldllfr.\,fed/d11~.29, 109-113. h~nwatcr. Ondtr~lt'JXX>fl /<Jumnl of \'eretin<u-,• Rns:md,, 5-1. 333-336.
6g Ol!J.M, ::;,1,., :..'6kVAJ.. R,A.• OONA~llh P,t.. k MAHAS, ~..<\1 .. l996.. 90 ou ~LE$$.JS, J.l.. f;- ~"~~, L,., 1!)81. rroblems \\1th tho lmcrpreuulon oi
l)emo:ntrndon ofvenkal tr.)r~nils.slon or CouvJriu rwnfhnmillm, the cpfdemfolog!eru data in heanwaier. a.studi•on 23 farm>. O,ulem,J>l)Qn
cauuti\·~ ngcm ofhcanwn1cr. from cows to their Cal\'CS.. V,ucrinary Journal o/Vc111rln11ry Rt$earr:h. 54, •127-433.
l'Mulrolo/0', 61. 119-132. 9l l>U 11LE)°'Sl:i, J,L, \',r\~ CA$1 ~ OUVll/A, JA h DP.ZU102NH0l1T, J.O.. 1989. Tht-
-;o o:xo!\~, fl.\.,., 1s9,L ttrnmv:ner exptrim~ms..-lgritulturnl JoutJml 0/11111 hcteroget1ldt}' of Cbti,virfri ru,ni111u11i,,n, stock.~ CTOS$·1mmunt~· and
Capco/0/Jod Hop,,, 12. cyl-7(i0, >erolo.;:i• In shocp and pathogcnlci,:y IQ mice. 011d•r.tte;,oor1 Jom1111/ of
;1 01xo:-:. ft,,,... 1899, Htartw:ue.rexprrimenis. A.griculrumlJoumtll ofthe l'ererlnmy Res,ar(/r, S6. 1~201.
Capo ofGood flOJII!, 14. 201,-207. 92 D0MU1t, p;.. UA.RBl'T, A.~•• Pfll(l;lift, C., C,.\SITTI. C..;\.. PAU!t"R, G.H,. R.-\\", S.C.,

t.-\ prelimmary1101eon
72 DV PUSSL.."-. J.L. 1970. Jmmunitytn hea.m,11tcr. FUl:JIJ(U, l'. k R~"OlR\\.'A, P.R •• 200J~ R~.rg.:mitntfon of Ct-n<!r.a In tht

the rote ofserum :i.mlbodlcs. Ondem,:poorrJournal o[V~rerinary Famll~ Rleke11slaceae •nd llnopl~,m~••eeattln the Ordor Rlcl.eu,l:,l1$.
Rt>seare/1, 31, 147-U9. Unific•tion or Some Specie, of Ehr/fchin "ith .'\J1aptasma. Co1tvlri11 \\ith
l!hrllch!ft. :1nd Ehrlichiil \\ith Neorlckeu•la: Oesc:rlptlonsof Fin, Xe,,
73 ou f'U!SSI~, f,t....1!,70, Pnthogent'fisofheanw,ner: l C(J1vtl.rin
S;x,..:le. CombinallOni: o.nd Dc.<lgnation or F.hrlicl,ia equl and 'HCE
rumlnnnllum in the.lymph nodes or d9mtt5tk n.im1nzuu.s. 011,lmteJ1QQrt
ag.,nt' ;a Subfettf1-e ~·ynonyms o( ehrllc///11 pha_gocy10~hlla
Journal ofl'rrerinary R~.,nn::h. 37, 89-95
1111,matfo11al Jourr.,1/ ofS.w1emmlc11ntl E1'0/ir1io1wr .11kmblofo[()',; I.
74 DU l'l.£ss1s. f,L, r975, £!ec1ron mletoscopj' of Q:m:dritz rumltumtii,m 21.;S-2l63
in!ccled retlculo,endo\ltelial cells of the manunaii:m hon.
93 M)ISG'O>:, , .. 1898, H~anworer. Agrlc11l11,ro/ Jtmm11/ oftlrtCap1•of<:,ood
O,ulew,poorr Jo1m1nl of Veterinllf)' Rewin::/~ 42, l-13,
/lope. 12. 7•18-754.
7.; DU PIISS!S, J.L, 1975. i!i.lropathologlc:ihmdle, on 1M pathogonesil of
~).l 111a. ,.11. a:r SC.Hu~s. ,,. 195:!. Heartwntcr intmun.is:ntion under field
heamvatos as manlfesred in mice infected 1,ith asmlln oi 0>1«/rlll
conditions jn$\,"tu.ll:ind. Journal ofrJw Sout/: Afrl~an \'t1w.rf1tllf)' .\lwlfr:nl
rumfr:mrdum. ~tMcd.Vc1. thesk. Unil'Cr$ilyof Pretoria.
A>s«latio,1. :?.3, 9-14,
ro ou Pussrs. J,L, 198L The,applkation orthe indir~t fluorcscenu1mib0dy
95 FLACH, t.J .. \\'OODrORl'>. J.D•. MO~J\ftlJ\. S.'P.. DOLt\N, T.T. & stfAMB\\',\.'\A, t..
test to 1he s~rology or heam,•a.1t1r. /11: ~,,-uTEHt.\JJ. c.e. 1,, 01aso~. J.u ..
1990. !dentitltetion of 8bhf':S1(1 boi,is and Cnuitlrfo r,1.mfrJanrium on the
(eds). Tick 8iolo,>•1wt Cn11tr/Jf: Pr0<ccdl11gs ofa11 l11umotlo,,/II
islondofUn[IU)n. 7.an,jbar. Tllo V~1erint11y/!,'<'Ord, l26 ;;.59 ,
Confcrcn,.c. Rhodes Unh•cnity. Crohom6tO\,n. South ~\!rice: Tlc:k
Be;,earch UniL 96 Cl'tAU, <;.~.. PERJl\.. s.o~. KATF.,~o~J.M., MCl.)':RMOTT, J,; •• MOR7•.-.R.U.. S,P, &

n DU Pu:ss1s, 1 L, 1981. Theinlluenecofdith,osem!carba,.onoon 1hc

or
\VU!'-0. A.S., lS97, The preva1tmcc :.crum antibodies :o uck•b9rnc.-
in(eclloM in c;ttle In smnl!hol!kr d:llry farms in Murang'a DtStrlCL
1mnmmty of $hccp to heart,,>altr. Ondenr~poort Journal oft'attrina,y
Kenya," c:rosMectional St\ldy. Prol'I!'""~ Ver,r/11111)• ,\ttdlc111e, 3-0.
Rus,orch. '>8, I iS-176.
9S-107
78 DU russ;s, 1 r,., 1982. Mice. infected wnh a Cor<'dr/n rumli:amlum-llke
97 Gfl.AO\\f;U.. O.\',, \·~~ !'\"TEXEIU:. ~- & JOU6F.RT, o.c.. 1976. Aucmpred
•gen< AU mod<I In 1hc study 0Ct1tanwater. OS.SC. thesis. Univer>ity of
oniBcial infection or impala, blue wildebce,1. bull'al<>, kudu. ginl.ffo and
Preiori111 South Africa.
wanhog\'.;th h~nwarer. Joumat o/1llc-S-0uth ti[rfr:an -''-'l£rimuy
79 DU PLESSIS, 1.1-, 1!135, A me1hc,d for determining the C111l'drlt1 ~-\sSOCintiOfl, 4i. 209-210.
nm1fn<111rtum in<ection rate u(,\mlJJ,,'Omr,u: hebm~wn: ~ffettS in m·icc
98 ,,w, " ·· 1m, Chomka! blood st11dles. lll. Comparative ,:udies on
Injected with tick homogon.ues. 011dcrsnPQOrt Jo1m1al af\lererinnf}'
'loked and 'unlal(ed' blood filrrnrcs of sheep In honlth and dunng
Re.««rch. 52. S;i-61.
'hc1)ftw-ltt'r· ~Rlckttrsi.t1 rumlnouti'1m inf...,--cuun) and bb,ae-wngue
8Q Du Pl>MI;. /,L, ~99. Ondcrstopoon Veterinary lns;irute, Prt,-a1e Bag XS, (cmarrhal (en.•r), Or:de.rs1epoort /oumal ofl1,rerinary Scien<Yand
Ondorstcpoon UI 10. :-outh Africa. ~e~onal eommunrcau<>n. Animcl lndu:suy, l, 285-334.
81 OU PU.SSl~ ,.i.... 8FRCHe. P. • \•A,-.:. GA.$, L. l991~ T C(!ll•medfatad lmmunJry 99 GROSs<DPr. r.,.\\., 19,$. Heanwa1er immunl:r1nion or cland
to 0>11rdria n,minanrittmin m1cc: the proltctke roleofLyt-2.. Tceu~. tT11111orr,;c/,r~, or;o: or;o;t, Palill!l).Jo11r11al ofrite Sourh Jlrla,11 Ve1erinao·
011dcrJ1PpoortJoum11/ ofVPteri,1nry R,,...rclt, 58. 171-l 79. Medicn/hsociarion, 29, 329-330.
82 DU PLESS1S, ,.t.. i, 8EZ\/1DENl(OlJT. J~D., 19i.9~ Tn•,'CS't'igarionson t.he nntur.il 100 c;,,u,;s. a.. 1981, A practical npproach to the control ofhea."twater In th<
•n<l acquired r.:sis1ance of ca,tle ,o nrlifici:il lnfecuon with co,.vlrlo Angora goat and ecriain ,hccp breeds in the Eastern Capo<:o;i;,al
ntmimmtlr,m,Joumal of the South A.frt°('.IJ.11 Veterinary ..\ss0¢i11rfort. SO. reglon. !ti: 1,Hmmw:>, c;.u. ., 01n.o;(),<, 1.1>.. (edsl. Tic,· Blologj' and
0

!34-338. C,,nrrol: Pro~dlt,g,ofu111111,r1wt/()11(l}Q,nfett11<e Rhode, Uni\'er.dry,

83 UV Pl.f...SSlS, J,L.. BezUlPl:i.~11<.)U r. J,I,), ~ U,Dt.MA."~· ,.J., 198~ The Grnhom~omt. South Afrlc-•1Tick Researcl1 Unit, pp. 135-136.
immun,zation or calves against heartW3lCr: subsequent immunity both 101 GUbVE. \., !OXQ:UA...~. F., M8£1'>~UE, M .. l>IOUf, A." UJLEt\'1ltr1'G, C. .. 1994.
in the absen~e ,ind presence of natural lick challenge. Omf-Orsr,,poorr E»al !Ur le 1erroln d'un 1'accln nu~nu~ cont re lac,"'driiise.R•1111~
Jounwl o/Vctar/11t11)• R11$MfCl1, 51. 193-1 !16. d'St,,vq,e ,r de ,\/i!dtcf11, Vclt&/nalra des Pn)'!. Tropl(nwr, 4i, ~Ol-l~.
OU PLn..ffl!i, r,L, (;RAY, C. & VAX S'n\lffl, ~.F.• 1992, f1m11 cytometdc:amtlysls
Ul? r.Uf1"1f, .\ . ,1 \RTt:o.:r.,. n.. M11Fwmr.. "'·". ,·u m;-. T.H,,., n1nUF. "" ,993,
ofT cell response in mice infected with Cou'Cirla, ruminnmium. l!pidimlol<>glc de In cowdrl<l,e ou S<!'n<!'gal. 2. R6suh.a" de suM,
o,ulemepoonJ011mol o/W1erinnr; Research, 59, 337-338. ,~ro,opidi!mlolog!ques. kcvutd'l!ll,i•llr••' de .\fidi,c/11, Wti'rirralr"dn
Ss ou PW-Sb, J.L. JA.NSl!.~. o.e. • PR07.ESJ..'Y, t.., 1g83. Heanwa:cr in Angora Pllys 1ropttaru; ~6. ¥19-45~.
goacs. I. lmmunit)' subsequent 10 anfficial Infection and IO)a1mcn1. 103 eo£>'1i.A.. MllnNour;. ,:. & n10u,.,1., 1,q93. !lpid,mlolog!c de lo cowdrtosc
Oml.!rJrtpoort /01tn:alofVeurrJnarlResea1ch. 50. l3i-t.;.3~ ou S<!r-ogal. I. !ltudc de la mu,smission N du lllW( d'infoction
So D\J PU.US.$. J.L &- KUMM, ~.h., 1,971, The p.·lssagc or O,wdrta mmintmtfum d~Amhlyommn i.•aril8(1tum (Fnbriclus. 1i9J) dan.s la t6'?tion des ~ia~·es.
In mite. Joumal of rile South Jlfr/MJI Ver<rlnary .\f~dical At.ioc/,u/on, 42, fl•vuttl'Ele,.,,gut tlll.\1dde,·in• V416ri,u1/r,dJ.~ Pa>~ Tropfr.aia. 46,
217-221. 441~147.
 I li?artwat<'r 531

10,; OUM.)JOW, fl.. 09£1\£\~ P.T, tt VA,-.: ,'-..lS'ff,t.. .s.. 1988. ~cw idt-as-Sn lht th~Sudan Bri:/s/1 Vr1erinaryJ011rnM, 116.105-114.
~mpu1m~tic 1roa1m-.n1 ofhe•t1w1>1~r. Pro<i!udi11p ofrh• s,,,ult .4frii:rm ~~5 ~AT'/... 1,D., t,; \RBtT, \.F., M.\IL\..~, S,M1-, )."'"\J}JDULA-. D,, l.o(KltAR:r, J,M J..L(L
\,e1wlllmyAm>cim/1J11 Bln1111ial Ca1111rm. P!ctfilm•ritzbu,~ South M.K .. tJ!\WSO~. J.E.. Ot..SOX. J,G. & •m•ts<.:. 5.A .. 1996. A FCC'nmbmara antigen
Alrlcn. 119-9;. from 1hc. hcarn\i'lt~r ngent (Cowdritt rumimmtlwn> rc.acti\'e \\i:h
105 1L\1G. o .., .. 1932.. Xote on the u.~~ oithe \\'hilt- mouse for the crnnspon of antibodies in ~me bOUthea'-tem Uo{ted.S1ntcs whht·Htlll!d deer
~lf:llns o! heonwatcr. Joumnl of the.Sm,1/J Afriran \"6l~rlna.ry .',Jt'lficnl (Od=lleu.s v/,gi11/111uu1. but r.01 rattlo, seru.Jo1m111/ oft\'J/d/lfe
,A$.J«fhtll:m. 23. 167-170. (Jl~'f!il$t',$, 32 • .;z;;-;30_

106 1tuc. o., 195;. Tukbome riokmt,iosc• In South .\fnca. ¾li-ancn in 126 ~U..kftM.-.\~. r~.• C'OE'f1Ht. ) •.\,\\. ft .SAUOt. -r.v.·~· 1988. 11lam PQ14Cnfog,1
V1wn11al)' ~ie11ci·1. 2, 307-325. onll ,\1iit."'.otoxft.'?s~s ofLic'ltSUitk {11 Suurht.'m t\frlro. Cape·rown: OMord
107 111.,r.. i,..,.,. ,\1..l>XA.'\OCR tt.A, a, \\'111.5$, l\.t.. 1954, f rtatmcnt o!hc:im,-ater Unl,·erslty Pr~<;
wuh t<?tr..m}'dn. Jo11mal of:hr So111/I Afrlc1111 \ ·e1,rt11llf)' ,\/rd/cal 127 J:OCA,...... Ii{.:\: .. llfZUU)l·~XHOU r, J,U. & lHKT, /lw l98?, Uhrastructu.raJ featurl"S
:Wocimfcm. 25, -4M8. of (.'(Ju.rd.Tin rumlnantium in midgut epi1hcliat ~ells. andsath.1~ glands
w. 1956. Alllm..al Dis<?lB>'l1~ ht Som/J Afrlctt South .Africa:
108 ilf.NNt~(;~ \S. of n1mi>hal Ambr~.,,mmu ilcbrat•mtt, Omft,rswpoort Jo.umot a/Tcrutnary
Central !':ew•,\gcney. Ud. RtiMteh. 54. Bi-92.

109 IIOLL,\:\"I>. C.J .. L()GA,;, L..J...... ~Hi.BUS, C..\. & kls'nC. )I. 198-;. The"S:CrOtogkal 1.2 8 1(0('.A,,, '--~1., .)10RJ.ARIA, S.P,. \"OICT, \\', P., XI.AR.IP.:, J• • UJ.\'lN", ,\,0., 196(,
relationship bcrwcen Qmxlrin rumuumrium and·ccrt3in member~ of IJtmoru.rration of cole>nfes of Cou'llrf11 rtm1i11a,1twm in rnidgu!
Chl•,gcnus F.btlichfo. O,ul~h,ft'pot)ft Jouruul of Vtmoriuary Rhrarch. 54. epithelial C'~lisofA,mblyommn 1•11ri~tum. Amt.•ricm1Jo11rmll t)f
~l- Vcn•rfnnry Rl!S,(lrr./!. 48. 3511-,360.
uo HOWE'I.L. O.j .. Pl':t'N't.Y, T.:-:. & HORAK. J.G•• 19$$9, The bOSt:Sta.uu of the ,29 ~(>t;\S, J-:,)1.. ~OR\~Al. A.A. & t>O~OVAN. r.L. 199.!). Dewtopmcm tl'ld
siriped mouse. Rhftl.xlomys 1nm1illo. in rotation co the uck \ l-c,on. of Lmnsmlsston c! Ct>1.ntrill non:nanffum by Amblyomma moles
hc:artwater in South :\!rlca., Ondcrsltpoort Joumal o/\lettrintlt)' tt:msfcrrcd from infected 10 susrtptfbluhecp. Tlo,11«•,1'Si"'1<,gt1J1 de
R~rd1, $. 28~291. ,\fMrt:illr \ "it1~"i11air-.,dtj Pn)'"$ trap(Mlt.'C, 46. l83-l88.
111 Huoso~. 1.R.4- ttt-.,"'01:.H..t;ON. A.)1 .. 1941.Somt• prulirnfno~ cxpcrimcnhon 130 1.\1.1/t', s.c.. Xl<X)U, ~. P.\..UOY\, [s.A. C\Rl\ C...\ L &: SUMrrio;,;. K.I, :99;., The
ll:lt ,·mv!,-at or heam,'Oter ·,,irus· In ra~. Jo11mal ofrite So111h Africa,r Cowdr/11 r<1111in1111ti«m groE opcron. Microbiology, l~l. 2091-2100.
\"~t~rl,wl)· .\1td/ro/ /\s$O<"{ntlon. 12, 39-49.
131 LAWHLl\"'Qi:, f,;\ .. TJOR~UiOJ. ~. WHfll:u\SO, ,\.1'. ft).APU\\,\. l";r,, 199,5. The
112 kllTOf>'OS. i,,. 1900, Hr<1oryof heanw~tct. Agrln.ltum//011r110/ ofth• serotOjtic:al mspm1tc 10 htart\\-.-ue.r immunlYJltion 111 C.."\ttte Is an
Cope<ifGa(Xl HlJfH', I,, .I)()...! 17. lndJea,or of pro~tiv-c immunity. Re1,11u d'Glttvtge ,ut,,.\f(vi«lt1•
113 1Ll!,1<)u..,0E• .\. , .. 1977. Study or hc;irr:wnu:rartd the cau-smh·e u.g,ent, Wtlril:alrt d,~ Pt1ys Tropl,11u.<. 48. G3-6S,
Cou'tlrla rumlnanrium (Cowdl) 1925), in ~1geria. Distrrttuitm :\bMmtlJ
132 1..o<iA!I:. t~L.. 1g.a;. Ccu:drict rumfmmtium: stability :,.nd preseJ'.\11.uon 01
lturmmio,,nl. 38 8. 5-Jl
1he org•n"m. 0111/mtepo,;rt Joumn/ l)fl'ettrlllal)' R,•.,.,,.n:lt, 54, ISi-191.
n-1 1U:\10uA1>t:. A.A. f# 111.01 Ki\.,u·, c •• 1978, Climco-p11th0Jog1c;al Mudy nf
l(J3 l.()(;,\r,;, L.t., UOl.LA.~D. (".J .. ~11:8US, C.A. le RISI I(;. ;\I,; 1986. Sem1og:tW
hrMl\\'.lll"'rin gn.lt~ Tm1m11r1wrll!':i1, um} Pnrn.~1·r11/ngi;, ;;>Cl, -1-:n
reJ:monshtp ht1wttn C.oJ«'tlrio nt.mimmrium and certain Ehrlirhftr. 111~
us ILliMOB1\Dt.. ,\,.\.,. ISLO'f"'1\).tlf. f., 19;s. Vresetvation of Couvtria \'c1mrt<1"}' Tltc>I'(/. I 1'. ;58-lS3.
n,mintu:rir,m ,u low wm1pcra.rur~. Ht•$Mf'C'lt in VnmJ:nry Sd,mcv. J9,
13,1 I.O(iA;:.., 1.C .., \\'11\' \fU), 1',C... QUISTl!.RO, J,C ~ MllBUS-, C'...A., t98;. 1'ht
337-338.
dth·elupmem orCt,u·,lr:'a n,mfllo11rwm in neucrophils. Ondrrs:ep00r:
116 uu1.EL,1.AN.A..&: DIW.\'t:R. G.., 1932.. lb.c use of dox~~ycline to coutcol Joumal o/Wc;ri11ary R~"'urh. 5-1. llli'-204.
~"1.rtwa,or In <het!p,Joun,al of1/t~So111/t \frlra11 \'mrhuir;· ,¼«int/011,
53. 23-24. l35 1.ou~-sJJVRY, c,P.. 1900. Tlck heurt,;.·m~r experiments••-\gric'ulturol Joumnl
l>{tl,. 01JJ<'O{G<H/fl 110/1'', 16. G82-687.
n; 10xcn,.si, F•• 1991. Protective in'tmunil)' lO hearr.\ta\er (Cowdri<,
nunlnm:ti1mi ln.fc-ctionJ Is acquired nJtervacdnMion \\ith In IJO litt.(;1<1I."'7.tE, f',ti~ k MC"IL\AD\'. s .. 1987. Cou...'(lrln mntltmmlum inf«-tion tn
Jdtm,tucenuall.td rkkeustne. h,jec1io11 anti lmm,mity. 59. ;29-i3J. 1he mouse: o rc\-i1,·•w. O,uk~U/XJOfl /ounl(JI ofVt'lcrinary Rt!searcl,. :>ot
267-269.
118 10:,_·c,EJA:\, F. &- 1,,._1<~ c..r.J•• 1&99. Cowdria rumiranrlum: tCC'ent
cieYclopmerns in dfa~no~c,c me1hc>ds, molecu.Inrcharactt.tn1..ation and 137 M,,e,n~'Zlr.. ,.~.1. & '"·"" mlO\'t."· ''-t .. 1!181. Th~ loolat{c,n :tnd cuhun- ur
,,icclnc$, "" MOULT, o. ~ll~OIJQU I, r .. (eds). fl/ckcusw and 1;1ick,usinl COlvdrln n1mi.'l<1nrlum tn nlbino mice. In: wmn-.1-tfAD, {.,ff..~ c:-1n~s. ,.u ..
lJlm1Jt~ m rhc 1'am of1/u; Thfr,I ,\llllcnlum. Purlt: ~~c\ltor, pp. 363 386. leds) .. Tick 81o'o,gv rut,1 c.,,mrol: Procrw/urgro/an J,u~matl()nnt
Jl9 10~:ti~IA..~. F .. ffllEU.\tAX"t.. M,J .. lil\1£RE, <'-~UH.flNtU:OAC, G.. 199t.Anugenlc Confor,11"~· Rhod•, Unh·ersiry, Omhoni•town, South Alrioa: T,clc
R,,search Unit pp.33-3,.
dtven,;fl) of Cow<lrilf nuni,rumium f'IOhU~ d~terinfnL-d b;· cro~-
l1nmunity. R,m:a-rrh In Vtterl11ary &l~nnt. 5 t. ~4-28. J38 ~1A.ll,\.X, S."\1., A.\'DflE\\, tt.R.. Tnr;JL, X,, UUIU\U\GE. :\I.J. It H.\RMtrr. \..F. l,SS,

12.Q J(>W}l:jJ, ~ , f._, 1'1tl~~1A."''S", ,1. 1.C.. L>ft.GftOOT. \L, \'A.'" ):OQn!.~. P,J.S.. & \"A."
lmmwiisat{on of sheep against hcanwmcr with inncd,1ntt.i-d ,m,'llrln
o•RZlr,sT. 8..\.M., 1991, Competiclw cn>.)'1tt~·lln~cd lmmuno,10rh<>nl rumlmmtitmt. Rcs,·mch m Vc"ltri,:m:r :;cluncq, 58, ,l&,.49.
:ss.say for htartw'1ter UIDtg monnclon:11 an.tlbodtr; to a Cow(lrfn 13iJ \t,\lt,\!"., S. \1,. \tC(:uuu;, T.C.. ~[l:)IU~ t,.)I., town:. M,V.... IU:<CfJA:\, 1'..
n,111/,inmlum-,pccific 32-kllodallon pt(ltcln. \'tll•r/11n,y ,\1/crob!o/og;·. RUR.\Xf;mw.,. s-.n. & BAIU:!trr,,\.r., 1994, Mot«ular cloning of o gene
28, 199-21 I. encoding ,he lmm\mogenfc2l ~Oa protrin or<:o:i.'tirl.a ruminomwm.
12\ JQSG.EJ:\!':, P .. UtLJ;;SfUiftG, G., FM?./r,~r.N, r,,r,,, ~lll\-"ll.A, II .S1FU\\'F:'1.11tlll$, J., Mi~,wlc,gv. 140,2135-2142,
1968. ,\nug('fllc ditrcrcncl!S ~,ween stocks orCowdri11 rrm1/Jta11tlum. 1110 ~IA.IL\!\', S. \1., l'l!t'FR. T,f., ~'1U, $,~1 .. $tM8i, Ii.IL. SUlt.\",'1,, 1\.1\, & &\SIBl'T.
Re.featth in Ve,enilnrykien~. -u. lSG-189. •·"" 1995. Ulborntorpc:ned Amblyomma helm";"m and .,111biromm11
l22 JOX(a'.J.\N. JI•• Z.\~'DSERGf;"X, T.1-\., VAS Ol! WTl!L. P_<\.•• Ot CAOOT M , & 11(4ric,garwn ,,ck:, differ tn 1h~1r ~useepubilily to in!ec:11on with Co:(.tfrltt
u1t1,~6tJftO. c .. 1991, Th&nck..bomi! RrrJ.•u111ihCooi·drf(1 rtm:tnanrittm has rrwu'uautium. Cpttl(mfolo1r-•a"'I l11ft<>ticm. 11 S, ,)iJS-JS.l
o chJrunydin-file dcvctopmi)m.al cycle. Omlt1rt1e1p,>0r1 Journal Qf 141 \li\JIAN, :.. "·· 1•n111. T.f .. .)l\H\1, tt,11, f; 6URllll>Gt., M,J., 1998. PCRde1ectfon
Vclc{rillc.tl).' Rtstarrlt, S8.. 227-237. o(Cowrlria ruml11cmtfum in(ec:cion in ticks aud unun:1J.s from
l23 ~'"YAM.I. P,w '$- >:AGIRA, ,•• 2000. l11e role or parnshk d!se3:SeS as cau.scJ hrom,-.1cr•cndcmk regloM of?.irnbobwl!. Annal, ofthe .Vcu Yurk
or mort3llty in aanle inn high potential uroo ofcentl'lll Kcnrn, • .Atoc/emyo{Scitm:.-,. 849.8~,.
quantitathl!'anal;'Sl:5. Omh!rstepoort /oumnl o/\JitfJnnof)' Ri!Si!fJrCh. 67. 142 MAH,\~, ~.)i.. 1•m-n. T,Ji .. 51)11i1, tl.Jt .. h't)ChN, " ·· CAMUS. t!,. s.,n,F.T, \,f. ¥
!S,-161. omm1~. >1,J, 2000, Compomon of efficacy of ,\merlrnn and Mri=
\U .,....,.., c... 196o. Rir'.on.slol lnfectio11 (hCilrrwo,c, 1<nsheep and go:us in Amblycmm<l ncks 3$ \'ecto~ ofh<'artwmc, (Ccfutlrit,, umitmmhlm\
 1tli<(<ion by 1nolcc1~ar au•l\'Ses Md u::mmhslon U1als /oum11/ of of an ~7.~1nt'•linlr:i.-d lmmunn~<1rl1-f>.Jlt ,\~y. 011tlrts1~J)O()n Journal Of
Ptlm.~tology. 86. 4.;....49. VN,•rir.nr,• Ri~Mr(h. SJ. 205-207.
143 !l,L,\J-L\,.~. "-M .. s,-srrH. G.t .. >:UMDVL.\, o,. 8Ufl$UO(;C. M ,J. & U.UtBUT, \,f,. :!001. 161 ...,.,,r ,. \\,O. 1!t,3S. l'hc blc.i"burt. U>nma/i5tu~ nlbifro,a~:ind thr Blad
Reductfon in mon;sbty from hcar1,,1m!r in c:ndc. ~hccp .JOd J?Oa1s \\ 1ldt.• SttsJ Qmodwetcs,;.:mll ;1..5 t{lrrtmuftw.anwarcr. Oud111°Sli1/KHJrl
expcsed to fie?d th11l.h.-ngc usini; nn inatti,.·ntt:-d vaccinc- \',wri11al)· Jmtrnnf aft',•:.•rmnr:r:.tt(!ntl!cmd.'\mmnl lmfu.ury. 5, 3HO.
Patt1S1tcl~•. 97, 29~8; 162. ~t=n 1. w.o J.9).~. Thr immun(ty il'l lwurt\,'..tH.~r. 01l.(1'"11ttJN>0n Jqumut n/
1-1,4 M,\H..\,',., s,,t., \\"A.('ol.lF.LA, ~.o.. \t(I,UfRt, l'.C'., 1'Ult,\.\;(i,fk\\' \, f .R., \\,\'is.IS>:, t ·turlrituy .'ir.tttt:,,•m1d Ammdf J,r((w1ry, 13. 24~283.
L\. "8..\RBE'i. \.F•• 1992. r\ cloned i)~,\ probe iOf r.rmdrm nm,mamium 163 :..tJ i.l. •.\' .0 .• 1~~0. Uhm,n in thl, ltt"".1tou.mt ofhl'Mf\,.~h:r. /om no/ oj thr
hytmdizes ,,,th eight hcmnwnccr .,..trains ;md dl'U.'(L'\.m!t'CJ.4:d ~het•p, Soutli AJrictw \ ~rrinary ,\lcdtct1l .-\s.~mw,1. 11. l.5.
/oumal o/C//11ical ,\/kroi,/o/ogy. 30, 981-986
16.i ~• n,. ',\·.o. 19,17 TI1c uun"\ffl~~ion of tw.in,\':&terb~·Ambl)Y>mmtr
Hi>tOnqu..: du pcuplcmont tw,in 1.11
1-;5 l\~AtLuH.o, J.t' • \ll\lU..\AD, ~ •• 1~98. ,,,am1,c,1m11, Oomu'.. 19:09. S<>ul11 ,Vrium MIA!urv. 1, 83.
d• l'lmroductlon de 13 lique t\mblyomma ran,gnmm dons lcs !!cs
16s ~urz. ,.,.o., 1•167 nw <pldrmlol!Jllk:tl p:onem ohinti, procoph~,.i nnd
fran,;alkS des .\ntllles: :,yruhh<) bihllr,gnpluqce. E1/rno:001«lmw. l,
p:oi01.oa.l r.oono~~ tn n:4mon io game- pre1cn11Uon m South A(nc;i,
19-36.
J11umol 11{1t,~,;o111l1.Mncair \ t/f'ft11nry Mrdlcnl Auoclorlon 3N. ll!l-141
i,.;~ "l<\.RTI~U. o,, 1997, :o\nal}.,f~ of tht.' imnlunu r~pon,t• nf rumin.ant~ H>
106 ,·1:1,:r~w.o.. 1q68. Hl!a.r'"-'ttlt. 8'1lft!llu dt I Offi,·t· lm,,r,:r,:fcmnl d,1s
Cou-dna ,-,,m,,wmimn lnf1.-ttJon. Ph.D. tH0>1>, Unh·~t>!t;· of Uuech,.
41,:.ootte, ;o, J:!!l-336.
Ucrec!u. The Xelhcrloncls
167 'l,"frt7. ,,\',O," .\U..'<,,\SOl.:R. n.,\, 19Jt. Th~ lmmonisa1fon ol r.:tl\·('S a~lln>1
u; ,um!<.L.7. 0 •• 2001. ClttAU·tM\'T. aP 5r5. ,;i6.; PQtntt- .a. Pi1rC" Ctideti..
bc,in,,:,1cr./uurnt1! u/1he Sm,th !tfrlr.·,m \ t•1,·nn<1() \frJlml.iluocimlo11.
Guadcloupt'. French \\e,t lnd1es. l\,r,on.il conimunir.mon 12' 10:l-t t I
ua .,um~£L o .. ~t\11.L\lll>, s.c.. co1sst. ~- SHt.nctaouoou. c & an~.,m. A.. 168 ~u1-z. w.o. ,\ll.XA.'\;Ul.R. J\,.\o,. ,0£.L\AH, r.f.. 19.1;, StuJ1C~ rm unmunay m
199-4. Protection ofgo:ns ngalrut hC3n\\-..tct ..1cqum.,d b>· immunt,nt(on hc:im,utc:-. Omi,·rsrtpc,on /cmm11/ r,f\'c•urtnal) ~ct1•11a-ond .tltimt1l
\'with .inac.ti,,;ut·d: ~le1ncmary h()dit\ of Coudrlu rumi(Umtbmt. lmlusrry. 21 z.;~?52.
Vtlbrl1u,,,-· Jmmunolom·mul lmmum>J}fltlu,fl)g;)·. 41. JS:~!63.
1fl9 ,OM\ \I. ff..\ .l , \."U,U-·h. II n. & \'t):,;).t,IL C,I. ll)90 fnfl:ctfon ra1c1; \\ith
u9 M.\RTIStt. o .• snr.nrnouoou. e.. <:OUl\AOo. P.O. & nt:ri.S.\10.•, .. l99l. tu 11i1ttJ Cou'dri.tr r;mwmunwn 01 n)·mph,, ,,nd atlull .. qr ch~ hunt U('X
lnf«tion of bo\inl' btufn c:.ndorhclit,I ccllti.-bv Cm,.,lrlcr rt,mfnm11I1uu. ·'1n'1{rmnma ht"hrcu.•um cOHtcu.-d fn lh"" h~?d m lunbo.bu....-, Vim·rlnm;,·
Rcsmrrl: In \ 'ert•ri110') Sclencc,.5S. 258-2.60.
0

Jlarmiralog, :kt. 2:'i- 283.

tSO '1ARTI.SEZ, o.• S\\'lSKhU,. , .. (A\tUS, L ~ 10~(;>:f--\ ,. f .. 19go. Comp.tr.lh()U '"" '-OKVAt.. 11. ,.L,. :.,w111.:-..:n. f. \...19i'9, 'T11t-(1)n1rol ofhc.1m\':lttrln
de trob nnttgt.tn~ pour le serodiag.no~tic 1,h.• la co\\dnose p.u 7Jmbai:n\'r." Rhodl.."'S1.1 . .l.(ml>t:lm.,c llltotlrsid -l;.1·1cu/tuml Juurnnl, -s.
lmmunonumescencc indlreete-. u.•,·w· dit,•raf,:c' 1.•1 ,i.- .\t«fecin, IGHE;.
1'4r/ri,r,;lmtl.s Pa;~ 1'ropi,m«. ,13. IS9-I '36
171 '\:VI~\..\.., U\AUCT ,\ .F. UUl~lOOl". '1,J. ,1. )1.AJJ..\.""i. ' · ~ J:001 0~\
151 .>:.i.,nir;:ROs, c .. 11.\RR.L s .. CA,.,1vs. c: • Goour r., 198;-, G.:m•uc ft!"S1stnncc or ,~ccuutio1~ with mnpt followc:d b\ protl~n t>c>Mt a\t.l;tnl'nL, ptotuctlon
Gu11U<.1Joupc mt(i\.i.,: gO."'lb.HJ lw~n,v;u"'· Ond1'Y'1f"i"lmtJrl /11,mutf ~{ "~Jn"1 chulfon.8..: with r.rtu.t.lrfll rammm1litut4. the U~t'i\\ ofh\'linh'Ottl!
1~,rtn111)• «=rch. S.I, JJ7-3•IO. \.~trlllt" 290-1 1.-11.
152 M80t01, M.,1., 8U:l\f.R, r... r., .. kttU(f\\\(1\'.;\ r .. C'.1UIX( R, ,,. 6l JO~C.l·J.\S, r., li':l. '\'VU~.\, \,, ~WtAX, ~.M. AVRPIOGt\ M,J,. ,\1(.(,UHU:. l,C,. kllMXGIRW.A, t1. "-
1!¥.19 Validation()( chc fndil\'Cl m<tpHI cnl,)'ITle•lln~ed ,mmvno,o, bcn, 0:-\.1HU l, \ ...... 1998, ,.\ D~:\ \ uCCUll~ IU(IU!·Clfi. mice 3g~1n~ the ric~tf~ic.t
a..~y for dlDgn0Sf5 n( l~pt:rhncntul Co1tvtrm mmmrmoum m(t'rtion 111 .\Ai"nt CmtYirw rutnlmmtium, Par1JSi1,, lnummoll)JD-. 20, i a 1-119.
smwl ruminruus. Ctmic'fll $: Vinguaslic; Labomlflt)' lmm,mo!o~·. 6. 17'J OIJti.Rt., :. r , . ;,. tnrz.umi:~HoUT. r.u.. 198;. l·lc-..1m·,a1('f in host'lj: 01hcr lhan
66-7?~ dom('~t(C' rnmlnnm~. r)oders1epoorr Jnurnol ,>f\',•1•:ri,wf') R.t•sfflrfh. 5•1.
tS3 Mi.J.J7..tR. M.t.. S-OM\'AL. R,,\, k 2>0XAClllt, r.L.. 1!!9.3, Eflecb of lJC'.k 27t-2i5.
lnfestanon and uc!.-bomc dlsc;ise Infection< lhe~m,ater. snopln.1110.,i~ rN ant Rt,1, 1•.r. "'" 1'1.Zt110t'\1tbu,. J.u.. 19K7. 111.,• produnio:i o(he..1nwatcr
and b1\be~io,-.h1 on lhl· J.tcwuun a.nd wc~h1 Ji:Ull of MU&honu. cuulc in \'tl«Jn~. (),ut,•r1:ttpa,ort /ournnl o/v1,c1.,•rl11nry Rrs,Ytn:11. ~ 48>-488.
wmh-easiem Zlmb;ihwe, I m/>fcttl Am mat 1/cr,/Ilr and Pr(l(/t1cuon. 27,
l#.$ Ol:ILft[M, P. t C,U\1.)JO\\'. D. 4.. \'.\S" A,\t\rt.L ~fl.. 1~88. \n alll1 rnnth'e
129-1•1·;~
u-.l3trncnt for hc:mwnll't. Prve~i.'aln~ ofth~SnutlI Afrim11 \ ~/i!ri,uuj·
154 \U.Tt:U.10.:o\.'111, S,C., UTT1L\UU(, C).. i\A\IQL R.J.,OUOl·t-.t\:," 4,. f,"U!,.:., B.,.
.AJA'tl('/uti<m. Bi"•im.cal CongrL~~ rh:h?tm11rit1.hur.;. SouuAfric-n. 82-88.
ig*'>. The omng.e,yclJowp~gm.C'm of he3m,"'ater lnf«ted ~om goats.. 0

C:,,mparar/:y RiO<'h<rmistry(lnd Plr)f'to/og.·, 8 · Compamrl"· lt:o<l11•mf'11)·.

176 OH\~111. ~ .. Pt~UU~'\. r .S USV[I~ ft.. :roo,, Anah'i\!i QI ltQttS('ripuooaU)·
..lC'fi\'<:- gi:nc clu-.:cn of mnJor om,.,. mumlJromc pro1i::in mul11~tn~ r,un1iy
81. 299-3o.l.
in l!.hrl:rlrw canh nnd £.. clrn/f..~11Sis. fnf~rwn nnd Jmmuni:)'. 69.
135 ,:o~HKO\:strr.1, s...n•• 19.r;. r.nm1nt.ont-. b) n?adt:f'. . sr1,11:t:t te>t>. &?. :!083-209J
1~6 C,ll"\U~. /\ G. Q')()l)\l,\X. LS.I; (:IL\11';\, ,\,. ,98(). fi11· Pls.nrm(tC'rlltJgiral 41 Rlbc11t11:tnuil RNA: tt key IO ph)·lo~'°ny.Ji,un:nl
ul!'-f:..... -..,9 wotsr, t.:., 19~3
&u/J of n,crap~urra. London: l!~llli~r• lindall u_r tiu• f,"(/~rt~t11m o/.;nic:ncm, Sacit'tia for E:rµ.:rim~11/JII B,otoKJ,', ;".
1Sj )IUl.'.IIDll ..\.\\'., CJJA.,.s.&OKO. T.• u't:...UL\GlL\~. c.,... Pntlt. l _r, J.:RU~hl\, n.t.., l 13-1?3.
\1EDL0. (;,F•• ~HA.~. $.~1. i PlJllR\' ft,D,. 1q99, .\n b<J:f?Ssmcnt of th4_) c·a tJ\\T\: . ,.c..
lmuum, . ., .. ~Ku<.1·11, 1.,,.•
1"'~MU,, 11 ,.. :!Fl,
cmno1nk ln\p.sot oflur~nwlltt:r ~Om'iltln rumtmmw,m tnfoclfon) and F1t~~fopatho!ugfcal k,tturesc of htar1wa~tr ut~h~jl, Jo;,rnnl ofti:.· ~u,lt
it~ conuot In Zi:rtbahwe. l;r,niemfl"t· Vt1t'J11:nry· .'-1.:idlrln,•~39, 173-189. .Vntan \ ,•rrrw.ary .~St>t·tatiou, 4·\, 3ffi'-tU.3
158 \l'\\·\....:at. t>,».. )1AJ.H...,, ~-, ~'iANIUt J.):., T,Ut\CIL\. r..L. 41 ,tatt.\'IJL D.1.. r.c. ~-,1c;:r:\\ .\ J' T 1- ;gg:
li'9 "" '\t-11, c-.H • tlO, t.,. U lAPfr A.t- .. '1f(•Ulftf,,
S!,9S. Jmmuni1.4rfon of rr.mic l>y infection \\ith 0,11.·drin rumlnanrfwu 11u~ 1mmun,11nrm.-cll\t- Anuptuinm mnromnf( m.iJo: '.\uriil<'t' prot('ln2 I~
dtcu-' T lpnp.hn~-u.~~ thilt ret·ogrh1.t-:iu1n!ogQui., mftee11."d '-'nt.lmhcl!ul enc~ b\· ~ polyrnorphic 11111!11l(tne farnll) /n/trtlQII and /mmmrlr;·,
c.US ~nd monoeyie;.111[..:rton aud Jm111<mll)·, &:>. 18:tS-1860. 62. 3H(>!!-1$11i,
r;g l'I.J .. ~\'.\!\}IJl. u ... fl \RBCT u -." '\I.\IIM.:. :> ,1..
\1\\.o\.'(Cil. n.,, .. ~:O.."!i.LY£H. 18o .rut.r.tFiJ:i.t . A.~ 19,nJ, Cht•mo1h1Jr.\P)~antl ddh·cf) ~v,re:u'!' ·
\998, Mafor entigenlc proa,hto. I J.nd? ('If <:u1etltr'o tJ1m{,:amlum Jtc h;u.1ntr1p:sra~iH.~ \ c:rrtnan· Ptutl.${tolog.•. 5.i, ?:?J-248
wgcu !or T•l)'mphOC) tU re:Sµoruc"' ol lmntunc t"3.ltle. •.\Jutal( of1114! 1\tN.1.t 181 l'm'IC T~J., &\.RBlT. \.f., ALU'\tA~•.\.fl..-St~Ult ,,tf., ftUlL-UOQl.. '!.J.S.
lark -~<ml,•1t1)'<>fxie11,,,,_ 11>!9. 372-37,; M ,1L\.,. ~M., 2000. Det<'tliun u( u,e
iutent of ht•1&nw~u:ir. Cott'tlrlo
160 '.'o.(.117. \,\\' .II .. \'ILJOCX, Gj .• ht1.Ul0l;SIIOUT J,l).. 08ln.!'-':, t• f.. \1Mtl.N. L h nuui1:,1mf11m. fn .1mt,J_mmmtt 1kk .... by PCR: \',t1fdaHou and jppl!<'"~t:on
\'fa\l.E.VU.~. X.)..1.f., 1986. Ot.U.:.'Cli<rn of CtJ:,'(/1/tt ruuwmmwm ontigl•n of thr ;L~q~· ro 11t·1U uch. Jm,r,u,t o/Cllninil .\fr'cMlliof~. 38. 1539-
and 3.nllbod)" during tho tours~ ofheart\',.ltt'f ~ < " in sh<!cp by mc:.1n.;. l ;.;-1
 Mennwaccr 533

182 P(Trtt, T f., tUtl~ON. '.\,ft,, P"JR.H..... n,o .. ()°CAt.1.\CitL\~. C,J.. \\U)U), (i.t,. H(ri·t•lts,n rr,mlnnma,m ·CaMlry, 1923) fn.he.lt1MUcrbrnln.,_, Th~
J. ~ \tiUIA.'., $,'-1,. l999,
SMITH. C.,£... MU.)180. C .• H()k,U;, f,G .• PUfCflll>vr \I \~<WrlmnyRm,td.J; 4?3-H4
Cowdtin n1mi11ttmium in(~1io11 in I ic:ks in lhe Kn~sN .'.\:,1.ionnl l>:trL 201 1•uns1:u. flt .• t!t8:- D\1,c.•Jopn,t•n1 of tl proph~inc:cic r.:glm\.' u~1ng
Tl~ t 'dr<'t111n11>· ,qi!,:ord, 14.11. 30-1-307. Tcmm)Yin, l.,\ IU ,"-.,i:tt m th.: lntroduc:ti<m or~u"'cfptib~ t :. mlc lmo
183 P"CT'Ek. T.F., 13\lftkrt>fit. ,u., }ii\lt\N, s.M .. 2000. Compcumce of the he~nwute.r ttndemic: are.1.~ of ,rnca. Ond,•r..r~1HJt1n }01,mal tJ/\l•ldrlm1ry
.Aft.ii:.t.n COrtOi1ie Ui.'.k •.·lmblyunmw urarmQttum r,\i;-,sri: IM>dida('t;, .a.1; a lku1atrl1. 5,1. ;()!1-512.
\'OC-tar of the og~m ofhconwuu:c {Q)rn/r,q rummrmtmm ·• /fJfm:nl of l<>:.t ruR'-:t.U. fl.t ~,u, tu,. l'.I> &- ~amoorn. t •• 108,tt. nc,•t.•lopinont ora
P<Uasiwlogr 86. ~38-411, prn1>h)'l,trti1,· :~im~ us.1ni:,JonR·Dt'tlnA cecr,.1<,.'\('hnc tor tht' corurot or
t811 PliER, ff-- r)t.l'I\I.~.t.., AARll-ft,.\.J.•• :,.:OIWAt.. A--\•• "1MIU, ·-H, 11:tLU , P,1, A. rcdw;urr :·md h,~artw:1.1t:r in ~u"("t,plihlt! nnh.• movt-d ln10 an endemk
,u11 ,s. s )1... 199;. Developmen1 and cw1u.uh,n or PCR •l~Y (or arc" 1ro;umr i1,11m11/ Hl'l•ltlo n,u/ 1'1od11c1fu,i, 21. t 1-19.
dtt~tion of low Je.,·d.,. uf Ctm.Ylr/11 rumwtmrium inft'"CtiOn in 203 kA.\US.:r-t. J • u,tt.,~£JIG, G., 1910. Prescn-;i1ion of a ~,rain of Comfrl(I
.~m~omma tick. not detrctcd h)' l>NAptob<.'. 1~11r11nl ofc:Jm1«1I n,mtnamlrmt b)· ftet"..im1t. Re:•ur tl'Etc,·nt:t<"I lli! .\fl'tltefnr \ '.!t~ttm,lrtda
,\/lcroblofogy, 33, t oo-1 ·2 PtQ'$ 7'tJJ/Jf(rWS. 23. 313-316.

185 PKnH. 1 f .• tl'"1n tt.0. 0 'C,-\IUGIL\ ' · c.,.

~,1:ou,·. G.1 •• MU\UIU. ft,. ~<n Ht'UU't foH ,SUL'\(>'1,' \ f'.H •• IL\K1U~os. K.U .. M,\UA:S. s ,, .• DtTAMOUE, Mr."
,_.,.J,Utn. ,\.F~.a.. ~.\H \S, S.M,, 1999. Prcv.ih:nc~ oi Ctmdrw r:.,n,uumrtum UARflr.T, .\.f. 1998.. 5icqucincc hc1c1ogrn~it~· of 1hc maJor ru11i,&~n.ic
bf.c1lon In .{1nblromm11 llr/,rntum ~tks fron1 h,;anw•t«·cn<ltmlc pmrefn l gl"n.~Jrom UJ:,vlrtn nummmmu11 Isolate$ from difftr('m
=• of Ziml>l\b1,e. Ep/J/1•m1~/IJ!lj•t11ul /11ft'fti1111. 123 &m-316 gc;-ogr,,phicaJ iireib. C:lintrtd m:d [Jf(lll.110J1ie Lal.N)mt<Jry /mmm1o!ogy. 3.
186 Fl!l"'U, T,l<.. "o,..~. 1 C,I. Rt:OIAV, \·.. 1~87 Thc~<olog:.·ofth• \fn.-an ~ 1;-122.

\'tttors<>fheanwaicr. whh panlcu1:n rtforc.nct- 10AmM],'Omma oos mmc "nux."'u. 0.L 198.1 Tr!oo II l'htlichitae Phll!p l9S,. ,,. KM1>.<,.
>1
licbrncum and Amblyomm~ ,-,a,lt-gatw,t. Omlf'rs.tc;HK>~ Jt1ut11al u/ ,.'ft .. tsOU, l.<,,.
cJ..,. &·rg...-, ·, \/umuilu/S,l~J«·m(1l" &,u,,.rw!(l,&l• 1°54,
r,,..nna,y Rt.warcl,. S-1 381-a!u. Rnlom,m:: \\10tum~ & \\ aU.:.m;
tS, PU:.~ uR.. J.<;.. l9i'O, F.lr.c1ron m:icroscop\' of Om·dritt Rld.:tttsin; :110 MO,:-<)µ\\. )t., ~,rn, \.,.\\,. Dt \\',\.\l. n.r. UIJ l•U 'S,'(I~. J.L. \',\;\' r;~,. L. ..
,Umlnamlum <Cowdty. 19:?Gl ln :hi: endothelial cell~ of the, t,.nt.•br.11r- o,u;rr, , i.•ft ldennfir.,1l"n ur th• an1i~enlc prot.in, ol C11i,-,J.;n
h0<L O,:d,r,r,p,xm founUJ.I o/Vcterl11D?' H,.,-..,,rclr 37 67-78. rumltttmtttmL (Jmlrnc,•µoort Joun:al oft"crh•rituuy H...-sw,rtlt 5':'. 21~
188 PlE-"AAR, l,C.. 8.A~S. r ..,. & \'AN OER M:E.IW-1:. I.L,..Ot 8., t966. Smdfcs on 1hr 221
pa tholo~· of hearn,•mer <O>wdrut tRicl."i."ttsia) 11,,,:i1:muium. Co\,dry ~o; nun. t.~ \L,UU.\J\D. l.t ('.\.\HJ< L. OU'ftU. tr... ~l>\llt(KO'\ c;; .. ,m.
19:?6). I, Ncuropttholo1,"cal changes. o,ulm:rpt;On ltmmal of 01:turtcn,c: or t•,1pri1W fouro~it.\ ;imiRcn~ ta.Ai in Creo!t1 t¢Clats
liwn·r:atj•RMl!ittrh. 33. I li-1:18. ,u1;.et1ptihlt? r~i)l.1.nt to hr;im,·nter. R'11 1m•1/'Pl,l~J;c r1 ,ti.• ,\:tidttin"
189 roou. 7.0.11 •• :.S:62. Flock 1mmun~1ion of .$ht..>cp auJ ~o;at!, aguifl:H \ ·c11~rl11n/url~, t•a.:. TropirrttLf 46, 20;;-21);,
ticanw,ner Port 1. Jn\-CS:tigatlun~ regarding rou;lnc: flock 1mmunl!hltion ~o8SC.t1X[1D(ft, 1., G2Ulf.ftf. !li,L•• 11.\.'X"-'X. C•.\t., O(OA.SO. P., Ptl.111.\Jtt. I;•• SHl:U,
of ,heep. Ja11m11t 0/1/1,som/1 Afrrc,111 l'etcrmary .\/;,,tcca/ .-mlon. 33, r.0-1 ••1.J.11.As,n ,1."' lfJLl ,.,·.. 1999. lndutliOn of C:1)8,. T C't-fis u.smg
3S-U hctcrolugou ... prlnt~boo!t immuni<n1lt,n "1r.negit...-;. tmmw,ofogfMf
i.so POOU J,D.n. 1962. Hoc,. tmmunl5.ttton ot sheep ~nd goo1t., ag~ln.~ R<'4•ft.•:c;. l 'i'O. ~38
henm,ilte-1. P.ln 1 Prctlm1t1ttf) c.x~rimcnb on r.oci.. fnunun1)41Linn or .209 'S"c.tl RUJVlm. 11.s, 198u. A ,,mplt' itthruque to, che tQllecticm of brain
goab. Jc,unml ofrht· Samh AJ1;cm, Ve,c:rht<uy .\ted,r.nl oW(>('w1lo1t, 3:t snmplrs, for,~ dlngilm-1s O! hc;mwnwr 7f'Qp m ,/ Amnwr tkohO tmd
?;il-:i62. / 110(/ut/l°IJn, l:f ?.i-291

i.91 rfU- rz~tA,:,., C., 11A,U'•lt1 D,. !i,.'rOl'll.fiRt), O.ft,, 1:UI K!•• • .>,,A, & Rt~Hl~A. ' ,1 ,995, :no , 1'!\tU. ~.M .. J'nl "· r I •f ,10 .." f.01 \'/\, u.. 11.\MET. h,P.. fU~Ul: I \.,;, •. ,.,
lti,S rR~i·\ gene .1c:qm·ncc of "t'i) Rid.'l•t1$/n lwlmintl1o«11 .ind th MA11Ai\, ....,,,. :001. ~\nuhod~· f'e\p0f1~K tu MAP l B 11ml uthcr C.(jwdrfn
phylogenc1lc alignment Mth meml1<>rs Qfthc ~cnw. EJ1rlirh/11. nm1lmm1fumnndgcns are do\l,'Tl n.,;:uluu..*d in c.1nle chullun~~d whh
/m,...1nali01Ull Jour,wl o/Sytremmlc /JactrrlolngJ•, -l5. 2{)7-:!1 t, IJfk·t~nsmlned hem,w:tt('J. C/1,ural qnd t)ingnostk Laboro1q~·
192 ~ROV().,"l . .... & SllUIUt-,H<"IUT 1.u.. 1~117. The h:\t0rlc:,J backi;round 11nd
lmnmuat~·. A, 388-39U
glub;d hnponancf." or hean\\:1ter. O,utrnr,•P(H)rt Joun:'11 OJ \'rrermmJ ~U :,.HAll'.(.'l•IAHI • \.;'I., Kl:Vl .k"!, t:•• VA~ .\M\11 L, ,-.IL, .,WA'., t. I .. N \ '.A' 01. '-: BtRi,,
11(,ffiir(h 3-1, 16S..169. 1.~., 1~8 Tivatm~m of he,,n\..-atcr. pou:n1inl ad\'tr~c: 1.1ffi."C. U of 0

rurosl!ml<lc: .t~m(nJ, muiun un rtrtnm hu111~>Sto1lc p•u-::tmeh,lf) in

193 rROlh~'t. t 198;. Dia~nO'$l:e.: ofhcartw:uc, at po,1~mo"ein In nimlnam~
11Qnnnl ~hccp. /r1rtmnl 0/1/fi' .,outl, "Vrit'dll V,'f<>ri,uuy .4J:socl'1l'i<>II, b9
tnd thecontirmi\tfOJ1 ofCtmHlria ,u,nlmurrium in mic.,... O;ut,.,-st"'J)('lfnl
1211-11,;
Joumnl "l Vct~mntuy He-,;rarch. ~4. au1-:-w:1.
:n2 irnl>'°S, n.r... usu,,, . \S~.• MOA.fl'I!-. J.N,. MUHllll!.\ll.1 ~•• J\OUJXK. \.
19.:1 fROZIU,~"\ L....198;~ He.krtw:ucr. fhc 11101·pho1og,y Couvlrw or PIUC lll\ff.U, ."1,0, :"IH,'-U \, II.Cf,. TIURLWl'!U. t,.,,"·• HUYffR, A.Cl .. MR.\UU:\. I..,
wm1r..tJmmn1 and ilS 5,ta1ning chn.ructl'ristlcs. in the- ,·crttbratc ho~t und
.-r t1l.. 1987. f'rottttfon of c~udc ng\\ln~1hi.::.1nwn1,·r in llcusw:in.i:
in l'(lfo, Ondir.sup,oarJ Jtmnull <>Jl',w~rlnnry R1,1~t1rcfl. S 1 l '73- 176.
comp.i:rnth·~ .a:uk.tcy of different mcthoch again~, n:uur.11 nnd
19a, Pk,0'1.,MS.'l. 1•• 198;. fh-' p:.\.lhology ofhe:1n,,-a.1C1". Ill. A n.'\icw. hlnod·d~rh,-d chall~ng"' 711,• Fr111rlnn')' R/Nord. 120, t3S..138.
01:dersM(){)Ort Jour,w/ aj'\'t•r,-rinnl)' Rei,•tuc/J. SJ. 281-286.
213 ;'11M~sos. t1.,,
"Wllf\'•.w. 19.;J. Exp<1rimon1s wi1h .iur~unycin
196 PROi':~A"\, L... Rl:.ZUIDf-~IU)lrT, ,.o.,.: PATrnsos. c L, 1.986. ft(>.J.rrw;u.1.---r: :\n h)'drodoride- fhc us~ of aureomycin In the lft'..tlmtJnt ot
m viuo s,ud~· of the ultrastrn.c ture or Cow,trra nmtma11ri1m1. tu.-am\·.itct•.-iJirtun/ Ri.•porl ofrlu· \"t•1tt1·Jnu')' Dt!µartm,•rtt ofKtnya for
011d.'tltel)OOrt Juum~I aflvwr111t1')' Ra.<eurt'/1, 53. 1113-!jS 1949.-27.
1.9; rROus,a. &..& nu PLLlil.!olS. t.L.1985, nu~ p:nhol~· of hl'am,.,.111:r. I .\ :.:w ~Milli, (;.t: , .,~a-LMSO:\', f...C., llt.rRfl.lO(tt. "'··~· 1•rrnm. l.J, tt M,.,\li,\'.',, ~.M ..
s1udy of mtcr inicc-tcd ,,,rh ,hr W,•lgc\'onden ~u;un of CtJUv!r,o 1998. GJU\\ th r.,f Co:,'ll1ta rumuuu:rlwn fn tissue c:uhurt, endoth.e1ktl cdl
tumin(lnrtmn. Or,d.~f'$Jrpo,orr Journal of\ ~,r~rtnr.n• f(~arth. .:;:?. :1--;"'9, lines from ,,ild .-\frican rn.1m1mJl!h)aumaJ o/lWltl/lfl] ()Ue1:.J11.'S.34
-.98 rnu,u:n.,'. t, • nu s•a t:,·J•,. M-, 198,~ The- p.11hulol!o'Y oCht·.1nhlllt-t. JJ_ \ 297-304
,tudy of 1he lung k,!on; u1 ,http and goa,s mkctcd \\1th 1hc Ball3 215 ,rttl'.Ut,. ,.. 19cw t1e-.;im,t11er moculntlonexp~iimmt'"' Agnculmral
strain of C.O"drta nmrfnamium. Otttli!T'Stqpoon Joumnl 11/\ ,•t1.7'1u1,y ft11m:,1/ oftlr,C,1p<1,/Goo<l 1/o~. ~~. •13.'l-<142.
R<brt1trl1. 5?. 8145, 21G . . .nuu. 1. 19:2. Heam«tor /01,mnl &ft/:.• /),:Jx,r1111cm of \g'",<'11/1111..:
~9 PR0-1.E:)J!Y, L'" ou ;•1.J-~,13,, u .. 1985. I 1e~u1,,:u.tr in Angor.t 8{1:Lb 11..l. Un/1111 o/~011111 /tfrt«r. 4 t.1r~z45,
pothDIC>'6tG>I ,iud) ofanWcl:,Jly ,nrct1ed. tn:atec ~d untreated go.11>. 2:1; ,rA.,tl'-\. !>.. J9G9, L~pt•tlt?ncti \\ith hc-am, ':ltl'f cr.1.d lotlon. PtM•i."(lfn~ o/
o,utmupaort Journ<1I of\"e1nim1ty R1.·~.-nr(h, 52. lJ-!9 titt Sym,,,,,~,,1111-tm Bit1lo~•4ud C(m111,I tJ/T1'i..·J..'$ l11 somlum, .,Vrlrf.l
we>P1J11CH.1>t, H.s .. :11-1;;. ~ ,imple ur.d rupld m~thod for dcmon.irating Rhude, Un1t,.,.lty. G111h,m,1nwn,Sn111h Alrlca 133-1.0.
53'-l ~cuos 1111<>.~ Rickettsial nndchlam)'dial disease~

218 snc,, w .. 1928, Pathologi(al studies on heanwa1er. !Jilt mu/ Nth u11~$1>Uchc de Cof<'rfrir, nm111111111/11m 1solr!e en Cuadelo11~ (t\mJIIM
R•,xms ofthe Di,·«tQr oflf(•1erinnry Ed1mmo,1 and /in.weir. 283-30S. Pr:,r1<;o1,..,,J, Re"" t1'£lt:"1g;· ,., de ,\fffd,:c/11, W11'ri>:t11udrs l'frJ'>
TroJ}irtJ.U.\·, 38, a,i--.;2.,
219 »':'<Gt. BA,. 19;8. Bruin biopsy ror 1he dia1;'Tto'1, oi hc,,m,·3ter. fropia,/
.~nim11/ IIM/rlt wul Prtx/11ctia11. IO. 4s-ll!. 238 \",\CIUt.il\. :\',, TtV..r. l., Ttrrrt, ti., :'ltM:.r TISf./, U, ~ tlf.~MIO, A, 1991\.

22o~E11•u"<:HL, H.J.1 .. 196!!, Orgnnopho.<phaie poisoning in form Mock in lnhlbl<ron ot MHC class I ond cl0$$ 11 ceU $Uc(ace expcessu,n on bornie
sou1hem Africa. /u11m<1/ ofthi Sotlllt Afrltntt V,wrlnnry .,Jfdlrnl cndo1ht:linJ C(.tils upon lnfrc,ion \\tftb ('.0:111/ria ru>nilu1r.rium. \lrtl'ri11ary•
A.ssocfmlon. 39. 69-72. lmnum<,l()~· «nd lmrmmopculwlogJ·, 61 3; "43,

:.!21 'Jl!R8!.A...W:H1.....,.,. & \IIS!\I". J.A,, 1968. Th!Qdftn poi.s.onlng of ('3ltle. A CO.Se
239 \'.\:< ·'"HLL. •.11 .. 196;. Th• u,>l or ele.:110,encct>h.al<>$n>phy and brain
r~pon. /¢r,r11ul of1tw So111h A{rlNtn V,:l(ll'f11/(nl .\Jrdical ,-Wodmioo~ 39. lnops)' In 1h• clinical dln1,111osis of heamratl'!. 011dn11,•;,oor1 Joumnl of
Vitcrinn')' 11,,,,,vc/1, 54, 2.95-29!1,
91-92.
'222 'ffi"EU.e-R .,,, 190:_ft. 'fhead"not:e or our kncrw1cd~c rel)Oning the stock
.i,:o , ..\., .,,1snt.. !Ui. t. oo(iJt.f,t, r.T_ 1~.a,. l'he tu!3UTIC'nt of he:irt\\O..tcr.
Onderutpoort)ounutl of\,..?teriutuy Restorch. 5·1, -lfs-li'J
d1Seascs oi S0u1h Africa. Ttm,s,xtal,!pir11fwrnllo«r11nl. 4. 69-80.
Zoll \'AS ,\)ISTEl-S.R., Lt8Et(fM. f'.T.. IJlllO;\tESICO, M.. K11U~·P~\11UCt:. R.D, &>
Zl;3 TIC:l. G.A•• nttr~O:\, S,tt .. STb'\\'ART, c.G., OU Pl,.ESS1~, 8, &:DE WA.U., ri.T•• 1998,
MAn1rL 1.. 1.988. Thil prrs.t.1'.'lce of ,mdoro,cin ucu\ity in Q.stlS of
Thenbsence of clinical dis~os,· in roule in <0rnniurn1I gI117,1ngoccas
expcrimem-all~··induced heartwnce.r in sheep. o,ul.ilr~1.1f}(J()rtfourr.nl of
where tarn:ien am cha.n~mg from an imea.\i\'e dipping i>rogrnmml." ,o
Ve1rrinnry R£'3't<utiz. S.S.. 211-220.
one or endemic s1ablht)' l() dck-borno diseases. 011dm1epoor1 /OllrttOI of
Wterl11QI)' /l;,,·,:,rclr. 6.<,. 16!)-I,~. .24l \'1'::0. \J.1Srt J... 5.8.• ~l, 1'.Jb, t·., (~HI tuUt;.. A.J,. 08blli: \ I. t', 1, • H.t.ltTSCJil~(.:t;H,
IL, 1~8a. Theclimcal pathology of hc,mwa,ec. I. rl"emocqlllfil' and blood
244 TOnT, P,, tl[!'\5.UO••\ ,, )S.\llJ\X,., S. ,1 .. \1Altn~fiX. I).-&< '1Cli:.tl\1l!R, 0,J,.. 1999,
chcmlmy, 011dmr;;))O()rrJoum/tl o[V,wrlnary llt.reilrci1. 5S, 3,-1S.
tmmunc response$ ,o Cowdrla n1minantlum in(ections. l'nras/10!010·
Totfa.1·, 1S, :?86--290. -:t.S3 VA~ A~~l'l.l.• .)..ft.. Rl'Y~M, . f,,.. \1'' 8UR(af ..... 19::-.,4. ChangL... 1n the

~5 roTn... P., llU.N'AA&HT. o.. Zll.l \fWABA4.\fiO. P. ti W'EIWN:'\(, r.. 1993,. blood-~ >13nu of~he,ip "i1h ••1>erin1emally indu«,d h,am,.1e,.
Q11dcr.UptXtrl /()11m11/ ofVe1rrl11tlry R(s,arclt, Ot. ~3-49.
Inhibition of Cou:dria mmbumtium infcttiou~ yfcld ~· fm.e.dfr'Ons
alpl,, and gamma In endorl1cllal c<>lb. Rl!t•wd'l!lei•(lgl!et de ,Vl,ld.c/ne :24.,i \'i\~.A,:.:,,Tl.t.. S..R., Rli\'!JK, , ... MVDut,nu. I!.., PRL'TOR!VS. C, J, <..\('A'S. P., 199.;.
W/.!ri1wire1l.,.: P<t)~ Trop/0011.t, 46, 189-194 The clfn1cal pathology of 1te:inw01or. tr!. Ch:uig~ In blood clq1ting.
blood c.skium, hl<1od prot~in, h:acm:nocrit and whio,•,ccU t'O\tnt'I'. in
z:.26 TOTTI!. P., 10!1:c.tJAN, r-•. nn c1tF,1 A.t.. & " 'au:.~"F J•• 199.~. p·roduc1fon or
olph~ iniorferonln Cowtlrltl rumittomlm::,ln(tl<:ted <aule and 11< clfoa <hc,,p wltl1 cxpetimoninll)' induced hoarcwarnr, O,ul,r~poort Journal of
on inle<:1cd endo,hcl!al cell cultur,,s. lnJ'«1lo11 nnd /mmrmir;•, 62. V111,rhJt1rt ReA"lrrth, 6t. 21-2i.
2600-UQ~. \n OVeT\-i~w nf
245 VAS UEft\PtKAMP. lf.L , .. Jt.lt()/.l·.~).;.\'. , ... 1987, Ht!itrt\\',lfC,r.

22:' f-oTff, P., \!CKEB\'f'Jl. O.• JO~Ctf,\.~, f- U.\OB'ET, ,\., ,\1 WA-~.~. \J,, "WASCt, o.
!he clilllcal •ign,, •=pubnlcy and dltTcn:nllal dfagnosc,. r,ithcd{,nu,
& en~·suo. ~-· 1998. AnnlysiS'Of Ct'.Uulnr respons~ to narlw and
In domM1!c rumln•nu Otrtlem,·~v,ori Jo11ri1nl of\:t1tri11"?' lk;«•rrh.
!,,;, ?SJ-266.
recomblnom proteins or Cmt.'liritJ rwninm11lu111..-tmu,ls 0/thct Ntw n,r.4"
Arad,m,l'o/S<te,m,s, 8"9, 155-160. 2,;G v,w nn MEJil:\'.'tL L 1979, l!-1eld experience \\'llh hu.:im\nter .Cowdrin
rt111:1mmrlum) in anl<-.Jouro11l ofrheS001h lj"nca,; l'tttrltrf1ry
:Y.?8 TOffE, P, • .)1C'IQ:E.VE.R. u.. MAlmNez. o. $- IU'.~S.AID, "" l!i97, t\n:tl~'Si$Of
T-cclJ t<-$P,Onsc-s ln cuttle imrnuniicd n-gainst heartwa1er i>r vaccln:ttion .woci't.tion. 50. 32~2.5.
wuh klUed e!Nnentnry bodies nr Cowdrin n1111fnautfum. Jnfcctlon twd 24i ''"~ oe'l MtfC:WEi L 198;. lhe inreclloo.and rttatme.nt mtthod of
/111m11nl1y. 65. 236-2,11. \'-:I.CC1nation a.ga:inst bti!.T1wat~r. Ondor$t<'P()Qrl Jnumal tJ/V"tterftuu·y
R,m'llrtlt. S.. •189-191,
2:!.9 TOTTE. P.. \',\OIIER\', :,.: •• M~,,rrrxu.. D.. 'Ul\JI, , .. UAW ,G.\LL )i.T.. -MA.CHI..IOll.
~.i>.. B~~s,.uo. A. &WEJ'CD:~t:. , .• 1996.. Recombinant bO\'ine intt!rftrcm 248 VA, KUT"f, \1., OU:\TER, S,J,0 )I \QUI IA:O:. If,, ,\U.SOPP, &A., Sl!K.\11, \'. R
g:unm:.i lr.hlbir.sthe gl'owth of Cou.'tirla rr:ml,umtilml but fQ!J; to l:\duct e.w,\~ w.c:.. :tooo. Identification of<:o,v,lrin rwnl!lwitiwn antigens
major histooompa1ibll£L>' oompfo:x clos~ ti ronowfn,g infl.~tion of rha, smnufa1ep1oliforo1ion ol lrmphocyu,s Crom c;,rlle 1mmunii~ by
cncfothcUal cells. \ ·crrrintrry /111m,molog.·n.1uJ /mmunopatho/ogy. 53. 1orcc,1<iin and 1rcarm.en1 or with biaCll\tnted organisms...lnfrttion mu:I
61-; J. lmn111ni1y. 68, 603-614
230 u11.t.~8w(<,, c .. 19;t. Studies on bt!M\,..-ate-rin Mndugnsear. Pun l. Rwt1e1 U9 \'i\.~ KUt:F. M .. ,-nt'Z.
A.\\', 4' DE WMl.. l),'f•• 19.93- lsolallor.md
d'l:'1t:1t1~-e. et de M,o~rl11e 1'1!11Jrinntrt! detPn;~ Troplca11.<. Z4 • .!39-2,19. ch.imC;en1.atfon ofnmlganfc pro1eins of Qnedrln nm1i,im1rfum, R~11,r
lf/J.l1W1l[;t Cl dt lltd1Jctt1e 1'(,l(:ri,1t>ir1• dt-. P11y, Tropiaw.,. 46. IS~-164,
:131 uiu.Nll'IIUt~c.. 1!)81. 01.f(.'fU(rsofCm1kin 1Ju, lr()pu:.'f. E«momu:wul
200,1r,1,c Rt:le;.:(mc,:, TIH.: Hnguc. Nc:hcrian~ '-1Q:ninus 'tijhoff :tSO \'A.._ \Ul:'T, \.H~, Ju~<;,-:y..x. F," \'A."' DtR U.IJST, B.A... 1992, p~ logenulic

Publish<>~. po,iJJun af Ct,wdrln rumirumtfum (RkLeusi:ile:,1 di•tem1fnt-d by

annly&bot amplified 16$ n'bosornnl l)1'Aseq11enet>s. /n:,·nU111011"/
232 uu.1,ss~RC:, e... 198!!. Dboo:;e pmhlems a.,sucioted "i<h the lnrroducdon
/u1mml ofS;s;emnllc &creri9/11gy ,12. ~94493.
of EurOj>!r.ln caulc ln rhe tropic:;, Pl(JCl!,v/fngs ofth, Tw,lftlt IVorld
(;011~$$PII /)fseru,,,ofCaul,. TI,c 1'cthcrta11ds. 102~1032. :151 VA~ \'UF.T, .,.11 .. 1oxcti"~· F.• \·A~ l>:t.r1r;r, lt. .&- ,-A~ nn1 ,r::,~f. ~.,\., 199,i,
\loJecular c!ouln11, S<"qucnco ~nalysis,a.lld uxprc<,lon cu 1he gene
-233 uiu:snER\., c;.. 1.9&2.. ltq>etl.lu erunl tnn.!i.nil~~ion or Caw,.lna
encoding the lmmunodomtnam 32-ldlodalton protein o!' Cou'tfr/u
ruml,urmium by the Gulf coa)l tlck AmlJJyommn mn(t1latwn: dnnger or
rtlml1:11mlum. lnftv:1fon n,ul Immunity, 62. 14.51-l.itSG.
in1rodudn11 lwam,·~t<'1' and benign ,\fiit1in 11wllerfn>l<on10 rht
Americun 1naJnla.nd. Am.:rirmt },mrnal o/V~ttr/n(lfy Ra..'Clrrh. 43, :$2 \'.ii." \'lJI."t. J\.U, \".\., DtR.2.UJST. 1t.A.. CAMUS, Ii., "1AJ.IA..~. ~'1.. '-L\.R'rc.xa. o.
1?79-1282. & JO"\GElAS. r.. 1995. u~r ofa spedftc' lmnmnc,genk re~on on the
Cowtfr;a r,unlntmti'!mt map I pro1d11 (n n -k•raI0311;:,l ~uq~·. Jrmmnl of
:l.34 UlU.~I.ES\G,C.. !983,. f te.1.rtw:ittr {(.i.mdrln nmwramium infect.ton):
Clf11IMI .\ficrobialogy, 3:i, 2~0;,-2410.
current miu~Adt'<11t(tl In Velerlnmy Srltn,.-..,dud ComJ)Omtit'l
, Mtd[(llJi'. 27. -!2i-160. 2S3 \'()(.fl.. 5.-W.. \lJS"KEk. C'.E. & HUR'<, Ii., 1~98, Al1C3ni)tS to irnmu11i7.(.' m;ce
wtth ino.C:li\~ CowilrUr rumillamlum. In: coos-& L. "Rtmtgcuu.o. :,.t••
.2Ji' UU..Ei~\:Bt;kG, c .. ,~. Mcthc'4.t; currtntly Uii.'.d for the tontrol of (~d,). ~ml Jm.-mntitmnf ConJrrwrct a11 Ticl:-bomi' J>nti1.t>gf11S nt ,11~
he:mw•rer: 1helr \''1lidl1y •nd prnpo,:,t, ro, fururc t'Ontrt>l '1r.>1~'1,"""· 1/091• \tutor lmMja~i•, n Gl"blll PiW1"'t.~1j11r, KruJ;Cr~allon'*I Park, South
Pattu.<ilo!!lgla. 32, 5;,-52 Africa 531-53..
2$16 U1U::..\'B1:R(:, G., &\Rn{.~ .. CAMUS. I.., UUHRl:,C.f \1 .J. £ (:~RRJ:C, G.1 •. t9fl.i. !S•l WA(H·lf.J..A. S,I)•• RUR;\~CUt\\'A, r.M., ,\1A1CA.~. S..M .. \ 'U~)o;t,t C.t., (.n,\\\T-ORO,
lfoanw;ncr In the Carlbbe:u\. Prw•11d,., \ eurf,:nry .llvdlt-/ne, 2
T .n., &UUU"1 .\.,J,, emc.Rmc;,.. \t.J. ~ MC(H)IKf'.. ·r.c,. 199L ,\ clonC'd DX/\
2.:;.s...2s, probe identifies Co1«"1rw ntmlmtntium lnJ'1mbl>Ynnma 4!f1rt,:gamm
rlc~.Jounu,.J ofCJmitnl MfrrolJiofogy. 29. 2s11-2s;i.
 lleanwnter 535

255 <,'.uUR. o.H. (, D1J>1LEM, J.s.. 1996. Eincrgcn« or ill• •h!tlch!o,C$ a, Ondent'i1-,x>ott /ollmttl of VtzNionf)' R,·~'l1rch, 63, 159-1 i'O.
huninn hc:1hh problems. Emerging lnfer:1iQ111 V/1t1UN. 2. lS-29. 261 \'UN~ll. c.L. t'l\'1to,1.. s 1. &t.'lU. s.... t988. Culth":ttlon or UJttYlrio
256 \\.\l,Xt-.11. ,.n. & otWAc.£. A•. 198;. Thb tick \t.-"("tors cf Cowdrln rwnmamlum nmwumrium ln bo,inc,·u...i;cu!J.r endothelial celb. 11w 1:1,:yn
ILSO<lllldra. !xodldac, g<mu,A111//fy(l1111>111J ond rbelr distribution. \#u,~rl,u,rl(,n, U. 12...16.
l)ll<lt•tfWJJOOrt Jo11n:al o/Veteri11qry 11,!/tfUC],. s.i. 353-379. 26S \'UN nit. C'!:.E,• .:or.AS, ... ~1.. !\'Or\\'AI.. M.A.. & 8URIUOGI?. M,I.. 198';', Drs1inC'ti\."C.
:5? w1::.,v£1t. c.r. & uxA.:-:U.IJ:, E.A .. 1990. The t.osrimula:ory ru.nction oi sl.linln,s of cok>nfM of 0,u'flrfn rumtnnmium 1n mldg111:\ ot
.anttg\•rt-ph."-.\~utln~ et=lh,.. hmmmuft,g.,• lWu)', l t. 19-S:i. Amblyomma hc,brM11m. Omttrsr.tpo(Jrt Journal of \lvw,111111)' &scmch,
z$8 Wl:l~ ~.t.• UAIG. o"". & Al.f:XA~Ol'R, R.,\ , l9~ Aureomrdn In the S.:. 183•18S.
treaunem of hc.1m,~ter. Oudcrstopoon /Qunwl v/Vetm'nnry Rffl'nrch, :66 zw&>'G '"'"'· "-" 11,~t"'-'·"" .u.. 20m.A chemfcull}•delu,cd medium for
:?3. ~1-30. tht growlh ofCowdnn rummantimn-. Ondersu,poor,· Journf11 of
2l9 \\'foO~Gt\, F.D.• )1U).tJL1.\'f... ~...... &cat()()T(~HUI~. J,, :N>OJ. Tran:"lm[S'>ion of \ 'et~riruu1· Rl•1,.•tuc11. 68. 3;-40.
Couvlrln rr,m11uuutum by Amblynmmn gfmma from in!~eted .Afncan ~67 7.Wf!'YG,,m, u."' sv~1,11t:-.'6, "·'·· a~,. Conrmuous. in vdro pn,p:!g.ttlon ot
buffalo (S)·11e,rus "'ff,r) and clund tT11urQ1rog11.10,yX1 10 ohoep. Trop/ru/ (.ou:dria nmm;antlum {Wtlgc\-onden .stock) in a canine
;\11/11111/ lioolrh ar:tl Pr<ld11ttio11. J3, 379-s:!90. mucrophugc,:nonoc:)10 C<'li line. 011dem,1""'fl /01mwl of\ ,,mnflr,•
~60 \\:!IMS, D,1). fl RJCH \RI>, k.l>., 198,1. lmtrcep;lon of n v~ctor ofheanwQu..tr, Ru:search. 68, ,SS-lSi'.
1lmblyommn lwbr<Wum K<><h ~-\<:or!: lsodinao) nn bla<k rhln=ros~ 2b8 7.-\\"EtGARTI-f, r,,., ,osr.,L\S-S. ,'\.t, & uoR:-., Tt., 199tt !)enun,trc,c mcdi.a for the
tmponcd lnto th,; United S101cs. Prt1«<"1/1,g, ofrl1,• lilgi1r;· dgli1h A11n11n/ 111 ,·,rmculovation of Coud1111 mminanrium.A1111alsof1la· \l'f,· York
\/,..,r111gof1h~ Ui:ittd Srm,;••~11im1tl I/Mltlt:\ssorf1t1io11. Tho H)'•II ilcadtn:,-ofS<i"•11res. ~9. 30;"-Jl?.
Rcg•ni:y Fon Wor-J, lio1d, Pon Wonh. Tc~<u. 303-311
261 \'Ousc;. t. ,.. !USSO~·. tt.A... 197.l, f ftJm\'lllN in 1hc:- tf,.md. J,aurnal of tht'.
:!"&J 7,\\'nCAJITlt, f" JOS£~1,A.V-.., ....... \IA~ ;.nu,,. r-.• \'A.S HfE.HDE, . II., \U!tOPP,
M.,r-E..P~ a :\t.LkWP. b..A., ~ i . The KLlmm i$tlluteof Ellflldtia
Somlt Afrfr:M \'e1erinaryA$SOCi1tll'tm. •l-t. l85-IS6. rumhu1n1ltmr In t1rro bolalion. prupagadon, and chura.ctC"Tl.'1tfort
26::: YU, ...x•• \JClUUOE. 1.w.. 7J1A~(';I x.~ WAUO,.R. 0.11 •• 2000, Chara.cu:rJzation ot Onc/enr,poor. /oumnl nfl'tt,•r/11ar; /lnrorr/1,69. 147-1$3.
rho complcu; mmscriptlonaJJy nnivu F,,Jtrltchm dmjft.--e1isu2a ~a omur !!70 i.wr~,'OMtTII; &. \'OC,iH.. '-.\\',, JC).,~\\.,\S~•.\.I .t HOftl'\, .... 1997. /n l'ilfO
memhranc protein multlgcrw lanllly. (~11e, Z•l8. 59-68. ~olluton :uid :uht1.·~tlon nrru1~tlrln n1mhum1i11m undct~rum·fH..'<"
26311,"x••· r..t.. 1996 Heam,..,cr In ,hetp ond soar,· a resiew. cuhurecanditla11, Hes<'ll«II 1,1 V~ttrlnn,ySdenre, 6'1. 161-164.
 Lesser-knom1 ricke11s1as 111tcct111g livc$tock 537

- .......... emphasize~ the imporuince of a cla~sification based on age-

netic b~is rather than the species affected. since apparenlly
the same granulocytotropic agent is a~~odated with disease
in dogs. cats. hor!>e~ and human~.6 1 with a role for small
mammals in infection of tick larvae.9 1 In addition. Lhe recog-
nition thai 11gcms closely related to J.,. µlwgocytophi/a a,e
•
.. found aero~~ 1he temperate broad-leaved and mixed forest
zone• of Lhe northern hcmi~phere within Licks of the genus
lxotles. resulting in infection in humans and livestock from
China to California, further indicates that we should expect
to find a \'Cry widc:,pread distribution ofebrlichias in na1ure.
All of the major gent•ra of ticks in the family lxodidae infest·
ing Jive~wck ha\e been ,hown to harbour ehrlichias. and we
shou ld therefore <'xpec1 that most. If not all, hard tick popu-
lations will be inYolved in the transmission of those ehrli·
chias with which the~ may have ccl-evol\'ed. A\'ailablc
...-...' cvidcncl' ror the well -studied ack-borne ehrlichaas IC. rumi-
;.
•.
,;~ 11m1ci11m. f:. phagocyrophiln, E. ami$ and £. chnffe,msi.<)

.':'RI:,,
. '
},\.

.
which have a tropism f1>r leukocytes. indicates they are each
transmitted by 0111\· a single genus of ticks, Although evi-
dence of natural infection in other species of ticks has been
Figure 41.1 E:eciron micrograph of Ehrlrchre (0,1oeceres1 pllagtXf'll1Phila reponcd for ~orne of these agems. only one genus of ticb
within an intr.:cv,co1as1nic ·,acuole in a grar.ulotVle of a sl'.ea~ appears 10 be compe1em as a vector of each. for example
lxodf's spp. rather chan Demmcc111or spp. for agems in the
iheclade,24 but retaining lhespecles nnme. Under this pro- 1:: phagocytopilila-group. 141 Evidence or ehrlichial infection
posal the three genogroups would become the genera in ticks does 1101 therefore indica1c 1hn1 the Lick is Lhc nawral
Neorickettsin. A11t1plas111a and 6hrlic/11a. ,\rguably, agents in vcccor. and care must be taken in interpretati1>n of the find·
genogroup Ill ~hould be Cowtlric1 species sine-.: the d iSCO\'ery ing lhat inonalarion of ground-up ticks can cause infection
of the agenc of hearrwater by Cowdry in 1925 predaws 1ha1 since this b) passes the salivary glands: transmission by rick
of canine ehrlichiosis. In this chapter, the gen11s names nsed reeding i~ require<). The geographical distribution of the
are 1hose In use by the overwhelming proportion oi authors le~scr-known chrlich ia,, probabl)' corresponds \\ith that of
during the period 1995 10 2001. Proposed re,isions based on the vector Lick species. Given 1haL rapid discovery of addi·
genogroups do not suppon a .g enus 111th a single member tional ehrlich1al agents has occurred since anemion has
(C. rrmri11a11Tiw11J. whlc:h 11•ould be replaced with Ehrlich/<1 been paid 10 tick.~ in temperate regions of the nonhem
rwni11amwm: but broadly suppon Foggie's19 pre\~ous pro- hemisphere and the determination 1hac b0th monocy'IOtro·
posal for a division of the tick-borne ehrlichia$ into two, pie and granulocytotropit agt•nts can cause human di,.ease.
1,~1h a genus comaining rickettsias parasltizing circulating we should .:xpec1 10 find many additional ehrlichias infocc-
mononuclear cells. and 1he 0 1her. a CyUiecetes genus. com- ing livestock in Lhe tropics. which will require our eoncepts
prising rickeusias which primarily parasitize granulocy1es. of ehrlichial infections orllvestock to be reviewed.
Considerable attention has been paid since Lhe early Studi<.'s 011 the lesser-known ehrlit:hias of livestock are
1990s 10 tick-bome ehrlichioses since 1he recognition of woefully inadequa1c: a genetic characterization oi the
human infections wi1h the monoc~ 101ropic E. c!Ulf!et>mis 11genb involwd in bo\·ine and ovinc ehrlichiosis in different
and with a granulocytouopic agcm closely reh11cd to continents hn, 1101 been made and ls rendered difficult by
E. phngocy1ophila. The unparalleled lewl of study has pro· 1hc extremely llmiled number or labora1ory isola1es of these
vided clear indications rorrcvised concept~ of classifica1io11. agen1s. Rather than perp1:1uate the many doubtful $pedes
transmission and epidemiology of ti1e les;;cr-ltnown Ehrli- names. and the pocemlal Yector associations of each. a pro-
cltia species. The mosr obvious mC'ssage is that a classifica- posed ca1egorization I$ made b<.>low. in 1he light of a re\·iew
tion of the ehrlichias based on 1he host species in which it is of the literature and current conccpcs in the rransmission of
diagnosed or recognized i~ mislead Ing. sincP disPaSP usu:illy th'1 ehrlichias Small and large n.iminant ehrlichiosp, h:wE'
occurs only in accidenial hos1s. Fun her. the life cycle of each resulted from l rnnsmission experiments. or been a;,sociatcd
of these Ehrlich in ·species' may involve different ·reservoir in na1ural outbreak~. with Amblyomma. 1-1,l'<llomma and
hosts· according to ecological situations. and for lnfeclion Rhipiceplwtm cicks. It is reasonable 10 suspect that the
of lan,ae and nymphs of 1icks. The extremely close genetic agents associaied with each tick genus are distinct. bin
relationship between some European and :--orth o\mcr· those u,uall) identified as 'E. /Jo11is' and "l:'hrlichia 011i11a' i11
ic:an isolates of granutocy101rop1c: t>hrlichias further large and small ruminants, respecti\"Clr, resulting from
 Lesser-known riclmn;;ias infocting live.stock 539

Table 41.1 Prove., or preSumed lick vectors for e!l'licn1a ,nfe.:;1~ of livestock. e~cluding Ehrlich/a (Ccwdna rumimm1,um infection

IXODIO GENUS IXODIO SPECIES EHRUCHIA SPEC1ES LEUKOCYTE EVIDENCE PROBABLE REFERENCES
ASSOCIATION !El DISTRIBUTION

Aml:lyomma Aameriranum E cl/afieeT1sis Monocyiic (M} E1, E5 North America 25

A caj~rmenSJJ 'f llov15 M El Central and South 59.eo
Ameiica
A. var,eg1Jtum 'E bov1s' M E3 Afnca 78
Rhip;cephal~s R. everrsi evensi 'E. ovma· M E1 Sub-Saharan Africa 65
R bursa E ovma· M Et North Alrtca, 53
Mediterranean
reg,on. Middle East
R appendiculaws £. bows· M E2 South and Central 9,62
Af1ica
R. haemapnysa/01aes E ow1a M c6 Sn Lanka. lno;an 90
subconuneot
Hyaiomma H. species E. bo'lis' M E2' Iran, Middle east 14
(derived from Iran) North Africa
H excaVc1rum f. l/0\7$ M E2 Iran, Middle East 65
(from Iran) and North Afr,ca
H arra1olicum 'f lxms' M El' India !4
anarolicum
H margtniftum 1saac, f O'llni! M E6 India 97
lxo(fes lxoaesspecies £ phaga1:y10:,h118 Granulocytes (G) E' Nonh Af11ca, Europe, 35
genogroup oareanic
Unknown - '£ J!hiJg«ytOP/lila G Habitats in Pon
possmly Fxodes genogroup Ehzaceth/
liumansdorp area,
South Africa
E onditi G High altitude
habitats in l:ast
Alnca

• 11cks are presumed to have acquired infection w'1ile fee-ling on iheiletia annu/ata infected calves
Kev: Ei = b1olog1cal :ransm1ss1on during feeding of ucks pre,•ious,y fed in the Pf8vious stage on e,perimemall, infected animals, E2 : bio!og1ca.
transm,ss1on dut,r.g ieedmg of ucks previously !ed in the prev,ous s,age on animals 10 the fle10, E3 = ,noccra,,on cf homogenates ol !Jcks ccllecte.l from
o:
the field: EJ = 1noculat1on homogenates coHetted o• tlC'<S previouslv fed on inlected animals. c5: ideni,f,cat,on of agent by molecular tests 1i t,c~s
collect~ from the fieid. ES• circumstantial evioell:e exists :o essociate infection w11h 11c~ feeding

membrane-lined incracytoplasmic vacuoles in monoc}'tes. Epidemiology

producing dense clusters of cocci. lniected monocytes,
however. are seldom seen in smear:. of peripheral bl<>od.14 Although oven tli,ease is rare, l:. /J()vis in£ec1ion~ are prob·
Splencctomy. 15 rindcrpest 18 and environmental stresses26 ablywidelydi,trlbu1ed in Africa, the Middle East, India and
exacerbate F., bol'is infectious. Given the likely occurrence Sri Lanka.48 • 39 In the l\ew World the infection has been sus·
of recrudescent infections.. it is possible that the appear- pectcd in caule in Guadeloupe2• (but wa~ later considered
ance of E bo1 1i$ in blood smears of cattle following their to be heartwater 1" 1) and identified in Bra7Jl.5 it In the USA.
laboratory infestation with ticks collected in the field as nalural infection of goats and deer with I:., rlwffeensf.s has
originally reponed in Sourh /\frica9 might be related to this been shown,22 and cross-reactions in caule sera with 5ero-
phenomenon rather than true transmission. :-.:evenheless, logical tests for Co11'drin suggest natural infection or cattle
similar results62 • 68 in Zimbabwe corroborate these find- with agems closely related to Cowdricr. Had the laner oc·
ings and do suggest that fl. appendiculaws is a \·ector in cu rred in orher concinems, it may weU have been recorded
soutbem. and probably East Africa. as 'E. bouis' or·£-:. 011i11n· infc-ctions.
In one trial. cross-proteccion between £. /Jovis and Ehrlichia /Jouis cause~ a persiste.m infection of low
C. ruminrmr/11111 did not occur in experimentally infected pathogenidry in heahhy cattle reared in tick-infested areas
and challenged calves, 1:; but subsequently It has been where the parasite is endemic, but when such animal> are
found that amibodies in sera of ca ale believed tO have had stressed. either by adverse weather or intercurrem disease.
bovine ehrlichiosis. cross-react with the Ki.irnm s1rain of severe clinical disease muy be evoked and death may
C. ruminamium. 19 follow. 26 Similarlr, oven disease and death mar occur in
540 » rno, 11111u: Hickeusial and chlamrdfal diseases

exotic caule afterthcir imroducrion into an endemic area i 8 mon. Furthermore. in the same coum11• Rloche and Bour·
or in naive indigenous cattle following the introduction of din 60 detected infected monocytes in a calf and a pig after
per$iStently infected c:auk•and their ricks into a previous!)' they had been inoculated wirh a pool of grow1d-up Rhipi-
disease-free area.~ 1 ceplialus and (mostly! 1-/_valomma ticks collected from a calf
Pre,iousl) infected caule resist challenge 10 1t,(ection by which had been kept in Lhe same pen as affected pigs. which
E. wvI.1.. ' ~ Relap,es occw autl ma)· be Jmlm.:ed by splcm,c· the~ took as evidc11ce of'E. buvis'. Thi! vector associated with
tomy. 33 th.e major outbreak reported by Pierre; 1 is uncertain, since a
Reports of thi, experimental transmission of ·c.. bovis' heavy infestation with both Am/Jlyomma and /iyalomma
conducted "ilh different isolates rrom separaie cont inents ticl<s occt1rred, and transmission may have inl'o!l'ed either
in which three different genera (A111blyo111111<1, Hya/0111111<1 or both of them.
and Rhipiceplwlus) of ticks ,vere implicated. are consistem
with the concept of the invoh•ement of distinct agents. In-
Clinical signs
fection with 'Ii. (1ovis' by inoculation or ground-up Amb(vo-
mma 1•11riega111m in West Arrica ' \\'aS reported. as was 'II is possible 1ha1 the difference in severiryoflhe disease rec-
transmission by A caje1me11se in Brazil.~9 Transstadial ognilled as ·nofel' in West Africa and the infection in Olher
transmission through nymphs has been shown to occur ex- pans of the world is related 10 different genetic types of·£.
perimentally in A. mjc1111e11se.~'9 :v!onocytotropic ehrlichias bo1 1it. \\1th th!.' ~uspicion that the A. z,ariegarw>z-Lnrnsmit·
may be widel}• distributed "~thin .~mblyomma spp. 1icks ted form can result in more severe disease. Gueye (cited by
since an agent with genetic identity to £. chajfe1msi:s in I Gs Ui!enbergl01 ) coruidercd the disease io be an important one
rR:--:A gene sequence wa.~ detected in adult.\. resrudi11amm of exotic cattle in Senegal. l lowe\·er, other factors. Including
in China. 5 Transmission b}' Hralomma ticks has been re- breed and effec1 of inrcrcurren1 infections (perhaps rrypa·
ported: the original report of Donatien and Lestoquard in nosomosl~. or dermatophilosis i11 West Africa), might also
1936 did not specify the species which had orfginated in be expected 10 affect the severity of 1he syndrome.
Iran: '.'ieitz reported transmission by Hralomma e.,·cai:0111111 In most countries E. l,011is is oot considered to cause se-
ticks from lran 65 but did nor Slate if transmission was by rious disease in caule or water buffalo, but in some circum·
feeding the ticks on ~uscep1ihle animals or by the inoc:ula· stance~ might e~acerbare o.ther infections. rncubation
tion of ground-up licks. Workers in India have reported periods after parenteral inoctda1ion of infected blood or tis·
1ra11,mission by 1-/yalomma a11(1(0/ic11m a11ncolkwn after sue suspensions range from one co two weeks. 15• 78• eo Feed·
tick feeding on TheilPria a111111lma-infected cattle·; 1 and by Ing Infected adult ricks on s usceptible canle induces di~ease
1-/yalomma margi11acw11 isuaci after lick feeding on T. a11111t- 2010 41 days laier.62 whereas the injection of a suspension of
/at<1-infec1ed calves.97 In these experiments the possibility cnished ticks produce$ a parasitaemia in 16 10 20 days.'8
of recrude,cencc of latem infection brought about b)' tick Two peracute primaT) cases in Sahiwal cattle imponed ro
feeding or T. tm11ulara infection musi be borne in mind. as Senegal have been recorded.7A Both animals exhibited se·
well as the difficulty of discriminating theilerial and ehrli· vere ner\'OUS signs and died within 12 hours.
chial infection of monocytes. Similar concerns can be raised (n areas where E bo1•is is endemic. primary in[e_ctions in
regarding the ex-perimems which incidentally recorded the indigenous calves are usually nm observed. Exacerbation of
potential role of R. t1pf)('1Ulic11/a111sin southern AfTica.62 De chronic: persls\em infections may. however. folio\\ stress.
Kock e1 al.~ reported 1ha1 £. bo11/s was deteeted in ca11le after producing clinical signs in 101050 per cem of the animals in
adult R. apr;e11dic11/aws collected from farms in the Lim- the herd, with case monality rates that range from 5co 25 per
popo Prol'inc:e had fed on them; sinc.;c ~ome a11imals on cem.71 Morbidity and monality raLes are higher in primary
which the ticks fed had pre\ious e~posure to ticks. a recru- infections in imported cattle or in virgin-soil epidemics with
descencecannot be ruled out. However, the finding was cor· case monaliry rates ranging from 28 to 50 per cent. •8
roborated b~· Maison 62 in Zimbabwe who recorded 13 cases In acute primary cases rectal 1emperamres tlucrua1e
of E. 1Jo11is infection after feeding adul1 R. apps11dicult1111s between 39 and --11 •c for710 10 days. Affected animals are
ticks collected in che field on high-grade dairy 1.--anle: The in· anorectic and have diarrhoea. They arc apat!1etic and che
fection was detected 20 10 41 days af1er tick infestation. and respiration and pulse rates are accelerated. :'\lany de,·elop
was accompanied b}' rever. Norval68 also reported from nervous signs such as h~1>crexcilabili~· and paralysis of the
Zimbabwe that nymphs or R. appe11d/c11lar11s. collected larynx which disappear on recovery. In Olhers. the nervous
~"hen engorged. transmit E. boi•is as adults. Circumsmntinl signs are \'cry severe, affected animals having bou,s of
e\'idence also suppon,; the role of Hyalomma-associazed '£. drowsiness interspersed 1\ith pcri<>ds of hyperexcitabllity
bo11is'; in Senegal£. bo11is was detected in blood smears at and convulsions during which they show galloping move-
the end oflhe rainy season but not ia the dry season in cattle ment:.. Severely affected animal~ usuallr <lie during a con·
maintained in areas where Amblyomma were scarce or ab- vulsion about eight day$ after the onsec of illncss."1
sem.38 but where /-ly(l/omma marginarwn rnfipes was the In some West African co untries the disease 'nofcl' oc·
predominant tick, and Rhi/Jiceplwlus e1,er1si e1,er1~i com- curs shordi after the onset of the rain,. in emaciated cattle
 Lessi,r-known rickemias infecting liwscock 541

e.~u!nSi\·ely afflicted witl1 dermatophilosi~ and se\·erely organ microscopically. }.·lany of me convoluted tubulb are
inlt>stcd with ticks. 1t is an exacerbmion or a !JilrSistcnt necrotic and are reduced 10 basal lamina filled \\~th karyorr-
£. bo~·is infection. characterized b)' enlarged superficial hectic nuclei o.ndcytoplasmicdebris. In comrast. changes in
lymph nodes and swollen drooping ears.3 ·' Affected ani- the central ne:-vous S}r:;tem ,m: not marked. ,md are limited
mals have fevers. congested mucous membranes. ocu lar 10 cong~"tion of the leptorneninges ~nd discrete ,,apillary
nncl nasal discharges. inappetence and constipation. \,Inn:· congestion and rare perh•ascu!ar haemorrhages arbing by
collapse and die ii not treated. In some animals in diapedesis.
which the only signs are lymph node enlargement and
inappeumce. recovery occurs within a week. 26 In l\lali.
Diagnosis and differential diagnosis
Rousselot83 obsen·ed a distinct seasonality 10 cases in 1942
and 1943, occurring during the rain)' period. with. for the The history. clini<:al sign, and lesions of canle with pritU;ll1'
combint?d years. six in June. 45 in July. 45 in Aug11st. 17 in infections are readily confused with those of heanwacer.
September. se,·en in October, and very few at other times I laematological examinations will aid differentiation
•
when tick numbers were least. Rousselo1 did not identify de~pite the difficulty of detecting E. bo11is in blood smears
the vector. but recorded tha t sick animals had A unrfllgn, because bizarre monoq'les do 1101 occur in hea.rtwarcr.;9
rum, R. evertsl and 'above all' l·/yafommn infc,,1ations. lihrlicliin boi1is in \Vest Africa is usually detectable m organ
which does not allow us 10 associate infection "ith any one smears, panitularly of the lung, taken lmmcdlatel)' aner
or these potential 'E. bovis' \'ectors. death. The parasite docs noc occur in endothelial ccUs. 711
It should be noted that monocytotropic ehrlichias can The A. C(ljem1e11se-,ra11smi11ed agent in Brazil and the
also cause human illness(£. chnjfeensis infection. A111bly- agent in somhem india!l.J have been maintaint!d in cultures of
omma-1ransmittedl. and therefore some of the '£ bol'is' monocyte.,. The agent in India was dc1ec1cd after 10 days of
agent~ may also present a risk of zoonotic infection. This culture of monocytes from a 1,-ater buffalo. It reacted 11ith
ma}' also be t.he case w11h Rhipicephalus- or Hyn/om111fl- £ rn11is amJserum.~3 The results suggest that tht.' agent may
1ransmitted agents. as well as those iransmiued by Am- be pan of the group Ill chrlichias. which is char.icterized by
/1/yommn. since macaque~ CMncaca inunJ reacted with f.. cnni;, since cro,;,;-reaction5 by in<lirn<·t fluorescent anti·
fever and rickettsaemia on inoculation \\'ith 'E. bo11is' in Al· btld)' test (IF.\Tl between agents typical of group II and Ill
geria. where Amblyomma vecrors are absem. us The /lhipi· ehrlichias are weak.'" n,e ability to culture the agent in
ceµhn/11s-1ransmitred £ ca11is can also c.ause human mono<..-ytes does suggest fixed monocyics could l>e evaluated
infocti()n.2~ appareml)' wilh few if any symptom$. for use as antigens in 11',\T. although specificity may prove to
be a problem when en tin.> organisms are used~ anngens.
Pathology
Control
Despite the difficulry in de1ec1ing E. bouis in blood smears
during ,he course of 1he disease. the haematological Treatment wiih a tetracycline during the early stages of the
changes are of diagnostic significance.18 The characteri.stics disease is usually success:ful. The es~emial preventive mea-
described for 1he Aml>fyomma-related '/;, bovis' in mono- s ure is tick controJ. !0 1 The infeciions probably phl)' a role in
cytcs appear 10 hold tn1e for the forms of· f.. bo1 ·is' observed complicating other haemopanisite infections. c~peciaJly ii
in other countries. and transmined by different tick genera. they are immuno-suppressi,·e. Since some of the \'ecton
The cytoplasm of infected monog1.es contain seYeral vacu- may also be in\'Ol\'cd in the transmission of Theiler/a pan'll.
oles - most of which appear empty but some \\ith ricken- T. m111ula1a. or F.. rumi11m11i11m, the use of acaricides for the
sias presem - and their nuclear membranes are distorted co11rrol of 1hese infeciions can be expected 10 aifcc1 the
and even shredded. Occasionally, a nucleus contains a vacu- exposure and immunity to E. bo11is agents transmitted by
ole. L)~nphocytc numbers are little affected but those of these ticks.
neucrophils and eosinophils decline. ;\!any inunarnre cells
enter the circulation and cells in mitosis are usually
pre,em.1'!1 There is, in addition. a Lhrombocyt0penia. tQ
Ovine ehrJ ichiosis
The conspieuous gross lesions obser\"ed in West Africa
ore hydropericardium and severe lung oedema roge1hcr
Introduction
with hydro1hora.-. and asd tcs. In somcca,-es the interlobular ,\)though Ehriic'1it1 01•i11a was among the flr$t mo1l0cytoph ilic
pulmonary oedema is so severe that ii mimics t.he soro- rickettsias to be recognized. and despite the passage of time.
fibrinous exudate found in the lun~ in acuie case~ of con- there is still not sufficient C\idence to consider it as a distinct
tagious bovine pleuropneumonia.;8 Tiie l}'mph nodes are species from the agent \\'hich Is inl'olved In bovine chrlichio·
enlarged. oedematous and, sometimes. congested. The sis. 101 TI1ere have been a number of serendipitous obsen·a-
spleen Is reacth·e and the kid neys are congested. Rioche79 tion, of monocytic ehrlichial infections in sheep. or goatS.
considered the kidneys to be the most severely affected made in the course or other invesrigatlons 15• 5?, ,,.;. ; •• ~which
542 "'WO~ nlhll: Rickc1tsial and chlatnydlal dis~ases

the aulhors ascribed to £. (or pre\'iously RickeusiaJ orti,w. In known as 'nofel' in some West African countries. These·
addition, the same or a similar rickeasia has been linked to a verity of disease in Senegal and Mali sugges1S that a similar
fatal disease in sheep in Namibia,MAlgeria and Turkey, 13 · 33 agent. or mechanism of inducing disease. is presem In
Senegal.41 '.\fali CRoussclot. l9-13, cited b)' Uilenberg101 ) and 'nofel' and o,ine ehrlichiosis in that region. although con·
Sri l..anka.'lll As in the case of£. bovis. there is reason lO sus- comicam hennwater needs 10 be considered. Five out of 18
!Jt:Ct that according 10 1he agt'nt idemifit:!tl in sheep a sp.,cies mortalities In the rainy seasun. and two out of sel'en mor-
of its own maybe misleading. and not indicative oflhe level of tal!tie.< in lhe dry season. in sheep from an ,\mblyomma,
dlversity 1,1lhin the species. Since .%ipicepllalus spp. have free region of Senegal, vaccinated against heartwater and
been shown 10 act as vectors, ,Uld possibly Jlya/011111111, ii is exposed to natural infection, were recorded as being du<.' ro
reasonable to su~pect that some Rhiµicephalus-transmined £. o/li11a41 and lhe authors suggested that this infection
E. bollislo11itw isolates 111ay be genelically homogeneous. and may also exacerbate Coll'riria infection. The sheep were
possibly also Hyalomm<Hransmitted £ bovislovina strains. exposed to ht>a\l)' infestations of .4mblyomma sp. and
Since E. chajJe.e11sis naturally infects goats.22 it is possible that R. evertsi evertsi and. given che possible infection of sheep
Aml:llyomma-transmitted ehrlichias also cause monocytic by E. boi,i;, i1 is possible either or both tick specie.s \\Cre
ehrlichiosis in small ruminnms in Africa. Ehrlichit1 011i11a is involved in rhe transmission.
known to cycle in sheep in southern Africa but ils economic Information on transmission is very scant, but what there is
importance i, unknown. bm is probably low. suggeslS that at least one fonn of£ 011i11a is 1rarnm1ined by Rhi-
plceµhalus ticks. Neftz 1'' reported that £. ot•i11a has been suc-
Aetiology cessfuliy uansmined with adult R. e1 ',l'rtsi that had been fed as
n}~nphs on infeeted sht-ep. and in lhe original srudy of Lesto,
Ehr/ichln 011i11a possesses rhe morphological characteristics quard nnd DonatienSJ an emulsion of ground-up Rhiptceph-
of F.. can is and E. bc11is and its mode and site of replication are a/us IJ11cy1 ticks infec1ed .sheep, indicating that these ticks were
the same. 16 The host range is not known. but since most of infected by, but not necessarily capable of transmitting, thb
the ruminam infective ehrlichias of ruminants are not spe· agent. Several other reports from Africa suggest that R. el'ertsi
cies-specific. it is likely that ti. 011i11a also infects other rumi- ewrtsi may be a vector of sheep-infective ehrlichias. An agem
nants and may be gene1ically identical to some £. uoPis transmiued by II. e1,er1si e11(lrt$/ from a heam,-a1er-frt>e fam1 in
strains. Phylogenetic analyses based on 16s rRNA gene Zimbabwe induced seroconvcrsion in sheep to C. mmi11a11-
sequences. or other genes. have nol been reported. and there- ri111n:~ this ma}' potentially be £. 011i11a or a related organism.
fore it is unclear to which gcnogroup of the Ehrllchieae the From the results obtained in another stud}' on cross-reactions
agent belongs. The amigenic cross-reacthiCJ,' of a Turkish iso- to C. rumi11ami11m In domestic livestock in heamvarer-free re-
late"610 Co1t•dria amigens suggests a lesser amigenicsimilar- gions of South 1\frica it was concluded that both Rhipiceph·
ity to Cowdria than occurs berween Cowrlria and E. cnnis. but a/us- and 1(1'alomnu1-reiated ehriichial infoctlons were
closer than with E. phagocycophila. presem: high proponions of seropositive animals were in areas
(pans of ;\lpumalilJlga Province) where R. e1·em.i et't!rtsi oc-
Epidemiology curred but not Hyalomma spp. or R. r1ppe11dlcuia111.s. and also
in areas (such as the Nonhem Cape Province) where Hy-
Uilenbergm1 and Nei1zb6 ha,·e re,~ewed the scanCJ,· lilera- alomma tnmcanim but nor rile other two species occllfrcd.z1
ture. £/Jrlichia 011i11<1 has been recorded in Algeria and Tur- Thi~ is consistem with the prol)abiliw that both genera of ticks
key.•3 Namibla.8 s South Africa.as French Sudan.66 Tran. 75 Sri maimain ehrlichial agems. mid is substantiated by the report
Lanka.90 Nigeria, 52 Senegal (Curasson. 1942, cited by Uilen- of Du Plessir0 that he had bolated ehrlichial agents infective 10
berg101 J and Mali (Rousselor. 1943, cited byUilenberg 101 J. It Shl!ep from both R. 1J11ertsi and H. mmcmum ticks. Ln Sene-
has been identified In blood smears from splenectomized gal?' £. ovi11a was detected in goats kept in areas \\11ere R.
sheep and goats in Senegal,3' and also in imac1 goats and eiiensi e11ertsi was the predominant Lick after they had been
sheep in different pans of1he s.ame coumry. sptenectomized. In soulhem India an£. o/.'i11a-iike agenl was
In epidemics in flocks in which morrali1ies occurred, found in sheep with heavy infestations of H. margi11acw11
the animal$ were frequently suffering from severe helm- isaaci.N lio\\'ever, it ii, possible that the appearance of the or-
inthosis. In addition. Schulz85 noted se1·ere tick infestation ganism was a recrudescence of a latent infection and that the
and malnutrition. In West Africa ,everal repons link che Hyalomma species were not lhe responsible vector. Recem
di~ease 10 movement of $heep from arid zones with a re- isolation ofehrlichialllgents from H. mmMn,m ticks, and from
stricted tick fauna to more humid areas. The low patho- R. e1,'!'r tsi and R. ap,-,entlicularns ricks is also consistem \\ith the
genicity of E. 01,i11a ror healthy sheep coupled with rare concept 1har ruminam,infeclive ehrlichias are rnaimained by
epidemics associated with monalities in flocks in poor the two-tick genera, alLhough the studies?O which indicated
condition in which affected animals develop heamvater- that p:uhogenicity of these agents for ruminamsincreascs after
like signs. is strikingly analogous to lhe epidemiological rransnnssion ,ia .-lrnbfyomma hebracum have 1101 been con·
panern of bovine ehrlichio~is and its fatal exacerbation fim,ed by others, and are subject 10 doubt.
 Le.~r--kno\\n rickettsias infecting liwstock 5 1l3

Clinical signs ment with 1e1;racycline dn1g~ is likely to be successful, and

should be used together with measures to reduce sue~, and
The incubation period in suscepti ble sheep follO\\ing infesia- improve the general health of the animals. Control may be-
tion with infected adult ticks ranges from 15 to 18 day\>.66 '111e come an issue in areas where tick control is rela.-<ed follow-
onset of illness is signalled by a sharp rise In the rectal tempera- ing Immunization against heartwater, and pathogenic:
ru re which persists ror 3 10 l.O days66 and i:, som<: case,; ror as forms of the infoction are encountered. a
long as 17 days}:; Affected animals nre lisUe~~ and stop
eating for three 10 rour days. Towards the end of the febrile
s tage. anaemia is evident Reoove1y is rapid. b11t the blood pic-
Porcine ehrlichiosis
rure takes four 10 six weeks to become nonnal. 66 Ehrlichia suis wa:. first described in Algeria. 1 1 lt was isolated
The clinical signs in the om brcak recorded by Schulz85 fortuitously when blood frnm a ~ick pig believed to be suffer-
were characterized by thei r sudden onset and short rourse. ing from ,\Irkan s\,ine fever was inoculated into a healthy
Alfected an imals are febrile and de\'elop signs of ata.~ia pig whi<:h sub~eque111l1 developed an irregular fe\•er and
before going into sternal a nd then Jaieral recumbenc) died 12 days later. The lesions mimicked those ofheartwater
which is followed by opistho1011us. unconsciousness and in Lhe sheep.12 The causative agent was passaged in pigs and
death. at the second passage ehrlichias were allegedly detected in
monoc}'les and in endothelial cells of blood vessels. The
onset of the parasitaemias coincided with the onset of fever
Pathology
and the ehrlichias evolved in mono<:ytes in a manner similar
Parasliaemia Is at its maximum two to three days after the to £. canis and E. bo1tis. The organisms in endothelial cells
onset of fever. 15• 66 Relapse parasitaemias may occur se,·cral a re very small.
weeks later.15 A delayed anaem ia, eosinopenia and mono- Rioohe and Bourdinl!O observed a l'atal outbrea!.. asso·
cytosis have also been reported.r;,;, 90 ciated with ehrllchial-like organisms in circulating mono·
The gross rhanges in fatal cases of E. oui11a infection are cytes in pigs. Thei· concluded that it was e&semially an
similar to those of heartwater and include hydropericar- ·£. bo11/s' infection in pigs. since a calf died 12 days after sub·
dium, hydrothora};. a.~cites, epi· and endocardial haemor- inoculation of blood from at\ affected pig. The studies can be
rhages. hyperaemia and oedem a of the lungs. splenomegaly taken as evidence that the agent im·olved was not host·
arid nephrosis, ts. 85 specific. and funher supports the use of genetic groupings
rather than host species in the nomenclature of Ellrflrhfa
spp. The tick vector was not identified. a!Lbough Lhe agent
Diagnosis and differential diagnosis
was seen in the blood of a pig and a calf after inoculation
Jr ma~, be difficult to differentiate o,~ne ehrlichiosis clini- with a homogenate of Hyalomma and Rhipicepha/11s ticks.
callj' from other febrile diseases66 and fatal cases of the dis· TI1cy suggested that stress played an important role in the
ease should not be con fused with hcanwater clinically and manifestation or the outbreak.
at necropsy. A diagnosis depends on detecting E. ovina
within monocytes but not in endo thelial cells. u Antisemm
produced 10 5. ovina (ofTu rkish origin) cross-reacted \\1th
Bovine petechial fever
Cowdrid amigens in the !FAT with moderate titres (to 640) Synonym: Ondiri disease
bm reacted only to the 32 kOa major antigenic protein
(:\.1.APl) of Cowdria.46 This su pports prc\ious findings from
Introduction
southern Africa that crOS$·reactions with mine ehrlichias
can give positi\'C reactions in serological tests for hearrwatcr Bo,•inc petechial fever (BPI'), or Ondiri d isease. occurs in
in Amb(vomma -free areas. A rise in antibody titre. for ex- exotic and cross-bred canle in the highlands of Kenya. 8
ample using cross-reacti\'C an tigens or C. rumi11amium Tanzania, and possibly other high -altitude parts of eastern
(rFATcould be u~ed'6 ) migh t Indicate a recem exposure but Africa. It was frequently reported in tbe past and the sub-
would not be of ,·aJue in animals with recrudescent infec- ject of considerable study 1n the 1960s and I 970s but is
tions. Polymerase chain reacLion (PCRJ and hybridiunion rarely reported now. allhovgh individual farmers in en·
using the approach of Allsopp er al., 1 and hequencing of the dem ic locations continue to observe the disease which has
I Gs £RNA gene would assist specific identification and been reviewed by Davies.a and Scott and Woldehe\\~1.6ij
placement of the agent \\1thin the 1rlbe Ehrlichieae. T he vector or APF is not known.

Control Aetiology
Infections with £. ouina are either so rare or so ubiquitous An ;imcialion of SPF witl1 a rickerisia-like parasite in granulo-
chat control measures hal'e never been formulated. Treat· ~'le.~was made in 1962 by Haig and Dansk.in.4l but Plowright7'.l
5-l·l .,. Ho,""= Riukcmt.il and chlumydlal diseases

and Brocklesby3 had earlier detected e,1dence of infection in oedema of the intermuscular connective tissues. lymph
his1opa1hological specimens and blood smears. Foll01,ing a nodes and the folds of the abomasal mucosa.-o
subsequc:-m demiled morphological stud}. the parasite was Fihrlichia o,uliri multiplies initially in the spleen and
named Cyt()l/(/etes ondiri.50 Ph}fogenetic analyses have not then, during th.c clinical reaction. It parasitize, neutrophils.
been conducted and all isolates of the organism app<?ar 10 have cosinophils and monoc)ics.91 Elt'Ctron microscopic swdics
been lo~t. bu L available eviuem:<: SU!,(l;<:Sts the age Ill c. dusely show. in 1u.lullio11. that tlu, u~a111s111111u) I.le rt~e in capillary
related 10 the granulocytotropic ehrlichias of the [;, phagai;yro- lurnens of the myocardium and in cytoplasmic \'acuoles in
pllila gT()up. with ,vhich it shares antigens. and may tentatively cndmhelial and Kupffer cells:1' 1
be considered as Ehrlichia ondiri. Sernfrom catde after experi-
mental infoction wid1 E. omlirl react to E. phagaqrophila in
Diagnosis and differential diagnosis
o,ine neutrophils, and in immunoblots to antigen~ of E/,rfi.
cilia equi. anocher member of the E. phagocrrophila genetic The importance of BPF is small. although should vaccina-
ctusier.103 Strain \'3riation is recognil.ed,88 in ,intlence and in tion for East Coast fever (T. panm infcc!ionl b<' introduced
protection again~t hcterologous strains on challenge. in these and tick control subsequently relaxed. cases might occur
respects the infection mirrors E. phagocrrophila infections of more frequently. The disease is rarely diagnosed in Kenya,
ruminants. but this may in part be the result of the widespread availabil·
iry of tetracycline drug~. which are effective in trcanncm and
commonly used to treat disease in cattle of unknown aeriol-
Epidemiology
ogy. a~ well a, the reduction in numbers and habitat for res-
The indicator host is exotic cattle. including Sahl\.\.il breeds, ervoir hosts such as the bushbuck. Differential diagnosis
wich .i syl\'atic cycle in wild unrelope. particularly bushbuck indudes other haemorrhagic diseases of cattle such as acu re
( Trnge/(lp/ws scriprus). The disease hal> onl~· been reported trypanosoinosis. acute tl1eileriosis. haemorrhagic septicae-
in highl) restricted areas of the East African highlands. at al- mia, bracken fern poisoning. Rift Valley re,·er and. possibly,
lirndes bemeen l 500 and 2 800 metre,. Common features hearcwater. Diagnosis is by careful examination oi Ciemsa-
of all sites surveyed by Walker10:; w,m: thick bush or under- staintd blood smear,. for infecrion in granulocytes. although
growth giving heal')' shade. where the soil had a 1hid.. li1ter the chrlichaemin 1nay be waning when clinical ~ign, arc
layer giving high humidity. and a resident populalion of most apparent.88 Serological diagnosis of exposure to £.
bushbuck a11d duikcr (S>1ll1ict1pra grimmia). !'he seasonal ondiri can be mode hy lFA1 or Western blotting again~t an-
prevalence is probably relawd to the use of particular high· tigens of the granulOC}10tropic ehrlichias such as £. pilago-
risk paddocks at the end oflhe dry season. The disease spo- cyrophiln or E. eqw. 101 to which significanl cross-reactions
radically ,pills over into domestic cattle gra1in~ fore1>1 edges occur.
or scrub.112 The \·cc1or is unknown. but may possibly be one
of the species of I.rodes ticks which is found in high-altitude
Control
regiom of East Africa. Recovered cattle may be carriers for
several months. and bushbuck for up to 21 momhs.;· a The Treatment with tetracydine drugs is effective if gil'en early.
infection has frequently been recovered from the spleens of since active multiplicmion occurs before or at the ons~t of
bushbuckshot in an endemic location. clinical sil,'lls,!'8 but onC'e marked clinical signs have developed.
anti-ricke11~iai dn1gs are oflittle use. The only available reliable
prophylacuc measure in endemic locations is t0 avoid grazing
Clinical signs
canle in fore~t edge, and in paddocks \\ilh J>atches of thick
Sheep and goats can be cxperimenrnllr infected. bul severe $Crub. 1he use of acar!cide ' as a preventive measure needs 10
disease is only seen in cartle. lt isc.:hamcieriZed bya high flue· be invesugated. e\'en though the vector is as ye1 ankno\\11.
ruating fever. lowered milk }ield and widespread petechiation
of 1isible mucous membmnl!l'.. A few severe cases devclbp a
unilateral conjunctivitis known as a 'poached-egg eye' in
Tick-borne feve r
which lhe conjunctiva! sacs are ~wollen and evem.>d around a Synonym: Pa~turc fe1er
tense and prorubemn1eyeball comaining a pool of blood in the
lower part of the aqueow; humour. Other clinical signs are ab-
lnu·oduction
sent and affected animals conlit1ue 10 ent. Pregnant cows may
abort. The case mortality rate may reach 50 per ce111.fi Tick-borne fever CJllFJ is a benign rickensiosisof ca11le. goats.
sheep and 1,ild ruminants in Europe in areas infested with
lhe vec:tor tick. b:odl!$ rici1111,;, wherea~ pasture fever h a
Pathology
minor epidemic <yndrome lhat occurs when dairY cows are
The conspicuous features at necropsy are widesp(ead turned out of their indoor winter quarters in die spring and
haemorrhages. lymphoid h}1>erpla~ia. and gelatinous earl) ~ummer omo tick-infested 1,astures.43• 98 Tiie infection
ma\ also be pt<'Scm in South Airka: a single seniencc in a uc111onstra1ed. ,:, There r,. no 1ransplaccntal t ransmissi<>n
paper by Retief L'I a/.;7 states that in 1969 :-icitz detected 1ick- in shecp.9.~ The presence nf infec1io11 in and transmis~ion
bome fever in shl'<'P on rarnis in the coa.,ial region or th!! Port via II number of fxotles , pp. from the temperate regions of
FJv.aheth and I lw11rUlsdorp districts of d,c Eastc1·n C'.ape Prov- Europe. :-.:ortb \melica. and eastern i\~io has been ,ho\,n.
u\C'c. 'This repon has. howe\·er. not bcC'n ~ubslal1ti3tt'd and the bu1 ahhough thcr,• are several reports or infected V,mna-
V<·cmr wa.-. not determined Since I.mt/I!, ticks are found in ""111,,r tides from -.:onh \mE'rica, thE'}' do nr)l appear cum-
Soulli Mrica and infos1 livestock, it would nm he 100 pe1cr11 for onward iransmission.101 The large nu mbC'T or
surprising if associated grru1tdocywm,pic chrlichias 1Jn·ur and report, ofidenli!icat 1011 of gra nulocytic chdichias in Jxodi!s
info<"ti,m, of li\estock arc occa~ionally dct~c1ed. 1ick5 from across the minhern hemisphere 5uggt">tS that
most ff nm all lxodeJ spp. will he shown to have in1·ol\'e·
mcnt with trltn!'llli,sicm ol <Jhrlichia:. 10 1heirna1ural ho,1s.
Aetiology
wilh occasional 7.0CH,otic or a1' imal hcall h conscqul'tlcl'.
Elufichia rformerl} C,1•1oece1e,<l ph11goQ'10philL1 was the Hecovered animals re;.ist mo,1 homologous reinfor1ions
first iden1ified agent of the granuloc}10tropk i!hrli¢hial but the duration of the acquired immunityvarie~. 11 ' Passh·e
species,35 and is considered type-~p~'Cil~ for theg,mi,1icduMer prowcticm oi the suckling young through the tran~for 01 an-
of organisms which no" includos I' er111i. and the agem of tibodies in I he colo,t rum does not occur. ,,.i \v11en pre,~ouslv
human grauulc,c) tic ehrlichio,is Cl !OF.). .111d the bu~CQning infccit:d animal~ reuct on re-expo,me 10 either ~ homo-
number of granulocy1otropic ehrlichia\ \\'hich are a,,,ociatl'<i log<>u~ or heterologous i,olate the clin ical rc$pon•e and
"~ch infocrion land a1 times. disease. of11>n called granuloC}~k paras1iaemia«re truncated.
ehrlichio,is, in horses. d<>'.;~. cats and wild n1mimu11~ and small
mammals across th<.> northern hemisplu:re. 11,e inlc<:1io11 has
Clinical signs
been rccemly re,iewetl"" following the dl;;ro\'(1ry 01 1he
1.ooalltic potential of granuioc)10tropic chrllt'hia, Tiw iniec- The• natural incubation period rnngc,; from 3 to n days."''
lion of ruminant~ ha, also been re\'iew~L 10 13 The causutive ;\los1 primary tick-borne fever infection, in young animals
ehrlkhia muhiplle., in mcmbrnne-lined \«cuoles m the nro- in 1he licld are as\mplomatic. In older animals the onset o(
pla<m of circulating granulocyte~, partkularl\ ncutrophil;. illm•s!- b marked bv the abrupt onset of a high fluctuating
and. le~s commoni}·, in monocytl.',. It, e\olutlon i~ 1)1lical or fovcr of one 10 two weeks' duration. In non-pregnant and
crlkhias ol the Ehrlichieae 1ribc. 1 " .~ll l:urope,u, and north non-lactating animals climcal signs are nuld and uon-
American isolat~ arc amigenicallr related hm the} are not !.peclfic.-;- 102 The prolonged fe1•ers are claimed to impair
identical. and considerable amigenic homology occrn, ~,ithin sperma1oge11flt,is transiemly in rams 111~ und bulb,.- Nai\'e
the f. 11lwgocy10philL1 group: cross-reacrions oc<"ur in ,ero- pregnam animals ahon while still parasiwcmic rwo Lo eight
logical 1e,1~ \\'ith 1he flGE age111 and [. t~/lli,n and \,il11 day!! aftc.>r the onsut of rever.87 Occ:;usionally. the rooms
E a1ulirl. •t ., Howc:\'er, the strains oforgan bill which can muse mummifies and i< 11xpclhid \,·eeks later
disenSt' in niminant, cau,-e Ihde if any e,it:l'nce of intec1ion in ·nw dominam dinkal sign, in lhc pasture :a.yndrame are
hon;c,.-, and 1•kc: l'erm. \trains from hnrst'> generated very Im, high fever and a drop in mill ~ield."" Affocted CO\\'~ are ~lightly
le\'cl, or infection without fever in lambs, and did nm pro wet depr<!Ssed. are partiully inappctent anti cough i11frcqucmly.
ogainsc cilallenge wich pathogenic I'. pllagocytophila. •, ln ad- ThC' imponancc> of lick-borne fever in non-pregnant ani-
dition. there is great genetic similarity in the horse, dog and mals li<'> 1n iL, ability 10 e.xacerbaw ot lwr concurre111 latent
human pathogenic i$olates, whew.is ruminan 1 ~,rain< are infection,.•• The mo<t troublesome sequel to nawral infec-
more dl\·erse. This sugge.:,ts that strains should better be con· tion in lambs 1~ tick p) aemia.' 110
~idl'l'ed as pathotyp~. and that thow \\~th ;,,ounotic potemial !11 contra,t. le p/1t1g<XJ'l1Jpliila ma> also mitigmc the
an: nm the ~amc a., 1host• commonly c:nu~:ng clli,ense In rumi- effect:. tlf a concurrcm infection, prob:1bly \ia immuno-
nants; it is nor known if the resenuir spl•des arc .tl~o diffC"rent. suppress1ve effect~ through altered granulocne1 and l) m-
phoc~1c1 1" (unctiQn~. For cxumplt!. L'/tl!ryrhrozocm 01·i, db-
uppears from the blood or shi!cp infected 1\ith f.'.
Epidemiology
phc1f{O<J'lllphiirl" and. similar!}, splcnccton1i1,ed cah·..,, in-
,h natural tran,mi~sion is by ixod id tic,($. there i, in F.ur- fecti:d 11ith both f: plwgoc-rw1,hfla and Babes/« t/i1•nge11,,
ope a <ca~onaJ prt>\'alcnce of the disea~e whirh i, linked to have Jes~ mar~ed ha1:ma1ologicnl chrmges than those h1o~·u-
the e;irly summer rise in 1he number~ of ques1inl! ticks.·- loted 1~ilh either pathogen separaiclv.11
Although £. 1>hagtJcycophi la allegedly mulllplic~ in 1id:s in
the ooplasm and mi1ochondrla of de\'eloping uocytes.~ 4
Pathology
tho: agencs recognized might be true Rickerrsit1 5pp. since
Ri<:k<'tl$ia /1t>b·<1tic11s has reccntlrh<·cn identified a, a com- lnr haematologkal change, arc diagnostic: at the onset of
mon inferrion of /. rici1111s tick, in Europ;:,.~, and only fever thcrc Is a profound but transiem lymphopenia anribttt·
trans,.iadial transmission or /:.. phngol)'rophiln has been ahlc w a significant decrease in peripheral 8 lympho<:)1es.~ In
546 ,,...·110~ nmn: Ricke11sinl and chlam, dial diseases

addition. the reactivity of the lymphocyt~ ii, reduced. 11~ Two Diagnosis
to three days later !.here is a profound and lasting neutropcnia
and often a total eosinopenia. l\lonocytes increase slightly in Outbreak, are ,ea,onal and follow the ari'h'al of naive ani•
number towards the end of the oven clinical re-.iction. 87 mals omo pasmres infes1ed with quesling tiC:<s. These fac-
Concurrently. !l1ere is thromboq1openia.~1 tors. together wi1h the clinical signs. warrant a pre:.'umptive
After experimcntnl infection b goats. a decrease. In diagnosb whtc:h may he confirmed by the demon~tration of
serum alkaline phospha1ase dt1ring the fe1·cr. n marked E. phagocy1ophilt1 in granuloq,ei; in Giemsa-•tained blood
decline In plasma. zinc and iron concentrations. and a smears. :,erological e\idence of expo~ure can be made by
slit:ht. $teady decrease in haemoglobin and haematocrit val- IFAT\\1d1 £ pllagor:yroµhila-infected neutrophlh. although
ue1> ha1·e been detecte(l. 1112 antibOd} levels moy rapidly wane.
Parasiraemias are patent throughout the fe,·er. being
most marked on the second day of rhe febrile reaction Control
when up to 70 10 80 per ccm of granulocytes may be in-
fected. lnfec:1ed neutrophils are functional!} impaircd.3i. Shon-acting teiracycllnes are considered effective 115 for
111 Monocytes are also para.~itized. but more rarely.lll, 15 treatment of cases. bm a situaiion of endemic siability
.Most affected animals become carriers and the carrier state should be encouraged 1,·herever possible through e;q>osure
persists for weeks, sometimes months. and occasionaUy, of young animals Cat least prior to breeding) 10 Infected
years. 28 Relapses are not uncommon: some relapsing ani- ticks. Early expo~urc. during the period when colostrum-
mals develop shon fevers but manr do not. 8 • derived immunity in young animal~ is still present, may also
Death from uncomplicated tick-borne fever or pasture ass!St in co111rol of tick-borne viral infci;rions. In contrast.
fever is rare. Splenomegaly, hydropericardillln, endocardial pregnant animals should never be moved from tick -free 10
haemorrhages, petechiation of the mucosa of the small in- 1kk-infe,ted areru,. If the prevalence of tick pyaemia b high.
testine and haemorrhagic colit is have been recorded. :IJ. 15· 6 ~ losses may be reduced by the use of an acaricidc and by in-
.-\ heifer that died after exposure lO 1kks had lesions reminis- jecting lhe arnmals prophylactically with a long-acting tetra-
cent of those seen in bovine pe1echial fe1·er. 109 cydine.11~ If pas1un.•5 are re-stocked \\ith aduh animals ,,ith
II has been reponcd 1ha1 the lymphoid rollicles in lhe no prior e1'po~ure to ticks. iliis should occur when they are
lymph nodes were drained or lymphoq'les:' 5 Examination not pregnant in order to reduce the occurrence of abonion.
of impression smears made from 1h~ cut surfa<"es of the Treatment \\ith 1c1racyclincs is noi warra11led in uncom-
spleen. liver and lungs will reveal the imraC}'lOpla~mic plicated tick-home fever in lambs but it~ use in Infected lnc-
presenc:e of E. phagocylQphlla. Occasional ehrlich!a~ may be 1a1ing cows reduces losses by abating lhe clinical sign~ and
obser1·ed in ah·eolar macrophages or the lungs.6 ·1 the parru.itaemia.''6

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0

p.tthrJg..'Hl" pou: l~ hucu(. Rullr-tm tltt In .'X>dh,~d,. f'mi,c;J~tr &ottqm·

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:?7iA-.?rRO.
1s no,:Ant , .., . .,. u,1-nq\lAM, r .. tt:i;. Et,11 :,cn,d di.!'<. (onna1:,,;.u1:e~ \Ur
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a "\\1£, G. 1~93, Borlnr pe1t-cl11:tl fe,-.r :Ondiri dls..-a;;• 1,.-,.rlnal)• r1 DOS",\Tw,, .,.• u~roc,u,Ho. r 19~0. Ou (}-de e\·olutl! dr qlit?que,
,\/1.-rohioloic,•.31, 103-121. r:da:tt,:a, .-tumn·> tit /'Jmtt1111 P(IJll''" <1·i-14:..,,w. IA, lo~:? t3
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1hr1knnh ln South Afrko "i1h ,pad,~ 1en,n~o tQ T11etl,tl11 mwm,,._ R. lxwu Rulbmn ,It, ""Xitr,• Pmholog(f! E.lOl111u,. 32. 2•t!>-2-t8.
 Les.<er-kno\'.rt rlckensla. Inferring livi,srock 547

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dornt~tk rumm..1.nt1> in r<!'i:tlon, uf S1>uth .-\l'rkn. whrh." AmMyonwu, 4.1 u .,1(,. n ..\. A nAx.,>:1-..;. n .. 1~62. The ~tttiofo-gy ot ho,·mt..' pctL'C'hl.tl ttwc,r

Jt.,t,rtt,:flm doc,> nui occur. Oml~t••tX)Urt Jaurnttl af\',wr/1111,y 'Reif.•.ard,. tOndlrJ disea~t",, R,•J.t."CJr(-t: in \"t!lt"'riuar;· Sdt11cc.•.3. J2'l... t38.
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lnf<Clinn ol dome$rlt 1to•t, ""h l!htllcllla clwffttn«•. Jour,u,J OJ CU111ml """"lrtra tn ca1H•"' {fldltm t (NrltW') fmmu1I, 70, 86.~~
,\llcrob/otpgy. 38. 448-1-19
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F.ltrl/J:hla plu1gocyropltit11. und humun gr:,nulocytlc cl,rllchlu. /m,mnt of
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:t.i OUMt.ER. U• 1.: \\',\l-t,."f..R, D,11 .. ~001. TIC'k•borne ehrhchll)(i(~ Lnntf.'l
nm1/111111rfum I,""""'"',! \\ithln 1he genu, Nulirltin R,·Mi<rl't'lt.·ag,.,
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/11fe,:rlous o,,...,,,.'$. 21-28 April
.fi l..t>t~HAJU. f,:'11, 0.\\JU!!O~. \', .SL O.\\\''i!OS, I L 6 ,a \I.I lr;\f'\.'.Jrr. O.L. I~~~
2.5 r,,·1~-r.. t,,,.,,_ o.,w~~. J,f ..m4,, .\..o\., ttA~-Dt. R-.\\ . , ., ,xsrn <:-~ .
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lransml._..;on of E.hrlirhtn rlwjjt1msis (Hr,k,•rrslaln: Fhr!:c1,,,--at'. ;unong
01,'(.("ll,"t!'.\·.!n.1 ~~124
\\hhc,1aHed deer by r\mill,1·onmraamcnrn;m,m CIIL-ar,; /zodrdntt,, Jnumal
,1fMed1ra/ Emomu/Q1,.'Y. 3:?. 363-374, ,18 ._,tf'lVo. ._,~. ts&. Rtr,;~u.-.i.tl m(ccttons in ;,nfmill\ A f\''\ 1
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2t n,nus P,1 t~~. ~ot,:, cl.inJqm.•. ftidtttl!'tfo5rt l Rlcl.·,,.:;du bot·u cm /nurtml o/t't'itrmmy S<it'rl(~. J2. za2-ao1
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1
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T'11/1/c<11uc. 11. 291-292. by tl1rllcl11t1 omlin 7111• t·ru•rl,rorv Rl"'Ctml, 11 l. 5~fJO.
27 Unc- cid.~tt,i1.ht.· bo,in~
JLOCH. JI., u.PPOSI. \1, .t C.tntlUlJ. f'.. 197,l :;o );lt,\U!o'J, IL O.\\~. r.n.• 0111.t.V.JO). 0 ..\, i, CCKWU(. ,., • t9-;".l. 1bc
prolMbh.• en Citl-'dl·l~up,,: diffkult(,,1.du dias;na~u·. Rnltrti11 ,t~ {(( ~n'lrJ morpholllll)' 0'1h~ cau~al 3St'tll oi bn>1ne prrech•nl fr«r. Jmtr11<1/ of
r:r1•t,1hnlogidiz,11f1111,, 6$. 19,1- 1~8. c:nmfJ{1ta1ilH! Pmlwlc;J.-."·· 8.2, :?-11-246.
~ J-OGQt,l\ I9.SL Srudi(~ un tic...,.._ py::emfaand liCk·bomo Cl',·,u.
51 IOU(C.. X,R.£, l-l.01 T. J.G., 191\,; . tre,s.t')':~ ,\4m,u11/ o/S)':itc!mR.Tlf Bac.'"rinl"K},
~'.l'IIIJ>osuun ofth•\ LootoJ:,lrat Sorit.tJ·ufl.oudun. fl St-sa
Yo!. 1. Bwlimon-< \\'llli:uns<> \\11kin,.
i'J 1(l(,...1f , 19>Q!!.. Studic~ Qn 1h~ tnfcctfous, ngcni of l!tk·bomc fi:vcr in
52 wn,.\SY, p . & 11.£.,:etMOt:, .\.--\.. m;;. Tick,horne d1\e:s$r.-5 o( dm:ic-.ur
sh,ep. Jqumnt ofP111/mlot;)'""" lklrterfq/r,KJ·, 6l. 1-15.
ooim:tl, in nonhi,•m ~~cri;i.11 Restl'~ch smnm.uy. 19&1 ,o 1976.
3r, u.1., a-so.i. ~1ud.f~one1perrthm1.nan infection in
ttx;f11L ,. t- ~,:.af-T. fropit(I/ .\mma.t I ltwlrh muf Pf):H/uaum. 9, 211-218.
~Np~Jo11nu1tofCm11p<Jrmiw PmJ,aflJfD,. ;-4, .is.-ot.
r1."l ttc>s,,n,:,,; ..\. l9J'Ei ~ur un.e nouv<.•lle rt.deu,1>1:! du
1.J!'1'0QVA.1Ul, r i-
Ji R.)!it"ftR. ,,•.,,.,1. &C.,MDOl'I.", A.t ., 1908. Thror11bo~1op..,~ntn u, .Sh<"up
ntouton. 8ulf,1w d1· ltt soc1,•1td~ ParJrolog,et fo-:Xatitfllr, :?9 105-lOS.
.l••Mleinu..•'d \\Ith ~'Per{mrntnl 1icJi:.h(1rnt, ft\·tr lnfccr!on. 1,,11111rll uf
Comp11ra1it.\.•l'bthol01:,.'l', 78 251 ..25,l;. >1 uw,,. o. t97:.1. Th,• di!1cc.:1tf)n o( ncken:sia•likt• miCrn•orgam,-na,\ithm
lhc Q\'ttrlCi o; lrm.ok JxQdP'S ttdnus ticks; Xvu~l,rt/1 I"' Pamsi1t•11kt111d~,
32 rosrtR. w.N.'.\t & C."'-"''ut()~. A.r-~, 1q;o. Obscl\·111fQ11~on iht- runctlonal
59, :lll:;-298,
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<f(:JJ,n/)a/tl:it, Pari11>IOlO'• 50. l87-l9l. JI, E.'°P~lmt:nt,
55 '\.\[Lf.00, r. 19Jtl, 'S)Ui;ilC!">Oll licl,1.•bomc (~•\'l'rof .:-h('Cp.

3J t0'1T1' \,·,.v t()("..l*ll "-"''-''HM,01 19M H~<'mn"hrt,-tlcrnteridsln

on uansmt~l<>n and dt..~mbtnton ni lhl! cflkast"- Pam$1ml<Jg)·. .:a.
.1211-ng.
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'l'.i i'.OR.fXJ,. \\ ~.. BROWXUf. \ .• \\"II.SO~. D.R. a. \t\Cll0D. , .. lf*-:31 Tld•bomr 57 ,uc:uoo. r, .,, ,.o,mo:o-, \\·, .... 1933. Studlc,,. Qn titli.-bomc (('1,'Cf of $htep.
f.01,~ :.- hitherto undt'4',Crlhl'J d.1¾"3.,;e Of~hc~pi. Jourtml ufC0111JJ(Jr'ntil11· t, T1om't.fJl1\$10n b~ the rlc~. l,odc... ricmus. m1h .a descnpuon of the
Pmltol°&'Md Tht•ru;tt.•Uli'CS4 -lS.301-309. di.'rl!a,c produ,C'd. Pm<t~lfoftJSo·. :!3, 2~.203.
3b C:.Ul.\1:.A•• MIU~St;U(. ,1 .. u1our,. ,\.~~~Yll. M .. ~986.11(1\lts.-ct sn ,u1{(".11rrn.:, 'N, 19~ ()rtgm and ,•votuliQn c>( tlw 1'.'hr!ich.leac-. Ill: 'fi\N
llcm1>1>nfi1$it0<0'>du bemil "" S...negal I. Lo ret1on d,':'-..:i•ycs. Rrvue ~Jll"n'L,<," >IIU ~•, 11. ,eds1 £1:<Jtoglrat JMmffJl>.Shlps 1111d Fr>o/11rio11 nf
D'f.4•i'iJgl~1 di ,\lt:iltt:im• V~t,•rr'tuu~ dt,s Pny, TrtJpfc111u. 39. 381-393. ,,.,. R1rket1$1M,· Vol. 2 ll ..ran Ruuge: CR(" Pr"*
~ f.Ul;'J1'1 \., 'C,.\.,nc.~ '·'- PIOUt •.\. "1 \IUN(i,lff.. \1 t~IJ"7. T1quc~ C'I 5tt \I \("'-.\JU) c. PL., .. N8,t. Fhtlttftifl b<tt:ls (Oon.nHm & J l'\(Oqunrd... !936 •
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Ptty1 J'n IU((llt.'(, 40, I 19-l'Z:i,
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betull .tu Scnrgal.111. La ,one uord·s<>udanlonne. /1,11,u, D'El,w1~~1 <I, om Gado de t.,tre no Bra,li ,1,q1,/1'1Jl t/11 U11/tV'f1.id11rh- r.1/tml '111ml do
M,'tfffl,t~ \ ~ittflrinflirrdt1 Pt1)~ Tm()iC:((llL .;z, 41 l-UO. Rio ,/1• /mh•i.ro, S. 2:-1;-219.
J!f C,111 \'l. ,\,••,1ae.<;u1. \! . IH011f-. \ £. VASSIU.\.011'. 1:., 19-$~• PJnpbyhudc dl· f11 M,\<.'-tJX<... R,r.. owr:--.~. ,.11 .• ,m,.s. o.., KF.Jiu, t,;:,u•• rr.n.ta,T,.• ,,.,
IJ. (O\,.:driose ttobwl'\·alion~ sur la pmhologhr o,inc.• d.uti .Ll ttglon <le, BIOfA.,OORff". r\., f'OUC,11LI~. 1,.T., IU1T1.. G..\. t,.,:\!Ull'1'tt\, C:,I.., 2000,
i\ta.yC"'- JU Sc.nt;gal. Rn•t,t D'F.lt•ivt~· l'I dl" ,\frd«irti' \ '<"tlfflnntrt' d,•$. P11.~~ 51.'<lu•nc. an,\:)'l>l' of tht onk gent'Of lJranulOC)'tic chrlkh1a,.Jo11mnl oj
Tlt)fJ/CJJu.t, 4:?• .;97--Sf.13. <:llnimt .11/crobio!oif>\ 38, _'!!! ;-in~
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62 \t.\tsus P:.A 1967. Th.cilcnosis in llhod~~ 1. A ,1udr at dt~no,uc 83 ~ou~no1. •· 195.,. XOtC$ d~ l'~rasholo,:1rs I ropicolo. Torn~ I. Ptlram,•.<
,p1.."(1fflcn1.,. O\'U ,,,t)
St..-a.\.tJrt" /m,mnt oftlli! somi: ..t,,friam \ ·,•1t•rinnry du snugm'1 ntlitnllUA' P:!CL<j: Vigo1 Frere:,.. p. 152
M,:ditt1l ASYJr1mfm1. 38. U.l- 1O:!.
811 !',\\LD\l.. l),C., ).-l:,:U ,. , •. , $. \UH.Al\, S,M,, 1'98. b1d~ni,:«.• IO 'ihO\\' 1hat-an
bl ,,cn,·t ,, A.O•• 19,.r:. Tick-lu)rnl! re\cl in )'\)UJ1P, rambs. 11: ...• \ ,:wrma'), :tKent that cros,,rcat&$ scrologicolly \\~Ch OJu.vlttu rum 11rm:1Jwn m
11,'<'ortU,9. 198-201 Zirt\lMb\\'C ~ Lr,,n.,mined hj lie-~. fap,•rim1mt11J cwtl .'4.pp/i4.•d .-l«tTOlo,01•

W :--1U~EU'J. ll.. HlJ,1tR,.\,ft,. MAt'1ri..L'.1,lt_. C., kMOL"-lUt~. D.\, t982.

22 111-122.
Pulmo~ry lesions. m )lh.1<:I) lulluw1n~ ~*nrtrent3l tnkt:1ion by 8; ,<:1tuu. 1.: •• 1939. ,\ nc'i...tcr...11hl, 1u.'\,• ra Suu1h AfrtC':I. Onlf1.•t~ri.'J10Dn
F.hrilr.1Jit111i11:~·rnpltil11 :,nd Cltlom)"tli11 ~llfm:,,. /mtr11,11J ,;,{ /otJr1utl \If \'<Wrim,ry•Sci<·1:ctom1 .·\11inml /ndt1.,1n·. 13.13:'-289.
Com parati"• Put/10/"8.)•. 92. 11;-129.
a6 sc:on r..H.• 19ln. \\'h;if,. n~Kln tft'k,bome ft.•ve1. ..1nmwl Pro:"'"'i'lit:C$t1/
,;~ ,t1 r,. i956. A coO\OUdiJtltln of our knO\\'ledg«: c,f tnm..mt,..~on t,J
\\.O,. ll~ Wrtt'I' \ 'rTtrill.M')° S()ril•I)•. 5, 13.
11ck-bomt" dh.t.la.ie,. (/ml,•t-sti-1}Ql)rl Jour,,al of\ f'l«"J'1t;11,:, R1'll"'1tl'l1, 27,
8; st<>'n v.11... iqa,., Tld,·bOrrH: ft!vL,r In 5h<,"Cp, Vt•/('ti11nn•~l,mual. 24.
11~163,
100-106
66 ~l n-i, ,, .o., 1968, EJrrlil.'ltia ,.wlm, fnfocllon. B11llt•t111 dr fl)f/tl't
80 M:OIT fbrt, " \U)UlUU•WU. , ... 1993, lk,\•Jne petc?Chial ft~\·ur /,1; WOLD£·
/111emnrio1:al tl,$1:plz,,.,//P,. ;o, 337...J>IO.
ltl'.\\lt.~ ~ ,u . . m. \t, l'd\l, 1rid:er1tilul muJ (Jilmuyrllnlrf.4wJSoof
ti7 ,it.:.<o!\ . ~.• 11,PQu1..s1, o .• uu, ,\,J., , \I,-.<~~. 1 .y, fl'IM \.'. r.. ,.. 11M11.,n,. DQl1tffllr .-\lunmls. P~rganu,n P~. p. 427.
t ~ . Rid:tllsut l,d,.,·tlru tn /~rul••.f rkfruu tick.'" in '\\\etf("n, Jm,rmll n/
~. ~t:\F.'\UHl\,\, "· • mr.\S\P.U.\, !,.U... 1963 1.:.htlJ<'lm, lKWb·lli<c o~anbm> in
C/i11ic«I './im,l1i<1/r,gi• ;1; IOIH03.
caul~ m C"J'lon, C.,-/011 i.wn11my/,mm,,I. 11. 101
6ft ~oK,·\I.. J\.i\.1 .. 19:'tl 1 h:~ in(t•,u,ttun"' ~ncf lick·bomc dlo,\•a.w.. w
~ .)iC~~" \T'-,\. t". i, Jl\t~Ul)tt,:O., \t. R t9Q7, l11c l)ft"X%\C'C vf rJ1rllc/1lJ:
l..lmb;:ib\,·1.· Rhodc.. ,a. Jrmmut r,; 1hr SO.Uri• .ifrlttm t't"it>tllWf\ .:w«imitm.
cwi,w·I:~c ur,ron-1,m., 1n lh,• n1emonudc:tr c,"('ll, io CC) lcn ~·lrJt1
50, 2119-292.
r,•,rrlr;no·Jm,rnnl, 15, 1-11
Gt! OGUI,- ,. , .H .. \\Otorlll\\'lrt, I. .. lfAttf, li,,\., 1':f!W. GranuJoc;~·dc: ,·hrtithlo~ii'
$1 ~'i'Ot)(..11.\.~~- t'H{., 1975- P.:uhog<."ne-Js of bu\1n" l1Cl~Ch1aJ fcv..·r. l..ih:nt
,n crnt.'fglng orte<ll.<c:ovcl\.-d tlck·bamo disco><?. f11umr,/ n/ 1/rolcal
,nfc(llon.., imrngn1tr .,nd tis..1ruc (fis1ritnui<m flf c:_wo,'<1"n'1- p,1dlri•
\l/croJnalQ8),
. H. •lili-162
Journal n/Compnrmlr•.- l>n1J,olog)', 851 52..1-5:to.
-o Pll'.J\O'. !t,£...1953,. Bovine lnfl·ulnw, petcch1•ll (e\·cr. Ea.•t..lftUm:
9l "-XQD(;Jeh,i,i, :t.ll', Mfl'f.\f), 1. ~ c.;OOVlN. 1,r... ~75,. lhe tOl\.1 Qf nild
.-IJ;rimln11al !011rru,I. rn. fii>-tll\.
rununilI\l, In the cp1dcmlotogy of l)onne potixhlal fc,r:. Jo11mnl OJ
71 rtt:RMt. J;., 1.983, I. '<'httkhi<™: ha\:in4• en U,tt! d'lvoirr. Ef'lk!1.•mla1~c lfJ'f(l!nr. Cnntbn<i£tl!, 8S. 2-i~250.
troJtmem-p:oph)'l,txie. a,a•ue ,r1:1.,,,'fl.8t' ~ de lrid«rn, \ .¾t!n11nu,· tlt'fl
93 ')JtlIJN:\1.\R. •• \:-..';\.,oo\, k., 8..'\J..\SUXD,\lt.\.M, !-. ~ JQ1t,, t,. ZOOO.
P0.\1 TtOJIICOIU, 36, 33;.3~ I
1Jct~c11on oi on Eltr/1'hia b,wt,.IJke org~nism rn cultur«I builolo
72 M.(m HH;tfl . " .• 1966. ~omc nou.•,. cm bovine pc.:t(biJJ fo\ct iC>nrJirt mn,wh'tf,., rttJpirdl .A11imttl Jfrultlr & PfOttu,·wm. 3~. &;"-i2~
disaa;c at \lugug,c 1(~11,.,., H111/r1111 ~/1/1, F.pt;;,x,1/c W.<,'OJ,•, 11/, lfri<TI,
10,.i~so,. ~.a STAh'«"lfm, t,;,C;., \1,\.~"'US~. "-fl,. ,\IJ\CJ.:MU',1.U, , .h•• IIUO, I,\'•• 1, \.':Ot:JtS()'Si,;
1.r•. 1999. tnrection w'uh .t~<Hltsorhu111an gr.inuloc~iit l>hrl£(hio~1!1. lyr11t
;'] VU1'~hU. K.1- YOlJ'l(;, kll., IHliH 1-1.L,tO' l),V. /.I Hl.;\l'U'l'. 0,1 .• 11,i';"':'~ 'rhe
dl~t·J~t. 4.ttd b.t~o,! ... in wilt! whh"·.roottod 1nicl.!' lPcttmlJ'ftlt.( lt!UCaµus
or
hocm•tolo~~ e,pcnmc1113llr ind11red B. 1/i1'<,g,•t1•'1tld 1-: in Connl6-Cticu1,... Jo1111u,t of0111/rol ~llttnUwfo-.t,,.",:, 37". 283i-2891
111tt1f,vxy1opl1ila h,rec,ion"I in .. p11.mc.~tomi-..1.-d c~k~ [lu, V,·1,~m:nr,,
q; sT.,:.1,.1.1.4,,\\\n l \...l~ Tkk,borru:fo\~r&;a.UU~l'ofobo.rtlonin
1/N:tJrd 100, - ·
sh\.-ep. n,l' \.~t,'rittiJ,:',' 11,...-ord. 62. 46,:;,-..tG,8.
:'-& VU'>nRt.\. '-:., PU<TLIU.\. J,H .. tm.,u,. u. •-UrrL H,, \8':.111, r~,wr1mc-n,-al
c,.,....... ,nit-'Ctfons withlilul,rltl11 r11t14,1rx, tt:111JifJ11 and hutnan gnsnule><.;1k !J6 STVC.\. ~ .\:n-JUR,ii.O'.'.. K.. tr.. Ot~~o~ r~ra·.\ti. t-. •• 1~98 E.'<J)l.,rf.m~n131
,•hthrh,a-llkt· •gent In co,-, and hu""'- Ti,~ l'Pl,'ri"III)' Rmmf. 1·1/i. Ini't.~tlvn of l.tmb~ v..ith an ~qui.rtt.' gnmulOt/ tk F.i1r1frlrJ1 s.ptdt.~
311-JH. rcsembllnt Ilic .,gent 1h01 t•u$r, humou !1f011uloq·tlc <hrllchlo•I•
if·IGE, .. lrtt, Vrterirmria &·m1,Jiumjrf(1, 39, ,191-,<Ji
i'5 ILAH1j A. 1> ,MA~I-L\.\U, r,,, l~U;(i. C:omribu1io:, :1 l'ckudr di" quc:l<IU'-"

p-.1r.1sith du ...:mg du mm1mn cl de lit <ht"\'TC en lran Cl dnn~ fe:. f\ilY-' 9i OIIMlllH ,u ,Atll,\.\', Jt., RAl\\'fl.lJ, , •. 6 -,s,'li.U\S. ft .. \S91L \ \'<'Cur
\'nt,10~. 811//£>1J11 ,trl'OJJi.;cr 1111,,mmloual ,It·· EvW,ot{e;. 6.'i. I i69-1783, potrntiollit:,· t>f llyalvmttut mmginmum utwci in &ltt• tr.1n~mi,,~fo11 ot
n1t1tl~(lh (ll1'1Ulllt11. t"l1tlr(ll1, 21. 10-12,
~ IUO, \' ' .\., ta.,IA'-ts\\'.\Mt, Jr,•.S .. f'oJflJU,..W·J• \, 0.1 .. IO~tP'U, '- \. ,1;
(j~\'!I.\.D.,M~ ,M, 1,1· 1~i.A report of coneunt.Ynth.wmop-:c.,to1.u.an und 9& 1110"11•. t~hi....,Ilda~ tn tpid~m101ogyofbo\1nc uciri·bomt- fe, e-r tn
nckcu'(lnl fn!tttlon, In ,hce1>. lttdiw, \ cwrt,:m:,·1,,umnl. l>8. 18i· Ui8. F1nlo1nd \\Sld" dinh:.;J J\....,crl1>tlo11 uf flcld c.,~ ;\tm<tk~ .\f,,(lictnai:-
nmo. G,I'.. ,Hu., w.o,. /i'.\t,,r.rAttLAsl, 1.,
i9-1. Ob~"-aunn~on lltl' /:;tp,·nmrnmli, rt R101ng:ar F~nofac. M, ~uppl. t;. 1--0"?.
~ffet't nf tltl-bpmc f,'Vl!r l(.)10l'i'l'I-,, /IIUlg<><,'Wphl/11 Fogs:fc 9;qJ 1Jn th1: r,~ TOC\'\JI, f. 1i,b; F..xp,f°ritncn1aJ :§tu<lic.,; Qn bnvln~ Ur.k.·bom~ k\ ~r.
>l"'tmatog~e.t~ofbuU.,. /mmml 1,fthr 5omh .-ifllctm Vcr,•rt,,nn· I, Cli111r:,J nod h;u:matofogkal ,fota, s.o1ne p:opc:rti~ ot the 4:.\u,._·ul,'='
.-w«;,,,;,m. 4!, ~21-325. agem c::1d homuloscnt.1s nnmunit}. Acta J~rlmlo,;tr:n i!l M1rrol1JQ!og:,·a
;s R1t><:11£. "·· 1966 I.a rlck<1"lnse gciru-r.ilc ·,,.,,tilt' 1.u Scn~al. Hri·11~ Sc'andil1m•ktt iO. 't:?'9-1-15.
d'/!.Ltt't1gNt de.\/itfn·i,w \'lffn,utir<'d,-s /'11,1"> Troµ!r,IIU<, IY.4~19l. 100 1111-\1\, r.1 '\S-\C:LfAS, ,.n. c;nuso, c.R. &1-1')1t8tts. t>.1t•• 1!:'9z. A case of
-:-, moc-:ur, •.t .. 196:-. Wjnn!, mkro~coptqu.:i~cfo la rk.i;.f'thia.\.l° s~ntrnlr. h1m1an thrhch1<»1• acqu,rt'd rn Mttli-<itn,rJl ond l;ihor.11ory nnding,.
ho, int- a Rlck~tt,;ia tF./111/r/r/11 /><11•u OanaMn ~t l.t\toquard 19Jf,), Hrrn~ :\ntt'l'i<M /Q11n1nl o[Trt>pia,/ M,vlit:i11,w1d l/,11/fiM,'. llx !61-1 &1
dtok1-oge,i d<' ,\f,'d,..f,11• Wlirlnnit,•d..s p,._., Tru1dnnu. iu. 41:;..1?~. 10: UfU:..'\'.DE!tO. ()., t~J. Oihl't\.'h.rlh:hJos.esof run11naot, 111: \\Ol.1'1Utt\\1T, z. &
tto ktor.rn., ,1. a. 11ou1to1,, r .. J961J. Rkl;l!u,;;ip)(" d~ m0no~1e( oh'io.(."f\.'t·t' au~nt ,1.. {l'd~ • Rirk,•usiul and Chlamydinl tliMVJit.•so/Domt,iofic
cht2 lt potc au sent-gal lltww, d'JJt·:.VJ.J;e c.'l dt' .\!ltl~tth"' \ irr,1111,u..•d~:t AnltntJIJ.. P-~~.)n10n Prt.!SS. p.. 127.
Pa)• Tr<>pTrmL~. 21.45~111. 102 \A" Mtarf \.S..J,l'.A,\1,. \ /I.,..._ DUl:-0.:, C."t,)1,. Sf.llOT\l<\~, \,VI, 4o fRA.~sst~.
1h hJ~11r. \s., tg86. l)crtlnl'hl charaecurh,1k( or lcuk~1k rfcic,.Ht,i:ie or ,_,_ i,1114 C:llt11cm, ha,ma1otoitcnlund blood biothem,col cha~ge, 111
hu.m:uisa.nd.antm<lb.Jtr. u1\·t.. L (~. Jfic,.obiof(JiO·- Wa:.hin_g1on. ~o=u.s .i.f:.C"r t.'XJ)dtmcnwt Ulfcctlort Wilh ud.. -born~ Ith~ \~urml11)'
•.\mt•rk.m S.odfty of ~·l iewhtulog}. PtlftUito/o~- 16. 22;..2s:t.

82 nJ:-- H ,, ,.1mxso1.L.o.1-1g.A,1.lribcll.Ehdi.dui:at?luhplsi111,r..,11J1r... 10~< \'a>, M.P ,,, ;999. l:.pidcm1olo~·.:11d molcciulJr chdmrtt•ri.....wlon of
'.'\..ll. ~ H.Ot.T t,G. tc.dSj &•rgt1•:•, Mt,vmnl ofS)'$ttmarft Bticr,..--rfnfOR)•, \'ol. I. Eltrllcl111J µhngrxwoJ)hlln in ,~lallon to emerging ohrllcht.t<. PhD thcsl>.
Bal Um 010: Willi~rn< & I rm.In,. Unh·ebH\' 01 EdinburJ.th.
 ~<er-known rlckett,fa~ infecting liwsmrk 549

IO,l \'((;SJ:;. f ,P.. U \ IS. ).f.L i, FbH. n .. 1.9~9. Comp:nau,-~ \"t"Cll>f c:umpi"tC!:'l('t.~ I11,• t>Jt~b. of utiA>omc fc\·cr on some h.i.ncuon§.
111 \\'(UJ)f'HI\\ l 1. ;,_. 19,8;
o! J.)(•nnt1t,1 nror l'l'ltit1hilis nnd t1.CH.lt.Ssrl1p11larki ,\t'ar-i. /:tl)(f/tfni• lot rh~ of pntymorphononl!(ICM cell" uf ,,hili:p. Jm,m.a.l oj Comr,nrnrli"f'
31t,n1 o/ hum.,n ~,.muJoe~~" ehrllchlo<h. Jo,mwl of \Jl\li.:ol P,ultol~·- ~• • ,81-485
Fotomolol{l: 36. 182-f85_
B:!: woL01:11mn. ,_, 1987. r:>,·pu·~iun of l\'mpho~ic, resp.orb~ tQ rmtu~~n,
10-:; W.uJJ:fl, "ll. tt)OJlut. I c • S~'OOOkA.~ D.R., l'lf.i. fnVl">iigauon .. mm tbc In "'hcup tr,h,•ttl'd \,;1h th:5..,homr f('1,:~r. Journ11I ,,,rn,m,,arat11>t
epu!cmloluj<;• of bonno f'<'ll'th•af rcwr in Kt·ni·• and the pot.,111ial of Parlw/qg;, 97 637 6.:3.
TIOl'tlbl< ulJIJ ntit1."'>.t'). \o\..'CWI),,. ] 1t1µlc.'C1l.iltll!lcJI /(,:rtllh "'"' ,,,,.,.tu( tr,m, o. us w1.-,1 DElll\\n.1,.,-,c1•n r:-.R.• 1•181 hnrmmoto¢cal srud1cs
193-190
on 1ftk.. honll" f<.'"'-'' in :ii.h1."'1p. Jor,111111 ofC.Ott,fHfmlll1.• Pml:(Jlo~ ,
106 \\',\tlcr:R. u.tr~ & uu~Ji..ut. I.$,, 199...,, lirm.irgcncc o( th~ 1:htltchto~ ,L"- 92. i57-1Gi'.
hum~n health prohlcm-.. Emerging /11fe,:1,011s !Jiffllll"t i 11·2!1
au r>f.KI\\Tt ,. • ,con. <~.l\ .. 1982. St;igcs or dc,-elnJ>mcru of C.).rwar~
\\'(>1
10; w, rw,. w , .• 196.,. lnfonlli1y In the nun ~«oct~tccl •.,11h tlck-bmnc ''""' phD$fXJ'l0f)hilli, ~ c.tusat1\.(' 3-ff;t"nl of tkk:·b(une feyer, Joumnl of
in!\."Ction n,,,
\!ctrrltm,,1R,~rd, 7G, 1131-J 136. Coml)lJrart,-r Pml1ol1>~·. 92. .;ss...r,.a.
LOS WJ'J!,.kUJt(.. W.G... ()ORS<>~. ,1.L-. !-\\1\JF.L J.L S)A.-.CII C, .., •• \(\I.L\\'tA, LP
," \\'Ot ~E. c.n.. J96"'
&\C..\. o .. :-.t\."'-DltLo. L.. s:~·co1nc~·.-. f.r... wnsc... t
a
u.s ,,rnl)F11n,1T.r_, (corr. G R ~93-Tlt.k~bomc {pit,IUW re,•c:. 111. ,,.t'llnr •
lll" n L" ru~-r1c ,t. l"d5 • Ru·J..-..,rulal a.ml ('Jdam;ltllhl dh·M:!tJ "I
Ph11ogono,I< dJ, or,lt) of th• /Uck;/t1,$/0e.Joum11l •J &ut<•r.oloic,·. 171_
1Jom<'$ttr A1Jf,r.nh. O.\ford· P(•tg.1mm1 Pre<,;, p. 427.
-1?02-1206.
uU vu. x.1,, 1.u.,.~<,. ,.,..... ,tttrhtur~J,\\·•• Y.HA'-(';, Y.J... ., \\ ,n, n, n.u :?-Ont.
109 \\ttsOS, J.C . f()(.(;11. "'-, CAR..\JJCtL\llL ,1...,\., 1961. Tlcfi. ·bflrnt.• fc\1.•t ;~ ;,1
Phylo~ent::t1c re!.1.1101~h1p~ of .1t1oplllsmn mar,:lm,!t•.i.t~d •f.J;,1£(/:UJ
a~e of abo~iOll one.I stlll-blnhs In t•nle. nw \ ,:,ri11my R,,·ord. ;5,
1081-IQll.l
11111~·>' to other ~~ctiiK dultrrmln~d h~· Cirnl!l i.tmi.no•acfd 1J.equ~ncc..\~
lnttmwtlo,"'I /uurnul ,,f~.\.,lt111wtie'1111d li.t,'()/U,io11tlf)• Mit.'l"Ubw/ote,. S1
110 ,vuwl:'.H1wa. >"... 1983, nck·homc fC\er. A N\'fm, . t i:11:n,u,,:i Rr.~...-'flrd, 1143--11-lb
Organisnll'on, 6. 1\13-14~.
 42
Chlamydiosis*

A A ANDERSEN

lntroduction birds is no1 dealt with in this chap1er; several reviews on the
avian disease and its diagnosis in birds nnd domestic fowl
Chlamydiosls in livestock is caused by a number of specie~ are available.1 3· 1. 108
or strains of bacteria in the familyChJamydiaceae. TI1is fam- Chlamydiosis was first recognized in 1879 as a disease
ily has recently been dlvided into 1wo genera. Chlamydia that could spread from parrots 10 humans. when a S\\iSS
and ChiC1m_1•dophila, and ninespecies. 29 In 1his chapter, the physician observed respiratory illness in a household with a
words chlamyctiosis and chlamydia(e) will be used to refer to sick parrot. Avian chlamydiosis gained world prominence
diseases and strains from both genera. except when a refer- during a pandemic in !929 10 1930 1hat involved over 12
ence to a specific species is nec.cs.~ary 10 a,vofd confusion. coumrles in Europe and Nor1h 1\merica. The outbreak led to
The disease c_blamydiosis is characterized b) the devel- strict regulations on the imponatlon of parrots from South
opment of a ,·ariety of dinical syndromes depending on the America. In 1930. Levemhal. A.C. Coles and Lillie indepen-
species and strain of Chlamytlir, or Chlamyr/ophi/a, the ilost demly observed for 1he first time very small basophilic
species. and the affected organ sys1em.A \\ide range ofhos1s bodies In !he tissues of infected blrd~ and humans. and
may be affected by chlamyctial diseases. including horses. snggesced char they were the causati\'e agem (cited in
co\,·s. pigs. sheep and goats. Chlam}'dial straim are fairly Pienaar er al. 66) . In that year, Bedson and Blalld (cited in
hos1 specific, each primarily infectL'lg a given species or iype Meyer55) established the aetiological relationship bem·een
of animal and causing specific clinical disease:;. These the basophilic bodies and the disease. It soon became clear
diseases ma~· affect the gastroimesrinal, respiratory. repro- that chlamydinl infections were 1101 limited to psinacine
ducrive and nervous systems, and joints and eyes. Disease birds, bOl that they were \\idespread in almost all avian spe-
severity can vary from life-threatening to asymptomatic, but cies, and that chlamydiae from other birds were uansmis-
the usual syndrome is a modarate to mild clinical disease sible tO humans. Chlamydia! infection in pigeons was first
leading to persistent infection. Recurrence of the disease reporred in Soui.h Africa in 1940 by J.D.W. Coles after a
and/or shedding of the organism can occur followingsrress pigeon fancier who bad beon experiencing monallties in
or physiologic.al changes such as oes1n1s. What is currently his nock sen1 two affec1ed birds to 1he Onderstepoon Vet-
tmkno\\~l is the exren1 of synergistic effects these persis1ent erinary lmaitute. 13
chlamydia! infections may have with concurrent infections Enzootic abonion in ewes was first shown to be caused
\\ith other pathoge11s. by a chlamydia! infection in 1950 by Stamp and h.is col-
A\'ian chlamydiosis is the disease caused b) Chlnmy- leagues.95 Since then chlamydia! aborci.on has been recog-
dophila psirraci in birds.'1 It has been called pslnacosis and nized worldwide. It was firs1 seen in South Africa, in 1972,
omithosis depending on the type of bird ii \\'m' infecting or when a ~eYere epidemic of abortions :.wept through lhe
the type of bird from which It was contracted. Psinacosis sheep-raising areas wi1hin a single lambing season, result-
a11d omithosis aJ'e now considered to be the same disease. mg in up to 60 per cent aborlions in some flocks.88 In can!e,
.the rerm a\'ian chl:imydiosis being preferred. However, the Schoop and Kauker6f> furnished serological and cytological
avian st-rains have been kno,vn ro iJ1fecr hwnans and other proof that incriminated chlamydia as 1he cause of abortions
mammals \,ith severe consequences. Transmission of lhese in a dairy herd in I 956. In California, Kennedy and co-work-
strains from animal to animal occurs readily. The disease in ers in 1960'5 described the pathological lesions of aboned

• For recem chang~ In the noml!n1:l~m1re ortht.":«' or~1,nJ,m.,.. rofor 10 rhe ln1roducrlon in ::W:cli()n 3 Rick~-lt,t;i:'11 ond chlttn.iydi~1 cli.<o:tSc,

550
 Ch!ainrdrosls 551

calves and, from some of them. Stor£ er al. in the ,arne been filled wi1h controversy.';u bl much of which subsided
year 100 isolated the: chlnmydla! agent. Because of the when Page proposed that all chlamydiae be grouped Imo
sporadic occurrence of outbreaks, the disca~e was named one genm (Cl1lamydia)611 coniaining two species re tra-
epizootic bo,ine abonion. The agent i~ similar co. or the cho111m/s and C. psitttlci J.»1 111e di\ision was based on su~-
same as. that causing abortion In sheep. in which rhe cept ibiliry 10 sulphadiazine and accumulation of glycogen
disca~c i,; called cpizootic obortion of ewes. in 1hc ,•acuolcs when gro\\11 in 1i~sues of vertebrate tissues.
Thereafter. chlamydiae in ruminant li\'est0ck were also It also effectlvel} sep:irated the known human strains from
fow1d to be in\'olved in enteritis, pneumonia. anhritis, con· the animal strains. with onl}·a few exceptions. At the time it
junctivicis, and cncephalomyelitis. These diseases are usually was recognized that there was great diversity in Cltlnmydill
caused by the Chlamydophi/r1 µecorum strains. which are dif- µsiuaci. This classification remained unchanged until 1989
ferent from the abortion strain. and outbreaks are usually spo- when Chlmnydin µnewnonlae was proposed as a rhird spe·
radic. primarily affecting young animals. :-torbidity may be cies.33 At that time it \\'as thought to be solely a human res-
high. but mortality is usually low. pirutory pathogen. However. isolates have since been made
from koalas C/>hascolarcws ci11ereus1:t:i 11 '1 and a horse.!16 In
1992. ChlamJ-rlia pecorum ,viis proposed Sl> a fourth spe-
Aetiology and life cyde
cies>11 ft was originally considered to inducfe isola;es from
Aniclesm the historical literature 011 chlamydta and its ra.x- caule and $heep that camed pneumonia. polyanhrlcls.
onomy contain many later-abandoned terms which are. encephalomyelitis. and enteritis. Since then, C. pecorwn
however, still imponam in order 10 unckrstand the li tera- strain~ have been i~olated or idemified from other rumi-
ture. Chlamydiae v.-ere once classified as ,·iruses and nants and from pigs8;and koalas.~· 113
grouped ,,ich other filterable organ isms that required The wcently proposed reclassification of th~ family
liVing cells In which to multiply. Some of the names in che Chlamydiaceae in 1999 is based on sequence analysis of its
liter.ature include the psinacosis-trmphogranuloma- 16$ and 23S rR:\A genes. 2q Analyses of orher genes yield
trachoma 1Pl.TJ group. miyagmvanella. bedsonia. prow- the same results. rhe re~ulting division shows a high de-
nzekia. rakeia. and trachoma inclusion conjuncthitls.~· 6(; gree of corn.Jacion with host range and disease syndrome.
However. the use of a chlamydia-like name dates back to and thus help, in 1he understanding of rhe epidemiolo~·of
190i. when Halbcrsteadwr and Von Prawazek (cited in the ,~.irious strain~ a[fec1ing livei;rock. The major change in
Storz98 ) described characteristic intrac)'toplasmic micro- the taxonomy l~ the formation of two genera which corre-
organisms ,,ithin vacuoles in conjunctiva) scrap-ings from la te with the iormer C. 1mc:homatis and C. psillaci. These
a human patient with trachoma. They thought that the or- genera, Clllamydla and Chlamyclophila, comain three and
ganisms were protoioa and used the family name Chlamy- six species respectively. The species C. pecorum and C.
doz.oaceac after the Grl'ek word 'chlamys', \\'hich means pne11111011ifle are included in the genus Chlnmydop!iila.
mant.le. The term Clltamrtlia was proposed in 1945 and This new classification, together with the currently known
lacer reaffirmed by Page. 00• r.i primary ho,1 ranges and di!ieasc syndromes. ls gh·en in
The early classification of the family Chlamydiaceae has Table 42.1 .

Table 42.1 R!vised class1f.::at1on of

3ENUS -'<ND SPi:CIES PRlll,IAAY HOST!S f'RIMARY OlSEASBSi
rne family et,.amvd1acew?
Hurrans Trachoma lvmp~ogra1u o;.,a
vener,um. conjunwvi~s
~eooatal pneurr.il~ia
Chlamydia muriaawm Mice, ham;iers Pneumonia. lier, s
Chl?.m)•dia SUIS Pigs Pneumonia. ain1uncti\ ,:,s.
eotent,s
Chlam\'tf-Of)hi!a pneumonise Humans Pneumonia, syste~ic disease
Ch/emydcphile pecorum Sheep.car-fe,sWtne! Wide range of d1sea~s3
C11l;;m1':Jophila caviae Guinea Digs Conjunctwius. gen«al rret:l
mfe<:tion
Chlom~ophi!e fel1s Cats Rhinitis. CQnJurn:tlv,:;1,
 .,._ rn , rttnH llicl<ensi:11 and chlamydia! dl$eaws

Following precedent. 1here is some co111mvNsy abom The devrtopmL•mal cycle 01 chlamydia co:1S1s1, 01 fini major
1he ne,, cla,~itkation. primarily co:1cerning 1he divbion of ph8~L"<:
the family Chlamydiaceae i1110 rwo genera and the use or the • 1he at:achmcm 10 ,ind pc1wmuion Into 1he ho~t
term Chltin1ydophi/(I for the second genus. It is feared 1ha1 cell by the l.1:1:
1he new tl'rm could create confusion. The two genem and • th" trunsi1iu11 vf the nwwbulicallr inen EB mto
tlw nine ~pueles. however, have merit not only In molecular the metabolically ac1tve RB:
,erms. but also in classifying host ranges and clinical dis- • muhiplica11011 or the· RB b, bina rY fission using
ea~e,. The terrm chlamydioNb and chlamydia(e) Mt> u:,ed cnerID from the ho~, cdl:
hl're .IS generic terms 10 rt•fcr to members of both and either • rnumration ol 1hr mel:ibolicalll aclive RB into
•
genera. Howe\'er, the new scientific names are used when 1lw srabl<" lnfoccious EB: and
referring to a specific chlamydJal ~pecie.s. • lhl' rclta~e of 1he l'B from che ho~t cell.;·· rn,
Chlamydiae arc obliga!C intraclllhJar bacteria that multiply
in cytoplasmic inclusions in the cyrn~lasm of euknryotic cells. A cri1ical s1ag<" in 1he chlamvdlal life cycle is che pha~c fol·
They ha,·e a unique growth cycle con:.iSting of two devdop- lo,,;ng cat!')' imo the ho,.1 cc,11. The EB mu~t tran,-fonn into
mental iorri1s. The elementary body 1EB! 11; a condensed l'onn the mcrnbolically active RR and e<.cape from t:lw host cell
of200 to 300 nm in diameter thac is suned for suni\'al omsidc defence, while cran~forming. The normal cell proces,el> en-
the hosL The reticulate body !RBI is 1he replicating form, rang· dosomc~ ~omaining foreign material b) ac!dilica1ion of the
ing In size from 500 to I 000 nm. and found in a (:10plasmic cndo)ome .mtJ fu~io11 nf the cndosome ,,·Ith the ly:so<omn.
inclusion \\ithin the host cell (Figures 42.1 and 42.2). Replica- '111e digestion and proce~~ing of foreign amigen;; by 1he !y·
tion is by bina11 fis:.ion iypical of other bacteria. with the ex- ,o•,omt> i, critical for the proci>.s!;ing of antigenr. durini the
ception that chlamydiae rely on the host ce.11 for onergy. irmnune r<·~p(,nsr. II ha~ twen sh011 n that lh·e chlamydiae
T111em1ediate fom1s between 1he RB and the EB are usually prl.'\'ent the acidilica1io11 ol the endo~omcs and the subse-
seen in the indu~ion. t ti quem tu,ion wi1h ly:.osomcs.";· 10•

Figure 42.1 lr,ra:ytoplasmlc vacuole co1nani~.;; cnla'l'ytf:as in an enterocvte of :.rte 1le1.-m o· 3 ~atura I~ mtecred p,g Tt•e vacuole co•:a ;i5 e,e~~ar,
>od•es. •e:rcufa:e bodies. and a small a:n01Jrt of gl•,coser. (Cou'11!$'/0f Douglas Rogers. L,r.coln. Nebras,.a USA)
 Chl~mydio,ls 553

-.

',,; .

l.
•
't-'
'
,
'... •
'
. .. • •
,

Figure 42.2 1",rae1,1co as:n c vacuole tonta,ning chla:nydia~ "a con1unttival epithelial tell cf 2 r.aturarly ,r.ietted p;g The var.vole tontair,s elementar.
and ,e· c 'ate .:1 ~s. ar,d a fo•se: amount of glyto;eo 1Ccv:e-,. ,, D!iuglas Roge,s, Lincoln, i\et7!S<a. us:.

Epidemiology koalas ,Plwsro/<1rt·ru, dnereus). springbok :\11tidorcas

1twrmpia//s), ibe)> Capm pyre11aka), and bighorn !.beep
Chlamydiae art? among the nrnre ,ul'cessfui pathCJg(lns C011is ca1wd,:,,:$is)."" :Vlolecular and serological typing now
known. le was once thought 1ha1 chlamydinl 1sol01es :ire all mnkeit possible 10 cll'termine whether lhe.,e and 01h.crwild
sirnilar ,md rhat each infetterJ a ,\ide ho,t range. :'\ew mo- animab are rc~ervoir, of chlamyd iae for dom~tic animals.
lecular and ,crological 1echn1qucs now ,how that thcrt' arc c,r whether these ~,wcies han' their ()Wll host-strain rela-
numerous identifiable chlam)'dial srrains and chm rhe iso- liori£hips. :vlolecular ,rndies of isolates from koalas ha\·e
late, from a panict1lar avian or mammalian ,specie~ 1<:nd 10 shown 1hese animal$ 10 harbour both C:h/11111_1"<icplliln
be of the same strain. This palll.!nl indicates a triangular re- 1mf'1111w11itrt: and c:11/m11yrtophilt1 /NCMt1111 s1rain,. 39 · ·,
lationship among the chlamydia! strain, Lhe hos1 and rhe These strains howc1·er. are d istinct from rhose found in
disca~c syndrome. Research i~ increasingly ~hawing that othCir animal, and hum:ins.J9 Thl' snow,hae hnre and
whtn a ,train infects ils natural host persistent infoctions muskrat isolate~ :ir,• identical and similar lo avian
are common .. : These persis1e111 i11foc1ions clearly piav a Slrains.~··· ,:i Conversely. this ;\156 strain found in snowshoe
roll: 1n the maintenance of the chlamydia! organism in na- hares and mu,krar, hn, ne\'cr bE'en isolated from birds. On
ture. It ,hould be remembered that a 11umber oi the a\fan the other hand. a C JNc:,,rr1111 lsolare from bighorn sheep
and mammalian ~,rains can infect mher hosrs. including app(•ars the same a~ rhe ,Lrain causing polyanhritis and
human,;, \\ith severe c1mscqu<•11i:-c$, bur s('condar) $pre.id conjuncthiris in dorn<istic sheep?
lrom the~e infeC"tions appears to be limited. Trnnsmi<,;ion of chlamydiae pmb,1bl) occurs \1a
The role played by wild animals in the transmission of several routes. ;1s chlumydiae have been found in all body
chlamydiae need, to be a~ll<:sst•d. :--atural infections have ~ecretion$. including raec-es. The fa11cal-orol route may be
been reported in oposs11ms (Di<ll.'lf'his p(lmq11<1J,Wl.tis). the mo~t common !or mo,t ~trains. 2"· , ; ,-\erosol and con-
nonhem fur seals (Collorhim,s 11rsin11s1, muskrnts 1 011- 1ac1 1ransmiss1on may be more <:ommon in respiratory
rlmra :::iberllica), snow~hoe har<t~ U.~pus amerirm111s). tract and eye infec1io111:,. respecrively. ln,cC't tran~mission.
554 Sfr"T1ns '11RU: Hickeusial and rhlamydial diseases

e,peciall) by flies. must be considered. The importance of teau shedding of \iable organbms have been identified
vertical transmission ls unkno\\11. bu1 generall) has noc follo\\ing chlamydiosis in humans. animals and birds.
been thought to be a major factor. 7 There are. however. lmmunosupprc<sion with corticosteroids frequently
increasing reports of persistent Infections or 1he male reac1il'a1es ,heoding in these ca,es.
acces.~ory sex glands and of 1he presence of chlamydiae in Chlamydiae are capable of infecting and multiplying
scmen. 99 In ewes that have aborted. chlamydiae have been within macrophaics. 1ltis ability has attracted aLtemion
isolated from vaginal swabs at 1he rime of oestrus for over because the chlamydia! inclusion is capable of m aiding
two years;'"· t..3 lymsomal fusion.~: 1115 which is necessary in the normal
pathways of processing an antigen in order to evoke an im-
mune wpo11.~e. It is apparent that live ch:amydiae are
Pathogenesis
needed 10 initiate tlle avoidance mechanism. as anacti\'ated
Chlamydia! infections are responsible for a wide variety of chlamydJae an, processed by macrophage~.
syndromes in domestic animals, including enteritis. pneu-
monia. polysero,itis, lymphadenopathy. kermoconjuncrivi-
Clinical signs and pathology
tis. masnus. polyanhritis. meningoencephalomyelitis.
epididymirls. <,rchitis, seminal vesiculith, and placenritis. Clinical signs and related pathology in affected animals ''81)'
I he strains of chlamydiae each have a prcdllccuon for differ- according to the stram of chlarn:,,dia and the si11dromes
ent tissues and therefore different disease si;11dromes. TI1e from which the animal.s are suffering. Information on the
rouce of infection may play a role in the cype of disease syn- prevalence, pathogenesis and pathology for man} or the dis·
drome; nevenheless, oral or parenteral inoculation of a given ease syndromes is limited. This discussion focuses primarily
strain usually results in the development of che ~me disease on those diseases considered 10 be of major imponance.
syndrome from which the strain wru originally Isolated.
The oral route of exposure Is considered the primary Sheep, goats and cattle
mean~ of transmission for most st.rains.?S. 2 ' It has been Chlamydiae in ruminams have been primaril~ associated
shown that most strains readily infccc the eplthelial cells of with abortions. polyarthdtis. conjunctivitis, encephalomy-
the ga,,troimcstlnal mucosa. The organisms locali7.c in the elitis, t!lllerilis, pneumonia, mas1iris. and genital infections
epithelial cells of the mucosa in the jejunum and ileum. and such as cpldidymitis and orchi.1is. Abortions are probably
then ~pruad via the blood and lympli vessels of the lamina the most dramatic clinical outcome. having bo1h economic
propria co the mesemeric lymph nodes and the liver. ·n,ere- and public health significance. They are usual.ly caused b)'
after. infection may be established in other organs or tissues. Chlnmydophila a/iortus, but an abonion in a cow caused by
\\1th abortion strains, e\~dence indicates that infection infection \\ith Chlamydopl1ila psittaci sero\'ar B has been
may be initiated either by the systemic route or b~ venereal documented. 1 ' This occurred in a dairy herd where pigeons
transmission. Systemicinfcction causes a chlarnydaemia that were common in the barnyard. In addition. a bo\ine abortion
results in spread to the placenta. C!lusing e.,1ensive caused by a chlamydia-like organism WSU86· 1044 has been
placental damage. fhe extent of place111al damage may deter- reported.22 111• ijO The C. abomlS strain may also oe involved in
mine whether the lamb is born apparently healthy or weak, or mastitl, 12 and other fom1s or genital infcction.102 IU'J. 116
i:; s1illborn or aborted. 11• n Placental infection can impair ma- Chlamytlophiln nborw:~ strains arc serologically the same
ternal-foetal transfer of muricms, resulting in impaired worldwide. with the exception of one isolarc from Euroµ,:, that
growth or death of the foetus. Placental Infection may also appean; to be serologically distlnct. 110 The polyarthritis. con·
alter placental steroid and prostaglanclin relea:,e. resulting in Juncti,itis. encephaloniyelitis. enteritis and pneumonia
premature labour.48 In some cases. a ewe bearing twins may disease s~i,dromes ate considered to be caused primarily b~
deliver both a weak and a healthy lamb, as premature labour ,11rious strains 01 Chlnmydophila pe<:orwn. There is some
may be prevemed If the healthy placenta produce~ suHkient indication that there are strain dift'erenc~ ,,ithin the
proges1erone to ma lntain pregnancy. C peror11111 group. and thnc each srrain tends 10 produce a
.·\bundant evidence existi. that most chlamydia! species specific 1ype of disease or disease syndrome.
are capable of producing persis1e111 infection that can en- Chlam>·diai abortions in sheep and goats, also known as
dure for months or years in the absence of treatment. 112 The epi.zootic abortion, can occur as an acute outbreak in flocks
mechanism of persistence is not known. For years it was or herds. or as an endemic infection. In endemically in-
\)'\ought that persi~tence was associated with lncomplece £e<:ted flocks and herds, abortions occur year afcer }'ear at
chlamydia! de\'elopment and only sporadic production of rates of up 10 5 10 10 per cent per annum. lt is not unusual
EBs. Howe,·er. the fact that EBs can frequently be recovered after three to live year» to have an explosi\-e ombreak of
from asymptomatic: animals using cell culture techniques, abortion in yearling~. When C. aborru$ i$ newly introduced.
and that EBs and chlamydia! inclusions are commonly abortion rates ofup to 30 per cent are common in sheep and
detected in clinical specimens. indk'ate that productive sometimes oi more than 60 per cent il1 gc>ats.7-
infecrions are the norm. Persistent infection~ wi1h lmennit- A number of humru1 abortions followed by se\'ere
 Chlamy,Uo,h 555

complicalions have been associaled with exposure 10 In- abortion. If rhe disease proct'ss is not extensive it may lead
fected placemas ofewes and does. 10• 1~- 11 The prevalence of 10 the premature birth of a wea k lamb lha1dies sh only after
such abonions is not known, as fnrectious causes a re often birth or to one chat may lil·e but remains umhrifty. Ewes are
not looked for in humans. Chlamydia! infections must, how- rarely affected clinically and remain fertile. Thi~ is in con-
ever. be considered an occupalional hazard for pregnant trast 10 the disease in goats and cows in which therl.' is ten -
women who come imo comact with domestic ruminants. dency for placenws to be retained .ind for 1he developme111
En:n ewes exposed to chlamydiae before they have be- of vaginitis and persistent endomctritls. I loving once
come pregnant may abort.r.i. 115 In one study. seven ewes .iborced. sheep are solidly immune and wtll noc abon again.
were exposed by rubbing Infected placemaJ material on even though they may remain persistently lnfected.63 · M ; 2
Lheir noses rwo days al'ler they had larnbed. 11 5 During the In c.11tle. chlamydiosls causes abortlons. stillbinhs, or
following year two or them aborted as a resulL of chlamydia! the birth of weak calves. 111 i o, 24 • 34 • ~;. 98 • 100 Abonions are
infection. It has bet!n suggeined thaL in ~uch coses infection u5uall~ sporadic. but ma~ reach 20 per cent in a herd and
is firsL l'Stablisbl.?d in the tonsils. from where it is dissemi- usually occur during the last crimester, affected cows show-
nated haematogenously to 01hcr organs.\.\~1etber the infec- ing little e,~dencc of disease. The abortion i~ caused by pla-
tion persists in the tonsils or in other organs ts not knowtl, centiti~. but the ;iboned foetus may manifest ascitcs,
but it is thought that a state of persistent infoction \\ith in- ~erolibrinous pleuritls aod peritonitis. an t>nlarged reddish-
termiuent low-grade chlamydae,ma evennrntly r!•sults in in- yellow ll\'er, and hyperplastlc lymph nodes. Call'es born
fection of the placenta. Research has sho1,,1 that placental weak but chat suc-.i\'!! usually remain unthrifty.
infection.~ usual!}' becc,mc established bcnvecn the 60111 and In the UK, a syndrome in cows characterized bya drop in
90th day of gei;ration. and that pathological changes do not milk production. mucoidi mucopurulcnt oculonasal dis-
de\'elop uncil the 90th day of gestation. 11• ;z charges. depression and .inorexla over an eight-week period
The me1hod by which chlamydiae cross the placenta is h,L~ been reported. Hcco,·ery is slow but appears 10 be aided
uncenain. A.fter about the 60th day of gestation there is a by administration of rntracyclines. Abortion is common
physiological im·asion of carunculae stoma by chorionic weeks to month$ after the event. The only consistem finding
\'illi. This roincides with haemorrhage from maternal \'CS· is tha1 chlamydia! anrigen and /or rise in antibody titre to
seh. resulting in I.he formation of haemacomas that have chlamydiae have been demonstrated in affected herds. 19• 20
been suggested as a tnt>ans by which chlamydiae in the ma- ,\ seminal vesiculitis S)11dromc (SVS) ha, been reported
ternal circulation make direct contacL with till' chorionic i11 bulls.ij~ II Is characteri7.ed as a chronic inflammation of
epithelium. ll the accessory sex glands. epididymes. and testicles. lnfected
fhe 30-day period bctw~n the fonnation of the hae- b111ls have inferior semen quality and may develop testicular
rnacomas and the first observation of pachological changes atrophy. The syndrome is thought to be widespread in the
in Lhe placenta raises the question as to whether endocrino- world. An isolace from a bull suffering from the syndrome
logical and/or immtmc-related changes play a roJe_ll was found to be indistinguishable from the EBA59-795 lso-
Chlamydia! inclusions and lesions can first be detected In lotc: of chlamydia. indicating 1ha1 the strain Is the same as
theplocencaon about the 90th dayofgestaLion. The lesions that causing abortion. The S\"S can possibly be regarded as
first in\'olve the limbus of the placentomas in the hilar re- the counterpart of the abortion syndrome in cows. Similar
gion. At this time local necrosis occurs and the infection disl'ases or S}11dromes ha\•e been reported in sheep.50 '"'
then spreads 10 Involve the coryledonary and intcrcoryle- in open cows. cervkitis and endomccmis with a non-
donary placenta and opposing endometrlum. A mixed-cell putrid discharge hove been associated with chlamydia!
Inflammatory Infiltrate Is present. U ndcrlying the lesions. infections. 1119• 1 1"
fibrin deposits and a nonpurulent aneritis are present in the Polyanhriti.s in sheep. goats and cat de may be caused uy
thickened placental me~enchyme. Chlamydia! inclusions strains of C pl.'corum.l 1 fhe disease can be readiir repro-
are pre.em in the epithelial cells of the endometrium "'here duced experimentally by inoculating lambs or calves by om!.
the affected chorion is in opposition to 1he maternal tissues. intramuseular. ~ubcutaneous. intravenous or intra-articular
Severe necrosis and :.loughing of rhe cndomeuial epithe- rouies. 1~- i; In the natural disease, the route of infection is
liurn occurs during later stages of pregnan~. thought to be oral. the organi$m mull iplylng in the mucosa
In sheep, infection or the foems is secondary 10 the of the small intesLine. Diarrhoea may be present during chis
placenti1is and is not a major factor in the disease. Focal phase. Follo1,ing multiplication nt the primary site. the or-
disseminated necro,is ma)' be evident in the liver and ganism spread~ h.1enm1ogeno11sly to 01her organ• :md ti,-
other organs of the foe1us and mar be embolic in origin. 11 sues of which the diarthrodJal joims arc usually tbe most
Chlamydia! antigen. if present in the foci of necrosis, severe!) affected (Figure 42.3). Lesions occur in boch the
usually occurs only ln small amounts. The rnesemcric and pcri-anicular and articular tissues of 1he j()ints:6 · 17 '10, n;
popliteal lymph nodes ma}' be enlarged. having a demar• which usually are gross!)' enlarged because of a localized sub·
cated cortex with several follicles and gem1inal centres. cutaneous oedema and an increase in the amount of fluid in
The placental lesions may lead 10 death of the foecus and the S}'!lo,ial sacs. The fluid is greenish-yellow and rurbid. In
 ren tly with central nervous system (C:S:$) uwolvement. I he
principal clinical signs include moderate fever, ~tiffnes:.. ln·
coordination and depression. Gross lesions areindicatJveof
a sysLemic infection and include seroflbrinous peritonitis.
pleurit!s, and sometime,; pericarditis. similar to those ~een
in other C. p,:,com/11 Infections. Gross lesions in the c:--, are
not as e~ident and ore often limited 10 meningeal conges-
tion and oed~11111. Histop,11hologically, the c:-.s lesion is a
diffuse nonpurulent meningoencephalomycliris whtch ls
especially marked in the cerc,bellum. The lesions are charac-
terized by peri\'ascular and focal infiltrations or mono-
nuclear cells. focal areas of encephalomalacin and neuronal
degeneratlon.3 ; In sheep. 1he clinical signs and lesions are
~imilar to, but generally milder than those in cattle.
Enteritis and imerstirial pneumonia caused by chlamydia!
infections have been reported in s heep and carrJe.2a n &J. 98
rile isolates that have been characterized 10 date from such
cast's ha\'e been identified as C. pi!comm. There appears to be
overlaps in the clinical signs and lesions with those occurring
i1\ Lhe palyarthrith,, conjunctivitis and SBE S}1ldromes, indi-
cating different tissue preclilt.'Ctions for tlie various strains of
C. 1wcomm. Both s~11dromcs primarily affect young animals.
·n,e emeritis ~yndrome is rhai·acterized by a mucoid 10
,,'(ltery or bloody diarrhoea in young calves and lambs. Thi'
onset or the disease is rapid and the severity depends on the
virulence of tJ1e strain and the degree of protcetion afforded
by colostral antibodies.25· • "''· 98 On necrop~)'. a focal to gen·
26

eralized peritonitis may be seen. Petechial haemorrhages in

Figure 42.3 Serol1brinous synavitis typical cf Chii,mvdo;ihila pectHum
the abomasum and intestine occur in infections by highly
µolyartrri,,s niecuon
,~nilem chlan1ydial srrains. I I )'pcraemia and congestion of
the mucosa are common, and enlargement of the l}111ph
ad,·anced ca,.es. fibrin plaques ma}' be pre$ent in the Joints.. node:. may be seen. The ileum is usually the most severely in -
In severe cases excess fluid of a similar namrc may be fected; lesion$, however. may occur throughout the intestinal
present in 1he tendon sheaths. Muscle in\'Oh'ement fs pri- tract.is 21,. 9ff
marily limited to mononuclear cell~ in pem:ascular areas l lis1ologically. chlamydia! inclusions are primaril} seen
near the point of tendinous auachmem. His1opa1hological in the \'illus emerocytes and occasionally in the cryptal en-
changes in th<- synovium. tendon sheaths. 11nd sub&y11ovial terocytcs. Villus atrophy and m ultifocal necrosis of the \illi
tissues include inlihration of neu1roph1i,;. macrophages and :ire ofrPn P>:rPnsl\'P PPrrr·~ pnrches may lose ,heir follitular
lymphoid cells. Changes in cells in the S}"TIOVial lining can architecture and contain necro1ic cencres.25 · 26 o;,; 93
,ary from necrosis to proliferaiion. 16· ,- lij Cal\'es and lambs affected with chlamydia! pneumonia
Conj11nc1lvitis frequent!)' accompanies polyarthritis i1\ have fe\'er and a serous to mucopurulent nasal discharge.
lambs and may be a prodromal sign of a developing ou1- The) are depressed, cough. and have signs or dyspnoea.
bre<1k of polyarthtitiS. Chlamydia! conjunctivitis is usually Older animals ,dll ha,·e similar clinical signs. Conjuncti~itis
bilateral, and clinical signs var} from a profuse watery 10 may be seen in acute cases. In severe cases, young animals
mucopurulem discharge whlch. in some. seals the eyelids become prostrate, have laboured respiration, and may die.
together. and hyperaemia of the conjuncti\'a.-'8 In severe Most cases arc of moderate severity. with reco,·er}· taking
cases. photo phobia and keratit!s occur. Secondarv bacterial one to two months.,;,;· 98
infections of 1he conjunc1.iva arc common. The acute pulmonary lesions are characterized by multi·
• Sporadic bovine encephalomyelitls (SB1:) is a disease focal gre:,ish-purple consolidated areas, u,uolly locared in
that ha< been reported worldvdde?5• 54 · 65· "7 including in the region of the hilus and the ventral portion~ of the cranial
South 1\frica. 107 It prim,trily affects young caJ,·es. but ha~ and middle lobes. (Figure 42.4). Serofibrinous pleuritis may
been 1-.nown to occur in adult cattle. and sheep. Generally. accompany these lesions. In severe cases foci of consolida-
the morbidity in cattle is under 10 10 15 per tem. with mor- tion eXtend throughout the lungs. The microscopical lesions
tality of up to 30 per cent in affected animals.,\ gencrali1.ed in acute lesions are characterized by serofibrinous pneumo-
chlamydia! infeetion usually precedes. or occurs concur- nia and nleuritis. and small haemorrhages are frequently
 Ghlamydiosi~ 55;

Figure42.4
sneeo nte,s:,· a.
pneumc~ a ci ·>i:
1.~mra1o::;n1on o·
;he c:a<"'. a1ar::
m,dct1e loties ~. e•cai
of Chia~ 1"COPl1i i
;::eccrutr ,•'szucn

present in the parenchyma of the lungs. A:. the lesions tomatic. Some ~mdh!, have indicated that C. suis b not a slg-
progress. moderate numbers of mononuclear cells, partic11- 11Ilica111 pathogen. ,b h can be i~otated from nomial and
larlr lymphocrtes. are found in che alveoli and intersmial diarrhue1c pigs at the same rate.s~. toJ On the other hand. C.
tissues. Perivascular and peribronchiolar lymphoid srlis was the onl} potential pathogen isolawd rrom four
hyperplasia, in which scnuered plasma cells. macrophages, 30-day-old pi~lets ~uiferi11g from <liarrhoea 08 In another
and a few nemrophils occur. are also evident.6,; trial, inoculation of gnotobiotic piglet~ \\ith C. s11isresulted
Chronic pulmonary lesions consist of small greyish-pink in diarrhoea and extensive villous a1roph) of che ileum and
areas of pneumonia, depressed slightly below the surface of distal jejunum;; (Figures 42.5 and 42.6) Inoculation of
the pleura. Histopachologically these areas are characcer- 21 ·day-old piglets raised convcntionall)' with die same iso·
ized by atelectasis. infiltration of,·ariablenumbersof mono- !ates produced only moderate villous acroph) in the absence
nuclear cells and neutrophils, and severe pcribronchiolar of clinical diarrhoea.-- The role that coloscral antibodie< 10
ly111phoid hyperplasia.\;6 98 C. si,is mav play in moderating intestinal disease in animals oi
:-1icroscopically. chlamydiae are found in monoC}ies that age is unkno\\11. and h is not clear what S}11ergfa1ic effect
and in the C\10plasm of epithelial cells, maintr of the bron- chlam)•dial inft:'ction mav hal'l' 1,ith other pathogen$. Rect•m
chioli and alveoli. Organism$ are frequently difficult to de· $tudie5 \\ith )rtlmo11elia rn1him11ri11111 and Eimeri<l srabn
lect. partlcularl}' in chronic lesions.r,,i have shown that the di~eases produced by those organisms
were more severe when chlamydia was prescm. ' 7 <,<\
Pigs The &amc C. s11is strain~ ha\'e been isolated from pig, SUJ·
Dlseasc s~·ndromes In pigs that are associated wi1h chlamy- fcrlng from pneumonia and conjunctivitis although ii is
dia! infecrions are those of conjunctivitis. pneumonia. en- difficult to establish their significa11ce as pathogen, be-
teritis. reproducrivc problems, pericarditis, polyanhricis cause they ha1·c al:10 been isolated from clini<:ally nonnal
and enccphalomyelitis. 5• 106 A number of distinct strains pigs. When a C. s11is isolate from nursery pigs 1,~th pneu-
have been identified. including those chat belong to monia \\'8S inoculated experimentally into gnotobiotic
Chlamydia s11is. Chlamyrlophi/a pecorum and Cltlamrdo- piglets, several of them became moribund or se\'erel} dys-
phi/(l.(lbor111s.i9. 13• ' 4 • 114·"5 Of these. C rnis appears 10 be the pnoeic. although the majority exhibitc,d only mild
most common. Cltlamytlin s1tis Strains have been isolated dyspnoea. 7~ The gross lung lesions at Ii to 21 days after
from the conjunctiva and respiratory and inte$tinal tracts of Infection were tj-pical of bronchopneumonio. 103· 1i:
pigs. where they were associated with conjuncti\'itis. p11eu- Similar!>·· C sui., ,trains ha\'e been isolated both from the
monia and enteritis. 59 73 At least 10 distinct strains of C. suis eyes of clinically normal pigs and from those of pigs suffer·
have been identified. ingfrom tonjunni\iti~? 1 16· " H In severe case~. l:,"lllphofol-
~Ian~ C. suis Infections are thought to be mild or as~111p- licular hyperpla~ia in 1he palpcbrnl conjuncti,·a I~ often
558 ,Fa10,. uu,..: Rickc1rsial and chlnmydial diSt>Mes

present (Figure 42.7). \'\'hen an isolate obtained from 1he

eye wa~ inoculated experimentally imo che coniunctiva of
normal pigs. they did n()t develop clinical signs. although
histopathological examination revealed mild conjuncthiris.
The mild conjunctivitis may have been the result or a first
infection, as is the case it\ humans In whom it Is thought that
repeated or chronic Infections are required for severe cases
co de\'elop. The piglets inoculared in the conjuncriva or era·
chea developed diarrhoea with vlllous atrophy similar co
that seen in the pigs that developed diarrhoea after being
inoculated orally.•6- •9
The epidemiology of C wls strains is of imerest bet.iust
they are ubiqui1ous in pig populations. Serological and
isolation studies indicate that most pigs become infected
before six weeks of age, and that virtually all pigs are infected
by one or more strains.s• Infection with one strain rnay not
protect against another. Many infections appear to be per·
sistent. permitting rhe chlamydiae to be passed on from the
sow to her young piglets follO\\ing farrowing. In the USA,
many of the C. suis strains arc resistant 10 sdphonamides
and tetracyclines, making creatrnem or control of che dis-
ease S)1ldromes very difficult.6
Other disease spidromes chat have been associated \\ilh
chlamydiae in pigs include pcricarditis. encephalomr1?litis,
polyarthricis and reproductive problems. These are pmb·
ably caused by C. peccrum strains:12. .;~
Chlnmydop/zlf(I (lbortus, of the same srrain a& that iso·
lated from sheep and cattle, was isolated from a c;ase of
Figure 42.6 Severe vilfous atrophy in the ileum of a gno:ob1011c abortion in a sow.2 and has been detected in retrospective
piglet tnfattEd 10 days previously With Chfa71ydia suis thaematol<.'(lin studies in other cases of abonion.84 · 85
and eosin stain)
 Chlaniydlosi< 559

·~ ,. ..
•
' 'I: •• : .. - ··
--
•
..
- . .,., ' Figure 42.7

·~
• >". .. ----::,...-_~
·.; . .. '
.. .....,i-._
Lvmpnofoll,cJ!a•
hyperp.as:a n .i

-· - . » seci,oo of pa:penral

- ..
•
•
-
; • ccoi~ntiM ·,om a
SO\', infe:::~ \', 1th
Chl11myd1a nis
lhaema1oxv1 n a~n
<

Horses lungs, and lht' central nervous sy~tem. Polyartl:ri1is In

Reports of chlamydiosis in horse., are limited. However. ll(lrses caused by a chlamydia! infection has been di2gnoscJd
there is sufficiem eddence 10 indicate 1ha1 chlamydia In lhe USA and Europo. 19· 51 During tl1e l9i2 10 1973 epi·
should be considered as a cause in the diffcremial diagnosis demic of chlamydia! abortion in sheep in S0u1h Africa.
of such S}'Odromes as abortion. polyar1hri1is. cnct-phaJomr· chla mydia was incrimim11ed as the cause or ahonion in
eli1is. and respirmory disease. Because many of lhe isola· mares and of korawconjunctivilis in foals.66 Abortion, in
tions of chlamydiae made from horses were obtained before mares due 10 chlamydia! infection, have also been rcporrcd
strains could be identified readily, It is not known whether in Spain 19 nnd Germany.6
more chan one strain of chlamydia infeclS horses.
An upper respiratory tract/pneumonia :,yndrome ma)
Diagnosis
be one of che more important chlamydia! diseases of
horses.9 • 5 1• s:i, 56 An isolate (N16} from horses was rccem ly A definitive diagnosis or chlam)'diosis requires the isola1ion
characterized a, belonging to Cltlamydophila pneumo - or 1he dernons1ra1ion or the chlamydia! agent rrom infected
niae.96 :'-lolecularly. there a re enough differences between 1lssues. together with rypical clinical signs and/ or pa1ho·
this isolate and 1he human sirain of C. pne11moniae LO logical findings. A retrospective diagnosis can also be
make il unlikely that transmission or C. p11eumoniae made sero!o~ical!~ Ir the netiological agent C"annot be dem -
occurs between horses and humans. Isolations ha,·c been onstrntctl. Clinical sign~ of the disease nre taken !1110 ac-
made from normal horses, and from those ,dth serous count together with a four-fold or greater rise in serum
nasal discharges. as well as from the lungs of a mare ~uffer- antibC1dy titre between acute anti c·onvalescent ;:.!ages of the
ing from pneumonia.9 · 51 5' · 5b. 114 /I.lost of the equine d isease.
strains that have been isolated are difficult to grow in cell Chlamydiae are usually isolawd in embryonated i;hicken
culrures and embryona1ed eggs. Ancmpts 10 infect other eggs or cell cultures. Becau~e or the ease of penom1lng !so·
horses ,,11h these strains have resulted onl~ in mild or lntion, in cell cultures and or demonstrating that inclusions
subclinical disease."!. "1 This is also clmracceris1ic of the are chlamydia. c.el! culture is the method of choice. Thb ap·
C. pm11m1011ial! strains occurring naturally in humans and pliM to isQ!:11es d~rh·ed from both bird~ and mammals.
koalas. Most i~olates 1,;11 grow in a range of commonly used cdl
Other diseases in horses that mar be caused b)' chlamy· lines. Sl1Ch as \'ero. BG and MK. 1• 3 with the exception of C.
diaeinclu<le a hepatoenc:ephalopathy ~yndrome reported in peeomm strain~ that grow better in embryonatcd eggs. Con-
Spain.4~ The syndrome is sporadic. appearing mainly ln 1h11 ventional cuhUie techniques are adequate but centrifuga-
winter. and aifeccs bolh very young and older hor,;es. II is tion Cat l 000-2000 Xg for liO minutes at close 10 37 •cJor the
systemic, im·o!ving most organ system~. including lhe liver. inoculum 01110 the cell mono!aycr increase, efficiency.' 3
560 ,,., ™" rnru : R1d:c11sial and chtam,dial ois,-ases

Blind passngil1gat four to six dnr~ lnrre,.hl.'~ eflkkncy of i,u 16$ or~~ rH:S:A are mor<' pmm• ro gilc a fal:,c-pMitilt' re-
lation. 11 I~ Important 10 examine cul1urc-s for inclusion~ m ~ull with otlwr Gram-negative micro-organi~ms Scn,ith it~
days th ref;' 10 six after infection oi cell culture~. IL ,,hould be can be incrcn<l•d h1 targeuns: a relatively short o:-:.\ ~eg-
borne in mind that chlamydiae are relmh cly ,cnsitiw 10 mcm '" r,r h~ the i1~e of ;i neMcd pmccdurc. rn 1 llmH·1·ei:. the
freezing; 1he common practice of frcczc-rhaw m di~rupt th~· use of the: luucr procedure \\ill incwase the putcntlal for
cell monolayers must be avoided. 1.aborawr. comamlnmlon. ,\norhcr problem h the need 10
Collection and handling of ~pcdmcn~ b unpor1a111 for de\'elop !):\,\ t"<lractlon pmcedurc~ that can be tbed for all
the succe,sful ic;olatlon or dcmon,1ra1lon of chlamydiae. t) pc~ of ,ample:,.
In the case ol abonion. the p!acema as 1he prderred tissue. Seroloi,•y as a diaj!nosnc: 1cs1 for chlanwdiosis has never
When the placenta is no1 nvailuble. vaginal ~\\'abs should bern u,ed cxwn~i\'el~ a~ ii require\ both acute and conva-
be used.8 :!. ·~ A11emp1s to i<olate rhe organism from the lc,eem pha~c ,era. ancl I~ 1hc·rcforc not cqmp31iblc 11ith 1hc,
foems arc often not successful. In other chl,m1ydial db· need for rapid dia~nosis and m•atmem. The complement
eases. swab, rrom thf;' inf('{'ted tissue~ or ex\1date~. or fixation W"t i~ the mo~t ccm,monl~ u,ed. 1 s. 4Q rhe u,e of
samples of Infected tissues can be u,ed Faecal ,amplt·~ El.IS.\ ha, Ileen wported. '" Both of 1he~c te<:ts detect anti·
can be used ir high levrls ol an11bio1ics are addL·cl 10 1hem. bod,· to lhe t.P~ or group-rcaniw mnigen. but 1twy are
A diluent consisting of sucrose-phosphate-glmamaw ha~ prone m pm,;de fahc-pr,sitin• rt">ults. a~ <ome other Gram-
proved to be sn1islac1oi:v a~ a 1ranspor1 medium lnr negati\'e orgnni,ms ha\·e th<' ~ame I I'S amigen epitopes and
chlamydia! field samples' .1. ~ 1 and it can also br used as u give low-lcYd po~ith e reactions. In recent yc•ar,. 1here have
diluent \\hen free7.ing culture~ for,torage ·\nribiotic, may hccn attempb 10 dl',ign ,erological tl',t,. iu which chlarny-
be added 10 it t0 rcducu contamin~tlon.' clial pro1<>ln, are us('d a, the antigcn.:!l C<1mpcti1in, ELI!>:\,
Direct staining of chlamydiae orchlam~ dial inclusions in u~ing scro,·ar-,penlk monoclonal antibodil·,arcalso being
tissue). orl:-mears of exudate, or in cell l·ulture cell monolay- de1elopt'd. t
ers, can be pe.rform.:d using hJstodwmical or immunohls· mt• mlcr11•immunoll11c,r<•sccnc·l' tcq (:'IIIH) h- lncrea,-
tochemical techniques. Some of the more commonly used ingly beini: employed in human mcdicmc. It is labour int,m·
stains are Giemsa. GimC.:ncz. Zichi· \:et•ben and .Macchia· she ,ind therefore e~pensi\·e. Thi.' \IIFf line, haw the
vcllo. 1 .J. "" ~Jan} of 1hes1? art> being r.:pla('cd 11·i1h spi;:clHt· ad1'llntnge 1ha1. ii knoll'n :.erotypcs ar(! uS<•d <I' antigen,. 1he
fluorescein-!abelled monoclonal "mibodies which huve a serotypf that infel'Wd 1hc animal can be d.eu.>rn11ned. II has
high ~ensicivit~ and spcclficit). Ileen used t<> ~hem that pig, 1~hh antibod}' 10 one strain ma}
In recent years. a number of manufnciurer.s have prn- not haw antihod,· 10 a11otlll'r ;,ttain,11!• and that ;.hecp anti·
duced cn7.\'Tnc-linkcd immuno,;orhemm,say {El.IS.\) kits for bodlc, in for1al lluid, an., to lh<· ovhw abortion ~train. 1
the detection of chlamydiae in human~. The$e l'8n ah.o bt•
used for the detection of other chlanwdinl' tier!\ ed from
Differential diagnosis
mammals although some mo) lack scnsirivity and specific·
iry. The,e kit~ derec1 tht• llpopoly,accharide (1.PSJ •.ind soml' ~!any o: the clinical ,lgnb :ind lesion, cau,~d h} chlanll'·
or the LPS epitopes are ~hnrc<l with other Gram-negat!Vt· or- diae ,ire also cau,ed bv other bacterinl and ,·iral agents.
ganisms. In addi11on. lhc EUS,\ kit• usually require 1hc pre.s· 1'vo1w of the le,ion~ or ,il{ns is pa1.hognomk. and 1herrforc
ence of a fow hundrt•d organism~ bcfort· a pm,itiw tc,1 is laboratol") cvnfirmalion i~ almost ahva}~ nl.'<'ded. In 1he
obrainetl. The gcnertll ml<' to follow i, 1ha1 if thC' animal con- case of ahonion. the other ori:anism, most commonly
cemed I< manifC1>1ing the clinical sign< of one of tlw chlmny, rncoumcrcd are Bruct'lln ,;pp. In ,meat, or ~ec1ions or
diosis S)11dromes and the El.l'iA i, ,1rong!, positive, the infectC'd tissul.',. It may hl' diflkult for inl.'Xp(!ri\'nred per-
result of the El.!SA 1es1 can be considered diagnostic :--ewer sonnl'I ,o dillerentiatt' bNwcen brucella. c~la1nydi(l and
ELIS,\ kits using i\ lABs ha,·c fewer cross-rcac1ion$; howC\cr. cox1clla morphologicall), making tht.> \!Sl' •Jf immunohis·
these t6ts ~till need lo be e,·aluatcd for.;p1:clfici1y and "'11· 1Clchern1cal wchnitJUt•, advbahlc.
siri,it) 1,'i1h each 1ypc of animal specimen
Pol)merasc chain reaction (PCR) techniques are n•lat·
Control
i1·ch ne1, and arc being used in a numberol diagno,tic laho-
ratories. but there are no commercial te,ts a,·ailahle> tor use C:hlamydi,!l' are ubiquitous in the liH",tock population,
in animals. although there are a number ol published which make<> l'accination and ;mtlblotac thE-rap,· rhe only t'f·
.primer ,eh-"0• ·";, 1~ 101 ~los1 primer ,e1, arc dc,i~nctl lO foc:1i10• 111<.'an, of c:omrol. Currcm va,·c:lne, are ,1rain-~pedlk
react \\ith eilher the I 6S-23S genes or the major omcr 111e111 and arc onl~ a1ai!ahle for the mine abonion strain :md rhe
brane protein (;\lOl\ lP) gene Some ur these 1cs1,,. when fl:'linl' pneumonatb stram. Hmh attenuated lil'e and ina('ti-
combined with r~triction t'rdgnwnt k11gth polymorphism \'all'<l Yaccinc, arc produced: a,ailabilit) dcpl'nd, on [he
(RfLP . will also differemime 1lw spcdes of chlamydiaf'fl cou1u11 concemed Onl~ 1he vaccines produced for the con-
The sensitivity and sp<.>cificily of these PCR tests still have trol or 01ine ch lam) dial abortion are di~,u~se<l bt•low.
not been established. In g,meral. tc5h designed 10 dctc<:1 the lnaciil'atctl \·acdnc< again~, mine ahwtion due 10
 Chlumydio,;i~ 561

chlamydia! infect ion a re usually formalin-inacti1·ated and neously with Brucella Rev l and the live Salmonella Rv 6
contain an oil-emulsion adjuvan1.r>G. 72 To be effective they vaccines. 91 When administered with 1he live Toxoplasma
requi re large amoums of a ntigen to be incorporated into the \'accine. it must be given at a different site. 1? The vaccine is
vaccine. ~1osr manufaciurers use the level of complement currenLly available only in certain coun1ries.
fixing amibodic;: induced in the sera of rabbits as the stan- Some antibiotics are effecth'e both prophylactically and
dard for judging potency. These antibodies are, however, thempcutically in controUing-chlamyd!os!s iii most ;inim:1ls.
not strain-specific, and a re also 1\0t considered protec1h·e. but 1hey must be admini$tered early in the course of the dis-
Because the strain specificity and antigen incorporated into ease and for extended periods to be effective. Tetracyclines
1·accir1es are often not controlled, variation in efficacy of this are the most "id<:ly u~ed antibiotics because of their rela-
type of vaccine is often a problem. Although most of these tively low cost and ease of administration. They have been
vaccines do reduce the abortion rate. they do nor prevent in- used prophylactically \\~lh success in ewes under field con-
fection or shedding.66· 72 An nual vaccination of ewes is rec- ditions to prevent abortions. For optimum results they need
ommended for these inactivated vaccines. co be given chroughout gestation. which may not be olher-
A vaccine containing a live anenuatedaborrion strain of wise desirable. &i; Chlonetracycline has been used in feeder
chlamydia has been developed. The strain ls a tempera- lamb~ to control polyarthrltis. Tylosin and long-accing 0:..1··
lllre-se1)sitive mutant of the <:;. abortus s1rain AB7 and is tetracycline have been used paremerall y.w ;>..1'os1 of these
designated I B.70• ' 1 It has reduced virulence for ewes. goa1s antimicrobials should be effective In controlling other
and mice. When administered to susceptible ewes or goats chlamydia! disease syndromes in cattle, sheep, goats and
before mating. it provides protection against abortion for horses. but their use in controlling S)'lldromes in pigs can
at leas:r three years and preven ts pos1,parturition shedding constitute a problem as manr of I.he current porcine isolate~
oi chlamydiae. 71 • • 3 • 7'1 It can be administered simulta- are resistant to tetracycline and sulphad iazine.6

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l,..'e:nncm Squeuc. r.\; Amcrico.n ,~O.Uoo of \-.i.o.n P4U'loftJ}?'.1!!11¾ ~~1.· llt'COl'(f. t< I. 63,,Gr.
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6 ,~n1tt5£..'I: ..\.A.. & ttC>Gl ~ u.o.. 19.98. R~is1anet.- to U!uaq·cUnc .:md Rffi'~rc/1, 35, 171-176.
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t>,,,~wtling, of tilt NimIr l11temol/Q11t1I S)'tn/KJ511111l <m Ht1111,ui
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SJ:\-538
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olprtogn;,n1 sh~p.}0111110/ o/Camparati,,'I! 1'111/io!og;•. !02. 221-237. bo,inc fotus.. Jcumol o/Cl111/Ct11 Mlaobio/JJgy, 28, 8l<l-al6.
564 s,;cr:o., maa: Rickeuslal and chlamyd.:al diseas~

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)CTIJm a-.."'ld SCffll\fl. Vererlh110· /mmunolo-zrand. lmmunQpathology. 13. m WARn. ~1.r .• 198.8~ Th~ ch13m)'tlhd dcvc1opmtnt3J cycle. In: e.,1t0;\' A..L,
2iJ.278. (ed . . ,\l/cro/1/ology o{ChfDm)>dfn. lloca Racon Fl.. CRC: Press.
103 Sl[R£D1. L, SCHllJ.f.R. 1.• Sl'D1-F.1l, r.. cusc;m,, J.., ltOS[.'-'., E., C.ORl'K)Z, L, 112 WARD '1,t , 1999. Mechunisms or C:hlamydla·induc..>d d1,e:ise. 111,
[G<;~SBDtCliJt, t .. 1o~r.s. C.E. '- PO""l5CIIII. ,., 1995, ln,estlnal C/1/amydln sn:PR<ss. Jt.S, <ed.) Cl1t;1mydia: /11rmctl/<1//lt Bioto/(',·, Pati10~11,sis, 011d
In finlsh,ng pig}, V,r,•rfnary Pnthofog,•, 33. 369-374. rmmuntry Washin~on. DC: Americ.un Society for ~t;c:ob1olog)•,
lO.: YAAASJUIL,\. L, IMAJ, Y., KAJUWA, 11, &- H.\S.m\tOTO. ~ .. 1996, Polymerase
113 \\'ARD!WP...... FOWU1'. "" 0°('.AU,.\{lH>\X, P.. (.IFf.\RD, , ..... nM~1S. , .. 1999.
ch3ln,reactlon for thedeurc:tlon or C/1/am;'ttla ptitracfin the raec<~ of Choracten.l.3Honof thc. koala biovnruf C/1/amy,/Ja 1mt'Ilmo11illea, fo-w
budgeripts. Mlcroblologieol lmmunofolJ>', 40. 21-26. gene 1~1-omp,W0.1. omp9. lf>S rRNA, woESl. sp•c•r region. l)c«;,marlr
10s TAPP&. ,. &:A....OERSu;, A,, 1.99J. lnhibit:ion of endo.wmal ocidificauon by A1,pl1td .,rtcrobiology. z2. 22-2i.
Chlom.1Tila psfrzacl. Ab.mac,s of 1/re 91s1G<!11e,a/ .\lec,1,ng ojrir,Amerirfl11 11.i WILUI,, f,M .. \\'AT~OS. C.i.. l,U..:,)H':n, M ,. ~tAJR., T .5,. wooo. Q. 4t RtCH,,o~n. $-.J..
s«~ey for .\1krobiok,g.), M•.)' 5-9. p. 103.. 1990. Charncterwuion of CJ1/amydia psfua<r isol4ted from~ ho~e.
10S TA'i'lOR, 0.1 ... 1999, Chlamydiae. Im STRAW~ o.r-. o',U.Wftfi. s•• M~GI.IXG. l'rrulnnry•,'1:erob1ulogy. 24, 11-19.
w..1... & TAVt.OR, o.J .. {eds). Ditetl.1&0/Su,tnt. Ames. Lo\; Iowa S'mt~ l lS WU£..,t<.1R.&, A-t .. ff,Afl.'itO.;\"'-. , ·." DA\\'S():O., M,, 193.;, E.."<J)erlrr.cnb (0
Univcrsir,,• Press. demon_muce route.of transmission or ovfne enioock •!)onion. Bnt/.#1
,07 TUSTL'<, a.<;.. MAAf. ,. & VAN l1EF.ll0Dl<, ... 1961. A dlse35<! of Cal\C$ l'etr.rlnary/oumal, HO, 38o-391.
resembling sponidlc bovine enccphalomrelltis. Joumnl 0/1/;e Soutlr Jl6 wrrrnxDRl~Jt:, M.M .• 1,.1IWAL G,. I MIU.[, I),, FISt:Hfl.R. D,. };A'\Ui~, II, 4'
,lfrlw, Ve1eri1:,uy M,-dlcal llss«ia.11011. 3?. t 17-123. •1,P1,t., w.. 1990. Dlttttlion M Ch/1u11y,lla f»lllati in vaginal dl...,hargo or
to& \'ASJWMP'AY, I), , OUC.\TELU:, Jl. & l tAt'..St.aR(IUt."'· JI,. 1995, Cl:lt'lfllJvlfn 00"" • n,-c,ss31}' cnlargomcnc or bacceriologlcdlagnosis forth•
1»/110.d lnft:ctions:a m1e\v ,,1th tmphasl$ on a,i3n cblrunydlo,i,. ctloloi,-rc c!3rltla1tlan l)f renllilycil,.order. in the remnlr cow. Jo11rna/ of
\feillrinaryM/crob,ology. 45, 93-118. Veierlnt1?' ,\ltal<mt (BJ. ~I, ~92-S03.
10g VEt~IA. 2.. XU\!)S.Eft. \',, C\SOl!EILE, 1,. M.A~OVA.. t.. S,V£C()VA, 0,. MSPJ~1l, L, ti? Wrnt.,n"1...:K, M,M .. W£.S, t.., UOlt,\lUt, :O.",wA,\ 'l'fSU.ER(;. G. 4 UL"-:Ot~. ~ •. 1991,
nmr.fKO\'A, ,. ~ Zl!Al.Y, A.. 1996. Detection of Chmm}'d13 fn lnOomma<lon Bakteriologhchc Un1•rsuohuni;cn wm vork,ommen von Chlt1m)dit1
ohhe cerylx and uterus In COW$, Vewlnnry ,11,t1re111, (Prnha/, 41, psfuncf in Orgnntn ,•on Sohwclncn und In ,\bonlcnen S<:hw•inOfctcn.
297-30-I. Jo1m1nl ofVfter/11t1?' ,lfe,f/(ine /B). 38, ~ 11-120.
566 ,rrno, TH• E: Rickettsfnl and chlamydial disease:;

Figure 43.1 Vege.ative tLJ and small. compac; elecuon-oensa (S) f01ms
ol Cox1ella bumetit (Mccaul and Williarr.s5'1t tReproouced tJf kind
perm,ss,on ;;1 McCaul and Williams and the Amer1caa Socie!I' for
M1cron,ologyl

Figure 43.3 O!velop,ng ooiar spore (El of Coiiella bumeMIMtCaul and

W1ll1arrsS., Ji'lepc-juced by ~Jnd permissior, o' McCaut and \Vill'ams and
itie Amer can Soc,er,· ·er M·c1ob,ologyJ

Phase II organisms are obtained following serial passage of

Phase I organisms in laboratory animals or cell cuhure and
are ml11lmally infectiVC'. Phase JI organisms revert rapidl~· to
Phase t organbm, during a si ngle passage in a \'ertebrate
ho:.1. Monoclonnl antibodies can be used 10 distinguish the
phase variams. 99
The genome of sLrains of C. burnerii varies from 1,5 10
2,'1 Mb ;md isolates can be placed in genomic groups based
on analysis by restriction fragment length polymorphism
(RFLP) and pulsed-field gel elecLrophoresis.62. 96· ,- Eleven
Figure 43.2 E,ewon m,crograph oi a~ intracyiop,as.,nic colony of Ccxiel/11 cl1romosomal genes have been cloned and expressed in
oumez,, Escherichia coli 58 and plasmids of 36 10 -12 kb have been
identified in the genome. 9 ' Recent srudies have indicauid
tJ1e baccerium. The cytoplasm ofcl1e host cell bccc:,mes filled tha t genomic strafn variation may be less lmponam than
\,i1h ,·acuoles loaded wh h organisms (Figure 43-~J. 'La rge host factors in ex'J)la ining the occurrence of acute and
s:ell ,·;iriams' undergo sporogenic difforen1iation and foon chronic Q fovcr i11 humans.47
resis1ant. spore-like fom1s of the b::tclerla. These de,·elop 10
SC\'s lhat are released when the cell~ lyse. Epidemiology
Co:o:iel/a b11merii undergoes ph~e variation,81 rnainlr
due 10 varin1ion~ in the lipopolysaccharide of the organism. Coxiella bur111.11ii has been fou nd world\\1de, with the not·
Phase J is the natural, highly inf~tious form of the organism able e.xcep1ion of :S.:ew Zealand.3.q ll occars in a large variety
which is found in !nfocted animals, anhropods and humans. of ticks. rodent,. birds. and wild and domestic mammals
 Q fever 567

and anti-C. b11me1ii antibodies have been found in snakes Large numbers of organisms are shed in the milk.of dairy
and tortoises in India.a Allhough pigs, horses, camels, Asian cows, 1>anicularly at the beginning oflnctation. Up tO 23 per
buffaloes (811bal11s bubalis), dogs. cats, chickens. ducks, cent of dairy cO\\'S ma)' be shedding organisms :n their
geese ar,d turkeys may be infecced, domestic ruminants are milk 1'1 and 21 per cem of bulk milk tanks In England and
1he major reservoirs for C. burnerii infections In humans. 6 Wales have been reported 10 h3\·e serological evidence ofin-
In southern Africa, serosurveys in cartle ha,·e shown tl1a1 feclion.64 Susp<mded in distilled water or sterile milk. the or-
C. Imme/Ii is endemic. Antibodies have been reported In ganism survh·es at room temperature for momhs and
cacde in ~lalawi (2 per cent),'; South Africa (8 per centJ, 35 somelimes years.!U Although people may develop Q fever br
7.amhia (1 co I 2 per ce111) 31 • 63 and Zimbabwe (33 to 75 per drinking infected raw milk•0 or eating infected millt
cent).42 • 68 Seropositive goats ([0 per cent) and dogs (15 per products,92 in mos1 cases ingestion of infected milk results
cem) were also found in Zimbabwe and seroposiuve dogs in serocon\'ersion without disease. 11 Current recommenda·
have also been reported in Senegal (l 2 per ceml and Ivory Lions for the pasteuri1.ation of milk (62.8 -C for 30 minutes or
Coast (8 per cent). i 6 71,7 •c for 15 seconds) arc based on temperatures required
Prevalences of antibodies ro C. bume1/i in humans ha,·e to inactivate C. bumetii.~~
been reported from the Cemral African Republic (16 per tn Nonb America, outbreaks of Q fever have resulted
cem).' 0 Egypt (16 to 25 per cem).17 Ethiopia (7 per cent), 1 from direct and indirect contact witl1 partutiem cats. 5~
Sudan (10 percenl).17 Somalia (37 per cem).1; Tani.ania (5 per Seropos.itive cats have been found In Zimbabwe ( 13 per
cem),.,Tunisia (26 per ce111)48 and Zimbabwe (37 per cent)..;.:? cenc} and South Africa (2 per cent) but they han? yet 10 be
Although cartle, sheep and goals may remain chronically implicated in human infociionsY
infected for years, the majority or infeccions are subclini· Outbreaks have also been reported in humans follo\,ing
cal.'6 lnfected animals. however, may shed large numbers of exposure to infected pigeon faeces. 78 Person-co-person
organisms in the amniotic Ouid. placenta (>109 / g), foetal transmission of C. b11rne1ii is rnre bm occurs transplacen-
membranes. faeces and urine.82 A major risk faC'tor for hu· rnlly and dunng blood transfusions and auwpsies.58 A
mans developing Q fever Is contact with infected domestic recent report describes sexual transmission in humans.';o
ll\'escock. Those at particularly high risk are farmers, abat·
10ir workers. hide and fleece processors. and animal health
Pathogenesis
workers. Infections of humans and domestic animals occur
principally by inhalation of lnrecced aerosols of the peripar- The target cells of C. bume1ii are monOC)'tes/macrophages.
turient Oulds witll only a single organism being sufficienr 10 Most infections occur via the respiratory route and involve
cause infection.&>The ei,macellular C. bumerii are extremely the alveolar macrophages.~5 The KupfTer cclls of the liver are
resistant 10 desiccation, low or high pH. dlsinfectams and the primary cells Infected in infections that take place \ia the
ultraviolet radiaiion.'4 The: organisms may remain infective digestive tract. Subscquemly there is a bactcraemia whkh,
in aerosols for up to two ,~eeks and in the soil for as long as in overt clinical cases in humans, coincides with fever.'
five months.49 Recent s11ldlcs have shown that amoebae Calves in endemic areas may beco1ne infected in the first
ma~ be infected with C. b11me1ii and this could f-urcher pro· years of life and either reco\·er completely or become la-
mote their survival in the environment.58 Humans coming 1entl)' infected. Hormonal changes in the final srages of
Imo contact \,ith an infected farm environmem are also at pregnanc~· in domestic mminants are thought to trigger
high risk of de,·eloping Q fever:115 outbreaks ha,·e occurred massive multiplication of C bttme1ii in the uterus, foetal
with exposure to manure.69 In some outbreaks, however. tissues and udder. ln the pregnam merus. a mild to se\'ere
there ls no direcr contact with infected farms or animals and placentitis may follow the massive mulliplicarion of
their products. In t.hcse outbreaks the source has been as- organ[sms. 56. S$ About 50 per cent of seropositive heifers
cribed 10 windbome spread of C. b11me1ii from infected shed C. bume1ii in the placenta, birth nuids and milk at calv-
areas In the viciniry.3 • ing. but the percentage drops as they get older. About 50 per
Over 40 tick species have been found to be naturally ir1- cent of chronically infected cov.'S cominue to shed organ-
fected with C. b11me1ii,6·"6 and transmission of the organism isms interminently in their milk in successive laciations.
may occur during feeding of ticks. Although tick bites ma)' After parturition. mo~t infected ewes shed C. b11rne1ii In
playa role in the uansmlssion ofC.b11me1ii ln ,did rod.ems. vaginal seC'retions for up co eight days and some (c. 12 per
lagomorphs and blrdsij 46 there is little co indicate they arc cent) ror about two months. 13 They only occasio11ally shed
lmponanr in the maimenan~e of infecrions in li\'e~ror.k or organi~m~ at suhse('.jnem parturirions.94 Ahhough infected
humans. The organism does. however. mulriply in the guc dait,• cows may excrete oq:anisms In their milk for many
cells of ticks and large numbers of C. burnetii are shed in tick years.3 '' ewes only shed organisms in their milk for a few
faeces. Contaminated hides and \\'OOI may be a source of days: 14 It is not clear \\'here C b11.r11e1ii persists in domestic
infection for humans80 either by direct contact or after the animals between pregnancies, 14 but in humans the D:'\A of
faeces have dried and been inhaled as airborne dust C bumetii can be detected in the bone marrow for several
particles. - years after acute infection.:16
 Qfe\'er 569

is essential for the early diagnosis and control of Q fe\·er

outbreaks.
The control of C. bumerii infections is djfficuh as its
... e>.'treme resi~tance to physical factors and chemical d!sin ·
fection makes elimination of the organism from the enn·
ronmcnt problcm:nic. 3 ,\ppropriatc tick control smnegies
and good hygiene practices can decrease en\'ironmental
,:· contamination. Infected foetal fluids and membranes.
aborted foe11.1ses and contaminated bedding should be in-
cinerated or buried. Infected animals should be removed
from herds or provided wirh separate conrainmenc facll·
ities in which to give birth. Detection of infected animals
may be difficult. however, as serology is not .reliable and
Figure 43.5 Impression smear of 1he placenra stal'led w1tn 1he modified the use of PCR is generally impractical.
Zien~Neelsen method. Note nume,ous Co~iella burneriigrganisms Inactivated vat'cines for domesric animals have been de-
veloped. A \'aCcine containing phase rr c;,rganiSms44 may be
or economic benerl1 by reducing lnfertlllty. abortions and
detecced with immunoperoxidase a1>Says 19• 811 and capmce low birth weights. Outbrea~ of Q fever in humans have,
enzyme-linked immuno~orbent assay (EIJSAl or EUFAe.1 howc\'er. been associated with vaccinated animals:ia as this
but these techniques require fresh tissue. Care should be vaccine does not prevent shedding of C. b11me1ii. parricu-
taken in imerpredngresults as C. bumetii may also be found larly in animals namrall)' infected before ,·accination. 18 71 In
in placentas or healthy sheep and goat..~.9- Slovakia, large-scale cattle \'aCcination programmes 1\ith a
Recenc sructies have sbown that C. bumecii can be de- single dose of \'accme contain.Ing phase I organisms have
tected b)' DNA probes29 and a number of polymerase chain been 3ssociated 1,i1h a decrease in the incidence of Q fever
reaction (PCR) assays have been described. A PCR wi1h in humans.' 5
primers derived from the lupAB-associatcd sequence can be The need for treatment of infected domesticated animals
used 10 diagnose Q fever in humans and has been used 10 seldom arises bm. if necessary. tetracyclines are the drugs of
detect C. bumetii in cow's milk.sz. wi. !IS genital S\\'(lbs, milk choice. However. antibiotic treatment of cattle chronically
and faecal samples from ewes. 12 ca1de sera 98 and dust from infected whh C. lmrne1ii does not appear m prevent shed·
barns.10n ding of organisms.9 Doxycycline. 100 mg twice a day for l4
In humans. Q fever is generally diagnosed using serology 10 21 days. ls the drug of choice for humans with acme Q
which enables di!Terentia1ion of acme and chronic infec· fever.58 Chronic Q fever in humans is difficult to treat: it is
dons based on amibody responses to phase I and phase 11 recommended that patients be tested \\1th do:1.·ycycline (1 00
organisms.8 ; Of the serological techniques available. the in· mg twice a day) and chloroquine (200 mg three times a day)
direct fluorescent antibody 1es1 is rhe reference technique for 18 months or with doxycydine ( l 00 mg twice a day) and
for diagnosis In humans: an ELISA has been reponed to be onoxacin (200 mg three times a day) for three years.08
useful for seroepidemiological studies." s; The character- Infection\ fn humans can be minimized by ensuring
istics and value of other serological techniques such as the 1ha1 workers in the animal industry are fully informed
microagglucination. complement fixation, radio-immuno abou t the risk factors of acquiring Q fever. Laboratories
assay. indirect haemolysis. ELISA, dot immunoblorting and should be provided \,ith appropriate safety facilities and
Wescern blouing. ha\'e been re~iewed recently.< 3• -ss equipmem. Vaccination of humans in high-risk occupa·
Serology mar be used to detect infected animals but false tions should be considered: in Australia, a formalin-inacti·
negative results may occur. 13 va1ed whole cell C. b11merfi (Hcnzerling s irain. phase 11
vaccine (Q-Vax, CSL) has proven highly effective in reduc·
ing Q fever in abattoir workers.2 Reducing exposure to raw
Control milk and promoting the use of pasteurized milk a11d its
The dm·elopmem of effective communication channels products will also contribute 10 lowering the prevalence of
between public health officials and animal health workers Q fever.
572 stcnoq111ITT; Rlckcusial and chlamrdJaJ dis~ascs

fi~<tt Q, R(!('l1til dr MMtttne Wtb"lna1rt 163. 19.$-19<1 92 \\'l'l"',,_"· L ,gr,:,, Die Q,f,-·er FOJ$Chung m de, ~hwelt m den.Jruuen
84 TiiJELl. o., li.,\HO, "· f.-1:AAUS.s, H.• 1992. '.\lonoclonal anlfbt1dy ba.~cd 194 7-19; I /,;,lrii"lzrift /111 Trof)(lmlltdf.in 1111d P<11r1.sl10/~gu.•. 3. 29;--30 I.
copmrc EUS.\1 Et.lFA for dij1<c11on of CCI.IC/ti/a bum,iiifn clinlc.J 93 wess, £ .. ,~131 Gro,\ih and ph-:ysfology orrickctulac. Bacu:moJogtcal
'Jl«lmens. f11roJ)ftlJ1 /oumal ofF.p11Jcmlo(ogy. 8, 563-51, Rt:•ltwt, 3i, 2!;!1-283.
8S THOM.i\S, DkR.., TRUW!r K. I... ~f.MOS, ~-l..., n:,.;cu. s._,, .. COUiMA.\, 'l',J,, 9.i ,,'nMJ, H,H.. /E."(4iC,.. r w. .tt Ut'\'l!lTI'., t..u .. 1959. Q rc,,er !ttudh.•:t, X.'X.
Mf..\DOWS:, O.. MORC'.,\."'tH::.\PNCR, I•. 4 CAIJL.. l,O,. l9'9S. The ri,ka( ~&quiring Compori>on of four ,..,,ological ttchniquc, ror ahe det,'<tlon and
Q 11,,~r on f11rms., a <trocpfdcmlologlcal s1udr. Om1pmfon..1find masurtm<nt a( antibody 10 <,<>.<1<1/ll /11111zc1i/ In nA1uroll)"e.'Posed
En1,/nmt1umtnl ,\1i!dlrln,• .52, 644-6-17 •h<,;p.,l11writ1ut /o:mu1/ of Hygie,ir, 70, 1-13,
8§ nGr.RlT. \\.U.. Bt::Stssos. :\.S. 4rl.'.flCIH.SUUR, \\'.S.., 1!'61 Airborne Q fc1.-..:r, t.r :9:;1, Q (f\·er ln
'95 \\IIU>C.. >Ul LE..~---En'f.. f?.,11- A.SlSA~n. F~R.. 6- \\1:-r,.:~.
8'1':rerlofogfeol Rtt·lcws. 2S. 285-293 California. IV Occutroaco of Co111blla bum,tll In tho plocecm.o of
8i Tl~Ol OUPOST. u.. T!-URJO>:, X, R I\AOULT, D.• ;9:94, Q fe,\'~r .stro1ogy: r:u,off'
1mural!r lnfoaod ,h••P· 1111/rttl S1a11'< Pub/It fJ.ot1/rh R,•pott,. S6.
d~rmina110n for microimmuno0uonl$C<na,. Clf11ieal nnd D1ngnof/ia 1473-U7i.
IAlx>ratt>l)• lmmwzol11g,•, I, 189-196. 96 WIU.f>IJ. "" UGF.~. ~- • OAIJeJ<, (;•• 1.998. Pll)'Sltal and ge,,e11c map of'""
88 TO, H,, HTW&..k,J:., "-'\KO. N., klM, 11.f.. Y,\'.\l,,\G.UCJll, T,. FU~Vsm, ti, .. lHRA1,
oblfg.ttt in,racellut:1.r bocterium ('.o..'(it.•lfn hur1Mtff,Jo1,rnn( of
or
"· ,998. i'rewlence CM/fl/a l,1,metii inlecuon m d3irr canf• with Baci,rioit>gY, 180.381&-3322.
rcproducuv• disorders. /011mol of 1',rm110,y .\le<licol Scic11r.•, ijO, 97 \\1U,E.\ri. U • TIIU!U P . hUkG!R, C., RITrn.R, ,1 .• 0$\\'.\IJl. \\'. & i(lt.\USS. n,.
859-861 1991>. Molerular biology or Caxt~lln bum,i/£ 111: >:AZAa, J. "tQ,t\.'<. R.,
8g \'AS \10~ I'. ft.-\VM(;o'\ffl-._l!R, \\',,'fl.SIU.'-' U. « 1v...,1atl:.S, 't, ;993
Ced$ Ric~<11sl(11mul Rickeusial Disffl= Br.uislo,'8. S1o\"1lcla; S!o,'111;
tmmuoo~1ochem,cal dcmorutrntion o(Coxielln Oum~il anti~e:n In the Academy of Scicn<lb. pp. ~78.
re,ol placcnt~ of naaur.Ur lnfcc1¢d ,hccp and c.in.!t /otJmnl of 98 \\'ltJ,l.'h, Hnuw,. D ,, Ht:Ol~C>MIITTfR. R.. fl )."lf.i\U'5, H.• :99.;. De:ection
ComJH)mth<t Patllo/Qgit, JO!I. WS-301. of Coxr,./la Jrum~1/iio cow's milk using the polymc~se cha,n reacdon.
90 \\',\I.OHAL\:, O,G., S'f()IU(:0-:l!fl, lt,c.,. Sl,'1,1Q~"", tu:. MTIIO\IA-.'• LA., HJ:'9, Z1mr,afb/a11 f11r \-'crrrimumtklfcwe. ,\I , .58()....587
Abonio11 ussocl3ted ,-ilh Coxi,lla buml'lilfnfe<itlon In coot,, /011ninl of 99 W'lLU-\\1.b. u:,, IOUM.TO~. M.11., PMCOC:li, M,C., nfO)tA~, LA,,.Sl"6W.\R't, ft.'-
tha Amentnn \leterlnat)' Medie11/ ,ts:s01:lntinn. 1;3_ 1580-1581 PORTih, f1L. igs.i.. ~tonoc1,mal amibod(l~ dbtm~ul,;h ph.1~,"3.riam.s or

91 \\'U~liUftC. \\',C .. D08SON, M.l!., S.\."un. U- o_\..S(:H. c..,\., ~l\t.lA\"$A.. LP..

CuxM/a btmmil. l11f,c1ia11 '1nd /mm1111lty. ~3. ·121-128.
SAC\, 0., ll..\.SDELCO, l., SF.('.:ttft.E.\"I". J.r.. \\'l,>' f-4;· \\OU[, C'..A,, t989, 100 \'.\,."\AU. T., MUAA.'l,••~U. Y.• IXOU\'1-..i I., ()XA&\Yi\\H1, l., UE~O, IL I, ;\10i.tTA.,
Ph)1og,netlc w,·ersitr or the Rlc~eu,1•• Journal of Bncw1olf©', I 7 t. c .• 199& 0<11!1:tlo11 of Cox/el/a b11rnnil from du,1 in a bam hous<ngdalry
,202--1206. r;:,ntc. .'.ficrol,lo!cgy«md Jmmu11ofogJ'. -1.2. 51-;13

,
 44
Eperythrozoonosis*
Synonym: F.pery1hrozco11 spp. infection

FT POTG I ETER

Introduction Eperytllrozoon wenyo11i. £. teganodes and £. momii are

generallr regarded a~ being non-patho.genic for cartle.
Epel}"lhrozoonosis is caused by Eperythro:.0011 spp.. which There is, however. a repon of fatal epef)'!hrozoonosis occur-
are rickensial organisms chat occur epicellularly on ef)'thro- ring in five- and six-month-old calvt~s.6
cytes and thrombocytes. as well as freely in the plasma of a Epef)'lhroioonosis of pigs and shce1l are probably the
~,1de variety of animal species including pigs. sheep. canle, only two disea:;es of veterinary importance In livestock to be
dogs, cats. mice and rats. Most infections are subclinical, caused by epel)'lhrozoa.N According to Jansen,n an epery,h-
but varying degrees of fever. anaemia and icterus occur rozoon had been identified in blood smears from pigs in Zaire
{partlcularly in immunocompromised or splcnect0mized by P.J. du Toit of the Onderstepoort Veterinary lnstlrute in
animals). Natural cases of eperythrozoonosis are general!)' South Africa as early as 1942. In the lJSA Ir was found that a
not of great di nical or economic significance in livestock. condition of pigs associated \\-ith iccerus and anaemia was
The genus Eperythrozoon wa.s established In 1928 by caused by an epe11'lhrozoon.M Spliner'H named chis parasite
Schilling~5 when he named the parasites he obseived in Epery1hrozoo11 suis and also described another non-patho-
mouse blood Epery1hrozoo11 coccoides.5 ln 1934 :S:eicz ec a/.63 genic variety, Epery1hrozoo11 µar1111m. The lauer parasite was
were che first co describe Epery1hrozoo110111s in sheep in South also described In pigs in South Africa by Jansen.44 Studies in
Africa. In this article a nocc by ~eicz and Quinlan was added. splcnectomized pigs revealed that £. parvum appears to be
which reported that they had also seen an Epery11lrozoo11 sp. more pathogenic chan had been originally thought.3 AJ.
in spleneccomized calves. In the same year. Adler and Ellen- though E. suis is regarded as an economically significant
boger11 described Eperythrouxm ivenyoni, which they had parasite in pigs. it is considered to be unimportant in the rela-
observed in blood smears of a splenec:tomized calf. tively small pig industry in southern Africa.
IIO}'"le36 found and named Eperyrhrozoon reganodu, an The aetiology of eperythro7.oonosis in sheep in South
organism that occurs free in the blood of caule. He also sug- Africa is well documemed. 59• 63 Extensive reviews have been
gested chat some highly pleomorphic species of Epery1/Jro- published on eperythrozoonosis and haemobanonellO$iS
.;;L)011 rna}' eventually prove to be mixed Infections of more comparing the aetiology and other aspects of these
than one species. Although many scientists have recognized infections.2•1-~6· 17
E. reganodes, • 24 • 26 · 27• 3 •· 50• 99 this organism Is not listed in Epery1hrowo11 01,is may play a role in the poorly under-
Bergey's .\1a11ual ofSystematic Bacceriology. 1* stood ill-thrift syndrome (Chapter 214) in sheep and goats
A third species in cattle. which seems co exclusively infect particularly in the Eastern Cape Pro,~nce of South ,\frica.
thrombOC)'tCS, "·as first reported as occurring in Madagascaf' Eperyclzrozoon spp. do have a nuisance value in splenec-
a:nd later in Finland.7· 98 Uilenberg100 named this organism tomlzed donors of live-blood vaccines nnd in e1'.-perirnencal
Eperyrhrozoon ruomii. and It wassubsequemly concluded that animals. Subclinical £perythrozao11 spp. infections in $UCh
this paras ice is a valid species. 105 All three species in cattle have animals may resulc in the occurrence of aberrations in e.x-
beenJrl,mtified in Argentina and in Gennany.?1. l6 Hm,·m·er. perimencal data. ?l probabl~· due co the reactivation oflatent
E. ruomii. which also occurs in South Africa, does not seem to infections. For example. sheep dosed ,,~th f>achysrigma spp.
have earned taxo11omic status. even after it\\'l!Sshown to have plants, the cause of ·gousiekte'. often develop clinical
no amigenic relationship to£. wenyoni.9 7 eperythrozoonosis.21

• Forrecent change$ m the oomencla1ure of these org11n,;111s. ,ere, to lhe ,nrroduc1ion 10 Section 3, Rlcl:eimal ond chtnmyd,al disc;,;e,
573
574 <tc:no-< nma: Rlckcusial ond chlam)dlal diseases

Aetiology and life cycle are situated in depressions of the s11rface ot erythrocytes.
'There are val)~ng degrees of int.ima<.-y in the association be-
Eperythrozoa are non -motile. prokaryorlc. parasitic micro- tween tl1e parasite and erythrocyte.:?JI Epe1y1hro:0011 suis, al-
organisms belongir1g Lo Lhe order Rickensiales, family though closely associated with the erythrocyte, is separated
Anaplasmataceae. Morphological differemiation between from the plasmalemma by a definile space. ioz the interaction
some Epe1yc/zrozoon and l-1aemobar1011ella spp. is difficulr.2·1• l>etwt:e11 pa,asites and the cells results i11 se\·~re deforrnation or
4
; Depending on the species. these parasites occur epicellu- the erythrocyte membrane. Epel) 11hrozoon 011i.< is anached to
larly on erythrocytes and 1l1rombocytes. as well as freely in the red blood cell by a thread-like strucrure, 39 whereas E.
the blood plasma. Their association with the eryL11rocyte 1i<enyo11i. which seems 10 lie in a slight depression on the'Sur-
membrane is used to differentiate between some species. face of the ery1hro1..·yte. has a IOO$e relationship with Lile cell
A summary of the morphological characteristics and lo - and the parasite apparently causes litLle hann to ilS host.~•
cation in the blood of the different organisms occurring in It appears that E. suis replicates in that small immature
domestic stock in South 1\frica is given in Table 44 .1. The forms bud from larger immature and mature forms which
characteristic ring forms of Eiperytllrozoon spp. seen in 1hin seem to correspond to the coccoid. discoid and ring forms of
blood smear preparations stained with Giemsa are probably E. suis. 102 Eperythro.:0011 011is also multipli~s by budding,
anefacrs formed by the spherical organisms during the dry- and in this wa}·, chains or clumps of organisms are formed in
ing process.2'· olli, 46 The proportions of the morphological animals with a se,ere parasitaemia. 25 Scanning elecuon mi-
types of E. 011is apparently change as the parasitaemia in- croscope swdies on E. we11yo11i showed lhat the parasites
creases.25 Both scanning and transmission elec1ron micro- occur alone or in chains, clusters. or pnirs, which suggests
scop}' have added considerably co our knowledge of their tha1 hoth budding and binary fission may take place. 45
morphology. mul1iplicalion and association becween them It has been suggested that the bone marrow mar be a pri-
and the cells the~rparasit_ize.25. •IS. n. 81. gs.102, 10s mary site for the multiplication of£. 01,is and that morpho-
There is little difference between the fine structure of ep- logic differences are due co the stage or multiplication. 39
erythrozoa25· 5 1. 69 • 9a. 1412• 105 and Haemobarro11el/a bo11is.lfZ The complete life cycle of Eperythro.0011 spp. i, still un-
The organisms consist of a single membrane enveloping an kno,,'Tl.
amorphous matrix containing small ribosome-like granules.
Vacuoles and delicate filaments are also sometimes seen
Epidemiology
\\i thin the macri,x. Most of these organisms seem to be sepa-
rated from ihe host cell membrane by a definite space £::perythro::0011 spp. are very successful parasites, being
(Figures 44.1 and 44.2). widely distributed throughout 1he world. causing liule
In the case of 5. 011is, rod- and spherical-shaped parasitei. disease and surviving indefinitelr in the host

Table 44.1 Eperyrhrozoon spo that infect comest1c stoci< in southern Africa

PARASITE HOST LOCALITY SIZE/DJAMETE!I MORPHOLOGY IN G1Ei\.1SA-STA1NED SMEARS·

E wenyoni Canle Ep,,eiywoevt c anll iree 0,3-1.5µm R,ngs (typical). discs aM coceo,ds g,e comm,n
!o:ms \·.t.lcll siain reddish-purple Paras',es iorm
clus1e1s arour>d circumference of .ry..,rocy;es
E 1e~='5s Canle Free ,n p'asma Rods and th·eads S!eic!e1 roes and threads are m1mcommon q,og;
0.3-3,5 pm leng:111 v.11/l rod or threac auached lf1Ving-oao oullmel are
0,2-0,3 ~m wid,h rare. out characteristic of spetie.s Stain
reddish-purple
E iuom11 Cattle Altached !O thromboc·(les, occasionally Rings Ring, ovoid and coccoid forms. Da!klv sta1red rods
free in plasma attached to 0.4-0,Sµm are cccasionally seen Parasites ctmoletew
granulocyies rexeeptionall surround tllrombOC'/le.s in severe tnfect:o.ris
f OWS Sheep, Ep1·el'{Jltot\'ilC 2/lll :ree ,n plasma 0,5-1,0~m Air.gs. reds and uregulartv sllapen paras,1as ma\·
goats tllver or form clusters around circJmference of red
blood cells. Stain pale purple or pin~1sh pu•ple
E suis Pigs Ep1-eiylhrocv11c and occas,onallv lree Norm.I Ring forms predominate (large r-ng 'or'.Tls nave
• in plasma 0.8-1.0 µm c,stoned appearance) Coctoid. disc and rod forms
Large forms 0.3 vm also occur. Stain pale-purple to 1ed1hsll-l)Urple
E.par>,um Pigs Ep_1,erytiuocvnc ano free in plasma 0.5--0,B~m Coccoid and occasional ring forms Paras,,es
accumulate in large numbers on 1:in•viduat red
blood cells Stain pale-purple :o re.ldlsh,pu:cle

• Ttie c1:1lour va,1es with 1he pH of the buffer and me catch of stain
 Epcrythrowonosl~ 575

Eperycllrozoon spp. It is generally believed thal the~· are

arthropod-borne; the pig louse (Haemmopinus silis) Is ca-
pable of tTansmitLing E. parv(mz43 and certain mosquitoes
(Cu/ex a111111/iros1ri; and tledes camptorllynchus) can trans-
mit E. 01iis mechanican~,. 13• :i.; Stable mes (Scomo:tys calci-
crans) ond licc CBo1iicolabo11isand Linog11a1husviw/fi 61 may
play an imponanc role in the transmission oi infecrion
amongst stabled animals.7° Eperychrozoon spp. can also be
easily transmitted iatrogenically by all paremeral routes
with infected blood or organ suspensions. Eperyrlirozoon
suis is tra.nsmined uansplacemally.4 /11 wero infection of
E. 011ishas not been demonstrated in sheep.15
·n,e role that 1·.ild animals may play as reservoir hosts
would seem to be unimponanr as it is believed that Epery-
rhrozoon spp. are limited in their host ranges.2'' Their host
specificity is poorly defined and this is one aspect which has
cast doubt 011 the taxonomic posinon of some of them.ia
Figure 44. 1 Eperych({)]ODf1 wenyonim close asso,:,a11on ,·.•th, bu, Neitz61 sho,,·ed that £. oPis and E. flle11yr111i are both host·
separated by a defin1ts ; pace from. the redblood cell memorane Note specific in domestic stock. Hoyte3' confinned this finding and
parasite free in p!asma (>< 50 000) also showed that E. ragenodes is not infective to sheep. How·
ever, F. 011is may be lnfeclive to goats. 16 narural Infections in
goats wil11 E. ouis ha\~ng been reported in both South Africa95
and Australia. 12 Latent infections have also been produced in
two blesbuck (Damaliscus nlbifrons)60 and an eland (Ta11rorrn-
g11s ol)ix) ro!lo,,ing artificial infection 1,ith E. Ollis. 18
t\:eirz59 reported that in sheep age makes no difference to
the susceptibility to infection wilh £. ovis in South Africa.
But in c;omrasr. workers in some countries. especially Aus-
tralia and New Zealand, describe various degrees of suscep·
tlbility among lambs. young sheep and adult sheep.
lnrectlon may result in subciinical infections, in an ill-thrift
syndrome. or In severe clinical disease with high morbidiry
and morraliry.8
No detailed studie!. have been made on cperythrozoo·
nosis of sheep and goats in southern Africa. According to
Neitz59 farmers in South Africa describe an anaemic condi·
tion in sheep 10 which the name 'bleel:sickre' (pallor disease)
has been given. The a,etiology of lhls condition remains
Figure 44. 2 Eperfthrozoon 1uomii· electron micrograph of a section obscure, and It is unknown to what exte111 venninosis and
through an ,ntected platelet showing association bew.een the organisms mal;mtrition or other contributory factors may be involved.
and :he platele: The epidemiology of E. m,is infections in sheep is probably
further complicated in endemic areas by other haemopara.-
sitic diseases such as anaplasmosis and theileriosis.
A variety of Epery1hrozoon spp. infect sheep, goats, ca1tle Once anfmaLs are infected they remain life-long carriers.
and pigs in southern Africa rrable 44. l) and are probably Sheep that recover from £. 011is infection develop a premu-
very prevalem. However. very lin!e information Is available nity; a,rtiliciaHi•infected sheep may be infected for up to ten
on their epidemiolog)' in thesubcontinenr. Jt is belie\'ed that and a half years. H AulOsteriliz.ation has not been reported ,
heartwater. babesiosis and anaplasmosis live-blood \'aC· b\lt chemosterili7.arion renders sheep fully susceptible to re-
cines. produced inadvertenlly from Epe1)'/lirczoo11 carriers, infec1ion.62· 76 Splenectomy or 1reatment \,ith dexametha-
unincenrionally caused the widespread distribution of this sone results in relapses and a resurgence of parasitaemia.Z5 •
organism in somhern Africa.70 5 1 59
• Lambs that suckle £. Ollis-carrier ewes appear to be pro-
Jn Gennany it has been shown in a sur\'ey of blood tected from patent infectlon. 15 Serum antibody titres [which
smears from 570 pigs that 5. 7 per cem of healthy pigs arc believed 10 be protective) of uninfec;red lambs of such
harboured E. suis? 1 ewes apparently decline to zero within three weeks af1er
Little is known about the na1ural mln~mlssion of weaning. 15
576 ,euu,, nw,.: I.Uckcusial and chlamydia) diseases

One of 1he biggest obstacles faced when producing red- roc~'les and Induce desmiction by phagocytosis, primarily
waier (babesiosis) live-blood vaccine for cattle at Ondersie· in the spleen, but also in lymph nodes. 6q /111iitroagglutina-
poort is 1he imerfcrence caused by concurrent E. we11yoni. Lion of sensili.7.ed ery throcytes occurs spomaneously at 25
E. ruomii and E. regrmodes infecclo1\S in splenec1omized and 4- °C respectively. IQ~ The aggfu1ination phenomenon
donor calves. These infections result In prolonged incuba- correlates with increases of serum amibodies {as measured
tion pcnods and in the suppression of parasiwemias cali.scd by 1hc indircet haemagglutinmion 1cst) and of 10ml scrum
by both .Babesia bo11is and 8. bige111i11a.-o lmerfe.rence be- globulin concentrations associated with lhe appearance of
t\\'een concurrem eperythrozoon and other haemoparasilic lgM ·cold' agglmin!ns. 104 Erythrocytes, lost ro che ·cold"ag-
infections. such as bt1tween E. wenyoni and Am;plasma ghninin phenomenon, .ire replaced by reticulocytell. As the
marginai<?-0 and between /:. 011is and Babesi(I 011is.23 have new red blood cells are not readily infected and the altered
a lso been reported. Eperyrhrozoon we11yoni infection not erythrocytes are phagocytosed, the amigenic stimulus re·
only causes extension of tht! incubation period in 8. big· quired for the maintenance of indirect haem agglutination
emina infection,:!$ but it also suppresses the parasi1aemia of titres and globulin concentrations is ell'ec1ively re-
57
A. margi11ale and B. bigemina infe'ctions. Eperyrhrozoo11 moved.103 11aemat0logical findings in splenectomized pig,;
tega11odes apparently suppressed the del·elopmem of A. indicate rhe dcvelopmem of a normochromic anaemia
margina/e in a splenectomized calf 2 - Complement fixation caused b}' ex1ravascular crythrolysis.28
1esung has shown that Anaplasma and Epef'}·tl1ro;:0011 spp. :--acural £ suis infections show no signs or penetrauon
share amigens. 16 However. the indirect fluorescent ami- and erosion of the red blood cell membrane comparable to
body test shows no cross-reactMty between£. ovis antigen those which occur in flaemobarro11ellnfelis infections. 69 Se-
and A11aplasma 011/s aniisera. 40 Even between members of vere deformation of the red blood cell membrane has. how-
the Eperythrozoon spp .. such as between £. womii and ever, been demonstrated in splenectornized pigs infecced
E. we11yo11i. there appear~ 10 be no anrigenic relarionshlp.97 ex'J)erimencally \\i th £ suis. 103 11is thought 1ha1 the humoral
Eperythrozoo11 suis and E. parvum infections result in a immune response is directed in part against these parasite-
subclinical carrier state which may be activated under altered eryihrocyt~s. 103 Autoamibodies to erythrocytes.
stress. There is no cross-immu nity between 1hese two spe- po<>Sible mechanisms involved in parasitic immunoregula-
cies.84 but infection of E. suis suppresses the development of tion. and misdirected immune responses have been impli-
parasimemia of£ parvum.85 Eperytlirozoonparvum also in- cated in £. suis infections. 103 Smith, as cited by Henry.30
hibits I.he developmem of parasilaemia of Babesia trrwt- believes :hat F.. •iui$ in pigs may affect all age groups and that
ma,ini infeciion.3 ils clinical manifostadon is not limited 10 the classic disease
of anaemia and icterus of feeder pigs under stress.
The only lnfonnation regarding the pathogenesis of
Pathogenesis
infections by E. wenycmi, B. ceganod~s and E. momif has
Litrle information is available on the pathogenesis of bee1\ obtained from s1Udies using sp!enectomized
Eperyrliro;:0011 spp. infections. animal5." 5· 93· 105 In none of chesc qudies were significant
The majorlt1• of Eperyrilrozoo11 spp. infections. including ultrastruc1ural changes of infected red blood cells or lhrom-
those due to more pathogenic species, are subclinical a11d boc~'tes observed:'•
may be exacerba1ed b),' stress. Diseases produced by
Epery•tllrozoon spp. are characterized by the desm1ccion of
Clinical signs
erythrocytes or. in the case of£. ruomii, thrombocytes.
IL hll$ been suggested that anaemia in ovine eperythro- Nei1Zand his co -workers"°· 62 · ,;. showed chat the incubarion
zoonosis is 1101 direcdy associated wilh lhe parasicaemia, period following anificial infection with £. ovis could be as
but that ii could have wha1 was called an allergic base.u-1 short as 4 10 15 days, but could sometimes extend to se\·eral
£:.perytlirozoon 011is also seems to affect glutath.ione, which is weeks. Parasltaemia appea rs concurrent!)• "rj1h the rise in
essential for nonnal ery1hrocyte imegrity. 2~· 92 Heinz bodies body temperamre and increases umil che first signs of
in erythrocytes of E. ovis-infecied sheep56· 67 • 9~ have also anaemia are de1,;c1ed five 10 eight days after the initial de-
been implicated in the pathogenesis of anaemia.56 tection of parashaemia. Parasl1aemia 1hen wanes abrup1iy:
Cenain aspects of E. ouis infections are consisrnm with few paTasites are seen when 1he anaemia is mosr severe.
those of a haemolytic anaemia. 19• ; g \\11ile Sutton90• 9 1 be- Recovered animals remain carriers and recrudescem
!Je,·es that intravascular haemolysis occurs. Sherilf8 con- para•ltaemias may occur at irregular imer\'ais [or six weeks
cludes that haemoglobinuria. as reported by Overas.GS is or longer.
rarely found in£. 01•is infection. which suppons the conten- The severity of the disease in sheep is e:1.uemely \'ariable
tion 1hat extravascular ery1hrophagoc~1osis rakes pince. and seems to depend on the degree of the parasilaemia and
The cause of anaemia in eperythrozoonosis in pigs has its duratlon. 1• A cominuous or imermiuem febrile reaction
been ascribed 10 autoimmune hacmolysis cause by 'cold' may be followed by \'ariable anaemia and sometimes
auto-antibodies.33 These amibodies are anached to eryth· lcterus. 63 Other signs include depression. anorexia, loss of
 Ep<·l)•throm<>no~is 5Ti

condition. debility. rernrdation of growth rme. lowered hae- lowish-brown enlarged liver. and splenomeguly. There may
mog!obin levels. a drop in wool production and decreased also be hydropericardium. a~cftes. and epicardial haemor-
exercise tolerance. 11 - 6 The pulse rate becomes rapid and rhages. Microscopic lesions include hyperplasia of rhe red
weak and the respiration accelerates. llaemoglobinuria oc- bone marro\\'. haemosiderosis in the spleen and li1·er. and
curs \·ery rare!y.59 ~tonality rate is usually very low.8 · ;i •• fatty c-hanges and necro~is of hepatocytes. 17· 32 38 · ,-
tn pigs clinical signs arc most likely 10 be observed ln Pcrtvascutar oedema and dlscre1e gllal moblilzation in the
young suckling piglets and sows shortly after farromng. !mun ha\'e been de~cribcd in E. s11is carrier pigs that had
Mos1 F.. mis infenion, result in a subclinical carrier stare been s11hmectomi,:cdY
which may lead 10 01·er1 disease when animal$ are Epwy1hrn:,xm wttnyoni lnfec1ions in splenectomlzed
stressed. 53 The early phase of the infection is characterized calves rcsulr iu lowered packed red blood ceU volumes. a rise in
by high fever, but in more advanced cases temperatures white blood cell counts and erytlTTophagocytosis. ijl 71 Changes
may be normal or 5ubnonnal, and the mucous membranes in splcncctomizcd 1·alws infected with E. wen)l()ni include an
may be pale and icteric. 01her signs include weakness. de- increase in lactic add. lowered blood pH, negative base excess
pression. loss of appetite, conslipmion. bile-Mained faeces and lowered mm<lard bicarbonate conu:m.106 i\ letabolk
and dyspnoea: 7 Some animals suffering from the dlsease acidosis has al<o been reponed. 101
may die. In animals that recover relapses mar occur. Eperythrozocm womii infecnon results In a prolonged
J;"peryrhrozoon wen.ro11i usuall) c:auses onl) mild fever blood-doning time and se\'erc reduction in the number or
and anaemia in lntac1 canle65 but ve,y occ-aslonally some af- thrombocytes and petechiae in the organs of sp!tmecto·
fe<:1ed ammals ma1iifest ~cvere clinical signs.-1· 73 Severe miied calves. 1i::;
anaemia and icterus occur in infected splenect0mizl!d
animals. 61 ·9""'
Diagnosis
Hoyte3" considered£. regnnodes to be of similar patho-
genicity 10 E. we11yo11i. /:'pe1y1hro::.oc111 111omii haYe been F.perycliro=11 ~pp. infocrions should be considered in
rnportcd to cause fever and anaemin in ~plencctomized anaemic animal~ e-Specially if they ha\'e recent!)' been
ca rrle.9b. 99. 100. 105 spienec1011u1.e<l or immunosuppres.5ed. In most case& epery-
thro:wonosi5 can1101 be diagnosed clinical!) until anaemia is
well aclvam:cd. If Lhe examination of blood smears fails 10 con-
Pathology
finn the diagnosis. subinoculation of blood imo susceptible
The major lesions cau&ed by eperythrozoa (except splenec1omi;:ed hosts ma~· be considered.
E. womii) are the consequence of eryihroc>rte destruction Successful di:;lgnosis of cperythrozoonosis is ofien depen-
resulting from erythrophagocytosis or imrava&eular dent upon the correct technique being used when making a
haemolysis and the stimulation of lhe immune system. blood smear. Dirty glas~ ~lides and stain deposits can hamper
which leads 10 hyperplasia of the reticulocndothelial cells. recognition of lhese delicate parasites. Blood smears should
In sheep. lowered haematocrit and haemoglobulin levels. be made from fresh blood preferobly collected direc1ly from
hypochromic anaemia. a rise in leukocyte counts and eryth· the live 3nimal. a,, the organism, tend to lose their srrucrural
rophagocytos-i• have been reported.~~. "-1 In some animals. integrit) and ma) nm be idcntifia.bie in smears made from
lowered haemoglobin and haematocrit leYels occur one week stored blood.- For ex.imple /;. w1•11yo11i'disappeared' from the
aftt>r pt>nk par:i~iiaemia.10 whereas in 111hers. rhPrr k a rrla- hlood nf a splt·nl.'rmmi;wd calf within four hours of death. ;o If
tionship between the severity of anaemia and peak stored blnod is us1•d, blood cnllected in EDT,\ is beuer for di·
parasila~mia.55 · 67 The ma111 chemical pathological fea1ures agnostic purpo~e,, than heparinized or citrated blood.8 It has
include an increase in hlood lactic acid ar.d a lowered blood also been demonstrated that F.. 011is is frequently absenc in
pl I.fl:". 88 P)•ru\·me and lactate concentrations reduce the smears made from the peripheral blood of sheep in which
blood pH 10 to 14 days after lnfection.'l Sutton87• 88 reported anaemia is ad1anced.s. 5 1 ,s. 79
that affected Cl'}1hrocytes have a high glycol}1ic acthity. On The 1l1ick blood smear te~hnique54 for Babe.<ia spµ. iden-
chc other hand. llemobade and Blotkamp ,i found no effect tification is extremely useful for making early diagno..~es of
on the con<:entration of blood gluco~e in infec1ed animal5. eperythrowonosis.;n Different scoring systems are used 10
Variable degrees or anaemia, iccerus, and fuuy dege11era- quanrit'y the parasitaemiac;.
1io11 of the liver have been reported in sheep?• 8 19 · u 51• 39 · The organisms appear bright orange when stained with
67 qo .:rhe spleen is enlarged and sho\\'li e reddish-brown 11c:ridinc orange ond \i~·wcd under o nuorcsccm micro
pulp and severe hyperplasia of the .\·lalpighian bodies. There scope.:i.i Thi, s1.uning technique is more reliable for demon•
may also be a marked hydropericardlum together with sub- st rating low infections than is the Giemsa stain.~;
cutaneous oedema a.long the \'Emtral aspect of the neck. Campbell er n.l.s- believe that E. 011/s infections result in a
Haemosidcrin Is present tn the kidneys. li\'e.r and spleen a1 charac1cris1il: type of :inacmia. apparently of diagnostic
peak and late stages of parasicaemia. value. which is predominamly either macro- or normoC)'lic
~ecropsy findings In pigs include anaemia. icterus. a ye!- and normochromic in nature. Hoffmann3 t contends that
5 78 ,,,nil'>''""'•; Ricktusial and chlamydia! disra~£>s

clinical signs in pigs are non-specific and that the phenom- Con trol
enon of instant agglutination of blood sample~ is important
in the dingnosis of E. sui.~. Different drugs hal'(: been tested for acciliiy agains1 epel}'·
A variety o( serological techniques can be used to dfag- rhrozoonosis.~ 4 Arsenicals and rerracyclines app<'ar 10 be
nose infection on a flock or herd basis. These tests include effective against mo~, infections/' 99• 100• in:;bm in spite of
direct lmmunonuorescence, '6 the modified amiglobulln their efficacy. Lhese drugs do not sterilize the mfecrion In
(Coombs) test.80 indirec1 haemagglutination, 82. 113 comple- most cases. 7 .;~ ;z Treatment of sheep infected wit.h £. 01•is
ment fixation, 10 indirect immunofluorescence40• 4 1. 0.1. 66 with spirotrypan sterili1.e~ the infection, and such animals
and enzyme-linked immunosorbe111 assay (ELISA):19 become fully susceptible 10 reinfection. 77
The arsenical drug, neosalvarsan, is effective in the treat-
ment of E. ol'is and E. 111e11yo11iinfectlons.6t. 63
Differential diagnosis
B01h arscnicals. which are becoming incro2singly difficult
Any condition that resu lts in anaei:nia and icterus i11 sheep, to obtain commercially. and OX)'tetracycllnes at a dosage rare
goats, caule and pigs may be confused \\ith epel}ihro- of IO mg/kg bodyweight administered inm\\'enously or intra-
1.oonosis. In sheep these include malnuultion. helmintho- muscularly, are regularlr and successfully used t0 treat £.
sis. and the syndrome known as Ill-thrift (Chapter 214 ). 111e11yo111. E. womii. E. rega11odes and E. ol'i~ infectious in sple·
Differentiation between anaemia caused by copper and nec1omiZed cattle and sheep a1 the Onders1epoon Veterinal}·
cobalt imbalances and£. ouis infections is difficuh.8 · ?b Re- lmtimie.70 In acme cases ii ma)' be necessary 10 repea1 the
cent work in Austr.ilia has shown that£ ovis is the major 1rea1ment if the febrile reaction and/or parasimemia persists.
cause of ieterus in lamb~ and that copper poisoning is and during parasitic relapses. According to Campbell el al ..8
much l&-s hwolved. 2 trcauncnt of affected sheep should be based on the reduction
The mncroscopic lesions caused by E. Ol'is in sheep of stress by good husbandry rather than by therapy.
closely r~semble those seen in 01•inc anaplasmosis60 or Tetracyclines appear to be the drugs of choice for rrearing
chronic copper 1>oisoning.z £. su/s infec1ions.w..JO. 38• 58• 71 F.pcrytlmnoon suis infections
ln South Africa. concurrent Anaplnsma oi,is and T/11.!ile- do not seem to have an appreciable effect on reproduclive
ria 011# (T. sef)amta?) infections ma~· aggravate E. ovis performance, 101 but chlonetracycline treatment in the diet of
infec1ions in sheep. sero-positlve sow~ results in lncre,Lwd productl,ity.;.;
The poisoning of goats wirh the pods of Acacu1 ni/otica Although not conclusively proven. the possible .role of
bmussinnn may result in the development of anaemia . eccoparasites in the transmission of the infettions should be
icterus, methaemoglobinaemia and haemoglobinaemia.95 considered. It is therefore recommended that animals
Babesia 1ramma1111i infections and lepcospirosis should be rreated for ettoparasites d11ring outbreaks. Care should be
be differenriated from £. $1/iS infections in pigs. 29 exercised 10 prevent iatrogenic transmission of infection.

References
\Ill.ER.~." uu;:s'Tl()(lr:,,, v., 1934 ••\ no,e on rwo new blocx! para,1t<~ o( moflai..tly 1n l~n~ ,n \'1c1oria 3:;50<'iated '",th F.pt•T)1h10;,xm ons.
ca11le. F.p,,ry1h'rr,;Q011 ond 8nrtondlo. /011mal o/Cim:pamri, ~ Parltolog;· A1uuol1,u1 \ '1•1.:mutr)' Jo:mwl. ,ii. 5,38-,5,11
(lftd n,~m,i.:ur;u.41.2J9-221.
9 rnuz. J;., UliaJ'.,U)('), I I 01•t7., H.. 1'01..'\',;~0. H •• U))fBILW. Jt.o. 4" R0D1UQUE7.
2 .-\RCHE1t. J,h" unu11011N~. 1,x., t.98, /;p(rythroU,011 oi•ls and copper o.:< .. 1;.85. 0,g,n p,\tholo~ in ~xperlmon1al p<>rclnc cp<:i;'thro~QOnosl!..
po'k-Onint,:; 11., c-.a~\~~ of jaundkc m lamh carc:isc:f ..-1.usrm/Jnn Vttnina,y Ru1 1J,ff« <Ir: ~lltul Anlmnl, 7. 59-GG.
/Ol(Cf!fl/, 61. 312-<!13,
10 UAl>l>cl\, ,c,.. ;9;-. ,, complcmc1H fixntlon tc-.).t for th~ !l(:U.'tli1>n of
3 u,us.,,.1·.,.,., 1963. Epcrythro1.oonosb in pig$ In i:,nya. R111/11i11 of E/J,!()'thro:oor: lnf,'Ctlon In ,lu:ep. A11Mralla11 I ,i,rlnai,·Jo11ma/, 53,
Ep;:0011, D1«tu.esof"1frlc11, 2. 18$-ISS. 139-1~3
.a Bf.iSUUl:R, 11.u." r.our.1. R 1,., 19p-1 F.pcr,.rthrotQonC™s tran.smiu.ed In llti!rtt ti Ep~f)'thro;,,,Km 01 1is- a C'QUit." or ~..acmia, reduced
tl,\J)l>(J\\ )(.'.\., 1,':'9,
trcm C':.ttritr SO\\'b co thuJr pi(;~. Jtmnw/ 0J1/:~ .'4mtrfmn Vct:trma,y p1od11alon and d,-rrcu><.'<l e><crchc 1olernnce In W<'<!p••{wtralion
\/e,/i,:a/.~-,orlm/1m, 124, 98-100. '"'t,•111ur7=: Jo11r1111I. 55, ·l3J-i:u
s sa.u...,.,.()(~Ht, s. • ,·.,.s~11 um,, 1•.. 1929- ConmbutTon a l'e:tudtt dt"S ,: uAr')1>(J\\ Iii.,:\" 11179. na• natural otcurrcnet• in il..l{\J,"JI o!an orgunh,m
tpc~1hro1c:,.,Jrcs coccoldc;., Annnlts dt Ynra,110/0B4· Humal1u: c, n..~mbllni f..P-'f)'Jltro:«m m·ls. \usrmlftm Ve:tlfm1,:. ,ountul, ».
Com,,..,01,.. \11.)5.1 .... 3';3. GO~,
r. ,,~:,; ,~>0. A •.Ul.,n:. o .•(.ur.uu.M1~0. It. -..nru, .\, u .. !s,&3 Scb.'lla!rudonr: 1,1 O\lU>CJ\\, 11,. ~ , 1980. t..:ulannnuli.10,tnS a'\ a.\ 1?1:tor of i:IN')·1'110:.00" 01•i1
di Epcrythrou,onoii In 1'11e1U. Sdw tdur drtllt1• fl1r T/nhrl/k11nd~. 12~. lnfeciion in ,hotp. v,wmflry 11ams11otogy, 1. 313-3!;,
3:.3-35,
1,i D.U>DO\,·, x.~ .. 19$1. The dumton of th~ eatr.e:r ~uucof F.~ty:;iro..:.oon
i C\UIJ\\, LL, 1984, Prmo:.ool n11d JUrk,1ttd11l P&•ase..<.. Vol. s. Canixrrn: o,t/sinfectto.n in ')h.t'~ AuµmUn11 Vvl1i.'rinary Joumal~ i7 49,
.\11$!rallan Sun-au of Anlmol H<•ahh ..~u.imllan c;o,~mnienc Publishing l5 UADD<:>1.\ ~~ .. 198:0. The prOh..'t.'UOJ\ ofhunbs fr01n ~pcr.1hr02oon
X°f\.1Ct'
ln!t.-c1ion whdo \Ucillnt( f;~ryrl,ro:oon 011is ranil!r cw~. \~1.-rf11tuy
a CA.\tl'Rl..il. 1'.W., SLOA~. t:.A, ~ 11-Annun. PI\., 19~. Ob.serv¢.don$ on Pnffl:lllnltJg;. 10 41-15.
u, DONl\'rfL'· ,. ~ 1tljTOQJJ,,.. n. .-., 193!-, E:d~1tnct.1 u·P.perytbmiotm O\i't e-r\ JnJ)llttfSt' Juurm1f fJj\·j,1nm1UJ?·X'l1•nt'1•, l·J, 127·-l&Z.
,\~Cnc BulltWJ dt In s«f,:,1,rli' l'mhola;ol E..w1iqu~~. 28. 4:!3-i2&. Ill 111 MQJl,\l)f ,, ... I.- Pt.onA'-!P, c •• 1g?8 Fpt'f)'flllrJ:AHHI uvt..:. t. ~n)h.>J;ical
Ii PlJSL\P, I,\, l'l7~ Pf0lQ7..(M, /tr: DU-'.\"Nt If,\.. , A t \H.,1 U)., cd": /)fl1•mt'J d1.,gnMI> 01 lnf«rlon lw 1ht tndin.•<:t innnu11ofhu,rt>ec111 .tntlt,,>d,· ,.-.,.
tJ/Swm., ..;th edn \mt.~, fow·J.: Iowa ~fj\C tJm1,,'1:n1ty fin:"'\,., r,,,,,.,nm1·di:1,, und Pmu.JW,l.ogic.•. 29...~tJ7,...J IO
111 1,1t;J,.,. 1. . 1g.,:z. Oic, Fmphtlghchk\-'ir d~1 r1cnanulopto fur Am,plmmn 01•li ..u 1u'.\Joaw1. .L,. &. .l!lll'>O..,.,,.,,, 1q7a. F.pt:'')'lhrt>:oun twls. JJ. Pnt,11lcnc1.·
und l:)X'f';i'thro.-:,"''' 01•!, l'J.•utU'h•· T1uJ,){'m,,INfl..:1mrt:hl /tw1rhrlft ..it, ~1utUt~ in .,,Ju.,t!p in '\~cti.1 U\inl,': th~ mdit«t lnununoOuorl·~<'n.t
18-:>l. anrrt>udy lc"-1. tmJ,h•m>h,-di=fn mu/ P«r(t.s/tO!ngit1• 29. 111-314
19 tCCiGU:. .\..,.,. sr~i;n u.1. t40.t Studi1$0n EJ)f)l)lhto::,H>,1 infcttfon in ~ 1tni0tHm. ·'""· lt a1 o'n:\)W, •••• lff7U. F.prrytlzro:Qon rwl..~. m Thr c.ffc"ci
.
sh.11_11,. Journal o.fComwra1it..: Ptulrolo~·· i'·l. ·15--61 of lnlt'<"Ut>n on blood pH. conccnti.u10n')of p~ i\l\-'3lt.1ac~lC .ir..d
20 rcnm LL, Ll"\"l', H,t.. 'TOR:DCR'J. 8.J. &,C.Xolt.,'8\', \ .. 1 19;i- fntcrfr.r~n(t! i;luco,c In blood, h,tcm~tolui:r,.. n.nd un tonturrl'IU and ~upethnpo"'-'d
r0•1ttwt)µlm111 llnr itJft'<tion hi <hl·ci,. Tff,/jr'1tm1vli:ln wul
bttWC<!n ttnnpht.t.:mol\f, .md eprt)-thro1.onnosf\ m ,plt."nc:,t0m11.c-d <:.:m.h.·.
•½ozqnmn fournul af l 't•u.~nmtry J/eSt.,nrcll 18 5Sb--:;:-;9,
l'oru1JJUJto;:1t•, 2~ . .;..;3-4so •

21 R>UlUE ,. • PRoU.:oa·. L.1~~- \"etcr,n:u1· RC's(':trch fn~tinnc, -0 1,,~'-1 "-~ 11 · l'l~:? Tut oc<:urrenc~ or l!.flll)'lhmuum J)ltr, um splntN,
Onderstepoon. Unpubll,hod dom I 930 In Suuth \fr'ol"An ~\,inc. OndcNlt·t1tXJfl Jt>t1rt1UI nf i ·,w,umry
R,•.v-,u'Ci,. 23 :r6..
22 Fl!IF.OHOFJ-. J;:t., 1 ~ Ricl:emolcninJ'<•ltiunC?n (f:hr,1,J,t,, E:>..'1)111tP:J>t)n.
Hn~mabatto,ullJi btl H.tu..,tU.'ft'n In O..-ur1oc:-hl.:nd FotlMhrittt' :/411 .,a.J JOU\\ 11 o 19b;, l!ptr,·1hr<,;1,on ,,,,(1 Ult\"Cuon m ;t lamb. ·'"w /,-1lm1rl
\'eterfn/1rmwl1cln, ~upplem,int m/.t:111rnlbla11 fur\ .,r,·rlr:tJrmh.lI:.m. lS, \'._•rffl11nry/mmu,I 15 4-:'-48.
20-Ht09. -'~ l.r1 ro,, .._~. u J\J:\, ,.c 1973 F.p,rry·1hm~r, cre11,1..,ni: A '.Wannms
2l AlntDlfOt!JI. i;•• ono,iMut. \\ "'wuu·u \fit,.,, .. ,9:,. tnrd.1ioncn mi: c-k~rrc,11 111:<n,,.~:upc- ..,uJ), Jo;.1tnal "'//fmu,iw/c,w·• .s~. 661-lfr :1
£«9•1Jtrrr.nou a,,is bt:i Sch.ifen 1n ~ord1.h:u1sth1o,nU Btrlf11t/t wtd ..ati J.HULn.. ,., .... tu!orn. ,, M8J. Morpho!.u~l1; Jnllgunfc .uid pa1hogenlc
M!tncJ:ent1r n~ra1cild1t~ l\'rxJu111.Sclmft. a.;, 361-380. rh1uut1crbllr""-u( f.~·1h,o:.nm1 m•b: :Jnd F.1><-*-l)'tl,ro.:oc,11 t,,_.,,.rom
u GOnt£. n. ~ ~Cf\. 1.r .. 19r,. tcg;'J}t/m,v.l/o. /..flt•t;.·rluo:.1"'111U1d Am,otU.\'UI Jm,nwf oj \;•r.- 1 ittm)· R,,~<'lm·I, U ,188-llOO.
H<U:molHm1JnrUa. /Ir. >.Ra£,., Ir•.. t1:tJ.1. PCJta.rtlJt Pro:tJ:,o..-, XC'\\ \'ork. -ti' t,,,HMI u, ,.t• 1.. R1rtu,, '.\I., t!11il:i, U:u·moh:1-n6nl"l1ui,,h~ Fpt~t}·thtQ,:oor\o.:i~.
S:in Frantileo ~nd London: A<adomlc Press Grahan1t"!!osl~and Erhhch1n,1 ..... 1n w,
1'-M.\X, u. • 1n~i.<.. ~, .. 1 ~ .
25 1,UU..\SZ), T,M,. OOXl!Y. t>,L ... :,tOTT, (i.ff. ttJ87 ChitnJ{l.njt morphol08)" of /nf,Ytinm 8/oo,/ DJJtwt•s ,Jj.'ifm: mul .•\ii/mots,. \e\, ):'otk.. lundon:
/;JKrj1hro~l)On 011,. 1/e;carrlr in \'ocm,1<77) \cil'I•<~- 13, 611-91. .\C':lth.•mu; Pf"-">

26 K..\D.\~'I• \,, DI!. It.\.\'\. L, UJUUl!l \lO'\.. ,. \ ~ ,\;\t, \'\I, O,'-, t~Ko. ~g ~R.ff-<.,, '\;.ll 6l JICIU. ·"· 1~8 I ~1lft'''J. \J11mwl tJ/S)'~lt-•hutt:( &u:lt·r:o!OJ{\'.
E?4, t}'lhrutoc,)n~ls. t'n ''-'meH>~ tn la :\tgt'nun.\. R,•i•L"'' 11,• Mrdtt:ittn
0
R,1ltlmm1.o: \\1Jtutn:,:ind \\'tll.ln."--
1;•1crt,u>rln. 61. 9-19. 19 L\Nc;. 1.~1 • 1 LRFJtn. , ... M.,. ~n.,mu.$, r .J•• 1987, Uct~tion or :tmlbodit~ to
!l.i" f(U),\~f. A, GU01 IU..\to,r \.,\, \.X,ll ,~,.. 0 < ... r,\llU!U,!\. ti, ,,;,,,(,UI n, "·· Fp,.·l)•thm:.omt n,:•l., b~ tht• us.eof ,tn cn~nt~·Unkctrl tmrnuno,nrhcn1
Of.J(~UCEY \.~ C,\.,108. t.C.. s, r.o,:r~\LI/ JJl 11:l(h. l.. ~82 A CJ...'-("' of a,ss3\·. /t1..•,i-t."<Jtt'h fo t ~ttr/l,£11y Sdc1m•t·, ..t1, ~.J<J-25~.
app::ue!nt ~u.ppre~~lon of /11u:µft1J11w uwr,1µ,u,t~ lnfot1tor1 b~ so 11111...,:-.c.- P. t973. l'rc.-;ilcnc(" and significanc<: Clf tkk-borne- cJi.~.J~t~ o!
cpct)'thro1.oonos1~ tE1~1)lltro:»fJtt tegrtn!Hlr$; \'ctt"rmury· PuutJilol0£0• dume!o11, 4-<0<'k in northc:rn '-,;1g~rta. /n: \\ 11 m ... 1.~.H 1ed,J. Tlck-bon1i'
9,267-287. Oi"!M;'fl.,~ mu.I rhuir \ tt.·um,. l!dinbm~fl; Uni"-'\"r~t)' Pr1.'\.!o.
23 Ulll~RITI!l. K., \\'[S'r7.. ,. ~ 80UW:'\UN, w,,lCJ8.i. Ha.c;n:uoto.cththt' Bmundt
~,1 1 m LI 1011:-.~ 1.tr 19•.\0 Fpt·')1hro1oonoii~i an .:ohrcp. Attstrallun
ba dcr nkuum F.ptr)'tbro1,oono""-• dcr Sch\,oine. Bt1rllt11.•t mul \lm,1/11(11)' /rmmal 36. i60-:U,5
..,:a11rlh1t1cr Ti<t(lnz.ltel:~ i'/(JC1'tn..<chrlfr, 9i, 40 1 107.
~ M,,u,,tF•.,., ., 11r.t;t.t M, ,u •41ll "· H,c:., 197;,. ~.:&.anmg and u:.:nsmb')•an
29 11r....,~1~0., '1,w•• 19.:;t,;.. A.nlmnl Diseases tu Smull .1tfnca f~etorfa.. ~u1h
t lt"C"trun m1crol!(op~- o( Jf(J1•nu1J,n1(tmrlln,"11ts ,uuJ 6/lt'f']. tlrf<i:<1m1 m 1f~,
Afric.i: tcmml ,c"s A~ency. Lid. Anwrim,r /oun111I oj \'f·1t·rlt1my ~r11•nn,. 34. 11 ~&-t 201.
ao m,:m·. •.c. 1979. C.link31 ob,~r1ontlon< on ,•po~1hrt11M1u),!< Jnim,nl ,,f " ""u,1,, l'l8h, f:pN)thtollMiJW~"-1 ho~~\fo -1:n o\'L'f'!'IN "})U!UU\~
\1
th• \mf!rkm1 Vc-:.orlttltl)'.\11•dlt1J./ Ai$!H'lhtlnn. J74 r.,01..JJOJ.
v,w1Mk Vi•1,1rl,·a,m11Ylkm, :m. 5---ti7
31 Ho,-,:,u,, 19,a.). S~-mrtnrn;11ol~gi~ und Dtat:.1111,tik dttr
ff .
1-1 1'-1,\ut,:\I"\, u.f. ·.,,Lt. pt., l'ilt1. 80\inc b~lods, Thkk blood film, for
Ept1l)thcot.oonosP lu ferkelt•u.eus~rbf!Utto~I\ T1t-11Urt.:Jleh~ Um.sc'l:mr,
1hc-d.,teclion fflp,1ra.,1t.1t~mb Ari6ittJlltm Vt·wrimuy/our,u,J. li, •14-47.
39. ..;;4-4,;q.
~$ Nl\11tl\:, lt.J., llkl~J', l I • \\'(R1U D,tl. ,-,KJ,Nt,IUI, n,\., 1986..
32 uo1•F\1t\ \:!\ n.. $- M.AUt 1.1>. "'·· 1977, Au$bnu:h ~mer F.p,·r)1hru,oc:,no~· in
E1>nytluo:btJ1101·,,. •1ht.•,llth rf<.k for rl•.-.CArch ,h<.-ep fAh~rnct,
eioem Schwcmenas1bcs1and. Of11,1r1,~11emr:~l/tJi, ll'o,J1m.<1·/:rlj1. 8-1.
I.Jlht,rn11Jry Ar1imal St·l1•nt·,•. 36. SSL
·.9.
33 HtlfT'\J.\:"o'-.. n. -.t:ttMlU, iu.1 . • 11(.111 \lt\S~·HZI fl. t•.. 1981, F.'}1hrot11c-
:;6 't.\'WJ\, n.w.. ,~-.:-.uu.. " .. l)\llJJOW. ~.S", ~ \\11'1, O.J • 19BO, J:per,•:t,ro:OQn
ontibodfo., on pornnc cpcrythroroono ..ls. \ ·,,r.~1r.ary lmm11no!<1,g.·n11d tu·,s ""r<>rdrd in Ta.,;;m,miu in as)ocf.ttion witb Hein?. bQd;' d'l.!\·t?CTpmt'r.i
Atl<C1uhat1 Vttumn~1 /orimnl. S7 -10~
lmnumoµmlm/o;;.-. 2. 111-11!1.
34 HOW.uu>. G.w.. l!r.5 "fhe ~rimen1Jl 1r,\n\ml<.,it,no1 E~ ri110:.1m11 .:;; '1t1t\n11v. "' Iii':",; f.l11mmuiun ur P.pN)·rluo:r1m1 '\pp. (com m~.M:d
cw/J b) mo,quirn~, Pa,111/tQ/ogi•. ;i n."11 ln(rcuon, with Hahr.<,« ~md.o\na11fnsmn. J,mmmrifmal /011nwlft1r
/'om.<llblo,ry, I. 10·-108.
JS 11!)\JF~ u.,,.n .. 19&1 lnilinl dcvcJQpmt.•m or inJctttOn't \\'hh 8<1brt,in
b.1gemin11 /uu-nu,J ojProwu:x,l()J.I\', 8, 462.-166 .lfl ,m11 Ii.fr. t-... • ,.-nnD,'fl:U T' G l9:"~- lft>rrythro:oon fn ,rlnen
l'erkclcu,ugclb;1rieb IJl ,~,ddr111~chlnr,d. Pmkti,-,/:, r,,,,,1r..1 60
J6 11anE, IL\t~D.• lg62,f°~l)'lhrt)Z001' lcrgtftl(J(/("S,p "',... (Rid,c1t,Ulfesl. 66~.!.&S.
p;ir~siuc in c:it~c. /'r1TM1cofog;., s2. S:?f>-S32.
:It '-rrtt~ w.c,,. J!tr. liV<"f)1hroni.nftn"'h, m "ht.."<-'?· t)mlt.•h-t'1J.'IX>tl Jo.•,r,wl IJj
3; HO\~.,, K.'.\U),r 1971. The in.(cx:tlviryor T11,•iltna mut1111r. Fi>t't) 1ftrozc,on
\'rNnr:m;, ~!1•11,:t"mul.~,u,rw/ lutlt1Jlf)', 9. ~JO.
Ut!IJ)Y.mi .'Ind £ ttg,1,1wkt tu toht:cp m,d of l:i~r.-t)1ro:JH111 n: u tu atd<".
Brlti.Jh \ N{!Tinm,· J1m11wl, 127", 1-3, 60 , .. 11,.
\\'.O~ 1939, Thr lmmuml)' tn ~a.,1\,'0(tr. ()1uk,su1KMrt Jriunwl af

\,(wrlm>ry' X'1t•,1Nn11d ,tmmnl Jndm,,,,•. 13, 245-2H..l.

38 HloU. f.~.. onr,;c, \\'~8.. 1-IU. su... (.;HOU. S'.\'.4. ~lHF.h, \\'.1 .. 1'98~ Ou1btta1,.;,
h!'ld tre.annttn; of tpB)ihro1ocmn~i) h, ,,, fne in Tn.h,,1n. Joumnl ()j"tltt 6l S"lJTZ. ,., ..c.. l9.$0.. l:pl't)'lhl0:t:OOlti:b.s~ ln C8nlt.~. Oml.,11')tt1p<J<Jrt /OJmal of
C/1£11<# :;,,, l,·n·o/ \';r,rt,.,,,,,s.·1,•11c,,. 11. 211-220. ~·t1'"•rJnnry S4:U"t~t:md .-\mmnl l,ttftJJ.try. l--1. 9-28.
J4J lCHIJO. ( .• l-lomC\W,\, s.~KI~!. D. II XO!\:ltlll, 1 .. 198..: Sc.mnfni;:i.ud 6'2 :<r.rTL \\.O ... t~. Ep.•,,·tl:rozoon orfJ. lnfto\.,inn. Uull,•ttn CJDiu
uan.,mk-..lon ckl.'.'tmn mlcro,copic ob-scn'JJfon of /:Wl)'lllro.;oott o:1u. lr1tt'fnU1lo11t1I ti1:~ F"pi:oo:lt"$, iO. ~73--178.
580 ,ocnox YH~re: Rickensinl nnd t·hlamydial di~eascs

6-J ~Ert7- w.o., AU)X.\.''()tH. R.A, & DU 1orr. P,1,, 1934.. Ep,vrythf'()ZQQllOViS tsp. Epcr")1hrozoonosism ~'\\10C.
86 SPUTit.R. F,J.~ \\'ILU\.,,bO!IJ, R.L•• J.9SO,
nov.) 1nfcraon in sheep. o,u/.l!r$ft1J>00rt Joumal o[Vi!t~rln(JIJ' St:f(!Jfte mu/ Jouma/ ojtlzeAmcnro,1 Ve1erl11al')' ,l/odl(11/ ,ls;oct«rlon, 116. 360-36-1.
.Animal /rrdustl)', 3. 263-271.
8; <Strrro~. K,H .. \9~- EJ}<'l')1hruZ<JQ/1 Qv/s: The dlf!'erence !n curbohydrate
6a P-1., 1986.A modified indlr-t.~t lrrunununuu~nt
'.1cnaLI.!).. T.r." VP..~ me,obol,~rn ben,·tc:n 1nfe<:itd and uninfected sheep e:ythrocyt~...
as~y tor the detectfon or antibody to s,~,1hro:oon 01,fs lit sheep. ...,,1x,;,
3.J, .;49-;;s,
AustmliM Vnurl11al')'/011mal. 63, 15i-1;9
Ha c~,·. h.Jt ., 1917, Th'"· t!t'fect of b'pcr}'thro:oon oi·/6. infaction ott thc-
60 oa,. r.u. I!> A:,;('IAA, v.o., 1980. I lac1na1ologka1 ,1udic, on domc•11c gluto>c k\'cl ~nd •omc ~dd·b~~ f,mors In 1he vcnuu, blood of1hr:c11.
anlmal5 in ;111gori4. lV. Cllnfco-haezna1o!ogical l<>aturesofbovlne ,1t1,1mlln11 1',t<rinol')' Joumnl. 53, 471H81
11ypanosozninsl<. 1ho!lcfio51s. an•pla,mo<l>, c))<ll}~hro,.oono>i< and
89 SUTIOl', 11.11 .. 19;8. b)'<'':'th1b~*" (>l'is: 'l he dlfftrcnce ,n c.orbQh)dme
helmfmhla\-fS. z~ntmfbln11fnr \frltrlnlinnedt:t,,, 27, ;,a9-;9;_
me1aboll1m b<!iwcea hucoeted and uninfected sheep C:)"<hroq-.ei,
66 ODll[l..ll, n .. 1967. SOme ob~rv::uions on Ep.tr,,rltrtJzoou in.fottion In A11¥lrnlfnn Jorm:n/ vf /f.xµ.•rtm,mnl Hiolo1,;·n1ul ,\/edtrrrl S.:/r111Y, S4
non-splrne<:1omlzcd shccp :llld the dou,c:tion of thr par0$ltes 1111d tl1elr .;49-4;a
ontlbod!os b)' lmmunofluor,:>;set,nc•. ZJ.•,Jra/blat: for Bok1eriologic,
Part1Ji1tnkwult. !n/ekrlanskrankllitfhm und 1/ygf-tt:~·. Etsrt'Abr«,illms
,o wno:,., 11.u•. 19;8. Obf<!C\'tltlon 0111hc pnthologr of EJ,..,:.1/zro.:.oo11 ,wit
infr:et.on in ~hc.'C-p.·1\'tu· /.t"tllmul Vttrrhmry Journnl.16. 229-23n,
Orgfnnlt, A 203. 391-10 I.
67 0\'1'.l<ils. 1•• 1959. Smdlcs on l:p,l)'tlircrJJoi: 011/$ lnfcaion in ,!rccp. ,lcrn 91 ~vrru~. R,tr., 19;8, The pmhology o/ 1i/><1')'tllrtW>1111 o,,is, Ye::- Zmla111/
\ 'etl!rlnnrlti Semuli11aulcll. Supplc:om,,mt. 28. l· 148, v,:,ri,urry/1,umnl. 2;, 18.

63 O\'E=, J.. 19&;. linemogfobinufio In eperythroz,>onosi, ,n shttp. n,, 9:? ~-unos, a.u.•• a~;-g. Meet of Epc:rytfJro:.oon ovis and the ruduc-tl\·e
pottntlaJ ofsheep 1."'11-ihroqie.s. v,~reri11nl')1 Parasirolopy. S. 11-15.
l'ertrinor)'Rtt<>rd.12l. 277.
69 PO!lPISCl{fU. .\, "IIOPl'>V.~N•••• 1982, EJJi'l')'tilro:.OOrt $11/s In n~tura11y g3 stmo....:.11.tL, OL.\.Rl.l:STOS-. W.A1C• • cows;., G.H ..19r;. F.p,:r,•thro:oon
Infected pigs: Alight w,d electron mkro<cnpic ,1\ldy. \'ncrlttol')' ll'l'U>'Om- ti bJ<>od parasi1eofcnnlc. ;Veu, Zealmut Vttttrlm11)-Journal.
Pat/zf>logy. 19, 651-657. 25.8-9.
;o POT(;JJftl..l(.. Y.T.. 1989. \r(;h!rin:ll'y RCSC!;:tf'C!i lnstitUIC, Ondc~~cpoon. 9-i s.urros. a.it. ,1;.·10u,-. rc.t>., 1.973. Brpcdmcmtal Eperyt//rozoou ovir
Unpublished dnta. infoe:1t0n ofSht.."ep. .\°flti Zealand l"earrintnJ,•Jouma/, 2 • 100-166.
71 PUHXELL. R...h.. BROCKU:.SSV, E.\\.'. "\'()1J~G. f;.R.., 1976,. Epet)'rhfQ't,(J(Jn 9.; ·ruun..\SCt:, \l,, l'Jf.NMH, l,G,. UUJALKE., R.. f< \'.\IHl,\IH.E.~- , .. 196;. M(lrfn
wt11yo11/, ~ possible couseof nnneni.13 In !lrltifli c:itllf. 'n1( \ 'r1rri11nr;• nr11Jtlm rL, 0&1. rob~p. J:rau~;mu, Ulcnth.) Unman.ma P<"l,onous
n,oord. 98, 411. ;,tan, in South A1nco. /011malof11te .Soutlt Afrfrx:n Vew-lm,r:, ,\/ed/i;o/
1:: PUk;-:IU.l.. ILtt.., Ui\\'I~ u., fl0l..."1A.'i, \l,fl. ti \OU~G. IUh, 19$1. IO\'C'St;g;tfons i\s.iodntio1t. 38. S7-63.
on• &rb,,,!o ii,ollltod from Scoubh sheep. Plfra.</rolugy, 83. :W,-3;6. 96 ru(.))11. J.. 1966, A mkro-01gani)lm (tffet:1ing bo\·me platelet.,, F...fJ}'l!flc1mt1,
73 QUl)<t.\.'<, JJ.• 198,;. Su,p<.'Cled ~perytl11oxoon<»l1. In duJry cm,~ /rlsll 22.~5H59.
~•t~rf11a,;• Jormzol, 39, 2-2;. 97 nt0'\11. f. 4,: r~,~s.K.,~f..~. n., 1980. Antigemc t1on-reh11tor;Ship o( l\\'O
;.1 aos.~Kk,\ss.. C.J•• 'l,Jss. f),\I, & 1i-;ssE. c.w.• 198,.:t. Chtoncn·.acycline as t1 bo\One ep•~·tluo,oa aeinons1ratcd by the 11nmunolluor<-s~c111 mo1hbd
tre-atm~nt against the~m·,c:. of epc~·throxoonos.isu.n.sow ptoduetl\'IL~·. Aero \ 1.•terin«ria Sc'<mdi11n..uim. 21, G99-i0 l.
Proci!i.~1ingto/1/Je Efglllll l111~mmio1wl Pitl/erorlntuy Soeltl)' C011gn'$$. 98 ruo,u. J. ~ ,·o~ no~-wosu-. f':.n., J967. UILru$LtUCtut~ ur a mtcro-orga.nJwn
27-31Augu<1198~. (;hen,. llcliium. assoc!J11ed "ith bo,fac platelets. F.xp,,rlcnrta, 23, 1-;
;s scwwxu. v.• 19:za. E~()·tllfo:.oa,, cocwuff!.s emc aeue. dun-h 99 un.i;s)ERG, G.. 1965. :-otC$ sur lco Epel')1l,ro:0011 do bo•ins •
Sptencctomie ak1ivierbarc OauNmrek'tjon dc-r ,,·elsse. Mous-, A7iniKlw Mad~!!!t\Car. Re1•11t<l'Elcr'tlg.,t de MM,,cin~ Wti!rinol~d~ Pay,
1Vorlwn$cltrij1. 7, 18i>l-l855. 1'1qpica11x. 18, 73-131.
;& .$HfHrn, n .. 1972.. otagnn•ds or Eperytlrro:.0011 o,,Jsinfocuondl1tJ'trallan U.')() UIUIS!fRC:, r,., 1967. F.pttiytllt():.()011 womu. n.sp. tRiCkt'tt.SiA!t-"il,
\'tUttfnary Juurm,I. 48. 128.
troM,m• espl:ce d't~ryrllrouxm dos bovlns ~lad•ga..,a:. R~m•
77 \H.J.::.tufF, u .. 19;-J... The ,,ffcct of SplrOlf)'P:'"m nn Epttry.·thrO~l)()tf od~ tn d'El<'l"S'l 'l rfe ,\·/i!drri,r~ Wt.<rinniredt.< Prrys Trupi,011.~. 20. 563-569.
~hcep. Tlzt Vtr~ri,uuy Rl/rlJrd. 93, 388-269.
101 \\IE,\:~!XG, T.H.. SC')U\\'E!\":\. t.;., ~CllOTMi\!'I.' , 1\.J.K., \!f:R:,,,'00,. J. & .I.WART, O••
;a sa•m~. o.. 19,8. Th~ paihology of Ept>l}11ir0J:0011 ovls..Ww Ztnla11d 1g7-4. Tht, crTra of E,~rythro=oon w,m,t·<mi un tht: glut'O~' !<:\'<.-! a.n-d
Vkl•m·nai,• Journal. 26. 315. acid-blli-'balontc of bo,i,,. blood in vu:oand fr• :•itro. Tlfd«lzrifr
;9 WtKlf... u .. CL\l'Jt, K,H, k- Kt.10. M.A .. 1966. J;"'jwr;.·11,ro~rm ,wl~fnfe<:·tion in Dicrg,11t.,,,k1111tlt. 99, 13f>-142.
South ,\ustmlla. A11Jrralia11 Vcwr/11111')' Jc11ma/, 42, J6$-l i6. 102 ~,..\OUR\, 1.,#..,.
tu.~~1 t.. r:.iJ., 1~0.5. E.rychtc.)C")rtC membran~ alterntfom~
80 SHFRlFF. o. "'<J~:musc:. >1.c.. l96!J, The andglobulin !Coombs) 1es1 in 3S-')Or:i.1.1L·d \\ ilh tho illtnc:h.m~nl nud repllcailon of Eperythto:oon suis:.\
F.~·tiaro.:mm ov/sinfec:tJnn in sheep. All$"llllnn \'rmrfna')'/oumt1l, ,:s. Hght :i.nd cJ(!etton mlcrosc-oplcsm<l)'. t~ttwrinn,:.•Ptt1/:ologJ•. 22, 16+-l iO.
505-50i 103 ?...AOIAR\-, J.F. & ~,1rn,. A.it.. 198·5. Expcrim~ntnl porcine
8t ~lMffO:>:, c.F. • LO\'U. J.!'r:,, J9iO, Fine structure of Hcwmob11ttonella bovU ijJi•l')~hro1.0on.o,,, T-lymph<,cytt suppression :ut<l nlisdlrttted
1n blood and Jivorof splnn.,.,1omized c>l,~s. ,ln,,r/ct111/011mal of lmmunt ~po.nscs.Amerfcau /ourne,J tJ/Ve1ttrlnn,y Ruto1':'h. 46.
Vett~n)ihf}' R4'.<lVUtlt, 31. 2.U'\-2:3 L 82J-830.
82 StSJ;. o.s.. COi£. J.8. t, ~·URNEU., A.R...
1980, Scrolo$iC inddtnc~ or 1()-J z-1...:x-. ~. \,,,Jf~;;J: o,t\'.,.. '()(')Bsn~. A.W,. 19'S3- E.ffe<t of ,,~ey'throiocmosis
•P•l'}'throzoonos!s in Genrgla ,,vine. 2311/ ,lw:uat Proc..,1/iu1,-1. College on SO\\' producci\it)·. Joumt1..l of1he Alm•rletm Vttc'rinary• ~\lrdlml
of\'eionnory Medicine. Unl\'el'$11y of Georgia. ,\lc<O<tnti~II. 182. 3~371.
sa s>1m<, ,.,.. "1t11Hs, R., 1975. An Indirect hemoggturlnotlon rest for 1hr tn.s zw.,,n o.. L.C.tlltAX-t.. 1•. "\'A,'' von~Tf.."1:80SC.11. c.,_,.11.\". 1969. SrudfCS on
dJ~o"' ofEpetytltru:tKJn sui.sUlfoct.ion in swine Amt'da,,, IDurn<tl of an £,-Rf)'tltro::oon i'H:SOciAtt9d wnh bovtn~ 1hromhocytns. Zt•mmlblmtfur
\ 'Ntrl'1:ar;•Rese,m-lr. 36. 1319-1321. Baktl'·r(ologft•. Ptlfll..tft41JJJ.111uW. bift1ktlo111krt1nkht!lt1111 !t't:d U~,1tlt1III!. Er.ere
80 ;,punu. e.1.. 19so. EJl<'rythro:0011 mis MP, and Ep;,ry~hro:t,011 pori,11111 Abteik1n~ Origin>!•. A 210. lr>-105.
n,<p.. t\,o m:w blood p:trJsit.c-s or ~wine.Scft•tt1,,Y. Ill. i'il--514. 106 ZWAfrr. o .. ~Cl10T)1A~. A,J,H. & S"l1C.W l ' ff. A,l,M .. 1970. Mo~'3.bohr changci.
as SPUTTEJt. E,J,.. J.$53. Epcry1hro~oo11 pon·1J•n a fihcro.bfe blood p3ttt~hc c,r during J;)N!fj.'iltro:obn fnfl!-c1lon in c:sule.. Httiit.>tlft:lt in \imTl11nry~f'i1J.nr-..t
!l.tdn~• .Vorura. 172, 40 . 11.105-lll.
 45
Bovine haemobartonellosis*
Synonym: llaemolx1rro11e/la /Jot•is infection

FT POTGJETER

Introduction species, which infect the mou~e. cat and dog respecti\•ely. are
curremlr included in the genus Haemobar1011el/a," The
Bovine haemobanonellosis is caused by Haem0Il(lrto11ella classllica1ion of mail) of 1hese organil.111s remains in cioubl.;
/Jo11is, which ls a rickcttsial-like organism that parasirizcs Extensive reviews h;we been published on eperythro-
erythrocytes. Most infections are not clinically apparent and zoonosis and haemobartoneHosis in which the aetiology
disease, characterized by fever and anaemia, is seen almost and other aspeclS of these infecrions are compared.2-1
invariably on ly in splencc1omized cattle. t\n elucidation of the complete life cycle of Haemobar-
Uaemolmrtonella bo11is was described by Do11aden and ro11etla spp. is still awaited.
Lestoquard in 1934. as cited by Gothe and Kreier. 2 and ib
ofter) encountered in blood smears of splenec1omfzed cattle
Epidemiology
ac the Onders1epoon Veterinary lnstilme in South Africa. 8
J\llhough 1he majorily of /laenw/;ar1011e/la ~pp. infoc1iot1~ Hal!mo/;art1.111elltt /;m,i, is p1ouably widely distribuwd ln
which occur in several domes!ic and wild animal species are southern Africa. fl is believed that live-blood vaccines, such
non-pathogenic. Haemolmrwnella fells. the cause of feline as those for anaplasmosis and babesiosis. produced from
infectious anaemia in the domest ic car, ls of considerable clinicall)' inapparent Jf. bo11is-infec1ed canle may be inad-
veterinary lmponance in southern Africa. vertently rcsponsibll' for rhc widespread distribution of
H. bovis in South :\frica.8
f/ae1110/Jano11ella bovis infection in cattle is not re-
Aetiology and life cycle
garded as of much clinical or economic significance and
Haemobarronella spp. are pleomorphic. rod· co coccoid· bas only a nuisance value in splenectoml7.ed donors of
shaped parasite~, 0,2 to 0.-15 µm in diameter (Figures 45. 1 live-blood vaccines and in experimental animals. lnappar-
and 45.2), belonging ro che order Rickensiales. fnmlly Ana· Cl1l Hat>mobr1rro11e/la Infections may result in aberrations
plasmacaceae. The organisms occur in indentations of the in e~1>erimcl1lal data and arc probably due 10 the acriva·
surface membrane of erythrocytes. and are more closely 1ion of latent infections.
associated with chis membrane rhan are £µe,yrhro:;oon spp. Littll' is known about the uamral mmsmission of
llaemo/Jar1011e/la /Jo1•i$ is separated from che plasmalemma /-1. bovis. but it is thought that rhe pa rasite, as is the case with
by a definite space.• The parasites are sometimes arranged Epe1y1hro:0011 spp.. mighc be arthropod -borne. lnfec1ion
ln chains. groups or pairs and have a limiring membrane. can easily b«:: uansmiued iauogenicaJl r by iuoculalion of in-
but no cell waU or nucleus. They arc rarely encountered free fected blood or organ suspensions. Unimentional mechani-
in the plasma. cal rransmission by all parenteral routes is possible.
Haenzolmrro11el/a stain well with Giemsa s tain and arc:>
Gram-negati\·e.
Pathogenesis, clinical signs and path ology
T~e genus /laemobar1011ella was established b, 1\-aer and
Weinman in 1939. Although 27 species of liaemobarwn.el/a No uhrastruclural evidence has been found to show that
have been named, and iumatned Haemabartonella spp. infec- haemolysis or ~,hroc)1es is associated with high parasi-
tions found in a further 11 species of animals,! only three taemias of H. bo11is in splenec1omized c:alvcs.9 Packed cell

• For tl-c:ent chang1..~ it\ the nomenclature or th~seorg;111bm"'. tdt't to the intro<foction to St:!ctlon 3. Rid.t:rsi!ll and C'hlnm)'\J1rJI discnso.s
581
 •

Figure 45.l f1e.;11on micrograph of a se:t on througl1 ar. er11hrocv1e Figure 45.2 L•Sht rnicrosco;>ic phOtograph or a Giemse-sta• 0clKI blond
aniected ,.. ,th Hilemobanone/la /:Jov,s Note tne Qaras;1e poslt1oneo 10 the smear ei a •alf •n!ec:ed •:mh Haemabanone/111 bov.sh, t 00,'.l/
1rn1antation ,n lllS cell and !he intimate asso: anon llet·:,-ae.~ tne para!lta
and e~/\ll!OC\~e memb•ane

volumes show a significant decrea,e after whith the parasi- anaemic caull'. <!specially if they have receml~ been sple-
1aemia subs.Ides, and It is conc:luded 1hm phagocyiosl, ls re- nectomized or immunosuppressed. Cllnkal ,ign$ are only
sponsible for the annemia. A decrease of blood glucose has evident in animals with advanced anaemia. !fthe <:xamina-
beet\ found 10 orcur c<.mcornitamly with the parni;icacmins rion of blood smears fails to confirm the diagno~is, blood of
in splenectomized calve~.b F.rythrocytes para~irized by an th1: ~uspected animal may bC! inoculatl.'d into ~u~ceptihle
!·!. bol'is-!ike organism show a l.69-fold increase in glucose splencctomized animals.
metabolism. 7 1Jaemol1t1rro11e/la /101,is has not been observed Depending on the level of parasitaemia, Ii (101•is can be
to be pathogenic in Australia I or South J\frlca.3 confused \\ilh 411aplasm11 spp. infections on blood smear
•.>,pan from causing mild anaemia and fever in splenecco- examination. pamcularl~· ofspleneccomized animals.
m!zed calves. Ii. l;o11is genernll) appear!. 10 be non-pa1ho-
genic.6· Y Kreier and Ristic 4 cite two reports oi natural
Control
outbreaks of severe disease caused by H. bouis, bu I rhese are
regarded as unique. Growth oi Hc1emobarro11ella spp. ls inhibited by arsenicals
Little information on the pathology of H. bo,,is infoction and m.yterracyclines. but not by penicillin or streptomycin.
is available. 1· 1;, 9 Gothe and Kreier' list drugs which have been tcsrcd
for acrhity agains1 epe1y1hro1.oono~i~ and haemoharro-
ncllu~is. Fur rnurt· i11fumm1Jun on the :,pt;;c;lfic trcaHncnt
Diagnosis and differential diagnosis
of hacmobartoncllosis. refer to Chapter 44: Eperythro-
A diagnosis of H. bo11is infection should be considered in i:oonosis.

References
c.\Ub\\. 1_1 l!Jd.; ·\nlnwl 1teulth in .\u..,trJll.t \otuin'-" S.. Pro1<1:0(2/ amt 1 t:RtH,, .... t.- ltt>:ll, 1.c. i~4, &·t,:.')·'s ,\/llntllll ,>j .,~f1•t1N:J(C BllCh'1'W(~gJ·.
llfrJ.:.::mal t)bn1.1,e, C.nnbttrn: .·\U)Oali"n Bu.t\.'au o: Anunal H~,hh. Bothimorc: \\ tman , Jtul \t"ilk11\f!.
.\u,tr.,llanGo,ernm~n1 l'l1bllshini;~,,n·,.,... pp ~211 6 LO\'l Ji"·"' ,1cl\\t'.\. 1 c,_ 11,;2. lI~·puW.)ccntl."l M-~OC1au~d \,ith
2' GOTIU fl.I. ):;kEl(.Ft,J.I'., 19r;.•1t&,'Y'/1tfrmrlfr.. l:prrytl11tl:<JOll3mJ. Hut•111dttu:01h•llt1·1lk(! lnf«titm fn ~ph.·h(!ciomu~..d rah,'t"t .·t1m•nr,1,,
Hn,•m11llmtontlli,. 111~ -..;11urn. ,.,.• (\'·d J. Parr&!iltr,· ProJO:O<J. ~cw York: Jcmrm,. of\ i"lt.•ri1U1()' Fl<'ffl1rrh. 1:t 2.tmi-2089
Acatkmlt l'IL'» 1 LO\'t.1., .• wn."iO'.\
. ,: fJ.. \tOMl:S,I , .... WJ\'C.\l.. G.,197:' '.\h::aboU,Tnnf
,
l imr.tr:tt. J,t•.~ RbTIL. ~1 .. 1963, Mmphufogl(, ,1.nUgcntr nnd p..,thtJJ;t!nh: 1tiuros~ :n Ha,•111ol"'rm11diu•lik,· ln(l'Cled ~')1hroC)WS :n ~plcm,c-
ch;uacter111ucsaf f:1H'ri1/1m;.DQn m·/11 :md EJ~'r,1i11~om, u'i.·>tyom . tnmwu:t t':;\J\'c'!' ,·\ntum•tm Jm,r,111/ r,f\f,•t.:rlnnry1 Rt1.);t'(lfti1. 38. ;39-j.; I,
..\mtn,,m Jou;,1111 t.if Vrwrfunr;.• R,•,'-,vud,, ?·f, >t88-430. a POT<Hh;tiJt. t T 1989. Onde~tcpoort \"rterinat)' ln-.ittutf. Unpubh~hcd
4 KflUfR. ,.~. R!~'rlf., M,. 19C..-8.11:.ir.mub.::.rtond!n"fs._ f.pcf1,thru• Qbj('f\.\llor,.,._.
:r.oonu""is.. GnhnrnrJI(')'\I"' wul Erhtkhl<H1t~~ In: ,,·l!.1s,1A, . n 1,- "'"-1Jr,. ,, .. !: ,t,1r:i.o,. ,_,.."' 10\T r , tg:,;,. rtneriuucturc of Jlnc"mt1!mr1011tllti 1101•/1
!eds;. /11fc1.:11ou1,R/r,c,d Ufrta.~·> n/Mtm rm,I .lc,:fmnl:s. \'('\\ Yar~. 111 blood :,ml liver 01 ,plcnccwmll!.l'CI ~,'t',. A,m•rlc,111 Jor,rna/ of
M~cl•n11c PIL'S> l t'tt>rln"'Y Hrtit1rth :s ~, 22>-.?JI
 46
Potomac horse fever*
Synonyms: Equine monoc.ytic chrlichiosi~. equine ehrlichial colitis,
equine ehrlichial abortion

I E PALMJ:R

Introduction cu.hurc media. the continuous murinc macrophage cell line

P388D I is frequencly used for isolation and propal(ation.
Potomac horse fever is an acute tmterocolitil> of equids The original )1aryland Isolate grow~ read ii}' in this cell line.
caused by a group of closely related !ntrnccllular-oocurring in iliallr appearing as a clusler of singly occurring elemen-
ehrlichias collectively known as Eltrlidiia risticii. first recog- tary bodies follow«:>d by the development of initial bodies
nized In 1979 along the Potomac Rh·er in ~laryland. l}SA.29 and later forming morulae (mature inclusions}. Heavily in-
Ehrlichia n~~ricii infecrion in horses has been referred to as fected cells evenruallydisincegrate, releasing loosely packed
Potomac horse fever. equine monocytic ehrllchiosis. and groups of orgamsms held together by cytoplasmic Slroma.,;,,
equine ehrlichial colitis. It has been recognized 10 occur TI1ere is considerable biologioal diversity of £. risricil
throughout :-onh America. The original term, 'Potomac isolates from clinical cases of Pmomac horse fever. ;1,torpho-
horse fever·. was coined by a television reporter covering the logically. some tbnn large cytoplasmic morulae (inclusions).
original epidemic along the Potomac Rl,:er. ,\!though it is \\'hcreas others form small morulae or are indMdually dis-
the lea,t descriptive term. its \~~de acceptance and its high persed in the c~1oplasm in murine P388DI cells.I I Similarly.
level of recognition ensures that it will remain the mosr pallerns of antigenic proteins may differ considerably
common term for the disease. Equine ehrlichial colitis is a between isolates. 11 The sequences of the !GS rR:--A genes of
more descriptive term for the oraginally described syndrome isolates ma; differ bet\,'een each other more than from the
of fever. anorexia. depression and diarrhoea. Even mild next mosl closely reiaced Ehrlichia sp .. E. sennetsu. It is e\i·
cases nm manifosring diarrhoea hal'e e,idence of colitis. <lem that Potomac horse fever is not caused by one Ehrlichia
Equine ehrlichial abortion is an appropriate umn for the sp. bul rather by several closely relnted lmt distinct J:.1rrlichia
abortion syndrome caused by E. risticii. spp. 11I Thi!> amigenic variaiion of isolates causing the same
clinical disease has resulted in incomplete protection from
1he first generation of vaccines which all originated from the
Aetiology
same type suain of E. risticii.
The causative organism of Potomac horse fe\·er is a member
of the tribe Ehrlichieae.:?3 Based on nucleorlde sequence. tl1e
Epide,niology
genus Ehrlichia is phylogenetically incoherent. Elirlichia
ris11cii is most closely related to £ilrlicl1ia se1111e1su and Potomac horse fever is confined to North America where it has
Neoric/.:errsia /11Jlmi1111toeca (levels of sequence similarity. l>een reported in a wide variecy of geographic regions in most
> 95 per cent).ss Although only definith·ely proven for N. USA states and Canada. Surveys in the USA ha,·e shom1 that 16
ltelmi111iloeca. it is likely that these three organisll)s also to 33 per cent of hOr$es with no history of illness have amibod·
share the unique property of being the only known obligate !es to E. risrici/ 18 3~· n5 and that these are seasonal lluciuations
intracellular bacteria that a re Lransmiued ,ia a helmimh with the highesc nwnber of seropositive horsos and highest
vector. clinical occurrence in July. August and September. 16· l!l Over a
Although £. rlsticii grows readily in a munbcr of tissue five-year peri()d 70 per cent of 900 clini~l cases recorded i11
culture lines as long as antimicrobials are not used in the Maryland occurred during the same rnomhs. 53

• Fo~ recent ch:inge, ln 1he nomencla1ure or these organism$. refer 10 1he intrcxh1cl!on to Sl!ctlon ~- Rlckeaslal and ch\;lrn:;dfal dl=s..'S
583
584 •«:1,,:,~ n mrr: Rickcttsinl and chlam) dial disease.,

Clinical disease shows an unusual sporadi(; panern with infeccitln produces clinical disease which is identical co
a low prevalence rate (< 5 per cent) on any one farm despite !hat seen when 1he organism is given inira,·enously.H If
its frequem occurrence in an endemic art>a. In fact, when horses arc exposed orally to a sufficient number or \'iable
multiple cases occur on a large farm. the pauern is also u~m- ehrlichias. infection \\ill occur. Large numbers of E. risticii
porally (within the senson of the discaseJ and geographically are shed into 1he lumen of the colon in exfoliating epithe-
(among the pastures and burn5 on lhe furm1 sporadic. Occo lioJ cells.ti~ Th1.1s large numbers of chrlichi:is arc present in
sionally an epidemic form occurs which is characterized by the faeces of lnfcc1cd horscs. 6• 38 l lowenir, direct comact
a h~h attack rate (20 to 50 per cent) on an mdividual farm. does not result in 1ransmlssion. 14 so ii seems unlikely that
The disease ma)' be concentrated on a particular fann or contaminated faeces play a direct role in the transmission
race track, resulting in an ou1bre::ik simultaneously involv- of the disease. Tndirec1 oral transmission may occur
ing a large number of horses. 1 he reason for this epidemio- through concenr ration of the ehrllchlas in a vector.
logical varialion is unkno\,~1. 15· 48 ,V11orii:ke11sin ht'fminthoem and ,Veorickl't1sinelokomi11icn.
The mode of transmission has been extcnsinily sn1died two ocher members of the tribe Ehrlic-hicac whfch share
but significant questions comlnud to exist. Direct contact common an1igens'q and ONA homology55 IVirh £. riscfcii,
transmission dbes not occur. 14 Tne seasonal occurrence are transmitted through ingesticm orinfcc1ed helminths.22•
leads to early investigation of a possible arthropod vector. n The metacercaria of the salmon fluke (.\'a11oph,1'<'rut
Common \·eccors of ocher ehrllchlal age111s are ticks.r,-:- The st1/111i11rolt1J carries these pathogens. Dogs are Infected by
only adult tick found feed,ing on horses along the Potomac ingestion offluke-infestedsalmon or trout. Similar()·. E. ri,t·
River during lhe original epidemic in the early !980~ was ricii may be conce111rated in infected helminths carried by
Dermacemor 1•arl11bilis.9• ;i fhe lar\'al and nymphal smges anhropods which are inadvenentl\' ingested. Elirlic/ii(I
of this cick feed on small ground mammals such as the se1111,•1s11. a close!~ related paihogen or h1rmans. has been
white-footed mouse (Perom.vsc11s leucopu,<)."I :>.lice are suspected of being 1ransmit1ed by ingestion of a fish nema-
highl)' susceptible 10 experimt'mal E. risricii lnfection. 24 code.2B The im·olvement of an arthropod vector or
HowC\·er. anempt.~ 10 transmit E. risricii to horses using helminth would be consistent with the seasonality of the
field-captured adult ticks (D. V(lriabilis) from endemic farms disease.
failed.:-:? Funhermore, larval and n)mphal stages of D. tiari- Recently a reservoir of E. risth:ii has been identified in
abills fed on ehrlichacmic mice did not rran,mit the organ- trematode cercariae 1virgulate cercariae) pnrasitlzing fresh-
Ism to other mice or to ponies.~'1• .su and "hite-footcd mice \\'ater stream snails and aquatic insects in two geographi-
infested 1vith immature D. l'ariabilis and lil•ing in endemic cally distinct areas of the USA. 2• 10· 25 • 56 · .;s The trematodes
areas of the disease have con~istently been found to he which can become infec1ecl with £. risricli appear to ha,·e a
seronega1ive. 9 Dogs. 1he primarv host of adult D. l'(lfiflbilis. broad Intermediate host range. ha,'ing been found In abun-
are unifom1ly seroncgativc on endemic farms53 despite dance in adult and immature forms of the follo\\;ng aquatic
being susceptible to infectior,.138 Demwce11wr uaria/Jilis is insects: caddisflies (Tricl1011rc,ra). mayflic, (Ephemerop1eml.
not the vector of £. ri$Iit:ii and auempts 10 transmit the damselflies COdo/1(1/(I, 7.ygoptera). dragonflies (Orlo11ara.
pathogen with other ticks including RhipiceplU1l11s sm1- ;\11isoprem). and sconemes (Plecopreml. iu In one swdy. the
g11i11e11s, lxod/!$ scap111rlris. and Amblyomma mnerict11111m prevalence or£ ri.<iicii wa~ 32 per cent in 13 of 17 aquatic
have also failed. 20 spt'cies of Insect~ acting as hosts for the trematodes. tn
Despite the \\'ide e:q>erimemal host susceptibiliry or R. ris- The discovery of an aquatic reservoir is not surprising con-
ridi.1 3. 2 •1 6jf, . : non-equine mammals in endemic area.< show ;i sidering the early observations that 1he disease outbreaks
surprisingly low fre<1ucnC)· of scroconversion.a· rn "'1 · 71 One centred along majorrivcrs. 1HThis finding could easily explain
investigation revealed that a high percemage of cats on some theobser,ation that rhe dhease appears to be associated \,ith
endemic fanns were seropositive.33 but tran~mission studie:. areas and nm ho~es in die areas. One of the original index
using the cat flea (Ctenocl'phalides felis) and the chigger ml1e farms along lhe Potomac !liver was abandoned and left with·
(Neorrombic11/a w/umom). both l'Ommon eccoparasites of t'3ts cnn horses for several years becuuse of the high pre\•alence of
on endemic fam1s, failed "hen E. ris;idi-infected hor.;es or the disease. It was then repopulated with a new herd, Poto-
mice \\'ere used as donors and mice were used as reeipien1$. 71 m(IC horse fever re-emerged on the fan11 during the first
The ehrlichaemia that occurs during the acute ~'tage of spring thereafter. a high mmck rate being recorded in rhe
the disease makes transmlssJon b)' haemawphagous mes newly introduced horses.
possible. ThP rolP of aciult stahll' Oil'< (S1mmn1~ ralri1rn11.~) The roru111c1ion hetwecn 1hi>aquati<' rl'<Pr\'oir and 1ran•-
has been investigated:8 despite demonstrating ~hrlichaemia mission to horses ha-snot been clearlycharacteriz;cd. Trans-
in donor horses. reci piem horses did 1101 seroconvert or de- mission attempts using naturallr infected trematodes
velop signs of Po1omac horse fever and reco\'ery of£. risticii through skin contact with wa1er harbouring E. r lsrlcii-
from the flies was not successful. infet·ted cercariae. through ingestion of ll'ater harbouring
All the e\'idence indicates tha: blood-sucking anhro- c. risricii-infecied cercariae or through lngcsrion of different
pods arc no1 Teliponsiblc for transmiuing £. risricii. Oral aquatic insects - caddiSO) lan,'l!e (Limnephllidael. adult
 P<>rc,mac hor5<' fever 585

caddisflies (l,eptoceridae. Llmnephilidael. n)1llph and adult system. probably rhrough modification by inflammatory
mayflies tHeptageniidea) and s1onefly n)1nphs {Perlodidael cytokines. Whedier or 1101 the immune depression is clink-
- harbouring f.. ri.~1icii,infec1ed me1acercariae were only 3lly imponant in horses is unclear. Infected mice shO\\
successful in one horse which received large numbers of evidence of significant depression of both humoral and eel·
adult caddisflies. Two other horses receiving fewer adult lular immunit~-62 and develop lymphoid depiction in lym-
cnddisfl ies did not dt-11elop disease, seroconvert. <!evelop phoid 1issues.!il Although infected horses have 3 1:ransien1
ehrlichaemia or shed ehrlichlas in their faetes.33 Consider· denease in antibody production 10 :md lymphoc)1C func-
ing chc susceptibility of horses to oral infections. it is sur- cion.6a no histologlc lesions develop in the l)111phoid
prising that it was not easier to transmit the organism S)'incm.12· 63 In cicke11~lal infections amibodies can be pro-
thrnugh ingestion of the rrcmatodes.'14 The failure ro do so ls tective when lhe) block the pathogen's anachmem 10 or
probably associated with Jack oJ sufficie111 viable ehrlichia~. penetration oihostcells.21 Antl-l:'. risricii equine lgG inhibits
How horses would iJl!,>eSt sufficient number, of these £. risricii imemalization and imerfert'S with lhe me1abolic
aqu,1tic Insects to re$ult in Infection has not been explored. act ivity of£. risticll. rendering them incapable of prolifera·
Caddisfliesspend rheirlife primarily on tlie water orin vcge· tion In host cells and resulting in the evemual destruction of
tation near water. They may be attracted by lights at night. 1he organisms.~6 Op~oni7.a1ion of £. risridi with ami-£.
When the ecosystem of horses and caddisflies. or other risticil serum renders J:: risticif more susceptible 10 macro-
aquatic Insect~. would allow for transmh~fun is unckar. r\n· phage.: des1ru.::1io11.e.1 :Vlice c,,n be p.i~>ivcly protected frum
other issue that needs resolu tion is the observation that E. risticii infection b) antibody transrer. 26 However. ehrli·
cases of Potomac horse fever can occur In horses at signifi- chias can be isolated from animals in the presence of a rising
cant distances from major waterways as well as in horses antibody titre,11 and presence of antibodies dOf.'$ not always
that have no apparenr access to streams or other aquatic en· correlate with dearance of ehrlichias or \\~th presence of
vlronmems. ii. ·15• ·18 protecllve immuniry.00 Effective neutralizing antibodies are
The faecal shedding of/:.'. ristic(i. which appears to be a the important subset of t.he general antibody response to
consistent finding In infected horses.bis an ideal \\"llrto seed ehrlichial infections. In horses. clinical disease occurs in the
aqua1ic emironmems "~th the organism, and introduction face of rising antibody titres. Modulation of the immune
of recently infected horses could conceivabl) convert a prc- response br the pathogen so that 1he initial antibodies arc
\iously disease.free area into one potential!) endemic for not pro1ccllve may be an important aspect of the pathogen·s
the disease, if aquatic environments do prove to harbour the virulence.
source of endemic disease in horses. It Is apparent that, al-
though a freshwater reservoir Is probably hnportant in the
Clinical signs
epidemiology of£ risiicii. many questions have been left
unanswered. There is considerable \·ariation in clinical manifestations of
Potomac ho~e fe\·er. The clinical signs Include feyer.
depression, anorexia. ileus. colic. diarrhoea and lamini·
Pathogenesis
tis. 1<u~Aa. 51 • 52• R9 11 is a common misconception 1ha1 typical
Although the mode of transmission is unkno\\11. it is clear cases show most of these signs. On 1he contrary. an> combi-
that within hours of infection E. risricii can be found in a nation of these signs may be present. Common 10 all t'ases are
subpopulation of blood monocytes. Although the pathogen the clinical manifes1a1ions of colitis which do not al•,,·a}'S r<.>·
is readily phagoc~1i1.ed by monocytes,s-: it appears 10 elude suh in colic or diarrhoea. Mild colitis can resul! in depression
the hest's defence mechanisms by lnhibiting lysosomal and anore>.ia. \,ith or \,ithout fe\'cr. Normally fom1ed faeces
fusion with phagosomes.lQ The ehrlichaemia persists ma>• be passed \\ithout signs of colic being present. Cases
throughout and beyond the clinical period. The pathogen with equivocal ~igns of gastrointestinal tract involYement
has a predilection for the mucosa or the caecum and large typically have. on careful auscultation of the abdomen, a re-
colon but is occasionally found ill the mucosa of 1he markable decrease in borborygmnl sounds, confimiing gas·
jejunum and small colon. M rmesrinal epithelial cells, mast 1roinws1inal disease and long periods of silence broken by
cells and macrophages are the targecs of infection. Lesions shon bursts of loud. high·pitchcd sounds produced by gas
are confined to the imestimll tract. 12 Th~ inflammatory re- passing through a tense intestine with a g,1s-tluid interface
sponse 10 c. risri.-iiinfection. which appears to ,,ary in !men· will be revealed. Simultaneous ausculta1ion and finger per-
sit)', is mediated through C}'lOkines. Preliminary studies In cussion often rnsult in a high·pitchcd resonance in the dorsal
mice suggest that increased macrophage product ion of abdomen. Simultaneous auscultation and ballouemem oi
imerleukln-1 alpha. but not tumour necrosis fac1or-alpha. the abdomen reveal splashing sounds confirming a gas-fluid
interleukin-6 or prostaglandin E2. may be primarily interface. These signs of ileus arc the: mosl consistent clinical
im·olved in thC' pathogenesis of the disease.7ll findings in all case:,, and mkc on added significance when
Ehrlichin ristlcii causes Si{lnificam immune depression they occur in the face of an absence of clinical sign~ or colic,
in mice and measurable alterations of the equine immune and I he passage of normally formed faeces.
586 ,,, , o, nn•n: Rickcusiol and chlamydia.! di!-t•a;cs

The fever I~ fre'quentl}' biphai.ic, although the first phase ,een in equine 53Jmonellosis may he pre,en1, but man}'
is often not deiected. The initial increased recml temper.i- horses han:• normal leukocyte counts despitr other ~fgns of
iure range~ from 39.4 ro 41.l •c but, since it is notoccompa· enteritis...\ marked leukocytosis can occur within a fe\\' days
nied by other signs. the pre~ence of thi::. initial fever will not of onset of clinical sign~ in some cases. A iotal \\'hite blood
be realized unless the horse's temperature i~ mcasurt!d fre- cell coun1 of 20 000 to 30 000 cells/ µI may be found after an
qucmly. h rypic.11ly resolws within hour., and is followed in initiul leukopcnia.Allhough thi!, profound lcukocyiosisdoe-,
lh ree 10 se\'en days by a more persistent t'ever accompanied not occur comistellll) with E. risticii infeccio11. it is a rare
by other clinical signs. finding in matuw horses suffering from most other diseases.
Although diarrhoea is often thought of as a cardinal sign , If it is detected in a horse ,,ii1h consistent clinical signs. sus-
it develops in less than 60 per cent of cases. 10 when present. picion of Powmac horse fever should be high.
faecal consistency can vary from cow-like 10 watery. pip~- Gross necropsy. even !11 fatal cases of Potomac horse
stream. but lhe diarrhoea is not a::. profuse as rhat associated fever. is gem:•rally unimpresslve. The most consistent post·
with emeric salmonellosis in horses. 111<' duration of diar- mortem findings are Increased :1mou111s of fluid comems in
rhoea may be as shon as one day and rarely Iasis as long a~ the caecum and large c;ol()n, and focal areas of mild hyper-
10 days. Chronic diarrhoea is not a manifestation of E. ris- aemia primarily in 1he mucosa of the caecum and large
ticii infection. colon but occasionall~•also in that of the small :mrstine. The
Decreased feed intake may be a pronuncm sign . .:,ome 1mesr1nal wa!J 1s not oedematous and the mucosa 1s intact.•?
horses may have complete anorexia for up to l Odays in the 8y light microscopy a colitis is usually e,iidem and ehrlichia,
ab:;ei1ce of other signs. The major manifestation in other can be idemified in tissue sections s11•jnecl with a silver
cases may be a profound ileus evident by severe colic, severe stain.'6 Ehrlichias may also be detected b)' elecuon micro-
toxaemia and dehydration. In some outbreaks severe scop~. and by a tissue polymerase chain reaction <PCRJ
laminitis in all four hooves ma~ occur in up to .io per cem of assay. 11 The histologic appearance is somewhat variable but
affected horses. but in most areas it is a rare sign. :o. 1•· ' 8 The generally includes mullifocal ne<>rosis limited to che mucosa
laminitis, when present. may be the major mani(estation of with a mL'<ed inflammatory infiltrate. The histologic lesion
disease and result in humane eutha~asia of the horse \,ith in often appears mild in relation 10 the Se\'ere clinical disease
days of 011sct. ~lore often. however, it occurs together \\'ith which has led 10 a fa1al outco111e.
other signs. is mild and completely re50Jve5 within three to In addition to the placentitis, histologic examination of
five days. A few cases develop chronic laminitis. foetal tissues reveals a remarkable colitis. periportal hepati-
The course of disease wilhom cherapeuuc intervention is tis and lymµhoid hyperplasia of che mesemeric lymph
usually 5 to 10 days. Clironic manifestatiom. \\ith the CKcep- nc>des and spleen.
tlon of laminllis, do not occur. The ca$e fiuality race of 1111·
trearccl clinical cases varies from 5 co 30 per cem. Some
Diagnosis
fatalitie~ are related to an apparent uncomrolled $epsis. lhe
horse suddenly dyi11g despite 1101 being se,·erely dehydrated Although lhe epidemiology, clinical presentation, haema-
or ha,mg enreme add/base or electrol)"le imbalances. This wlogy 5ndings and response to treatment may all be sug·
may be an extreme inflanmrnto~ response orchestrated by gestive of Potomac horse fe,-er, It~ diagno5I~ is based on
a modified systemic inllammawry response S>i1drome scrolOID', PCR and/or isolation of 1:. risricif in cultures. Ai-
response. Other horses may have to be euthanased because thou~h t.he organism does parasiU.ze circulating monocvtes
or severe laminitis. 41' :md can be seen by light tnicroscopy of blood smears stained
E.hrlichi(l ri$ticii can cause abortion. Experlmentally. with common haematology srains. only a small percencagl.'
nai,'ll pre1,,nam mares infected bec\\'een 100 and 160 days or or circulating monocytes are infected; this renders 1he ex-
gestation aborted between 190 and 250 days of gestation. amination oi peripheral smears a not ve~· rewardmg diag-
The mares developed typical signs of Potomac horse fever nostic procedure.
and appeared to have recovered fulty before aborting. Abor- Ehrlichia ristitii can be isolated from monocyre cuhure.s
tion is accompanied by placC'ntitJs. detectable hhtologi- oi blood ,,ithin day,. of experimental iniecnon and consis-
cally. and a retained placenta in most case>'. Foals whkh are tently umil30 days after lnft'ctlon. 11 F.hrlic/1ia rl$ricii may also
carried 10 rerm mayor may not have a presuck titre of senun be cultured from affected foetal 1issues. especially when the
antibodies 10 E. risricii. in eirher case full-term foals are period between infection and abortion is long. The placema
31
.helllilw. and are 1101 ehrlichaemic. l5. 3? i~ usual!~ retained and bacterial contamina1ionofil interferes
\\ith the isolation of E riscicii from it. Howe\'er. isolation and
growth oft: rislicii in antibiotic-free tissue cuhure (co-culli-
Pathology
varion oi tissue or blood monocytes with muriue !'38801
The leukoc)'te count during the early phases of clinical signs cells) are research techniques not easily adap1cd lO diagnos-
rn11~· be variable. Leukopenin with a neuuopenia and lym- tic laboratories . .\!though a definitive diagnosis of Potomac
phopenla producing a haemograrn similar to 1J1at typically horse fever requires that r:. risticii be Isolated. in practice
 Po1onmc hur,;<1 fever 58i

reliance is placed on serology or PCR for its diagnosis. Serol· have difficulcy w1lh the procedure for Potomac horse fever.
ogy 1s not a suaightfon,'ard diagnostic technique because of Inconsistent antigen preparations, improper use of posicive
Lhe som,·what unfque kinetics of amibodyproduclion ~timu- and negative controls. c:onfu,ing nonspecific lluorcscence.
lated by E. ristlcii. Infected hm-ses have a rapid rise in nuores- and inexperience with identifying specitlc nuorescence can
ccm amlbod)' ticre, which level out within a few days. In the all lead to erroneous resulcs. The most consistem results will
absence of a blood <ample for serology taken in the early be <>btained from laboratories which frequently run the
stage~ 01 the disease. detecting a significanc rise in amibody assay and take great cure in nmning both negative and pos! ·
titre may be impossible. E,·en samples taken \\ithin hours of tive conrrols.
tlul onsec o f clinical signs can &how fluorescent amibodr ti· A recent study ha~ shown that even when performed by
tres oi 1,2 560 or higher. As a re~ult a decrease in titre is as experienced research laborawries. the serologic tesi results
common as a rise In litre and must be caken into account ma~· not be accurate. Serum samples l'rom suspecc c~sesand
when antibody titres in acute ancl convalescent senun a sample frr,m an experimentally- infected horse were sent
samples are compared. Thus II four-fold or greater increase or blindly to several tuuversity aJld private laboratories. ~lost
decrease in fluorescent amibody tirre can lie considered diag· laboratories reponed high titres in many of the sample,.
nostic. Approximately half of acutely affec1ed clinical case1> Close scnuiny of the s.amples by Western blot anal)'Sis
will ha\'e no significant change in tim: because the antibody showed that che onlv true positive sample came from the ex·
levt:l ha. rt:dchcd a plateau before the fir»t ~ample is dtd\\ 11."' pt.:rlmomtall) infockcl hor:..:. Th" ru,1 wet<' fal,c po,itivcs. ~•
The determination or antibody ri1res in single ~crum This suggests chat caution should be used in Interpreting lhe
sample~ serves no purpose. Although clinical cases will have resu hs of serological tests when used for diagnostic
titres greater than I:80, there b no correlation between purposes. Serology should be abandoned in fa\'our of more
height of tlcre and lilwlihoocl or acute dis1:ase. Antibody accurate dlagnostic methods. as they become available.
titres in horses tha1 have been exposed 10 Potomac horse A PCR technique ha, been adapce~l co detect F.. ris-
fever for longer than a year beforl:! antibody determination~ cicii.3· ;;. 6• 38 • ~J 3 ' The organism can be idencified in blood
are done may be as high as 1:5 120. On t:ie other hand, If a samples and cis,ues~ 1 whhout the need for culluring. and it
horse is found 10 have a negative tit.re afrer clinical signs de· takes less than a day. Although false negacive and false
velop, it is probable mac it is not suffering from the disease positive results5 1 can occur. wi1b proper cumrolo the PCR
and other diagnoses should be pursued. rrcaunem 11ith OlC)' · assay should be a rapid, definicive diagnostic test and
ierracycline early in the clinical course or the disease does should replace serology as 1he preterred clinical diagnos1ic
not affect lhe development of high antibody lel'els. Horses method. This ,cs ting procedure should not he \iewed a~ a
which ha,·e been ,:accinared with a Potomac horse fever ·gold' standard since doing so mny lead to unnccessaf')
,oaccine ma,· ha,·e titres as high as 1:640. In general. titres morbidicy and monality from unnecessary antimicrobial
conbistently decrease after vacdnaclon and after six to nine therap~ resulting in antimicrobial-induced complications,
months ha,-e disappeared. When l'accinated animals such as antimicrobial-associated enteritis nnd renal
develop che clinical disease (e.g. when a \'accme 'break' disease. as ha~ recemlv been reponed in human medicine
occurs} antibody th-res ma}' l'iSe drnmatically. In such cases associated wich the diagnosb of Lrme borreliosb.:': 50
amibodpitres are often over I: I 00 000 despite the presence F.hrlichiaJ abortion can be tentatively diagnosed by his-
of clinical sign,; of the disease.:ir.. au, 70 cologic examination of the focral colon. In fact. the colonic
l'he first blood sample for diagnostic serum antibodr lesions are as helpful in establishing a diagno~b of E. ristlc-ii
determinations should be taken early in the course of the abortion as the foetal liver lesions are in equid herpeS\i rus
clinical disease and the second live co sel'en days after the abortion. Confirmation is possible through a PCR assay of
fir~t to maximize 1he chance of seeing a significant change. foetal tissue~. 11
l'his shon clme imen-al between che ftrsl and second anti-
body determinations is con tr.try to what is generally the case
Differential diagnosis
for rno~t other infeccious diseases in which. for diagnoscic
purposes. rtllance is also placed on the difference• In anti· The differenrial diagnosis of Potomac horse fever includes
body levels in acme and convalescem phase sera. thl~ latter salmonellosis. intcscinal closLridiosis. nonsteroidal ru1ti·
being detennined two to three weeks or longer after 1he innammato~ drug toxichy, antimicrobial-associated en-
fonner. In Potomac horse fever this longer time imerval has terlcis. coliris x. fungal colitis. and toxic enteritis.
no ad,-amnge. le is of uunost importance co have 1.hc anti·
bod) levels in both the acute and chronic phase sera decer-
Control
mined simultaneously in the same laboratorr.
Great care must be taker, in choosing a laboratory 10 Elrrlicflin risricii is scnsirwe in uiuo to demeciocycline,
assay serum samples. The most common technique used i, doxycycline. cxyttitrncycline and minocycllnll.'" Ox1-1e,ra-
an indirect fluorescent amibody test. Laboratories adept at cycline is the drug of choice for treating E. ri~riciiinkction in
running other indirect fluorescent amibody assays ma)' still horses. The organism is an obl.igace intracellular parasite
588 ~t<:110" rniur. Ricl;ettsial and ch!am) dial disease~

thlll survives within phagosomes In macrophages by inhib· As in any horse ~uffering from colitis. the administration
iting phagosome-lysosome fosiorl.11? This inhibition is re- ofnuids is an important consideration. Fiuid and elenrol)'le
versed il1 virro by ox-ytetracycline, resulting in de.:.tniction of replaceml'l1! therap~ should be tailored to suit the indi-
the ehrlichias by the host cell.32 Oxytetracyc-line may ,1dual case..\J any or the clinical signs in acute colitis cases
interfere \,·ith the production c,i a cell wall protein Lhat is re- can be a1tribu1ed to inflammatory Cytokinc responses.
spon$ibll! for the phagosome-Jys,:,~ome fu~ion lnhibirion. These ca,es may benefit from 1herap)'Wilh anti-inflamma-
Oxyretracycli ne administered intravenously at the d<>S- tory mediators. When choosing s11ch therapy, It should be
age rote of 6,6 mg/kg once a da) for five dars Is a very effec- borne In mind that experimental evidence suggests lmcr-
tive trea1mem for Potomac horse fe,er when given early in leuk.in-l alpha may be much more important than lUmour
the course of the disease. A response co 1reaunem is seen necrosis facror alpha. intcrleukin-6 or prostaglandins in the
within 12 ho\1rs as noted by a decrease in rectal tempera- inflammatory response caused by £. risricii 79 Secondar}
ture. followed by an improvement in appetite. habitus and salmonellosis is not uncommon in horses suffering from Po-
borborygmal sounds.4' Diarrhoea. If present. docs not re- tomac horse fc\'er and this pt>ssibility should not be O\'Cr·
spond to oxytetracycline therapy:· Since destroying the looked. J=orrhis reason. and because a definitive diagnosis is
pathogen would be e:1.-pecied 10 take longer I han 12 hours, or1en dlfficuh. it l5 felt that horses suspected of having Poto-
the rapid clinical response to treaunem implies that meta- mac horse fever should be isolared despite the Jack of direct
bolic b) ·product~ acti11~ through the p1oductio11 of .:ytuk· .:umact tran~Jllis~ion.
il1es re~ult in chcsys1emic inf!ammatOT}' response s~·ndrome U111il the mode of transmis·s ion is clarified and the vector
which may be respoi1sible for many of lhc clinical signs. unequivocally identified. anempts to prevent exposure are
Once treatment is begun, progression of the dise~e stops. unlikely to succeed. Although aquatic e11\ironmentS appear
The response 10 0:1.~'tetracycline therapy is so dramatic tha t 10 be a loca1ion of one rcsetvoir for the organism. there is no
it can be used 10 strengthen a clinical diagnosis. If the diag- current e\idencc that the a,•oidance of such areas \\ill pre-
nosis is c-orrect. a resp(mse to therapy ~hou Id be clearly seen vent infection since the disease also occurs in horses which
\\ithin l2 hours (with the exception of a change in have h.id no direct comact with them.
diarrhoea). If no improvement occurs \\~thin 24 hours. the After infection "ith E. n'sticii. protective immunity devel-
diagnosis should be reconsidered. If therapy is begun early ops LO ihc infecting strain as evidenced by clinical protec-
In the eourse or the clinical disease. clinical sign~ resolve by llon to reinfection for as long as 20 month~. 16 Protccti\'e
the third day or treatment and no more than fiye days of immunity can be passively transferred.!!A suggesting that it fs
therapy are needed. 17 If a relapse occurs after discontinui ng primaril)' humoral. :--eutraliz.ing amibodies 10 specilic pro-
therapy. a second course of OX)'te1racycline is usually as tccti1·e antigens appears 10 be an important a~pect in ihe
successful as the first. induction of an effective lmmu nity.1;, .,.,, •.,. 31 These ami-
When treatment is beg\ln earl~ in the cour~c of some rick- bodi~ appear both to block internalization of£ rimdi and
etrsial diseases. relapses mar occur.2 ~ • 5. "fl Relapses are 10 interfere wit.h ics metabolic activity in host cells. render-
thought to be due 10 Jack of sufficient lime for adequate Im- ing it incapable ofintracellular proliferation. and result in its
mune stimulation oflhe host resulting in a tack of protection.;5 evemual desiruction.36 ,:hus it should be possible 10 in<l11ce
In e:1.1>etime111al E. risticii infection. relapses do not occur when Immunity by the ad111i11is1ra1ion of a ,11ccine.
Lhcrapy is initiated early in 1he clinical course:"; llo\\'ever, A number of Inactivated, partially purified. whole cell
treatment during Lhe incubation period merely prolongs ci1e vaccines an! commercially available in the USA. Vaccina-
incubation period but does not prcnmt clinical disease from tion has been reported co protect 78 per cent of expcrimen-
developing.~~ Early ueaanem docs not intt?rfere with the sero- Lally infected horses from all clinical manitestations of
logic rt'Sponse or the development of protection:'; Potomac horse fe,·erexcept. in most cases. the fever (22 per
/1. combination of erythromycin (25 mg/kg) and rifampin cent of them had no clinical signs. 56 per cent de,·eloped
(IO mg/kgJ administered orally twice a day is also effective ill fe\'er only. 22 per cem developed fc,,er and other signs of
treating Potomac horse fever when given early in the course disease ,.7° \ 'accination -induced protection is Incomplete
of clinical diseasc:13 The response to this treatmem is similar at best. which may be associa1ed wrth poor ~1imul111ion of
to that ro ollly1etracycline. wiLh 1he exception that there is a neutralizing arniboclies by the .vaccine.17 n,e protection
slightly longer delay before the animal becomes afebrile (24 induced by vaccina1 i9n wanes much more rapidly than
hours vs. 12 hours).43• 47 Such a delay in the induction ofapy· that induced br experimenta l infec1ion. Only50 per cent of
!exia has also been observed when rifampin has been used 10 vaccinates are fully protected at six months after ,,accina-
tri?at ric,keusial infections in humans. TI1e time period be- rion ,n and protection decrease..~ to 33 per cent after nine
tween Initiation of therapy and rerum to nom1nl of 01her months,' Based on this evidence, re\'accination at four-
physical findings is similar for both neatmem regimes:13• 4 ' month inten-als appears to be necessary to maintain a rea-
Allhough no experimental C\idencc is a,-ailable in horses. it is sonable likelihood of protection. In most endemic areas u,e
considered that doxye,-ycline admi11is1ered orall~ should also disease occurs ~easonally. In these areas. if any of the cur-
be effective in the therapy of the disease.62 rently available inac1ivated vaccines are used. horsrs should
 Potomac hOr$~ fc,·cr 589

be. vaccinated in the spring one momh before the fi~t cases expenses are not decreased by vaccinaiion. 1 The reason for
nre expected and agaih four momhs later, if easel> are srill OC· rhese vac.c;ine iailure, may be associated wi1h inconsi!ltent
curring in the area. antigen preparation or. more likely. il is associated 1,ith the
During the past few years, protection by vaccination apperuance ofse,·eral genetically dis1inc1 strains of E. ris1icii
seems to ha,·e become less efficacious. lt1itially. after the irt· which vary in their abilicy to induce 1he formation of protec-
rroduct!on of the ,·accines. vaccinated animals which did dvc amtbodies.' 10 "'' \II vaccines currently available con -
develop disease appeared 10 have a mild form ofil. '.\tore re- tain only 1he strain that was originally isolated. Polyvalem
cently. howe,•er, some vaccinated horses have deyeloped recombinant 1•accines containing amlgens which e:icit pro-
the severe natural disease from wliich some died despite the tective antibody formation may. in future. become avail-
fact Lhat they were apparemly adequately vaccinated. In able. These should increase the efficacy of protection
some areas, the prevalence rate of the disease and treatment afforded hy \'accina1ion.

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.,1w1u.. LR.~ MOHA\1.;\H-l), n o.. 1996, 6\7'.h.unfon or \ll.\~c1notioo 01 hor,1;1 .. 13 um L\.,--o. ( .r,. wmn.on:. R.H. & ,r-~to:,.i;, 1.. t988.
D;\\\'Ml'· J .t,. 'IIJ'tTIC, !i.t.
as a i.ttialC'&\' 10 conuo1 Q<luinc mc;>nocvtk ehrUch1011J, Jcmrnal nftht· lll(>luuon o( Ehrllthur rn•(,t,i ttom a fo:-u~ or .s mai e \\'ilh Potomae horst:
N,h·rirtm VNtri,100· .\1t.odit:"I .'4.ss.<x·,l,t,on, 208. 1290-1294. re\'Cr, Jn P,~>fYWiir.B> ;,om" :;ympq... f11m on Poromac iJor1-t 1-t,'.ir,
2 llARLOV<JH. J.e. R!:tJ~l!l.., C.:,H,, }1..\0IC"~,"• l,t... \'N.tOt\'Ufl, 1-.l-,, \III-U.ft. P.l,...f.·
Loui.ville. Jll' 10;- I 11
IWJHIAA. ,.., 19!!8 Deiect,on or Ehrllrlrlt> rlmrl/. agent of Potomac •h• 16 IIUT'U. "' ~ \UITl \ t.l \, U,t. fo Ml.\.."'"-,\K.,,PJ",\• u,. tg.89. ,\ntibody r~Jltm"il'
ho~ r.:,-er, in !wshv."Bt~r '.U"t::tm :-cn,1lh (Ph:uroo:nd:1~ Jt,stt ~pp. co EJ:riicJ1ia ru1tdt J.nd antibody reach\ ll\' 10th"• coinpontnc inug1,-ru. in
from northcm californla.A11p//<tt/ ,'I, E111·,rQ11111c11ta/ \/ftroh/n(o~·· 6-1. horitcs \\ith 1rduct"<I Po1ontai: ho~c (e\·tr. l11/~·ti(J11 ~ lrnm,mltj·. 5';',
2888-2893, 2'J59-296Z.
3 8,\J\LOUCll, J.t .• RtJJHlSA.. \. 4 '1.{AL)IC,,.:--. J f,. ,997. ~C,q\!'d
polymcra.~ 17 nurn. 'i.t,.., n ....1uur\l.u. ft, 4,: ,u,w,,. a.. 1998, Assod.iUon ot dcfkitncy
chain r<>o«lon for det~crlon M Eltrlk/11,1 risrlrli G<nnmk D'IA 1n In anliboll) ,.,_pon,c 10 ,•ocint• ,md ht1,•rogc11,hy of J:hr/iclrl~ rim<li
lnfc:t·tNI horM.~. ,,.,nerfmu~· Parasftt>t,,~·. 68. Jff:"-373. ~rain, wich l'o:om,1c hor,,e r,.,..,
,,,·cine (nllu"• In hor,,c;. Jormuil ul
"'"""'· U,G., 1~S9 $chcring•Plou~h Animal H•alth Co:por.,uon, P.O. !!ox
Clfolr.ul .\llrrnbtul,>g:•.16. 506-512.
3182. !Inion, 'l,J. 07083-19R2. l'Cll!On•I corrunu,11ca1ion. 18 r.01111., I.L. lliHJ..\.""t>. C.l,, IJ\\\',C.ns, ,., •., JU!t-Tlt,, M.. s.1r.111n1. ~ .. ~UU.\l'\,

5 8IS\\'A5'. tt.. )IUKHJ:Jll£1-, D.., \l,\'rTIS'ffi.\'•~,\PfF!R, B,L 4,. nun.,,, s.~. 1991. X.G.. 10UN1'0~. P,,J ... ~IHl:.frEJt. n.,. £: l),\~I K (';,J .. 1989. \tunchl~· iJfC\'"41l!lltt

O•nl'n('I.Wc nppl:<;at.!on.of pl')Jymer.ut' chl.\U'l r~{'ctcon for dt:ttt:to.n of ,in 1986: or an1lbod)· thrl!S .,g;,ln<t equine n1onol.;11c chrhchlosfs in
Elir/ltl11a ri,tlcir in equine mon0<")1k ,11rl!ch10SJ.s !PotomJc horse re,,.,,. oppattnd: ht'ahh) hDTSt...., in lllinnh. /\mtrit"lln Jo11nu1I ,,f \l te:n'>14r,,
1

1/,•ioortlr. 50, !936-19.1~.

Joumui ofC/ft1f(,il Mfcrlllriol,111>·· 29, 2228-2233.
8 Jl'5" .,~ a. \.L\ tUL..\PALU R." O\fflA. s.;,:,, 1994 O!!'ll'(IIOn of fJrrh,iua J!J C..OJU)()'.\_ J.(.: atc.11,,at..,L~.. J,L. Jo,\n&\A. \\ , f~TU. \\", PA.m,0%~ .)l. &,
l:OH~. J.., 198$. F.pidemio!oglc Jn,estlg>llon ol .ill',>ct<>d funn,
-~u1Joi 1rom ft.'CC'.5 of inft>cu.>d horses by 1mmunom.ag:ncdc st'pmanon
>nd PCR,Jo11malofCl/rrlt:l1I ,\t,crohtol,,ro•.32. ZJ.17-:/lol. Pmi•Miff1g,. o.fu $),•mpo.rt11m <m Pt,Mmnr lltJt,,, J,;,,,,,r. l..otl.l:i\1J:tt, "T.
Uv\. pp. 21-2S.
-: tll~\\',\.\-1\., \'F,\IULAPU.U, R, 4 Ul.lrrA. ).~•• 1998 • .\loh:ntl.u b11,Js {o:
antlgcnfv varintion of a pro1c<:cive ~trnfn-;)pccltic- and.gen ot F.11rlfchl11 ?0 11.\H~. S'.l!., H,rTCltAt. M Jttct, ft~\ l .. t:nC\."1~ l.. \I., 11,.\:\"M·.S . J.\\'. ff.\111. t ..\ ..
1t,\1\ttft. n.". & PERR\~1,1.0,~ 1990. Am!mptl.-d U'3Jbnlh.,tor) of EJ:r/ieltla
,;,rir/i. l11femo11 ,'< lmm11nll)•, fi6. 3662-3638.
rlstlcli, cau~lh-·e nf(t\m ot Potomac hor<e Ccvtr. b~ lht1 ticks
8 IIURt•. ,.(;.... M()lllJIT'r-.••\,\\',. \\1LI.L\'l"•i. S,N .. PO\\'l: 11 D,(;,, $. '-~.u•v f..\\,,.
(J,•nnm;~trt1Jrt1ar!ttbJ/i.~~ Rl,/pJr41phnh1.t..sa11J:11hwu-1,, b(.H/t"> ft'PPIIIIUU
1990,\ncmptl!d min~mls.lon of ldlir/iclrffl n~idl ,Ritk~ml~c;,nc; "11h
and AmMyou;ma am,•r(<~rnum. F.'<J)c!rim,mltll &;\pp/11,I .J.carol11~·. 8
S1omwcysatlci1m11s ID!pwm· ~luscl!la<·)./orrma/ of.Wediral
•11-SO.
Emomol"to~-. 27 8i.;-.e77.
21 UAt"(.sO..._ o.A. 1~83 Ffft:ct of tmmune scru.m 011 lnfe<t{vat~ oi RlcS:tms!n
9 CAftltOl..t, ,.t .. ~C.UMlOntAS:r>.. !LT. & ruu, H..M •• l9ts$. \\'h.ite-foat~-d. mle~~ u,usuglWWj/,i, Jnftet1011 .S· l111111tmlty. 42. 341 ...349.
Tfck burdsms u.nd role in the ~pfzontlulogy c,f Pmumac hm"''" !ttwr fr.
\la.l)1•nd. /011m,1/ nfWl/dllf,• f)L-,. ZS. 397..;oo, 22 mBtu,,s.c tto;,,;""· ' D. ~ <'llf'E.''C. c:.f•• 19io. n1..i.e11>illl ln1ec:lons rn
dojt, part lll. >almon d,,.ea.e complex 11nd h:>crnobanoncl!OS!S.
10 (th[. r.s.. PUMU\l.,\, s .. Jnlt.N'SOS. t; .. Dr!.k()C)., E.. UWLEA-, s..r•.,k.!\t,\Oltas.
Comp,•ndium o[Com1m,1,1s !Me1cu11011 ;,, \ e.•itrlnar;· l1tt'Klit,•. 8. :!E 1-
J.t:.. 2.oon. lnfectio11 of aqua11c insl~·t~ whh rrt-m.,todt- mctJtcrt"llrlae
256-
nffltii. tho caUS(I or P<.nom:ic hor~c !eve:- Jou mat of
CtUT)il'lg E.htl.ich/11
\/,-dict1/ E111omol1J~·. 37. 619-625 23 IIQl't. J,(l. "- J;;!!.f£C, S,A,. l90.:. l'ri~ FJrr/1(/)J~(lt_•. /11: Btrgi4/S ,\.l(lfltl~'ll nf
S)'Stmnntlt 8<1,Mio/f)/(1' Baltimoro, \\'lllinm, ,1nd Wilkin,. pp. ;o.;-; 11
11 OLUf'JlA.."<A$11UWI ~YA, w .. nu.;.nnSA, \"., \',:\l\j..\MOtO. s.. HUD,~••
C"AA\\."fOJtO. T..9., f'[fUt'flL\,"i: t .f_,S P.41..\l~R, (;.U.~ 1~9J- AJU~~tn1c. ::t.i 11.t.':)r;.1~~. s., .. ,o, u.:,; K.~ JL~~'l. 1,.1 .. 1985. Po1011lat hors~ !t',d:agt-·n: iJl
marpho10$tlt'• .ind mofc.tC'uh,r chnrottcrt,~tlon or new Ehrlidw1 rimhl m:c:e. T11r \ ;.~,.•ri,ml'\' Hc,rortf. 117, 5Sf)--S:,7
i~ol,\lCS, Journ11l ofClinirol,\ firml)ic>l"lr)•, 32. 302ti-.l033. 25 1>:.',11:ft. :\f,, Man, J.. ()ttA1<i1H. :\,. t-lULU. tt.. KHU,:,.., U:O., \'.~-: It- ftl~Jllb.\, \.,
l!1 CORl)f.~ 1),0,, PLJUlY. l\.D,. tmmUS.\., Y.$,CIIIC~atlX(; W.R,. 1936. ~o.,\noly<J>ofl&S rR:\,\ nnd Sl·kllodillton omlgengoncand
6nierocolltls ,au$1XI b)' /:1,rltc/lfa $/>. 111 the ho"", l'otoinoc horse fe\'cr). tran)mi.:s,.1011 in mice or FhrtfC'i,m n(-11t1f in \1fR'Uh.ue l\"Cm.1.wdc~ fmm
\"c"r1.?f1mr., Pmlwtv~J. 2'3, 471-rtt. IJ.lfmm llt'l.'A', r1..,.,nAU1t m Ohio, frmmal o/<'J{u,,:o/ \11a'1i11olo~·. lfl,
0

3:l-1!\.3.l.511,
l3 '.)~\\'SO:\. 11 , \11'1,V~A\\'Altl)l'"(,\, I., 1(0l1.Ai\'U, C.f.• 'IUl.."-1 "1,1,)l. l,J rJl'i.Ttr.:.
,1., 1988. Susc-e-ptlbllltrof cois to Infection with UJrlidua n's1irii. ~6 "-"t\"LOtl. P ',., CK,\\\'J"OtlU, 1,b. \10.1\\' "'• T.f, 4oi l'.\LUDf. C,.U., l~J. ?.J.,,.t\•e,
cau),mi\'c agent o! equim• mono<11ic l htiichlo,t.., .-1.mmcnn /oumal of
0
crans!cr o( an:Ibody tb f l,rlfrhln ri.Jtlt:fi J>TOhtch rnk<" from chrhchio.. j...
Vcreri11nry R,·s,-arc/1. 49, 2119\,-2 100. /nfe<r(Q,r 011d lmnu1t1lf), :,9. 2056-201>2.

U 01'\\'M>S. J.1-... N f\\'1SC, ~I\.. 199~ Su,cep1ibill~ ,.. rdt,~ to infonmn \\ith :r'." ~L'-"\1::t'.\:. lll.H 6-l'EOtA~l.~ JR, cf 198.o, Jmmunl! tt-spunf..t!I- to Ri'-'1<tll>in
FJttlirhia rltnffi"<"t14U. causariv~ :1gcm or hum:.tn cbrlkhlo:--~ ..\mm1:a11 n~·nn in(L.~don fn C'l?nS'-""ltclly ~lh)'fflfC" nmh• nuce. J11fil\-1Iou tmd
/011mnl of\'eu•riru,ry, l!esoorch. 53. 13:!2-1327 lmnmnlr;, 21. 310-313.
590 ,tr.:~" m•u. llickcnsial and chlamydia! disease~

28 KIT \0. r .. rw.nu.1. ILi.:.& n1l:WU-\, T.. 19·8. O~tt~tfflilinion of'-::ilmtJt'I \f'(1uhtt" L-hrUdu:ll culitf~ in ponic."\, ,tm~>Jitm1Joumato( t ,•1,trl,m,:,
pQl,ontng.dis.e,1~"' unrl £1okomln Hult (l"'\ ~·r F'h.1.ou-tccm uncibod\· Ri'S~1r.·l1. S..l. 2J0()-2J.!l4.
$.NdiC\ ,..,ith
Rf<k,•us.lu ::rnnl!Ull, Amrrfcm, /burnnl of\'tt«'riunry (8 l'Al.\l[R J.C... "\\lll"Jt.OC1r.:. IUI. l!,, m ~.so:-... LL, 19MG. Equin\•chrti('-hi.'!1 collti~
HNwtrch. 3'1. ~.-n2s. 1 i;tntt'lm~c hot<i• fl"\'t·r1: R1\~nh1on Oflhi: dl-.1'"8~<'! in renn,c,yl.....m!A. ~t•\,·
a9 ,:sow,.ts. tu:. ,~utnsos. c.w.. SIIIPU\". \\ t> •• \\'lfrTIIXI._ JI II. l•f.ftftY. Jrr"-i:r ~t\o,: \'or.k Ohio Idaho. .imJ Conn\!ttkut Joum'1l of thi• i{uu.•riu.m
a o. ,. n.-.,,mos. ,.1•., 198l. Acurr (,Qui.nu dlMthcal.:t.~ 111.lromc ·\EOSi~ .i_ t·~rL•n,m,:t ,\t,.odfca/.~Uwo,,. 189, 197-l~J.
1>rdhnt:1ary rcpon. Ptt>e.1'.•11/11,c:,.,• oflltt" :t$Jtfl Amr•r:cnn r..»tw:rmton of .;9 ru ,11 n. 1.[,, w1nnoc-K, ft.It M Jtt.'-&ON. c.c.. soas. F.qui11\1 ehrlh:hfnJ col Im•·
IItlui,w Pr.ttctltfmwrs. La... Vt.•t,:::is. 'tii.'V, US.\29 l:>3--:t>:
1."ff"OO or oxyu:maC'\cline :rc.-ant"nl during lht" i11cuhation J'('riod of
JO U\1S[.. rs.. ll.\'\", .-..s:.r.., ~J(]l0L$Qf\, \\ ... & c;.\r.l':M $l.8. 1.qq-0, .'\lti.'Ulpl(d llJrlkhfD n.stini 1nf«ti.C>n in poni~Joumal bf rlit ..~mrrir,,n \ t1trw.ory•
f.llr/fc/tlr.. nsllc1i mm~mis."ion \Vithl)qnmc\•nror rnriabH/1 t,cari: ,\IL'thc~! .-b...'<OC}t,tlr.>tr. 192. 343 ...J.$£i.
1xod1dae1. /1J11m,1J uf \1Nlin1/ f 111011u,/"1:!', 2;, 931-9.13. ;iQ ··,\11 f, rt•• (1flOU1>, ).,1 tJ}\\.J\.kO"t, \\.1),. WTlll,1111, ,\ f. 4: M-".IIM ~ii: s.,i .• ~uuu.
31 w~c;, -,.,.t \,OtT.I., T.r..... KMW,tr\, t .. W11n1'1,.t\', ,u~. ltlClr., 'Tr IIOUA.~&). th"ath from i.niJpJm>pnr.ie tht•rnp~· frtr Ly1m.• c.tl~w.e. Climc:111 ln/..'ClltJUf
c.1 •• tOKt.,ti\..,, 1.11 c.J .. 1!)Sj. E\ .titration o!(e1a1l lnfe,;tinn and
& fl\l.T..fl., Vt•w.--,s,31. TI0~-1109.
abomon in pn"WJ3.ni pomes t+x1acmmcnudly mfcc:1Nf with l:'ltrlklm, SI Pf NH\, fi,l).., P.\t \tr-JC, J E., BUN H, 1,11., ,1,\(;Nlt->-,.t::\, t\ \.0 'M'MHl,-. 1),0. I.
rt.ftfril A.m,rican Jouma/ <1J Vr1erlrmry· Tw.~arch. 50. 130:- ll 16.
Tnmn, 11.,.. l~.& 1£pidtmfolog1t:tJ charact~r,1~.ulon or lln ..a<Ulr
32 LQ'I;(.., M,T., G()tT/. l,L. \,,rm lrt, 11 C.. ._,\lrl.\~.,-1i\. l, (.-1.0U"i T.t .. lffl diarrhoea '.\~Tldromt,: The rnw-con1ml nppronch. Pror,•rdmg:s of rh~
ldcnlifkatlun ol Ehr/1(/1/a mtic/1 ;u 111• c.,u..ith'1l a~ttnt of two equine ~nc.·1tl)10J ,·m,~mmry F.pu/rm,o{~· urul Pr~·1·1•nt.aw·f .\1'-fl1rm,;,
abortion\ roao"1ns na1urnl maternal ln!ec-tlnn./ourna/ of \·(;r,irfonry· Ld,nburgil. pp. 1-18-lil
Vlagn.o.sti('J1Jt'(r$logar1,m. ;, 201-205. ~ Pftmv. 11.0... PAI \!J n. J,F~. nmUT. tt.h, nnu:o. J.• ,a·>1n1.:~. n .• E.Hk1r.Jt. ,,,.,._
33 ,\1,'.blGA.'=. , ...... PUSTfJtl.A, ;\,1 iOll~'\.(JS", L, c.u.u. ,...... ,u,nJtu. us .• lq.~ .-\ cast·C''>JUtnl ,tud~ of Pn1<Jmac honc fr•, er
Jtt.l\llU.~\. l'..
OE~ t'.. , uw1 l:lt. -c:....s•--"()(HJ. Tran,mt,<lon of Ellrllrhftl riJ.Jidl, 1ht..• Pr,,JWrfUll(-e \ ~'tcri,:ur;.· .\lf!1i,,i,,.-. 4, 6:~2..
agent of PolQJn.1.c hor)C l~,cr. o..ing na1uMU~ ln!1..)('1td aqvmfc im,,;..'('1, 53 1•1'1Ut\. 8,D.. ~ltMJtJUt,\SS', t I ., Htr.f, f\.M, , HM~'-!~. J.\\'., l 1.ET(.OtR. ,1.•
and h.clmtnth \"t.'(tON-! prcHmln.ac')' rep<>n. 1;·,,umtf l',•r.,rimu)· /<mmal. 32. TOftSfH. 1:.,..
hOJU. ,,.r•. '- 11.o\11"\, ~.r .. t~tt. l:pidc1nfQIO~ nf Pnimnac
2¥:>-279.
ho!'$• ,,~er An Jn,..,.tJg•1lnn ln:o rho ~;lblcrole ofncn·oqulno
3,1 .'-!<\nlG,\S, J.L, m...1HJ<.,, "·· 1.Al.\f[h, 1.1£., Of.1\0(J., c. e. )~On., .. ~q~
1
m.m1m,ts. Tl:r \ ·,·t,•murr;• Rtt.,,t,I. I ?5. 83~HG.
E-,d,na, for n high rn<r ofraL,r•110>1<1"~ mule, with the tndiroci SiS Pt.rK:Jsc:. u.11.. 1993:.,\mpliftc:uion 1>mduc1 toofumfno.tlo:\conuul Jn.
fluor~nt am,b!ldy 1cs1 for lilrrll.-hio ri.<tldi antibody in har1e:.. M lbJSti, O.H u-,{J\'IJl'.J.·,, 'i.MUU l.~• .&,\\'ltllt t .. ·ed~ DitlJpWJt{~
Jounrnl of t/111J\m,riro11 l'Nrri11nry· \/trllro/ .J.,Jffri11(1011, 20:. 14~1}-l•L~l .\Jo/t'(·n!ar .\lt<robfo!ogJ~ l'rinrifJl<':i am/ -~Pt>lf(m/011. µp to;-??!~
3S. "[1WR5,1,-,.:s,, "-1.. P"'-'lt._ J.I . OIJU..<. J.A <WN.l!. H,ll., Dl',O,. C.f, •
55 Nll.T'l..,.'t\~. (, .. M..UPH, n .. ~·,ontARt>, P.l-t., FUUl'iT. ,...... & numuu ..... tlt.91,
,,urn.oc.:, R.H., J!}8lt ~roloft)• for thl• diatcnu<ti,-ot ~u1n'-" ehrliehtnl I~ rfl"..\ g.t11'.• M:4ue11C(I' of XcorlcketJsi,, h,.'lm1111h0tYAand us
<"Olfris-. Proc~,"J{ng, of<r Symi>o#mn dn Po:tJmm• H~m J.'~'""'·
t..ouit.villr. J>h\ lot,:.tnt-th: n1iWtna.•nt ,1o.ith nwmhc"' af Uw 1,1,tmtJ.;; £Julkl1iu
!.'1. llS \. lli ~I•)• 1987. pp. 3.1-15, /111t.wu:tfut:nl Joum(I/ of System111lc &1ctc.~iol08)·, -$5 207-21 l
lfi ,:f:!t~tCJ..'.. r.s.1- H:Km1M. Y- l99.L lnh.ibiuonofbu\din~. c..n,f1: or
56 VU:.T[RL-\.:,,; rnttsso~· I ntA[ J.,rUSHRl..\,l.lC. OUU)(tr;,(,& '1.\0tG\S,
ln1raci,1lu1Dr prol1fcm1Ian or Fl1rllch/11 rm:cu In 1'38801 coll> by onti·li,
1-1-:. :woo. tnreruun nne of t:lrrllclna ristrtli. the 8-g<mt u! rotollU'lc ho~,·
fflttril serum. hnmu.nogJobulln <.i. or Fab fr.ll,,'fltcmt. hJft,·llrm t1ntl
(tc-shwo.t'"'' ~tn~ •mails t/l,g" _\'f'1•kat1ui.1 from n1>rthern
fC'\'i'T. m
Immunity. 62. 3156--3161
C..Ufomia. I tttriirnry Pnui,uolol{I', 92, 151-136.
3; \IOLLO\·; l'.J,, Pl ilSl:\Ci. n.H f, urRAtU>I, \",i' . .i:uut. Faisc'-po~h1\~ i6i1JhS of
Si 1•U~f1Jll.\, "\ .. U-11 f'(:S(C'".(',t-1(, t:,:o.t., ,1f,1'!1itf, ft1, <..1L\t. f .S.. L:Jil.11, +
l'CR ,osung for 1.~1ne dlse;,sr. C//11iral 111/trtlt11J.< f)/.<NJx·.<. 31. •I 12-.113.
MMHG.\'.\, J.t:•• :?oun. [)(l,ect1on tind quumimuon of Ehrl,rltu1 rt,ncu
3-8 \10l'r. f., ru.JalU"-, ,., ,.u.\.,r., \., M~O. ~M 4
,v. ("_\,.1997. Ounp:ui&on or g.crnornk Di\r\ 1n mfoctC'd ho~ :mdsmiUs h) rcr.tJ,umc pc:R t n.f,,_-r:naf)·
l'CR i\nd culture 10 the lndlrc<I Ouor~ccrt·anrlbodr lc\t for dlagno,;, rormMlaic,·. 90, i?9-l3S.
of Po1omorhorse f.,..e,./01,mnl o{Cl111/c11l ,\ftcmblolog, ~S.121S-Ul9.
58 n..etmtt., c,K.. lf.\Pl.f>lJf1H. J,(•, t; '1AmG.~"\, 1. r ,, 1~'8 Produi.hon and
3! OI..CUO\W, T.W. "1t<. 'f,11.,. ,wr lT('A.. t.\\,, IY'IO. Scrodl:\gnm.i" of <tquirH· Ch:lt\"t('~l?.aCiOl'l Of /:.hrl(Clziu r;.'it,Ci/, lhl' Ql,i\.'nl o( POloll"WC hor-.c fcvl!!.
monocytlr thrUchlu,1.> ln>t'l«'tod gn:,up,ur hnrst', ln ~llnn=ia. from ""ruill" Plr11mandt1r. Ju&w ~p.) 111 o1,1uunun1 tulEL re• and geneu·c
/aumnl OJ 1h,.~111eri~n11 W11•ri1int;r .\/edic111.1,..,.,,w1/011. !96. 1%7-19i0. comp~ri-ion :o c-quine ~rrun" Jo11r11t1l ofl1muyu ',tfcroi,i,,1,,~. 3i.
,JO PAL\\~. IL. 1987. Poto1n.:1c-hor~.:, fever. Jn MD.bt"-(O~. s.c_ (i!d.J. c,,rn•nt 1501-Ull.
Equi,:e 77t,,rapy2 Phll•delphlJJ:\\l'B.SmmdN< Company. pl). 92-93. 59 101r:u11~, \',. 1,at. Cu.>*·f1:-'Uing :anugcn., bcl\\ccn Si:or...·J,...:11,m
41 '"' MtR. J.E... 1~2. N~,· Bohon Center. UntvorsH) ot Prnn....yl\,mua. lwlmm:ho«n and F.hrls,h/1; .,pC'dt~. .>hnwn hr 1mmuno0uorh4.·~nc'°
i-enncn Sq=. P,\ 193411. un1mblished d•1a nnd \\'es,cm <nur.unohlonlng. Jm,mnl oJ Clmlrol ,\ffcrw,q/1>g,1·, ~9.
202/4-2029.
-4::! r.,t,Mm. 1.1;., t9.93. Po1umnchonw rc~r.
\(A•rlmTl}'Oint«ofNorrh
.1mericn - /;q11/11e P'rac1/c,'. 9,.399-110. llo rosuus.•. ,., 1991 1 h~ 1ribc Ehrl/"11/,'il~ and •hrllchl3l dl,eo,.,.. Cllnlrnl
.\l/aob,11logJ Ht•:•l(w1. .;, 281>--308.
.:3, V\1..\IE~. IL-f. flf,"l(ON,t;..l.• l~9.:! f!fttt aru.:ooumtnt \\l:h c.•~thl"Om)·tin
and rifamptn during 1hc acu1e -.1.agc,, of e>.pl'mn,r.iall, induC<?<i <'(tuine Gt 10~1111~,. v. t.• Jl""·
11 \I. isas..
In ,,tr,o~u~epublh1i¢t.. at l.il1r1tehln rl.uFnl
uhrlldlial wlith in pon~e~• ..inu,rriru11 /Qur'1t11l o/Vett>ritmf)· Hrsttm:h, !l:I, t<.t cl!tht J.nubiotk"\- Antlmi.:1oltful Af""" nml t.h1miorlti•<'1tf'.'. :l2.
20il-20i6. 986-~1
.;.a P.,1..,:r.A. Mi,,., aL,~x. c.L. 199..;. Studil'$c,n or.1J ,ran,-mb-sion or 6:2 Ru:.1111s.\. \·.""' Jl\Wt. B."-.. 198<t uren of .11uibio1ic"' on dinic.,L p~1hologic
1'01<,mnc horse (C\·cr.Joum(I/ o/\le1eri11nr, Tm,m11/ .1/,dki,:r. a. 87-92. nm.J lmmunnln~1'- r""'°pon~c,r. ln muri1Ht J>utormic lum~e !c\<-r~ µronicrtw
.;.s r1 ,1~,1tft. J.L. Rt.Ssh:\', c.c 1r, wmt1LK:X'. n.n 1986. Potomac horse fu\.'N: e1rcc1< <if do,,•~ycline. Vt1r,/r111ry Mitm/1/n/1,w. 19. 253-262.
~ Di.1gnosuc rt11rr1:i and tC!('ow11rion ort.ndcmJC' arca4\. Proct"t.·11/111.So/1/tc· 03 n.1,m1tS1\. \·.. 1ou,.!'O,. c.c. &- tttJfl(\(ft. c.1.. t<Ut:. Rtdurl"d unmunc
S.-t01ul S;_vmpos:lum mt &Juin,• CoUr Rrrt1J~~·h, \thtn\. GA. 157-160. rc-,.poml\L'nrJs :!l.nd lpnphotd 1.foplt.'1k1n {n mke infl"CH.'d \\1th Ehrlfrltm
rl~tlrif. Jnf«tion ttrt(I lmm1111iry. 5), :?2 l:i-2.222
..:6 l'AL)U:JL 1.k., tlJ ssos. C..il.. ti WIIITLQ(:!'. n..11 .• 1990, Hesis.,.inc.·c 10
<kvrlopment uf rqul11c tl!rllthlol c,oli11, in CX1><rimc11m!I,· ,noculntcd ij4 KISllll!I.I. Y.. P>RR\, ~.o. & '""1'~"'· t>.U.. l98S. 1Jhr,1"1J'UC!Urul ,wdy or
ho:'$6Qnd porues:. ,·\1mrrit,m f<mntnloft·ai~ln.ftt)· R,~arrlt. 51, 763- eh.rlichrtt.l org:mL,ms in 1ne l>l'tl" colons of 1101,i<S inrec1,-<1 ""h Po1om11<
;,;s. hor~ i('!n,-r ftlA..:tttJIJ and lmmm:tty• .J9. so:;...s I:!.
•,; PAt '4tll. 1.r. nr,-sos, C.6. ~ WHITI oc~. A,n•• 1992.. 6:fcct or 1reannrn1 will1 us HU.>flllSA. '{. kU II, '-)f,. MM:- N~A .• GOitO()N. l,<:•• 4, rkctl'\f.\~ r I., \9$0,
or
oxytttracydln~ during th\" .lcmc .stages (-Xpimmenmll~· mdut't!d Sc:,rasur\'tvof hnr,;;c,; wi1h- t:·vJd(•nt'.C- Q( tq\lff)(" monOC) u.:- chthc?uOS.lS.
 Potomac horse fe\'~r 391

Journal q/th, ·lmmrmr l'4!terr11ary,\ll'tlict1I Assotfa11011. 1~7. 11?7•13:!2.. L'.x('lOscd in hyper<'11dt.•m1c :\ft:\!' of scrnb typhus,.A.m('f1crm /1mrtwl oJ
I 1,vgll!m•. 50. ';'5-9 I
66 """'""'· Y.. w.,i,.,, n.. •• m,. u, "YMO.Moro,, • 1q,:i, n,v.lupmcnt of
ncurrn!Jzing nnnb<•d) In hot>t'S info<:H-<l with !:lrrl/(lrlti r,mcrl. 76 ,,u,u-, t.t., minn•~·'· ,. ,..,,,.,lM'-A.,t., tftllh. /;J,rllchi11 orf101um,,t hor..,.
1'.-1,rlr.nry· .\llcro/1/11/og;•. :u;, l!.\9-14- fo\'cr idrmOfied \\ilh J. ~\·c, 5ta&n. i furinmy Parht,/t,g:,·. ?3. S~l-533.
G';' ll~"'Tlc. M.. l9$G.. Pt•tUntnt chnt~!neri,tit'i orteukocytic RJc~<"tl~i.\t." of , , \li.PttF,:t.nx. LU .. z.9qo. r-...,p,,rtmcntnl ch,li~hu,,i, in nonhuma.,i
human,and nnim•l, In, ,\f/emln'o/01,."I, pp. 182-111;. primau.~. In, Elrr1tr:unJ.i$. .\ L'«lor,(u,m.,,t/Jvaw ofnnimtrll and
Jmmm,1, CUllf'nt TtJpia i" l'rt..•rlnnl}· .\1nllcim,111ut Amm11t ~:,mt'V'. 54,
b8 ms-nc. \! .. OAWSO:,.:, J,. ftO l l.'\.,'«0~ <:.J.& JP,NNY. " ·· 1"988. SuM.'t'pttbUiC}'O(
dois to infection \-.ith l:."hrlichin ri$flril. C-:H1$Uli\~-ag.cnt ofequine 9!Hl9.
mon~c,.'tic ehrlirh10,t~ ,Potomac horse fl."\'Ni ...l»:c.•rfcan Jm,rm,t tJf $8 1"W.\R"r7_. J.C !'>lllRAt, .\.. ,llX\MAJU,C .. ~\UNU.EK.\. l,f•.. HUXSUU.. D.L, &
i·mmnm,- R,.•UtJft'it. 49. 149>-1500. OKO\'t'- \J.C... 1082. Do")~ dlru: p:01,h)'l:ucl, for hmn:in ~n1h t\l'Jht:>
Gu mrnc ~f., nou...t..,n. r..1., r.,.,wsns, ,.r1., st,...i()'.\:', J. "pu.,,1c_n, J,L. 1~ft6. /orm:n/ ofl11j,,:1/01u D~<M>c;.146.811-818.
D1a14no~i!- ot ettu1nc nlQn{>e~·tic thrlkhfo~i"' f Potomnc hon,4.• f1t\cr1 by 79 \ '.\,~ IIUOJ:RL~ ••\.M Jllt-:110::,.,\. v.• PAJU.\ ,. "fl'IITU. Jt., 1993
Tumor
lndir«.t JmmunoOuor~«.nc;e-. /oumal oJ tht'i\Jliilrrrou V,m.•rimuy rux:roS-is factor u.tpha. mu!d~uki.n·l atph..l, imericuJdn -6, and
Medfr11/ Nsoc:imiQII. l89, 39-46, plllStJgfandln F.2 produwon ln murllie pcrittmca! macroph>I\O'
70 ftf)TI<::, :.: .. Hou.,,0. c.J. a. <;on 1,, r.e....1988, E\.aluauon n( a \-:t«im· for lnfc<:tcd whh Ehrlirhln ristitii. h1ftv:t(tm flml /mmunJC).', 61. 43l3-IJ3:;.,
equine mon<JC)ttc ~hrllchlu•b. Pror,,.~/111g,oft1 ~>"'"°""'"'on Puromac 80 \'E)Hn.w.uu.. R..., lll"t\\·.\.,. IL "nU"ITA,.!-,>-..19-9:5, l'athop:eruc. unmunolC'it'IL·
Hor.wf,,¥r.1.oumillo, f..T. USA. pp. 89-100. ;m<J mo1C'(;talar d1(u.,111n.;i-s brrweon t\\'U £hrlit.:hf11 ri.s:tclistrall~, Jo11mol
7l "iCIL\UOn:.\~"·· E.r.. RIC.I., R.!\1 •• ROW.t. \1,r;. 4, ,\l.\O, \.F•• t9._S,•atrh fo; Q[Clinfr11f ,11,vo/>10/og,•, 33, 2~8.-2~93.
.n anhropod, ,-c,or ot El,r/1,·lrill rl11icll. Prott,•du1g:sof11 S,1·mposf11111011 u 1 Vl· :\1Ut..\f'A.W.... bl~\\',\.'- ts. A,• UUTTA, ~.~•• 199». Studlc:i,; wfth tc!COmbmolnt
Potomtu: J/ol"~· Fet·cr. Loul>vtllc, 1'"1. USA. pp. !\-16. 111ut~Hl$nr Elr•//(/1111 risrlcit ldrntllkali<)n af struln,,p<!clflc an1tgM a.•
i,l SCHM!Ot'M,\.~S'. E.T•• ~oau ~f.G." CA.MOU.. 1.r... 1986. J\t<c-mp1ed pro1<-c1h-e ;mtigen I rwn11ory Pnrn,/10/"IIY• i6. 18~-i02.
ttansmis,fon of Elrrllt/Jfn rtstlalb) treld·capruml ()er,11t1cc11101 8: Wt.U.~, \; ' ·'- itUlilHIS..\, ,·. tglJS, ~ck on~onml Ni.ion \\iCh
I.VlrlttbilUt,ktfrl: l,u>ditfw:, .•A.mcrim,, /tmrnal o/r.."~:1Jrint1ry Rt•M11rtll• .a;. ph•gt),Q""-'5 cont:1lnln1tf11rl/drfa ri>ticrl In P338DI c-,U,
2.!393-2395. Abrogatlon ofinhlb!t,on With o~~iltmcyclln•. /nfi,:tiort amt ll•m11111r,·.
73 iussro~s. J.F- s, DAWS<>~, 1 1988. Producl ovalmuion: ~1n.ryland field [10, 3209-3215
t\·.1.h.z.atJon 011hr -PotQmac ho~ fcvr.r \11tclne. f:11uim· Pm,·rfn•. to. --12 llJ wr...:. , .. J,tlli:IHI'-\, ~ •• FUi:R'il. v ,A. ,I..QWCll'~A,lto\\UR.\\"A," ., t9~.
74 )H,\.,J:,\llAPPA. 8.. 0\111',\, '- ~ Jt l\t\Ttl~W. ,··~Al'ltl\. a... l992. ldtr.Ufit.atlon 01.-.~i~· ofl6S rR:-.A ll<fl<.., of nr,w Eltrllrlrlrt ,1rJ1n, lsolaivd hom
of \ht! protcc:dvc4,;•kilt>dalton rt-cnmb1n;tm :tnt~tn of E/JrUthw rL~tf{U. h(,rl>b \\Uh dln:r.1t ,,gn.,. o(Pocomac horw fNer. h,remmirm.fJ/J<mn;nl
Jnf.x-tfon(lnd fmm:uti:)1, GO. 612~17 t1/li.1'$WllllliC &1,·1Mo/og;·. ~•. 31.i--318.
i.i, ,,t.\!HiL f..l••• TR.\Uh, 11.. t rl". 11,L, MU..'Wllk~. J,I'.. ftUX.~UU. D,L.,.. hM()\'[.~ a., r1uf,,·i,·. P.1 .. 1993,.. In rJi,ru killing ut Ellrllrhlu rlstt(U by
\\'11.u,\..w,... ~.~1 • .,
·,,.G. 1950. Olloromph~nkol (chl<lr()m)'t••lnl in tho ch~moprophyl>Xi$ atth~11:(J and immom,. mom,c prrhnnf:\l m,u;ruph.lg~. fnf«rltmtmrl
of ,aul> typhu, 11"~u1,usomu,hf di.co«!) 11 RH\11,. with ,olunttcr, /mmrmlt; 61,861-867,
 ANAPLASMOSES

1\naplasmosis is an arthropod-borne disease of cartle, diales. As a resuh of these changes the genus Am111lr1$ma now
sheep. goats and some wild nuninam species caused by comains several species, pre,fouslr included in the genus
obligllte intra-erythrocytic rickeu~ial organisms of the Ehrlichit1. which are not intra-erythrocytic.3 The obligately
genus A1u1pla.min in 1he family ,\naplasmataceae·. order Jn1ra-cl)~hroc.1~ic .411nplasm11 species arc notv known as the
Rickeusiales. The parasites are biologically transmitted by el)'throcytic A11ap/asmn species? 111c name Annplasma
lick~. bur mechanical 1ran$mission by hae,matophagoi1s in- mesaeu.rn1m was proposed for an apparently di~1inc1 ery1.hro-
sects and iatrogenic means also occur. Clinical!~ . .-\110- cytic)l11aplt1.<11Ul infective for sh<'ep and goa1s6 but as yet there
plasma spp. ittfections in mammalian hosis may range from are no phylogene1ic drua for this organism so its provenance is
inapparem infection to severe disease and mor1allt)·. Gener- uncertain. rn
ally. the disease is characterized b~ le\er. progressive A varie~· 01 .\h'kan and '.\orth Amencan \did rummam
anaemia and ictcrus. Following subclinical infection ";111 species have been found to be suscep1ible to ..1. marginale
Anaplasma, or recove1y from the disease, infected animals and /or A ouis infection. bu1 anaplasmosis in these hosts is
usually remain carriers of the parasite for life. usually mild orsubclinical. and very Huie. is known about the
Differentiation of species within the genus A11ap/asma wa,, role that wild ruminants ma)' play in the epidemiology of
traditionally based on one or more of1he follo1dng criteria: dif- anaplasmosi~ 1n domestic animals. Uncharacterized A11n-
ferences In hos1 range; location within erythrocr1e:s: morphol- pla.<11w spp. have been obserYed in. nnd in some insrances
ogyof1he inclusion bodies; and, 1.0 some exiem. palhogenicity isolated from. a number of wild ruminam species in Africa,
of !he organism and immunological diffetences.s Under the and 1here is~erologicat e\·idence of1\1wplasma infections ln
traditional Sy-seem only four specles ofJ\11r,pl(1smr1 were recog- an oven ,,1dl'r hosi range.0• a Giraffe lGirajfa camelo/1tirdnlis)
nized. three of which CA mnrginale. A. cenrrale and A. cnudn- is ,he only wild rurninanr in which anaplasmosb has been
rum!) occur in c.itde, while the 01her (A. Ot•is)occurs in sheep associated with serious disease and mom1licy.2· 11 A11aµlas111n
and goats. The changes which have occurred in bac1e1ial clas- 111mgi11nllr4 and also uncharacterized Jlnaplasma infec-
sification O\llt the last decade. as a rcsuh of die application of 1ions1 have been found in :-:onh American white-tailed deer
phylogenetic techniques. have been noted in 1he lntroduc1ion (Odocoilem ,,irgi11ia1111s) bu1 there is no evidence that these
to the section on diseases caused by Rickensiales and Chlamy- animals arc an imponam disease reservoir for caule:'

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AM~~. ,1.Q.. unu1 I.L. A.M .. t\VI ~'\IS(.,(; , (..,unn:r.,ui r• ..tt"~Ul, "·· s 1uu u n. 1.P, • R1..,r-,r ,, , 11t&3,. An,\pla\n\011,is. Morphalogh: ,:h~r2Cltcrf'>t1n
.WM'\l.JII.. r \·t, c:<ntt.R, J•., .. t,uu.ut. ,. ""'qoRcu.c- \ 200s nett.~donc>f nr the p~:;i.,lte m the bin<><! ofcal, "' infecu:d with ow Ortgon strain QI
f.11r1tcl11n !>pp. In the blood of Mid \\hl<.Mnllc<l deer In \li,,oon b)' P(;R ..t.,1n11lm.mn trU.IJlr.tlli• .-tnu•rlrmr Journnt ()j \'tJeru:nn· H(1$,'f'lt'Clt, 2,;.
a<,.S)' ,111d ,i,erologlt •oal)',l<./011rm1/ <ifCllnlc,1/ .\1/cruh/olo;..'V, ~ t, 676-68,.
1263-5, 6 n tIA. J.. 1 tw8,.J . An.npltzt1r.n fnft"Ctit>JU:. 1n \'1.ihJ ..rtid d('lmKIJC
1,;11

:.! AUtiUSn"\, ).',J. 4 !U(,,U-Xti, Jt.O.. 19;z.Ann11lttt,1fl/1 i~h.'(!tff)n in .:. ~ua«~. ruminant, ,, tt\iL'" Jouriw/ oft\'il,111[• DI«,;,,,.. 10. l2-20.
O,.<f,•rnnpoor1 Jn11rm1l nf\',•1rr/11ary H1•xwrrh. 39, 29. 7 u•w \.E..(i,UI lti:,ft.,MISOll'\;,C.:OJ,.~JIK,\J',\'. lkflJ:1 \\ h.\Lf-1,l .. ZOOJ,
3 DUML.Ul. I~. 8.\ffl:tl"T \..f .• ttD:~tlt. C,. DAf:iCII, C":,..,,\.., l',,\ UllR, G.H .• JlAl', $ C•• Ph}logrne:lc anal}'"' of<ht<'l)ihro~~tc.ina1J/,w11a<pcae, ha~ on
Rrorgant,·....,tJon oi Crnl'r.t ,n tlw
R31-:ottSi\, Y. 1- -MUMA-"Clftw,. F. it.. 2001. l6S rO:xA ,llld GroEl. l tsrso, ~u\lnct.."!t ofA marginal,·, A r4:11trnf,-,
Famm,:r.rud,:c•uM:i.c-t-ac nind Anapln~m!u~c~e in chc Ordt-r Ric-kctrsl~lc,! ~itc.f ..;. nvi1 nnd ihess,«iric dttCClion of.A. c.rn1mf~\-acc1ne s:.rain,
CnJfia,fon of Somtl' Specie~ of £h1/khln ,,;1.1, ..i,mvtasm11 Cotnlr..o,,ith V,!1,'Tlna,y ,\1/t"obfol~·. 92. 145-160~
El1rJkl;w. and Phrlithiawith \feorid:1.•11sln:. Descripdon~cfFiu· ~e,., a 1011N. >,.~f ">JlfU.. tt., 1973". Ga.nu: 1.u1npJu,;mo~tt.. The !'°l:uton a(
:,Pede< C,,mbJnations-: and Oesignotinn of F.hrllth!n cqu1 and HGE ;\nnpla.wia OfiJ.OJ:5'ffi1. (mrn am elope:. J!ci1srl1ri/t fr,r Trop,mmt•dl;Jr, 1md
~~tnt' u., Subjt-cth·c ~ynt,.n}·mfi nr l!J:rllrhia 1,h,1gcxJ1oplal11 Paraslro!ngh·. 2-t. 192-~9;
t,irer1111tronal}o1~ rmrl of!ly1r.1mru("nml F.i-olutffJttllfJ.' .\lftrobfo/~1• 51.
9 tu,qnc.. ,, "'Jillt1Hl, t,r .• \98,:. FamU} Anapl.1MtUUGc:t.·J~ ,,,. ""'111l(';, ~ "· e.
214a.11w
um T, 1,c.. 1,..i, &>'liar> ,,:a1111ul ofSrttN1111rlr /1t1ct<'1lo!ag:.-, 1·o1 l
..: li(St..t.. ,u.:.. corr, \\',E... II DA\'IIJliiC)X, W .R. 199.5. An O~-mtrU oi thc- Doitlmorc and Lond~n· \11Uf3m< t. Wilkins
roh.1 oi wh:tt·tul?ed dttt in thu epizon11u10tn' -0f a.n.apl~-m<bh in
10 UILl:~HfRO. (;. \'~~ \OKSrF.,uo.sor. C.J .\,U.V. I, ruu( K~t .• t~7'9. BtooJ
1h~ ~0111.hei'l..,tt•m Unih?d ~tt~lt.!1 Joumnl of'rVild/lf;t Ois.tttY.c. '.ii ..
p:iru,-hn of ~http fn the ~c1hcrl:lnds. I•• \nbptu.,mo mL'$0.t•1t.•n,m sp,n.
3?8-85.
UUckertsftth.~ . .\n:tpfn,maract-acl. \ ttt1rlru,ry· Qt1,1rwrly. I H-22,
0

•Fora classllicntion rHer 10 tho intr<><luction 10 Sl't'tlon 3. Rlc:ketts13l and chlamydia! d~ascs
593
 47
Bovine anaplasmosis
Synonyms: Gallsickness. galsiekte (Afrik.J

FT POTGIETE R ANO \<V H STOLTSZ

Introduction tions of Babe.sin bi3emi11a and A margina/1!, which frequently

occur. led earlier workers to believe that these ·coccus-like
Anaplasmosis in cattle is an arthropod-borne disease bodies' represented a stage in the life cycle of B. bigemina.~ 18
caused predominantly by Anaplll$mll marginnle, and is Theiler in 19 I I231 referred to the follo\,/4ng serendipitou~ dis-
generally characterized by fever, progressive anaemia and covery as a ·tuck)• coincidence'. In 1908. cattle due to be im-
icterus. Two species of A11aplasm11 - A. margiJiale and A. ported from England to South Africa were vacdnated against
cenrra/e - are known to infect cattle in sour hem Africa. B. bigemina ,,ith a South African isolate or this parasite. On
Anaplnsmn celllmle generally produces mild disease, and, their arrival in the then Transvaal. Theiler challenged them
as cross•immunit)• between it and ,I. marginnle exists. II. with blood from immune local cattle, Fifty per cem of chese
ce111m/e is used in a live-blood vaccine 10 protect canle animals died. having shown fever, icu1rus, sen•re anaemia.
against the more ,irulent A. marginale. Colostral and non- ·marginal bodies or points' in the erythrocytes but no haemo-
speclfic Immunity largely protects cal\'es up to the age of globinuria. and this led him to believe that thesl' deaths were
about nine momhs from the clinical etTects of illi'ecrion. due to a different cause. He concluded that the ·marginal
Susceptibility co severe disease increases with age. points' \\'ere independem organisms responsible for the
'Peripheral coccus-like bodies· and ·marginal points' were acute signs obser\'ed and called them anaplasmata b"cau&e
obser\'ed in erythrocytes of canle by Theiler in 1910, which of their lack of C)10plasm, and the disease caused by them
was loog before he identified these inclusion bodies as being anaplasmosis. The specific organisms were called Anaplasma
the causative organism of the specific disease known by the marginale because of tJ,e marginal position (Figure 47. ll
local fa.rmers at the time as gallsickness,:m-230 ~1ixed infec- most of them occupied in the erythrocyte5. -

•
• •
•
•
•

•
Figure 47.1 Anap/es,m; margm;J/e: blood smear figure 47.2 /.naplasma cenuale. blood smear
594
 Bovine anaplasmosis 595

TheUer23 1 in 1911 discovered what pro,·ed to be invalu- produces mild to severe anaplasmosis in 1',onh America.
able in the control of anaplasmosis. This was a different va - There is no small laboratol)· animal k1101,11 robe suscept-
riety of the organism. which he called ,I. mnrginale (var. ible to Ana/JUISmn, and the absence of a suitable laboracory
centrale) and which was subsequently named A cemmle animal model.as well as the lack of a long-cerm in rdtrocul-
(Figure 47.2). He based the distinction between A. margi- turc system, have placed severe cons1raints on the progress
nale and A cenrrale on: of anaplasmosis research. Namral infection in the African
rhe different position taken up by the cwo parasi1es giam rat (Criceiomys gambia11us) in ;>;igeria has been re-
\\ithin the erythrocyte, the presence of slightly smaller porced,55 buc could not be confirmed in giant racs of che
sized organisms in A. ce111rale infections, the lower viru- same species rrapped in t11e Mpumalanga Province lowveld
lence of A. cencrale. and the incomplete cross-immunity In South Africa. 170 In addition, all auempts 10 infect intact
to ..i. margina/e demonstrated in animals recovered from and splenectomized giant rats br inoculation of A cmrra/e-
A centrnle lnfociions. and .-l. marginale-infected bovine blood. and by rick crans-
mission, using A. marginale-infected R. simus nymphae.
Theiler immediately realized the trementlous potential of have failed.
this organism oflower virulence for use in a \·accine, and the In Giemsa-stained blood smears, 1hc intra-erythrocytic
difference in the lengths of the incubation periods of B. big- organisms appear as dense. deeply purple. roundish inclu-
emina and A. cemrale infections made It possible and prac- sions 0,3 ro I µm in diameter (Figures 47. 1 and 47.2). The
tical to vaccinate imported cattle wich a combined redwacer organisms of the three bovine A11aplc1sma spp .. as well as
and anaplasmosis vaccine. wh ich was sold for a shilling. 232 A. 011is in sheep. are morphologically indistinguishable in
The first A. ce11tr(l/e isolate made by Theiler was prob- Giemsa-scained blood smears and are locaced predomi-
ably unique. Since this isolation was made i1 has been, and nantly marginally in the erythrocyte, except for A. i:enrmle
still is, extensively used in a Jive-blood vaccine in southern where the majority of organisms are located in a more cen-
Africa, as well as in otl1er parts of che world. including Aus- md position, as the name indicates. Anaµlasmn caudawm
tralia, Israel and Sourh Amel'ica. Therefore, the discinct and some A. marginale isolates have so-called inclusion
possibility exists that any subsequent isolations of:\. cen· bodr appendages. These are thought to be a product of bo-
rrale in chese countries may represem reisolacion of che vine host el)1hrocyre.s as they do no.t occur in parasiti7.ed
original isolate of Theiler. crythroC}'les of deer. 40 The appendages in :'forth Ar.lerican
Amore detailed account of the history of anaplasmosis In isolates of ti. marghtale are amlgenic 101 and their presence
the ~ourhem African region aJ1d elsewhere ls gi1·en by is thought co be linked 10 tick transmissibili{) of the iso-
Henning.'~ late, 9• Similar inclusion body appendages occur in ana-
An analysis of the sales records of che .-l cenrra/evaccine plasma parasites in the midgut tissues of D1m11ace11tor
produced at the Onderstepoor1 Veterinary lnstituce 176 a11dersoni. which is one of che vectors of A marglnrtle. Their
serves as a rough indirect paramerer of Lhe prevalence and a uachment to midgm epithelial cells nnd intimate associa-
importance of clinical anaplasmosis in South i\fl'ica. Judging tion with cercnin fonn~ of the parasite known as small par-
by the demand for the \'accioe and che 1,ide distribution of ticles, has led to the belief that these structures may be
the tick vectors oi the disease identified thus far,,-., bol'ioe involved in the infection of rick midgur cells. ;\'evenheless.
anaplasmosis is the mos1 widespread. economically import· some isolates from the USA that possess appendages are not
am tick-borne disease of can le in South Africa. The fact rhac 1rnnsmissible by ticks.z•a
1he disease is also transm!ned mechanically increases irs Uhrastrncrurally. initial bodies have omer and inner cell
potential distribution to areas which are free of rhe 1ick vec- membranes V\ith a macrix fn between. A filamentous pro10-
tors, :-io srudies. however. have been undertaken in south- plasmic network Is attached to the organismic membrane
ern Africa co decern1ine its relative economic tmponance in and is continuous \\itl1 dense chromatin clumps and aggre-
the cattle industry. The severity of the d;scase is, amongsc gations.'00
01her factors, probably strain-dependent and is not re- The development oi ...\. mnrgi11ale in the Yenebrate host
garded as being or equal lmpommce in different coumries. was first studied in deraii by Riscic and Wairach in 1963.:?00 Its
Con,rary ro the situation in southern 1\frlca. certain South complere deve!opmenral cycle occurs in mature erythro-
American counrries and Asia, anaplasmosis in Australia is a c~'les. Inirlal bodies released from parasitized erythrocytes
relatively mild dise.ase.~ enter other erythrocytes by invagina1ion of their cytoplasmic
membranes 10 form ,·a cu oles containing the organism. 65 · 197
The initial bodies then w1dergo a series of binary fissions co
Aetiology and life cycle
form ~trucmres known as inclusion bodies. which ate made
Of the !hree Anaplasma spp. that are recognized in cattle, 196 up of berween four and eight initial bodies (Figure 47 .3).
A. marginale is patl1oge11ic in cattle, .4. ce111rnlecausesa rela- \'\lhen ery1.hroc)1es containing inclusion bodies are dis-
Lively mild form of the disease, and A. c·n11darmn, 103 which rupted. initial bodies are released which invade other el)"lll·
has been isolated from mixed lnfeetions 1•.i!h A. mnrgi11ale, rOC)1CS.
596 ~r.--r:a., 1111111: Hickensfal and ~hlamydinl di,mnscs

' This has also been shown ex'Pcrimen1ally in the five dilferent
tick speci~ (8. tlecolomms. B. microp/us, R. e. eL'el'lsi. R.
simus and 1f. m. rufipes) common!}' found on canle in South
'
,\frica 175 and in male D. mulcrso11i ticks.20 '
/\ot all A margi11ale isolates are equally \irulem.
Kutder.'M usi11g Theile, ·s A. ce11ua/e bola1c In a ,tutl}' or
' 1he comparative ,irulence of ,I. margi,ur/e and A. ('enrrnle,
ob~erved only minor differences between 1he latter organ -
Ism and 1wo .\mcrican isolates of A. mnrg/11nle.
/111tirro cul1iva1ion of A1u111/asma hat- been the objecth·e
t of numerous srudies. ; 9 Only shon-cc.mi erythrocyte c-ullu re:;
have been successful, and i n these it was sho\\11 lhal cell-to·
cell transmhsron does,ocnir.90 Succesliful cultivation of the
organism in a 1ick cell line derived Crom l)ennacemor ,,,1ri·
at1ills embryos has been reponed.80 ~lore recently continu-
ous cuhi\'arion of the organism in uitro was achieved using a
tick cell line derived from lx()des sca1mlnris. 153 The mor-
phology and developmem cycle ofA margil/alein a tick cell
line cuhure were found to be similar to tho~e observed in
naturally infccled Oermt1Ci't1lor spp. tick.~.!4 • 93 Such a sys-
tem offers considerable advamages 10 future investigations
into the biolog) of Anaplnsnw spp.
Figure 47.3 U!trastructural morphology of Anapli!SfTII! 1718tg,nafil
Epidenliology
rxodicl ticks are the principal biological vectors of anapla,.
mosis, but 1he argasid tick Omithodoros sa1,jg11yi, can alm Anaplasmosis has a \'ery wide dis1ribu1ion in the world and
transmit A mnrgi11r1le. 11111 Several smdies have been under- appears to be endemic In mnst of the cattle-farming areas in
taken 10 determine the developmemal stages and morpho- ~omhern Africa. The occurrence of anaplasmosis is s1ill
logical chnt11c1eris1ics of A. marginflle in adult o. f111tlerso11i spreuding, and in count rics such as France it is a no1iliabiii
, ... 161 ~(1
ticks. ' Small electron-dense forms. larger reticulated disease. It is not endemic in Canada bu1 occasional out·
fom1s. pleomorphic fonns and small particles have been brea~ do occur, follo1,1ng which ii is apparently success·
identified in 11\'e morphologically differenr colony types in I.he fully eradicated. a1 high cost. by 1h.:, slaughter or
midgul of these ticks.00 Similar developmental smges have seroloiit'ally positive ca1tle. 1ua
been recoi,rnir.ed in Rhlpicep/1a/11.~sim11s infected with a South tn South \frlca. 1he role 1ha1 specific ticks play in the
.\frican isolate of A. 111argi11ale17Y and. althoug/1 the complete transmission of the disease has not been e~umsh·ely stud·
life cycle of the parasite and i1s mechanism of transmission br ied. The one-host tick B. deco/omr11s has been incriminated
ticks are no1 fully understood. a comple:x developmental cycle. circumstantially as being the most important vector.2 1 sim·
bas<.>d on these sn1dics. involving several morphological stages ply because occurrence of 1he disease is thought 10 correlate
ha., been proposed.~1 with che dis1ribu1ion of this 1iek. However. four other tick
Theiler in 1911231 was 1he first 10 report on tick transmis· species ;8. 111icropl11s, R. simu,. R. e. e1·errsi and H. m. mfipes)
sion of bo\1ne anaplasmosis. I le ~howed chat 1rnnsovarial have been shown eiq,erimentnlly 10 be capable of transmit·
transmissron of I.be infection by one-hos1 and three-host ling the infection,174 • 175 and their distribution largely over·
ticks could take place, although subsequently. the majority laps that oi 8. decolomws. 81 These rick:. are therefore also
of a11emp1s 10 substantiate this observation. including more considered 10 be involved in the epidemiology of anaplas·
recent anempts involving Boophilus micropu,.s in Australia. mos is in Soltl h Africa.
D. m1derso11f in the USA. and five differem South African tick In Australia. 1he epidemiology of the dlsease is less com·
species CB0011hilus decoloratus, B. microplus, Rhipicep/ialus plex; only a single vecror. 8. microµ/11~-. is recognized.
e1,ercsi ernr1si. R. simusand Hyalomma margiuawm rnfipes) The transmission of anaplasmosis in somhem Africa is
.have fail11d. 11 98• 17~• i,;;. 166 The few ,;ucc:essful auemp1s re- apparently comple~ in the s<m~I! thnt sc,me of the tick vec
ported in 1hc Hieraturc are generally regarded as being ex- 1ors studied. such as the three-host rlck R. sim11s, which has
ceptional.10 It is bclie,•ed that transmission mainly 1akes been identified as being an efficiem transstadial \'ec1or of
place transsiadially, although inirasladial transmission by both A. mnrginflle and A. ce111m/e in the laboratory, do not
inurrhosr transfer ri.e., normal feeding l)eing in1errup1ed on feed on caule under narurnl conditions except in the aduh
one host and cominucd on another) has also been shown to s1agc. Hyalomma m. mfipes 1.aivae and nymphae will not
take plate in the one-hos1 tick Dermflctlllor nlbipicws. 222 feed on cattle at all. yer intrastadial 1ransmi&.ion br adults
 Bovine .anaplasmosis 59i

has been demonstrated. ·\1temp1s to prove thm transstadial sites may remain \iable and infective in arthropods for at
transmission occurs in the two-host tick. R. e. e1•ert${, have least two to three dars after ingestion.201
failed. but again imrastadial transm ission with aduh ticks Some insects that have been incrimin ated as mechanical
removed from clinical field cases has been successfully ac- vectors include biting 11)' specie~ of the genera T11ba1111~.
compli~hed in the laboratory. The two one-host ticks B. <ie- Chrrsops. and Siphom and mosquitoes of the genus P,mo-
colorarus and I). mit:roplus. contrary to what would be 11lwrt1.19~ It has also been shown cxpcrimcniall) the.t calves
expected. are incapable of transovarial rran~mhsion, but can be infected wi1ht1 margina/evia the ocular rome.~8 and
both transsiadial and imrastadial transmission are It has been suggested that eye gnat:. (Hippfllaws spp.J may
possible:12, 1,a. t~. tis&. 21~ rrnnsmit Arl(lp/asma-mfectcd b lood to the eye of a susccpt·
The transmission of anapla~mosis b)' the imerhost min~- ible animal after they have visired bloody horse fly
fer of larvae and males or 8. microplus In the field may be wounds-48 !n parts of southern Africa bleeding skin ,vound,
importam139 because B. microplus males may surv1\·e on caused by the filarial worm Pnmjllaria bo1•icola a11d the be-
cattle for periods excet!ding two momhs. u; Rhipice11lltl/11s haviour of face flies. such as .\/usca xa111homeles. •\I. lusorta
simus males (infected as nymphae) wer., s°hown to have re- a nd 1\ L nevi/Ii, which lap tl1eir meal$. and also possibly other
tained the infection after havi ng fed for nine days on a bo- /\.Jusca spp.. may be significant in rhis regard (see Chapter 3:
vine.18J Male ticks of D. a1uierso11i. fed on a chronic carrier Vectors, Muscidae).
cow, subsequently transmitted the infec1ion lntrastadially In the high semi-arid region of eastem Oregon rn the
and biologically 10 susceptible cattle. :i.;~ USA. D. m1ders.011i has been sho\,11 to be the only wctor of
The time required by feed ing tick$ to transmit the infec- a naplasmosis. The disease was not mmsrnitted from carrier
tion seems to vary between species. Adult D. 1111derso11i have cattle by any of tht' haematophagous insects which arc
to Feed at least si., or seven days before transmission takes presen t there. 17 1 In Mississippi. however. Culicidaeand Ta·
p lace.~ while aduh R. simus may transmit the infection banidae were shown to be potemial vectors in the absence
within 24 hours afcer infestation. 185 of tlcks.m Observations m;ide on a dairy farm in Tanzania
.~ot all Isolates of A. nwrgi11a/e are readily tick 1Tansmis- incriminated Tabmws f(le11iolaas the principal mechanical
sible. The A. ctmtra/e isol111e contained in the commercial vector.?• 1 but. iu addition. frequent outbreaks of annplas-
va~cine produced by the Onderstepoon Veterinary Institute mosi~ were apparent)) caused by needle transmissio1i.
t'11n be transmi tred b>' R. si11111s, but not by 8. 111icrop/11s or There is linle e,idence to suggest that anaplasmosis is corn·
8. dl?CO/oraws. 186 whereas both the lauer species are ca- manly transmiued bi• biting flies in Australiai 1"
pable of transmining A. marginale. In soutl1em Africa. the mechanical tm1isn1ission oi t111a-
Extracts oi ground-up infected ticks from a ,c1riery of plasm11 spp. has not been studied in any detail. The seasonal
tick species and salivary gland e.xrracts. bu1 not salivary occurrence of the disease and the prC\'llhmce of Lhe cattle
gland secretions. are infective when inoculated into louse fly (Hipp/Jbosca rufipes) in Namibia suggests that thi~
susceptible animals.99 species may be a mechanical transm itter of anaplasmosis.
:-.1either the precise mecha nism involved in the transmis- However. while a laboratory investigation failed to demon-
~ion of the infection by ticks nor the Infective stage of rhc straie 1hat this fl) can transmit the infection, a similar in-
parasite has been identified with certainty. Oe\'elopmemal \'CSt igation with the smble Oy (Sromo.\)'$ C(l/citmns) S\IC·
colonies of A. marginale ha\·e been identified in the sali\'ru')' cecded.111:! In Zimbabwe It is believed that mechanical
glands of three Dermacen tor spp. by fluorescent antibody transmission of anaplasmosis. afrer its introduction b)' in·
techniques and electron microscopy.99· to.1. a23 rccied ticks, may pl;iy a significant role in the e 0lidemiology
Anaplasmosis is relatively e:isily transmitted mechanic - of Lh is disease.1~;; 111 South Africa it i:; bcliPVCd that close so·
ally by needle passage or infected blood. and, in such cases. cial association of cattle, as in dairy herds and feedlots. pro-
the prepatent period or the infection is probably dose - motes mechamcal transmission - especially by stable tlie,.
related.195 Circumstantial evidence indicates that some once a tick-transmim:d outbreak has occurred. Other hae-
anaplasmosis outbreaks have followed vacdnation pro- matophagous insects may be involved in this region. but all
grammes in Somh Africa. incriminating needle transmis- of them probably play a lesser role than do licks.
sion between animals. Instruments used in veterinary Losses as a res uh of anaplasmosis occur under both ex-
procedures. such as castration. dehoming and ear-tagging tensive and intensive farming conditions. but dairy cattle
operations. which are contaminated with fresh blood. seem to be more prone to serious disease outbreaks. This
may also be responsible for the mechanical transfer of the may be due to the clo.se dail) observarion of dairy cat1le
infection.53 which makes this difference more apparent than real.
Reviews on the mechnnical transmission of anaplasmo- Caule reared in endemic .ireas develop a naturally ac-
sis by blood -sucking arthropods have been published.5 ~· in . quired immunity to the disease. Calve~ under the age of 5ix
2.3-i, 256 IL is be!ie,, ed lhat successful mechanical transmission months. irrespective of the immune status of their dams, are
can only be achieved where there is minimal time lapse (a resi.s tant to ~e\'ere clinical dl.$case. 1~7 Cross-bred rattle in-
few minutes) between consecucive feeds. although the para· fected during the first year of life suffer none oi the clin ical
598 =os rtt11£>: Riekeusial and chlamrdial diseases

effects of irifection. 169 but the risk of clinical anaplasmosis Africa since I912"3? is believed to have played a major role in
increases \,ith age.85 Caule raised in endemic areas there- the countf)Wide distriburion of this organism. It was also re-
fore have the opportun ity to develop immunity to the dis- cently shown that this isolate is readily transmissible by
ease before the age when oli nical effects become severe. ticks. even after 70 years of re.gular needle passage in the
Bos indicus cattle breeds appear to possess a greater re- laboratory for the purpose of vaccine producrion. 186 This
sisrancc to A. mnrginale infection rhan Bos uwrus breeds. mildlr pathogenic organ ism competes with :he more ,iru·
but ,·ariatfon of resistance of ind ividuals \\ithin breeds of lenr A. margin11/e In in\'ettebrate and vertebrate hosis. and
both species may well occur.'6 Cattle of the Boron breed possibly play,; a significant role in the epidemiology of ana-
have been found to be more resistant than Ayrshires main- plasmosis.
tained under rhe same experimemal conditions. 130 al- In endemic areas, clinical disease in herds general!~· de-
though there is a na tural variation in indh"Jdual resistance velops only in the older animals that. for whate,·er reason,
becween animals, even amongst those of the same escaped Infection as calves. Furthermore. the movement of
163. 19,
b ree d. cattle oiten leads co the unintentional inrroduction of sus-
The nutritional stams of animals may play a role in the ceprible ca11le onto propenies where. because oi the exisr-
severicy oi the cli nical disease. Catlle on a reduced energy ence of a stable or near-stable disease sin:arion. clinical
i make apparently show less severe clinical reactions than do anaplasmosis is never observed or diagnosed. and the prop-
those receiving a normal plane of nutrition.= while those erties are therefore thought to be anaplasmosis-free.
on a higher plane may. in general. be most severely It is frequently the case that cauJe of unknown immune
affected.4 status introduced into a 'clean' herd can be a source of in-
In one laboratory study, n 15,6 per cenc prevalence of in fected ticks or can act as reservoirs of the infection.
utero transmission of A. 111argi11aleandA. ce,ura/e infections The role that factors such as differences in parhogenicity
occurred in 77 cakes born of75 infecred cows. 187 ::-o clinical and immunogeniclty of different strains of Anap/asma spp..
signs of disease were obsen-ed in any of !he calves prior to stress. and concurrent infections p lay in the development
splenectomy. which was performed when they were a few and manifestations of clinical disease has 1101 been ad-
months old. It is probable that the frequency of in 1uero equately studied.
transmission is underestimated. Investigation into l11e role that possible reservoir hosts,
Animals which recover from anaplasmosis usually be- orhcr than cattle, play in the epidemiology. has a greater po-
come life-long carriers of the parasire.33·S9· 172 Autosterili7.a- tential in ~ubtropical regions with abundant game. as in
tlon tenders animals susceptible to infection again \\ithin pares of southern Africa. than elsewhere. It has been shown
several months.n because the clearance of parasites from that biesbok (Dama/lscus dorct1s philllpsi , common duiker
the bloodstream is followed by the disappearance of anti- (Syhticapra grimmia L), and black \~11debeht CC01111ochaetes
bodies to the parasice. 82 grrou) are susceptible to experimental infection wilh A. mnr-
n,e level of immunity which de,·elops after pfimary in- gi11ale. but that iniections are subclinical. t35• 158 Blesbok are
fection apparently depends on the degree of the antige1tic also susceptible to A. c:enn-ale infec1ion.' 06• 158 Based on
stimulus during !he primary reaction;253 the response is, lhese obsen1a1ions. it was concluded that anaplasmosis is
however. inconsistent. 1&1· 245 Anaplasma ce111rale does not primarily a disease of amelope. which serve as resen·oir
afford complete protection against A, mnrginale, io;, lM, 21 0, hosts for A11aplasma spp.
2$3 and there arc antigenic diITerences between some iso- A11aplasma spp. ha\'e been recorded in a variety of African
lates of A. margi11ale. to3 game species. such as giraffe (Gimffn came/opardalis), 1..3. 3 1.
Natural infection involving all animals during calfhood i:is sable amelope (Hipporragus 11iger).z33 buffalo (S)'IIWIIS
in endemic areas will ensure an ·endemic stable disease cajfer), and black \\~ldebeest. 177
situation·. However. low or fluctuating vector populations
and limited parasite reservoirs cause va11~ng numbers or
Pathogenesis
susceprible individuals in different age groups 10 be present
within a herd, resulting in 'disease O( endemic instability', After primary mfect!on. there is a prepatent period that ter-
Disease stability under these conditions can be achie,·ed by minates when the first infected eryrhrocytes are detected in
a long-cerm commitment 10 the \'accinarion of all calves blood smears. This is followed by a phase during which
born in the herd as well as any cattle thar are introduced. there is increasing parasitaemia and anaemia. and iliac ends
• In geographically defined areas that are marginal for either in death (preceded by e,~dence of severe anaemia and
vector-tick survival, changing climallc condirions that en- other dinical 5ignsJ or convalescence and the de\'elopmem
able successful. but often o nly temporary. establishment of of a carrier state.
vectors may lead to disease outbreaks. E.-,ceptionally wet ~eit:ler the minimal infective dose nor the location of
\Veather condirions may lead 10 epidemic spread of anaplas- the parasite during the lncubarion or prepat.enl period has
mosis and oilier tick-borne diseases. 181 been determined. The duration of lhe prepacem period
The use of the A ce111rnle live-blood vaccine in South seems to be directly related to the number of organisms in
 Bovine anaplasmosis 599

the inoculum. body weight of the host and route of fected and altered erythrocytes sensitiZ"e them, thus aiding
infection. Generally. prepatent periods are 5hortest after in- their removal b}' the RE S)'Stem. 1o.i. 1' 4 Atnohaemagglutina·
travenous inoculation. longer after imramuscuJar inocula- tion in blood specimens of Anaplasma-infected aanle is ef-
tion. and even longer following subcutaneous llljection. fected by imm;.moglobullns. which react with e~"l.hrocytes
Artificial needle-induced Intravenous infection~ using large but not with Anapfasnw antigens. 138 Anti-e~1hro<:}1e anti-
volu:nes [4 litres) of infected blood can result In parasi- 1.>utlies am pn,sent in the cin:ul11.1iu11 m, 1.>u!l1 t:f) lhrocyte·
taemias of up 10 88 per cent within four days. The prepatent bound and free antibodies. 211 Apparently, bound auco-
period can be as long as three 10 five wecks 13 ; and. in excep- a.ntibodies are able to attach 10 nonnal uninfected
tional cases, it may exceed three months. The course of che erythrocy1es. '" 211 • 21 z The discrepancy between the num-
infection is not affected by the size of the inoculum. 106 but ber of infected el')-rhrocytes lost and the total loss oi e~1-hro-
prepatency may val)' even between animals which receive cytes ca1-1 probably be explained by an autoimmune
the same Infective doses. 175 In spite of this. the number of mechanism whereby normal erythrocyteS" are also re-
t\naplnsma organisms in live-blood vaccines is manipulated moved.211 Free antibodies developing during the course of
in such a way that the desired prepatency is achieved so that che disease do not opsonlze er)•throc~-ies and their signifi-
a reasonable synchronization of reactions results In those cance remains unclear in the pathogenesis of anaemia.213 A
animals that will react clinically co the vaccine. Similarly. in factor that stimulates phagocytosis nonspecifically. and that
the manufacture of frozen vaccine, adjuscmems can be is not produced in the spleen. has been detected in cattle.
made co the number of organisms per vaccine dose co com- This factor increases fh·e· to ten-fold at the time of the
pensate for loss in viability after Freezing and tha\\ing of the anaemic crisis in anaplasmo~is. 62 It has been suggested chat
infected blood.17J Prepatent periods may be prolonged after the phagocytosis of infected e~'lhrocytes is limited by the
administra1ion of retracyclines.114 clearance capacity of 1he RE system. 1~ 85
The in.tensity and duration of parasitaemia and the re· The rate of e~'!hro~,e loss and the level of parasitaemia
suiting anaemia are direct!}• related to the age of the host. 85 during the period of increasing anaemia prior to the detec-
Increasing anaemia may develop over -1 co 15 days, and a11i- tion of reticulocytes in the circulation are usuallr closely re-
mals may lose up to 70 per cem or their circulating erythro- lated. faidence of a haemopoietic response in the
cytes. ) laximal anaemia usually occurs one 10 six days af,er peripheral ciraula1ion, which is a favourable prognostic
the peak parasitaemia. In animals chat survive the patent sign, is first apparent Just before the occurrence o( ma,i·
phase, convalescence begins with e\·idence of ef)1-hropoie- mum anaemia 85
sis. Com·alescence usually lasts one to two momhs. but in A11n,,las111n margi1111/e-infected bovine crythroc)1-es have
aged cattle as well as in splenectomized animals it may be altered phosphofn1ccokJnase activity and vef)' low levels of
prolonged {three or more months) due to recrudescence of adenosine trlphosphate (ATP), r:r. although the giycol~-iic
parasiraemia. pmhway does not $Cem to be affected. These phenomena
Although the manner b~· which erythrocytes become in· may indicate that Anaplasma is reliant on ef)1-hroq1e ATP
fected ~ not dear, an electron microscopic study has re- for its sun~val and muhiplica1ion. As ATP Is critical in the
vealed how initial bodies penetrate the erythroqiic physiology or ery1hroc~1es, its deplerion would cause severe
membrane.200 1t hai; been suggested that a direct transfer of dismrbanc:es to their functions.
initial bodies between erythrocytes takes place.200 In· The membranes ofinfec1ed erythrocytes are altered \\ith
cren,Pd nsmmic fragiii1y. pm<~rlP.d h}•a d~rrease in erythro- regard to their protein and glyrnprotein cqmponenis.159
cytic acety1cholinesterase has been observed In bovine which funher lndica1es that the metabolism of infected
anaplasmosis. 216• 2 47 This indicates that changes in rhe ace- e111hroC)1es is impaired. Lectins form cross-linkages with
rylcholinesterase activiry ma)' be associated with increased receptor molecules of cell membrane!> leading 10 agglutina-
cell membrane permeability, which in tum facilitates the tion of in[ecred as weU as non-infected cells. but agglutina-
entry of A. marginale organisms into red blood ceJJ~. 216 tion is greater\\ith the former.- 3 This observation supports
The pathogenesis of the anaemia in anaplasmosis was the fact that 1he erythroc~'le membi:ane is altered b> 1he in-
originally thought co be due to destruction of the erythro· teraction 1,ith the parasite.
cytes by the parasites. However. the absence of haemoglobi- The nature of the resistance of calves to anaplasmosis Is
naemia and haemoglobinuria indicates that imravascular unknown. but it can be overcome by splenectomy. 50 lt ap·
haemolysis ~ not a feature of the disease.84 The anaemia is pears that the competence of the RE system and immuno·
caused by extensive ei:ythrophagocy1osis initiated by para- logical competeneeofindiViduni animals arc the facrors 1hn1
site-induced red blood cell damage and the role of anti· govern the outcome of the patenr stage of anaplasmosis in
erythrocytic auroantfbodies. 14• 27• 104 Er)"l.hrocytes thaE adult anlmals. 197 Greater immune comperence and the
become physically and chemically altered during the course presence of foetal haemoglobu Iin have been sua.,gested as
of the disease34 are removed by 1he teticuloendothelial {RE) possible factors associated wicll higher resi!>rance in young
cells in the spleen. Liver, lungs and associated lymph cauJe.9 !\ge-associated resistance in Bos indicu.~ cattle is
nodes. 2 B In addition, antibodies produced against the in· similar 10 that described for Bos taurus breeds. 163
600 ,(010, TI!flu: Rickcrtslal and chlrimydial dl~a~

Cmde infected with A. 111argi11ale.31! ~ and those to Although A marginale may cause abortions and neona-
which an inacti\'ated vaccine has been administered.' 9 · 2ij tal anaplasmo,i,.Z3 · 2 ~,s these arc rare. 18; Foetuses between
develop both humornl and cell-medlated immunity (CM!J. 100 and 140 days of gestation a re capable of p=oducing anli-
As C:'-1 1 is probably of panicular imponance in the protec- ~\naplasnw complemem fi~atio n antibody.23f
tive immune response. 18· 36• 39 and as ciremically modified
A11t1ptr1Sma antlgens stimulate mainly C:\11, this lauer
Clinical signs
method of producing inactivated vaccine has been pursued
in countries where the use of live ,•accines is prohibiled. TI1e se,·crity of anaplasmosis Is direcLI)• related to the age of
Primary exposure of 1be host",; RE system ro replicating the animal: in animals less than one year old It is usually
parasites provides the antigenic stimulus for the simulta· subclh1ical: in yearlings and rwo-year-olds it is moderately
neous induction of both CMI and humoral responses. 197 severe: and in older cattle it is severe and often fatal.$-! Dif-
Subsequent low-level parnsilaemia of the infection ensures ferences in virulence betwee11 A11ap/asmtl strains and the
continuous amigenic stimulation. l n the case of re-exposure leval and duration of the parasitaemia also plar a role in rhe
to the parasite. the memory 1·-lymphocyt~ are stimulated severity oi the clinical manifestatiOnb.6 Generally. cases are
10 proliferate and afford a specific protecth·e response. presemcd lor irea1me111 when the disease has ad\'anced 10
The current model for \·accine-lnduced Immunity the sragc of the anaemic crisis. Anaemia is usually nor cllni·
against A. marginale 1b• propose~ that clearance of rhe or- cally apparem until a loss of about -lO 10 50 per cent of red
ganism is effected by antibody against surface cpitopes in blood cells has occurred.86
combination \\tjth macrophage ac:th·ation for enhanced AJter a nacural prepatem period. which generally , 1aries
phagoc:y1osis and ki lling. Central 10 this model is 1he co4· between 15 and 36 days (\>\~th an average of 26 days). al-
T· cell expression of interferon ganuna (IFN··1J. which en - though It may be as long as I 00 dars, peracute. acute or
hances synthesis of the predominant opsoni1.ing bovine lgG chronic anaplasmosis mar follow.
subclass. fgG2. concomitantly acth-ating macrophages 10 Peracute anaplasmosis is the most severe fom1 of the dis-
increase receptor expression. phagocyiosis. phagolrsoso· ease. It I, rare but usually fatal. and It occurs most irequenti)·
ma! fusion and release of ricketLSiacid.al nitric oxide. in purebred animals and high-producing dairy cows. 195 Ani-
Carrier animals cleared of the infection b:, chemo- mals succumb within a few hours or the onset of clinical
therapy are apparently resL~tam to clinical disease following signs. in addition to anaemia, milk production ceases, and
challenge ex1iosure fo r periods of variable duration, which there is excessive salivation, rapid respiration. irrational be·
may be as long as 30 months.53 • 133 haviour and signs of nervousness.
Hu moral immunity may c:on1ribu1e 10 the maintenance The most oumandiog fearures or the acute form include
of die state of equilibrium In the host- parasite relation- pallor of mucous membranes. depression. inappetence. a de-
ship. and suppression of the humoral immune re.\ponse crease in milk production, general ,,·eakncss, and a rapid,ly Tis·
alters the course and ou tcome of infection in ,\1wp/as111a ing parasitaemia. In some animals. fever provides initial and
carrlers::s Rec rudescence of clinical anaplasmosis is well persistent clinical evidence of anaplasmosis:· s.. bm this is not
recogn ized. especially in dairy caule: age. con.dirion of the consistent, 1'"' as in others the recta l temperature ma}' remain
animal. environmental s tressors and the effects of hor- be!ow 40 •c throughout a fat.al am1c1<.·1a As the disease pro·
mones during pregnancy ;md lactation may be imporram gresses. constipation. rumen stasis. weight loss. dehydration.
in this regard De.~ame1hazone, a synthetic glucocorticoid. ic,erus. laboured respiration. a 1humping heanbeai and
suppresses acquired resistance 10 anapla,mosis and muscle rremors nre observed in severely affected animals.
causes recrudescence in carrier calves. n 8 It also sup- The faeces are frequently bile-struned. Diarrhoea may be
presses amibody production. and treated animab show manifested in the initial stages of the disease.llll Abortions may
more pronounced parasltaemia and anaemia than d<> occur in severe ca~es.23 • 43• ""· l&i. l~ij t\b11om1ally aggressive
comrols. 59 beha~iour. probably resulcing from the anaemia and resultant
Recrudescent parasilaemias are accompanied by an an- cerebral hypoxia. is also manifested hy ,ome animals. The ap-
amnestic responst' in both lg,\ I and lgG fractions. suggest- pearance oficterus indicates that the acute phase has passed
ing lhat the maintenance of carrier mfecdons is primarily and that convalescence has begun.36 but no Lall icteric animals
controlli:d by antibody response~. 2.;o Paradoxical!~. such reco,-er. The course of the acute dis<'.ase i$ generally prorracred
persistence of!nfectlon occurs in the face ofa protectil·e in)- and mar last for t\\'O weeks or more before there is any evi-
,mune response. suggesting u mechanism by which Lhc para- dence ofimprovemcm in the animal's condition.
site evades the immu ne system. Low parasitaemias in Chronic anaplasmosis is manife.,ccd by slo\\' recovery fol·
chronically infected cau le are known to fluctuate in ,;equen- lowingncute disca~e. and it may persi~t fora period ofberween
tial cycles 69 • 90 and recent evidence favours the mechanism two weeks and three monchs. 19j It is characterized by poor ap·
of antigenic variation.60• 00 rather than any significant wan· petite. lo!» of \\'eight and ,-arying degret-;, of dehydration.
Ing of immunir)', in explaining rickettsaemic cycles in per- anaemia and icceru~ which are usu all) milder than in the acute
sistent lnfect.ions.167 disease.
 Con\'alescence follo"~ngan auack of anaplasmosis is al· damage. haemosiderosis. erythrophago<"ytosis by the RE
ways slow, and a recovered animal may ta.kc several months system and hyperplasia of erythropoietic elemen1s in the
to regain its previous condition. Relapses associated \\ith bone marrow. The latter may show depletion in c:hrnnic
anaemic changes and fever may occur a rew weeks af1er ap· cases.84 The hepatic lesions consist offany infiltration. cen-
parent recovery.;-; trilobular and paracemral degeneration and necrosi< of
/\naplasma margi11alc infections moy Induce temporary hcpotocytcs. bile sro.sls. and proliferation and haemosidetin
infertility in bulls.8" anoestrus in heifer:;, 227 and abor1ions pigmentation of Kupffer cells. Haemosiderin may al~o be
and neonatal anaplasmosis following foetal iniection.2 ~· 160 present in macrophages of the lungs and lymph nodes.24n
The mortality rate, particularly in adult cattle of exotic Varying degrees of degeneration. necrosis and pigmentation
breeds, may exceed 50 per cen1.n (bilirnbin and haemosiderin) or tubular cells in the kidneys
may be presenr.l ll, 2 10 It is believed tha t the parench~mal
cell damage ln the liver and kidneys is attributable l O tissue
Pathology
hypoxia resulting from the severe anaemia;;· zis
Increases in le,·ets ofsemm urea nitrogei1 {SU~). asparrate The spleen shows congestion, hyperplasia of the red
aminotransferase (ASTI, total billrubin (TBIUJ and alkaline pulp, and an inccease in the number of plasma celb and
phosphatase (ALPJ occur at the onset of parashaernia in haemosiderin-bearing RE cells.it~. ion
acute infections. and these may be related tot.he destrucuon
of Cl)'lhocyte$ by the RE s~-stern and cellular damage in the
Diagnosis
liver and kidneys.5 · ; , 8 · 75 On 1he other hand, !c I~ thought
that the Increases of AST. ALP and direct bilirubin (DBILIJ The diagnosis of anaplasmosis requires a comprehension of
that occur or continue after peak parasitaemia has been not only the pathogenesis of the disease. but also its clinical,
reached indicate liver parenchymal cell damage, and thar pathological and epidemiological features.
large quantities of erythrocyte cataboli tes, probably in addi- A varie~· of diagnostic methods is available for rhe con-
tion to hypoxia, lead co liver cell degeneration and other he· firmation of the clinical diagnosis. In most cases confirma-
patic changes.8 A smdy on A mnrg/11ale- infected cal\'es has tion of a diagnosis depends on lhe microscopic examination
shown that the rapid destn1ction of erythrocytes, and the of convemionally stained thin blood smears In order to
concomitant rapid relea~e ()!' their contents could lead to demonsuate :\naplasma organisms in the Cl)'throcy1e, of an
imbalances in blood pH and electrolytes.;, Dramatic reduc- affected animal. However. microscopic examination of
tions in pH and conccntTation of HCO~ during the anaemic blood smears from sui.pecced cases of anaplasmosis is usu ·
crisis ha\'e been obseffcd in calves that have died of the all)' gerfom1cd at a time when the cllnical signs are most
disease. pronounced (I.e. at the height of the anaemic crisis) and by
Little has been added to the comprehensive description this time 1hc parnsimemia may be very low because of the
of the lesions made by Theiler in 19 I0.~8 The lesions are removal of most of the infected erytbrocyccs from the circu-
rypical of those occurring in animals that have sulfored from lation. In addition. the presence of a regenerat!'·e annemia
anaemia due to el)'throphagocyiosis. Prominem changes in requires careful examination of blood smears LO distinguish
fatal ca:.es include $evere anaemia. icterus. splenomegaly A11aplasma inclusion bodies from basophilic stippling in
and hepa1omegaly. Lymph nodes ma~· be enlarged, the car- reticulocytes. Ho\\'ell-Jollr bodies, and possibly other
cass may be dehydrated and, depending on the duration of srained particulate maner in ery1hrocyres. A posim·e diag·
the course of the disease. a lso em aciated with serous arro- nosis can be made if morphologically d istinct organisms are
phy of perirenal and cardiac far.. The sptenic pulp is dark red observed in the mature erythrocytes which are still prcsenr.
and has a meaty consistency. while the cut surface of the Subclinical carriers are difficuh to identify by this method
liver varies in colour from orange-brown co mottled yellow because of ,heir low or. more frequently, undetectable levels
and shows e\'idence of cemrilobular degeneration. The gall of parashaem!a.
bladder is distended and contains inspissated thick brown In a light microscopic demonstration of the intrae~"thro-
or yellowish-bro,,~, bile. There are serous effusions in body cytic parasltt:s by the examination of stained blood films,
c;a\'ities. pulmonary oedema, pctechial haemorrhages in rhe personal preference and experience play a role in che choice
epi, and endocardium, and often evidence of severe gas- of stain and staining procedure. Giemsa stain is very reliable
trointestinal stasis especially of the rumen. omasum and and is commonly used in laboratories. The commercially
colon. The omasal contents are dry and impacted, and lhe available Ditl,Quick stain ser has been found to be superior
colon contains hard. di)·, often bile-stained faecal balls. A to Giemsa and 1,·right-Giemsa staining methods in Lha: it
variable degree of red marrow hyperplasia may be present in takes only a very shon time co stain a blood smear and
long bones. The urine is yellow to dark brO\\'n due to lhe apparent!)' allows more accurate parasite coums.'16 Other
presence ofbilirubin. stains that may be used include coluidine blue,209 new me-
There are ie,,· reports on the histopathology of anaplas- thylene blue. 113 and acridinc orange (which requir~ the u~e
mosis. I listologically there Is evidence of hepatic and renal of an ultraviolet-light mic.roscopeJ .7°
602 ><CllO~ m~r., Ricken.slal and ,hlamydial di£eases

A direct Ouorescem antibody staining technique has !.he detection ofhumoral antibodies 10 A11aplasma, an agree-
beeo iound 10 be superior to the Giemsa staining method for ment or only 86 per cent was obtained. which was considered
the diagnosis of anaplasmosis at necropsy.Ill iliadequate.255 It is belicvGd by some that the CA test is the
Several serological rests have been developed for the simplest and most reliable test. whereas the CF remains use·
study of A mnrgi11ale infections. Their principal objective ful in experimemal siruariom;.m The Cl' test detects
has been 10 identify carrier cattle, mainly for regulawry rca antibodic.s sooner than the CA test. bm it does not detect
sons. but also for disease control. especially in chemothera- carrier,; for as long.
peutic eradication programmes (see Con1rol). Serology is Radioimmunoassa) (RIM for the detection of A. mnrgi-
?Jso \\idely used in epidemiological studie~ and 10 sneen 11all! antibodies. has been used. 21 ~ Optimal sensithiiy and
experimental animals and potential !he-blood vaccine do- specificity are obtained in lh is cest by the use of an ·I. margi-
nors for susceprlbllity to Annplasma Infections. in Somh Af- nale antigen and an erythrocyte antigen originating from an
rica the complemem fixation (CF) rest. and subsequently uninfected calf rn this test cross-rettctions may occur In
the rapid card agglutination (CA} 1es1, ha, e been used in caulc that ha\'e suffered from a B. bo11is infection. but ii the
anaplasmosis studies. while in ~aniibia a modified indirect serum samples 10 be tested are pre-absorbed \\/ith an extract
immunofluorescent antibody test (!FAT) was adopted in of 8. lioi•is-iniected el)'lbroc)1es. the number of cross-reac-
Studies on the epidemioloID of the disease.:?.?· ~:t 1ions is reduced. The use of the two -amigen test system is
Bo!.h soluble and particulate Anaplasma anrigens have the major iact0r m redu('fng problems associ111ed with
been isolated from infected animals and used in various se- cross-reactlvlty. \\'hen purified A marginale antigens be-
rological tests. Despite problems of non-specific cross-reac- come a\11ilable. cro-ss-reactions of !his narure should no
thity and lack of sensitivity. reasonably reliable tC5tS are longer occur.
available to diagnose anaplasmosis. u 3 • 131 :\ high degree of Species-specific D}U probes for A. mnrginaleand A. ce11-
correlation exists between results of the Cl' and the capillary 1rale have been de,·eloped2~3 for the detection of A. margi-
tube CCTJ agglutination rests. as well as between those of the 11ale infections in carrier cattle61 and in ticks,'• but ha,•e
CF and CA tests. Both the CF and CA tests are recognized in thus far been used mainly for experimental purposes.
the USA as official tests for the interstate movemem of caule
where regulations require a pre-shipment negative resi.
Differential diagnosis
Enzyme-linked immunosorbent as.say •ELISA) is al'CU·
rate and can differemiate caule free of the haemoparasite The clinical signs of anaplasmosis should be differentiated
from those with:\. mt1rgi11ale infcctions. 18 False positive re- from infectious and non-infectious condilior.s preseming
sults. however, hal'e been obtained from cattle that have ,dch fe,·er. anaemia. icterus. depression. inappera.nce. and a
been Inoculated twice with Bn/Jesia bo!'is-infected erythro- reduction in milk )~Cid. Similarly, on post-mortem examina·
cytes. 111e cross-reactions arc attributed to antibodies pro- tion, anaplasmosi,should be differentiated from conditions
duced against erythrocyte amigen,. 18 which primarily cause lesions attributable to anaemia re-
:.tore recently. a competith,• Inhibition enzyme-linked sulting from e~1hrophagocy1osis or to lcterus.
immunosorbem assay (Cl-ELIS:\) based on recombinam In southern :\irica, anaplasmosis is mosr c:ommonly con-
major surface protein 5 (MSP 5) which is conserved among fused clinically and pa1hologically with bovine babesiosls
...\nap/asma species244 has been del'eloped and shown to re- (redwaterJ caused by Bal,e.lia bigemi11a or 8. l101,is. Many of
liably detect both acute and chronic infections of A. 111argi- the epidemiological factors of thenvo diseases are vervsimi-
11ale in caule.49• 92 · isi. 189 Results suggested that the Iar - both di$eases are tick-borne, the respective causatl\'e
sensitil~ry and specificity of 1he test was high49 • 9 ~· 1:.1 and organisms share common tick vectors. and animals of dif·
highly comparable LO those obtained \\ith the C:\ iesr. 15 1 Cerent age groups show a simllar variation in susceptibility
The IFAT. adapted for use with microfluorometry, has Lo both diseases. Although the distribution of anaplasmosis
been successfully used for the diagnosis of..t marginale in- in southern Africa largely corresponds to that of babesiosis,
fections.-Z Its accuracy exceeds visual determinations and is anaplasmosis generally enjoys a far wider distribution. One
more sensitive than bo1h the CF and C:\ test, In identifying of the main distinguishing clinical feawres in babesiosis.
infected cattle. whiC'h is absem in anaplasmosis, is the presence of haemo-
The simple latex agglmina1ion (LAJ test. suitable for ~loblnuria in many affected animals. Haemoglobinuria.
practical field use. shows a high degree of specificity and however. is not a constant feature ofbabesiosis. parricularly
S!nsitivity when compared to the CA teSL IS? Culture-de- during the early stage of B. bo11is infection. ,\lthough dia-
rived soluble L.\. test amigens arc highly specific when corn· rrhoea, which commonly occurs in babesiosis. may be
pared 10 antigens derived from l:1fected cattle. In tests present in acute anaplasmosis, constipation and nnnen sta-
conducted \\"ith the laller a relatively large proportion of sis are more regular findings. The course of the clinical di'S·
non-specific reactions, which arc probably due to the pres· ease in anaplasmosis is often longer than that of habesiosis.
ence of bo,ine erythrocyuc stroma, are encountered. 152 Care should be taken as outbreaks of anaplasmosis and
In a comparison of the CA, lFAT. CF and ELISA tests for babe5iosis may oocur concurrently.
 Anaem ia ,md p rogressive weight loss are also clinical followed by the implementation and long-term maintl)·
signs often associated with Trypano.<oma spp. infections. nance of strict tick control programmes (to control it and
Trypanosomosis (nagana), however. general!)' occurs ocher endemic tick-borne diseases. such as hearll\'llter. ba·
\\~thin well-defined 1setse-infested areas. besiosis and anaplasmosis) is 1101 generallr recommended.
Some forms of rhclleriosis should. in some circum- Furthermore, eradication of anaplasmosis on the subcon·
stances, be differentiated from anaplasmosis Theileriosis tinem and mo,;r other parts of ..\frica is not practically po,-
can usually be distinguished by the presence of enlarged Sible for Ob\faus reasons. Stringent tJck comrol would
lymph nodes and the microscopic demonstration of render the cartle population sus,erHible co several other
schizoncs in l~•mph node biopsy smears. High piroplasm tick-borne diseases with po1emially cal!lstrophic conse·
parasitaemlas and anaemia are most commonly associated quences (see Chapter I: \lecrors : Ticks). as has been expe-
with clinical disease caused by Theileria nwmns. rienced in Zimbabwc. 126 The development of acaricide
Anaemia and icterus may a lso be encountered in cattle resistance by ticks is common and the cost im·olved In
suffering from lep1ospirosfs, pos1-parrnrient haemoglobin· high-frequency dipping programmes cannot be afforded
uria. bacilliary haemoglobinuria.89 chronic copper poison- by many farmers in the region. Further complicating fac-
ing and intoxications with Qrassica and Allium spp.ss. 89 In tors are rhat populations of \\ild ruminants ~u~laln rick
these conditions. haemoglobinaem ia and haemoglobinuria populations and that some are probably reset'\·oir hosts.
a re usually also presem. 89 Causes ofhepatotoxlclry. such as but much pertinent lnformailon concerning these aspects
Lantana camam and Mit:rocystis aerogino.sa poisonings. in is stlll laeking.
which icterus is usually one of the principal signs. should For these reasons it Is generally recommended that a
also be considered. stable disease situation be pursued in the southern African
Changes in behaviour. such as aggression, occasionally region by allowing natural exposure of cal\'es 10 tick-borne
manifested by cattle suffering from anaplasmosis may also diseases. including anaplasmosis, during the period when
be part of the ctlui.cal signs associa ted wtth several olher in- they are naturally resisrant or protected by passively ac-
fectious dbeases (e.g. heartwater. cerebral babesiosis. cere· quired maternal antibody. :\s far as anaplasmosi; is con-
bra! thellcriosis and rabies) and non-infectious conditions cerned. the existence of epidemiological factors which
(e.g. lead poisoning). inhibit the exposure of calves to infected tkks \\~thin the
relevant period can be compensated for by the administra-
tion of the A cemrale ll\'.e-blood vaccine. If this comrol
Control
strategy is followed. namral or art.lficial manipuiation of
Corurol strategies vary greatl)' In clilierem coumries and tick populations. or any other factor which may lead 10 an
geographic regions. Generally, the effectiveness of a strategy unstable disease situation. \\111 generally not result in dis-
in a specific region depends on a thorough knowledge of the ease outbreaks.
epidemiology of the disease. hue it is [requen rly rhe case that Vaccination strategics differ in accordance with \'Cteri-
the scientifically based measures are constrained by such nary policies and avallabillt}' of vaccines. In some states ln
factors as legislation. available infras1rucrure. funding. and the US,\. for example. chc use of live-blood \'accines is pro•
the le\·el of management. for these reasons no univer~al hibited. hut inacti\'8ted \'8Ccine is available as an allema-
comm! programme can be advocared. 1lve. Cercnin other states. however. hm·c been granted
Following the imroduc.ion of anaplasmosis. whether by special licences to use live ancnuated A. margina(e\·accines
infected vector ticks. cattle or other reservoir hosts, into re- for the conirol of anaplasmosis.
gions complerely free of the disease and where an epidemic
amongst I.he susceptible reside111 cartle populations could Immunization
be expecred. eradicarion of the disease is Indicated. This is The oldest and probably still the most effectl\'e and \,idely
achieved by identifying i.nfec1ed caule by microseopic ex- used method oi prophylaxis against anaplasmo$is is \'acci-
amination of blood smears or serologically. Such animals nation \\ith Theile(sA. centmle isolate. 18'1• 208 · z; 3 However,
are then either sterilized by ime nsi\'e drug treatment. or are A cemrale. although generally a mild pathogen. can cause
slaughtered. Concurrem Stringent vector control measures severe clinical reactions follo"~ng vaccinacion.io
to ensure eradication of lnfecied ricks and 10 pre,·em me· b1 South Africa the original isolate of A cemrale, identi-
chanical spread of the infection must be implemented. In fied and used by Theiler as a vaccine for the first time. is still
order co eradicate the disease successfully. a i"Ophistieared used to produce a live-blood \'BCCine which is excensilely
veterinary infrastructu re, the support of releY,1111 legislation used. For many decades this vaccine was only avaiiable as a
and the a\'ailability of ample funding are essential. Regula- chilled (4 'C) \'l!Ccine of which the dose was 2 ml per anunaJ
lOI)' measures should also be implemented 10 prevem the regardless of age and was administered subcutaneously.
subsequent reintroduction of the disease. However. this chilled vaccine has a limited shelf-lifeY~
In southern Africa. where a naplasmosis is endemic in which prohibits its use in countries or regions that lack an
many of the important cattle farming areas, its eradicarlon efficienc cold-chain distribution network.
604 <1r.no, ruw: Rickeusial and (hlmnydial d~cas\!..

The limhed shelf-lire of a live tl1111plasm11-blood vaccine :.heep has been accomplihhed. 1119 Thb strain. which hll$ a
has prompted the development Qf fro,.en vaccines (stored at \'irulencesimllar to 1hat ofA. (.t>ntrtlle and produces a higher
temperarures lower 1han -80 •con dry-ice or in liquid n.itro- level or immunity agains1 \irulen1 A. 111arg11wle challenge
gen) 1,·ith extended periods of usefulnCS$. 140· 173 This offers a than does A. cemrale, 111 has been successfully used for vac-
practical method of ensuring vaccine a\'ailabiliry in regions ~ination purpuses in cattle in Mexico, 1~ Colombia{ 1 2~ 5 and
or countries where continuous producrion of chc chilled C:alifornia. 78 rloweYCr, in /\rgcntim1 11A. nmm,le js preferred
product is labour intensh e and no1 cost effective. Infected because of se\·ere vaccine react ions experienced with the at-
blood cryoprcserved with glycerol remains highl)' Infective tenuated A. mnrginale vaccine.
after tha•.,1ng. despite dilut ion and incubation for eight TI1e prolonged majmenance of a virulent A. mnrginafl'
hours at 30 ' C.87 field isolare in a splenectomized white-tail deer (Odocoi/eus
In South ,\frica the use of the chilled \'accinc has bcc.n re- vfrginlanus) has apparently hecn sufficient ro induce at ·
placed by the use of a deep frozen vaccine that is distributed tenuati011 oft he organism for ca ule. 123 Vaccination of ca Hie
on dry-ice and administered intramuscularly at a dose of with this attenuated parasite protects agaim:t a \'irulent het-
I ml irrespccrive oi the age of rhe ahimal. This deep froien erologous ..t marginnle challenge.
vaccine may be used for up 10 four hours after slo\\ thal\ing Similarly. some ..\m1plnsma isolates from other wild n1-
in melting ice. minant, may also produce only mild infections in cattle.
A lyophi!Jz.ed, expenmemal live-blood ,·accinc has been suggesting a loss of virulence afl<:r prolonged mamtenam:c
developed in Sou1h Africa. which is more siable than the fro- of A. marginale in the wild hosts. 11 '· l?9 There is some e,;.
zen \·accinc and has the advantage that ii can be stored at dence to sugge:.1 that such auenuated A. marginale isola1cs
-20 •c for extended periods \\1thout loss of effecthity. 1:"11 For may regain virulence upon passage in caule. I.!9ou the other
pracric-dl purposes this vaccine unfortunately still has to be hand. cattle arc frequcndy not susceptible to infection with
distributed on dry-ice or in liquid nitrogen. anaplasms from \1ild ruminants. or the)' develop onl) ~ub-
As young cal\'es do not react severely. it is generally nd- dinical infections that do nm p rovide immunity 10 \'irulent
\~Sed that they are vaccinated before or at approxumucly six .-\. 11111rgi11nle challenge. indica!lng the possible existence of
months or age. Jn older a nimals more severe reactions ca11 unidentified A1111plasma spp. that arc host-specific or that
be expected and dose supen.ision of them is indicated . are possibly more closely rel:ited to A ovi.s. 11•
Pregnant cows should preferably not be l'accinated a, the}' Subsequent 10 the establishment of long-term cominu-
may abort. The incubation period following ,•accination ous in 11ilro cultil'ation or A. mnrgi11nle in a tick cell line, 1• 3 it
usually varies from five 10 seven weeks. Vaccinated animals ,vas shown that culture-derived organisms retained all cur-
..,hould be clo.ely observed for clinical signs ofanaplasmosJs rent candidate diagnostic and vaccine antigen~as well as in·
during this period. and should. if n<>cessary, be speclfically fecth;ir ior cattle. thus providing a \'aluable altema1ive
and symp1omatically treated. In vie\\' of the possible reac- source of organisms for the de\'elopment of diagnostic tests
tion to vaccinat ion in old er animals, on ly a limited number and vacdnes. 15
of animals ~hould be immuniz~d at any given time. fhe current anaplasmosis vaccines rate poor!)' against
The use of an inactivated .-l margi1wle ,·accine prevents the concept of an ideal vaccinc. 1il-l By definition the laner
serious clinical disease.26 However, the need for t\\'O initial sl\ould be a so-called dead vaccine. have no attendant risk or
inoculations, followed by periodic booster injec1ions. may inducing disease. afford lil'c-long prolcnion. neutralize
result in fatal neonatal isoeMhrolvsis in cal\'es born to vac- 11a1ural infcc1ion. and prevent a carrier s tate from de\'elop-
cinated cows.5 1 It has been sugge~ted that rhe use or an in- ing, thereby removing the ~ou rte ofinfection for \'ectors. In
activated vaccine of sheep origin may avoid eliciting 1he addition, ease or production. long shelf- and stoTage-life.
neonaial isoerythrolysissyndrome.:r. Caule vaccina ted with and affordable cos1 are hnponant elemen1s which ,,ill also
this vaccine are still highly susceptible 10 cl1allenge \\~th het- determine the success of such a vaccine.
erologous strains.124 Such vaccines are not 3\-ailable in Advances in molecular biological techniques have
southern Africa. s1imulated auernpts 10 produce a subunit vaccine for ana-
A method by which a minimal infective dose ofA. margi- plasmosis ba~cd on the presence of an antigen common 10
11ale (obtained by diluting heavily infected frozen blood sta- all Anap/asma isolates. Such a common antigen. referred
bllates after thawing and prior to inoculation] is used to 10 as the major surface protein complex MSP-1 (prel'iously
reduce the severity of the clinical reaction follo\,ing vacci- designated AMlOS). compo~cd of t\\'o protl'ins. ,\~1Fl05
'!adon has been successfu lly applie<l under experimental :rnd Al\1!'100 .pre\iouslydesignared AM 105U andA:'.\11051~
condirions.n;. ; 4 ln field trials it was established that the respectively), has been shown to immunize cattle agai11J.t
minimum infective dose for A. mnrginale is 106 parasites homologous and heterologous A 111argi11ale challenge. H I
follo11ing Intravenous inoculation; the same dO$C i,.how- 16~ The genes coding for both proteins have been cloned
ever. not iniecth>e when administered subcurnneously. and expressed in Escherichia coli and arc being characcer·
The attenuation of A. 111argi11nll' by irradiation of a viru- teed for possible fumre c:ons1ruc1ion or a recombinant or
lent srrain and its subsequent serial passage in deer and s~111he1ic replica of ;>..fSP-l which could be used as a
 Bo\·ine anaplasmosis 605

vac<"ine.16· Hl A second potential subunit \·acdne candi- other hand, ha1 e been approved for commercial sale !n some
date. :'\·tSP-2 Cprn\iously designated A>-136), ha-; also been countries. These two drugs are primarily used as antibabe-
identified and this similarly proceccs cattle against both ho- sials.2·:i· t ts. :ua. 238 The carbanilides are usuallr administered
mologous and heterologous challenge.166 parenterally. and at normally recommencll!d therapeutic
dosage le\'els do not srerllize A11nplasma infections in
Treatment cattle. z. ~ :u. 5? !O'l t 2. 1,n
1
:

The trearment of clinical anaplasmosb in cattle requires. There is some controvers) about the desirability of treat-
firstly. the suppression or elimination of the parasite (spe- ing acute anaplasmosis al the time of maximum anaemia.1°
cific treaunentl and. secondly, the alleviation or prevention Much of this controversy is related to whether or not an ani-
of secondary complications and rhe hastening of recovery mal wlll recover spontaneously, and the conflicting opin-
(symptomatic or supportive treaimem).81 ion$ which have been expressed range from those who
Secondary complications. generally the result of tissue consider treatmem to be beneficial at this time to those who
hypoxia. are a frequem sequel to anaemi~ and in acuce ana· feel that the excitement and exertion incident to treatment
plasmosis the se\·erity of this anaemia .~. to some extem, are more likelr to be detrimental to the animal. At the time
proponional 10 the extem of the pamsitacmia_lQ. M.ns How- when increased erythropoiesis becomes e\'idem. spt!cific
ever. e\·en in animals \,~ch relativelr low-peak parasitaemias treatment probabl) has little. if any, effect on the course of
(below 10 per cent). or in animals \\ith persistent low 1he disease. since at this s tage thepara~itaemia 1s in any case
parasitaemias CO.I to 2 per ceml. a significant anaemia may rapidly decreasing as a result of the phagocytic removal of
develop if they are not treated.:n ~3 ' rherefore, specific Infected eC}1hroC)1es, and spontaneous recover) is
treatment early in the cour~c of infection is desirablc,32· ,;;. frequent.30 Specific treatment with chemotherapeutic dmgs
63, 8 1, us. l<6. 147 t,:n. 196• 197 and such treatment prior to the is therefore probably only useful when the parasilaemia is
development of a high parashaemia or the onset of severe incre-ising.
anaemia is considered essential for a favourable The tetracycline drugs cffcctil'el)' inhlhh the multiplica-
outcomc.115· 250 Oiten. however. by the time that acute ana- tion ofA11aplas111a in e1ythroc)1es and ha\'C been \\idelyused
plasmo~is is diagnosed the level of parasitaemia b already for the treatment of anaplasrnosis. 2IHIO. 47• bJ. 10, 14- 1~ 197
declining asa resull of the phagocrtic removal of parasitizcd Single imravei1ou$ administrations of chlorcetracydine m a
erychrocytes. and the animals are ~uffering from severe dosage rate of 11 mg'kg body weight. or single parenteral
anaemia. 30 NevertheleS!t. cvrn when rrcarmem is slight!~ (subcutaneou,. fmramuscular. or intravenous) administra-
delayed, specific rreatme111 i$ more beneficial than support- tions of ox) tetracycline at dosage raws ra11ging from 2,210 6.6
ive treatment.147 mg/ kg bod~ weight have proved effective in reducing ,.\11a-
plasma parasitacmia and alle\iating clinical signs.1:i. 14:. Ha
Specific treaunent I lowever, when cmplo~1ng a standard or short-acrin.g ,SA)
Prior to the development of the cctracycline drugs. a variecy oxywtracycline fomrnlation (contalning 50 10 150 mg/ml
of chemo1herapcutic agenrs. including ar.,,enicals, antima- concentration, or the drug), tetracycline or chlorcerracrcline
larials. antimony deri\'lltives and dyes. were used to treat for the 1Tea1mem of clinical anaplasmosis, two to three injec-
acute anaplasmosis. but these had little. if any, chemothera- tions administered intra\·enousl)• or imramuscula.riy at the
peutic effect on :111aplasma infection,.66· --. 1• 0 i;;o, 168. 197 rate of6.6 cc 11 mglkgai 24-hourintei,·als are usuallyrecom-
In many countries the tetracyclines (chlortetracydine. tet- mended.84 ! i; The usual dosage rate of oxyumacyciine (SA)
racycline, OX}'letracycline, rolitetracycline, doxycycline and used under fieid conditions is 10 mg/kg. 208 • 2.'.!I
minocydine) are the only effective chemotherapeutic agents The chemo1herapeurlc effect of doxyc:ycline, rol:te!Tacy-
approved for the treaunemofa.naplasmosis. 114 11 •· u 4• l!lii, 19:- cline and minocycline in the trearment of acute anaplasmo-
0f these. chlonetracycline, tetracycline and o:q1etracycline sis is similar to that of oi-·y1e1racycline (S1\ ). \\ith cwo to three
are the most commonly used drugs for the treatment of acute administrations at a race of 4 to 10 mg/kg being recom-
and chronic carrier infections. 112· 114 1i,;. 208 ..inaplasma in- mended at 24-hour intervals, 120, 140, 250
fections iu ca11le are not sterilized at the usual recommended Single intramuscular administrations oflong-acting (I.A)
therapeutic doses of the tetracycline drug;.1 20• 1,17 . 221 formulations of oxytetracycline (200 mg/ml cc,ncemration),
Gloxazone (alpha-ethoxyethylglroxal dithiosemicarba- at the rate of20 mg/kg body weight appear 10 be equally ef-
zone) and two derivatives of carbanilide. amicarbalide (aml- fective in moderating lhe course of A mariinale infection~
carbalide isethionate) and imidocarb (3,3'bi•-(2-imidazoli n· in canle.120• 221 • 2J 5 TI,c ui.e or a single 1reatmer11 of o~;:1e1ra-
2-ylJ-carbanilide dihydrochloride (or dipropiona1e)l also cycliue (L~) for acute anaplasmosis has an ob\'ious practical
have specific chemotherapeutic effects on Anap/a.;ma infec- ad,'l!mage in that ca11le receil'ing mulliple Injections or 0:-.1•-
tions.'- aa. ~2. 1oa-110. 112, 121. M3. 1-11. 20-, Howher. gloxazone tetracycline (SA) . or an~· of the other tetracycline drugs. need
hos not been made commercially a"ailable because ofirs un- to be restrained repeatedly and receive larger (two 10 six
acceptabt~, severe toxic effects in lactating cows. 142 The car- times) total volumes of the drug fonnulation. 122· u.:;. 235
banilides, particularly amicarbalide and imidocarb. on the lmidocarb hydrochloride 76 and its t.lipropionate sah 1·1'1
606 ,,er..,, 111...,,: Rickcusfal nnd chlamydia! di~~es

both ha\'e a marked inhibi10ry effect on A. marginak. clinical signs of anaplasmosis.are seldom observed. These
Therapeutic dosage levels ofimidocarb for the treacmem of mild relapse reactions usually do not require treatment. :?.S7
acute anaplasmosis in ca1lle vary from 2 co 5 mg/kg body Relapse reactions in splenectomized caule, on the oiher
weight208 and should be administered subcmaneously or hand. are frequent!)' as severe as, or only slightly less severe
intramuscularly. 112. 111 It has been found that a single sub- than. the primary reactions3 2 s:!. tzo and frequently require
cutaneous administration of imidocarb at a dosage rate of trearmem.
2.5 mg/kg body weigh! is as effective as higher ra1es varying The: duration of rhe relapse interval may reflecc the effi·
from 5 co 10 mg/kg in che conirol of acme anaplasmosis, 203 cacy or the drug and the 1reatmem regimen used. as a longer
and Lhat a dosage race of 2,5 mg/kg body weight is the mos1 interval between relapses indicates greater suppression of
efficacious under prnc1ical field conditions.2 140 tb.e parasite.an In one investigation it was fom1d that the
Administration of imidocarb at recommended therapeu- mean duration of 1he relapse inierval in um:reared control
tic dosage races usually causes rransient side-effects.13• IH. caule was approximately 18 days. compared to 36 days in
203
These include a iocaliled swelling at the injection site animals receiving two treaunems of 01'"}'1etracycline SA (at a
(following subcutaneous adminis1iation) and systemic cf· rate of 10 mg/kg. 2~ hours aparr). and 56 days in animals re-
fects, characteriz.<'d by increased rnllvation, lachrymation ceiving a single treatmcm of oxytecracycline L~ (at a rate of
and nasal discharge.:!. I I:!. Z<l3 The severlry of l11ese signs is 20 mg/ kf!.).221 Similarly, three treatments 11ith OX}'terrac)'·
dose-related. and when doses are repeated or increased 10 cline SA 11 mg/ kgl, combined rreaanents of o;,..yterracy-
eliminate Infections (see Control: Chemos rerili.7..ation). cline and gloxazone. or three trearmenrs of gloxazone also
more severe clinical signs or lo~;city including diarrhoea. resulted in longer relapse intervals. 108-110
dyspnoea and depre."5ion frequently occur. 1 Since intrave- Alier a single administration ofimidocarb (2.5 to 3.5 mg/
nous inoculation similarly results in more se\·ete slgns of kg), relapses occur approxhnatcly 1hree to four weeks after
toxidty 112• 11-1. i.,,
238 and even monalityatrates of3 mg/kg
treaunt>m.1 03· llC while at lower dosage rates. treated ani-
or higher. 11 z. m this route of adminis1ra1ion is 1101 recom- mal~ relapse earlier. >. second 1rearmem with imidocarb ad-
mended. ministered at the onset of a relapse delays aJld reduces the
Ir has been found that im.idocnrb administered once at a severity of the relapse. However. when the second dose is
dosage rate of 3 mg/kg body weigh1 is a more effective che- administered before a relapse occurs. its onset is delayed
motherapeutic agem than oxytecracycline, tetracycline or bu t its severity is not reduced .2. tCMf--l lO. 14<
chlorretrac:>·cline, uven when multiple administrations of
the latter drugs are used. 150 Apart from only a single dose Supportive t reatme nt
being required, imidocarb has the added advantageofbeing Fol101dng treatment with an effective chemotherapeutic
highly effective against Bti/x,sia spp.. 2. 3 • 113 and the neccesity compound. the Anapla$111a parasitaemia declines over ape-
of making a differenriaJ diagnosis in areas in which both riod of se\·eral days, reaching levels of 1 per cem or lower
anaplasmosis and babesiosis occur is obviated.1 14 204 How- only after appro!'.imarely one week. The decline in parasi-
ever. this latter practice is not ge11erally recommended since taem ia is usually accompanied b) a reduction ln packed cell
a specific diagnosis is essential for the implemenuuion of volume over a simflar period of time. 11 is iherefore impor-
appropriate control measures for the rwo diseases. Further- tant to bear in mind that rhc clinical condition of an animal
more. the do~age le\·el of imidocaro required for the treat· may deteriorate considerably as a result of progressive
ment of anaplasmosis in caule results in tlie sierili7.3ti<m of anaPmin non,1thsrnnding s11c·rP,,f11I ~plldfk rhrmn-
Babesi<1 infections. and possibly renders the animals 5US· 1herapy. and that supportive S)'mptOmatic treatment ma}'
ceptible 10 babesiosis on re-exposure. 2 • 3 be required to prevent monality.30
One or lWO subcurnneous administrations of am icar- The adminis1ra1fan of blood transfusions 10 can le suffer-
balide at a dosage rate of 1Omg/kg bodyweight have a thera- ing from acute anaemia is advocaled in the treatment of
peu!ic effect on A marginale infections comparable to two anaplasmosis,G6. 77• l3:!. 135 but the recumn1endat!ons for the
intravenous admini~1ra1ions of o>.ycetra1.·ycline CSA) a1 a acmal 1-olume of blood 10 be used for this purpose '111"}'
dosage rate oi IO mg/kg.52 greatly. To be effec1h·e. it would appear that large volwues !2
Following successful treatment of a primary .-\nap!asma to IO lltres or more. depending on the siz.e of the animal and
infection 1,i1h any of the abovc-menr!oned dru~. the para- the extent of the e1;1thrOC)1C Joss) need 10 be transfused.6-l l lli
sitaemia decreases to l per cent or lower within four to As a general guide, 15 ml of whole blood per kilogram body
seven days, 303· :!50 but there are frequent rclop;c~ (recrudes- weight hti,, been recommcndcd. 1: 16 Howc\"er, the results fro111
cences of parasitaemia) Of \"ar),dllg imenstty at \"ariable lnter- such administrations are often disappointing.&. For reasons
1·als rhereafter. 2• ~?.. 52 • 112• 1'1'1· 221 The severity of this relapse that are 1101 quite clear. blood transfusions ma~ also lead 10 a
parasimemia appears 10 depend on tbe 1irulence of the rapid rise of .-l margi11(1/e parasitaemia in can!e. 8-1. 252 but
parasite strain and the suscepribiiiry of individual animals. generally the)" lead to an improvement in rhe condltion or
In non-splenec10mi1,ed caule. the relapse reaction is gener- anaemic animals. particularly those animal, with severe
ally much Jess se1·ere 1han was the primary reaction and anaemia in which increased haema10poiesis is not yet
 Bovine anaplasmosis 607

e\~dent.132 Despite the possible benefirs 10 be derived from Control of vaccine reactions
blood transfusion. the technique does not lend itself 10 large- Following the adminis1ra1ion or live-blood \'accine,. some
scale application under field conditions and should probably animals may develop clinical anaplasmosis. For all practical
be reserved fur ve1y valuable animals. \\'here blood transfu- purposes these cases should be appropriately treated speci-
sions are indicated, blood should be administered slowly, fically and symptomarically as though they were naturally
keeping in mind the risks of vascular overload and lhe possi- acquired clinical infections.
bilicr of anaphylaxis due 10 incompatible blood groups. al-
though the latter apparently seldom occurs. The donor Chemoprophylaxi.s
animal should also be carefully selected so as 10 pre\·ent the No chemopro;:,hyla.cdc drugs. such as those used for the
possible transmission of other blood-borne parasites.n. 132 control of bovine babesiosis, are available at presen: for the
Because of the accelerated removal of erythrocytes in cattle control of anaplasmosis in canle. The chemoprophylaxis of
with anaplasmosis. 14· 202 the benefits 10 be derived from blood anaplasmosis is, therefore. largely based on the constant or
transfusion are of short duration (18 10 48 hours). a.nd repeated administration of chemotherapeutic drug$ to con-
transfusions should therefore be repeated: preferabl} within trol Anapla,m" parasitaemias in naturally i11fec1ed cattle.
24 bours.a.: Blood transfusion is contra-indicated in animals Generally. chemotherapy during the prepatem period
that are restless and difficult ro restrain, or in those "~th ict · merel)' results in the prolongation or the prepatcnt period and
erus.84 The cransfusion and excirement incident to crea.tment detays the on~et of clinical •igns.~7• ro. 127 With repeated
may c-ause stress and the deach of che a.nimal.30· 84 administrations of oxytetracyclincl.A (at a rate or20 mg/kg at
Because oi the risk of mechanical transmission of Ana- seven-day inte:\11ls). commencing seven days after infection.
plasma. affected cattle tespecially those 1,ith high para6i· the prepatent period ls extended by approximately che same
taemias) are a major source or infection ro uninfected time-span as the duration of u:eatment.57 12-7 Although a.U
susceptible cattle in the herd and should preferably be animals thus treated remain infected. fewer animals develop
i,eparated from the healthy animals. Discretion should be clinical signs cf anaplasmosis as the number of treacrnents
applied in deciding whether to remove the healthy animals increases.
or the diseased ones. as ~everely affected animals could For animals ex-posed to natural infection, a treatment
succumb as a result of exertion. When isolating diseased regimen consisting of!mramuscular administrations oi oxy-
animals, consideration must be given to the pro~sion of tetracycline SA lat a rate of 6.6 to 11 mglkgor Li\. at a rate of
adequate protection against tl1e elements, easy access 10 20 mg/kg at 21- ro 28-day intervals) t0 all susceptib,e canlc
palatable feed and water, as well as facilities for continuing throughout the season when vectors are most active and
observation of the animals and for rhe administration of abundant. and continued for a further one or two months
any additional chemotherapy, which would require the after its end. have been employed or recommended.84 • 111•
128 190
minimum a.mount of exertion on the part of the animals.77 • This miatment regimen should prevent clin'cal dis-
Additional S}'lllptomatic o·eatment may be beneficial ease in most animals that become infected. but l\ill notster-
and is usually recommended. Although haematinics such a5 lllze Annp/asma Infections. Thus. infected animals become
cobalt and iron probably do not have adequate time to carriers and subsequently de\•elop prcmunity.
stimulate eryrhropoiesis sufficiently ro influence whether or Similar prorection against clinical disease is afforded b}
not an animal will recover.3() these and liver-supportive prolonged oral adminls1ra1ion of chlon:erracycline through·
drugs may r-educe the time required for com·ale5cence. out the vector season. or1hroughou1 the year. 114 · 190. 193
Furthermore, conection of any gasttoimestinal distur· The use of s1.1S1ained-release oxytetra.cyeline boluses has
bances such as ruminal stasis. constipa1ion or lack of ap- also shown co:isiderable promise in the chemoprnphyla.xis
petite by the oral or parenteral administration of of bovine anaptasmosis.3:l..J• This inerhod should impose
rumenotorics. ruminal fluid, mild laxatiVCli and appetite fewer management and labour constraints relating 10
stimulants. may help prevem additional .econdary compli- parenteral administrations ofterracyclines. and should pro-
cations and promote a more rapid recovery. vide more predictable and consistent drug levels than is
Longer-term control of anaplasmosi, can be achieved possible with oral administration of chlonetrnc~·cline in
by a variety of methods. which may include one or more of medicated feed or Licks.
the following that Involve the use of chemorherapeutic Vl'hen 1errac~•clines have been used LO prevent clinical
drugs: anaplasmosis, animals should continue to be closely ob-
• the use of chemotherapy to comrol primary \'accine re- served for clinical signs of the dise;ise after wi1.hdrawal of
actions following the a.dn1i11istration of live vaccines (in· medication.
fection and \reaunem method ofvaccina1ionJ:
• pre\·endon of clinical disease in susceptible cattle by the Chemosterilization
prolonged administration of chemotherapeutic drugs Anaplasma infections in canle can be srerilized by a variety of
dun rig e>.-posure (chemoprophylaxis); and drugs and treatment regimens (Tables 47.J and 47.2). The
• elimination of carrier infections (chemosterilization). methods used 10 de1ern1ine whether successful $ter'Jization
608 Sff'TI()~ 111w:: R!ckem;lal and chlamydia! dlse,ts~,

Table 47.1 Successful ueatment

DRUG RATE OF ROUTE I NUMBER Of INTERVAL
regimer.s for :he sterilization of
ADMINISTRATION TREATMENTS
Anep/a;ma in!ec110r.s with lhe
ietracyclins d•ugs. lModif,ed lrom Te!ratVc!ine 11 mg/Kg 1\lor,M 10 Daily
Kuttler ·97g"·•1 Teiracyetine 11 mg/l<g IV or IM 10 Oaity·
Tetracvclme 11 mg/kg lVorlM !i Dail\
0.\)'letlacVtline 11mgl~g IVorlM 10-14 Dail;
Ch1011e::w,chne 33 mg ltg IV 16 oa,,y
Cnlortetratycline 2.2 mg/kg Orally 41 Oa,1y
Chlllltetiat'ichne 5,5 rng/kg Ora,lv 45 Dai~,·
Ch1or1etracycline 1,1 mg/kg Orahy 120 Dady..
Ch1or1e1raC'1clina 11 mg/kg Oral!v 30-00 Da ll\'
c~:011e1~cychne 3.3--5.5 'Tl{! xg Crally 60 ca,\y
~1orteuacvclme 11 mg/kg Ora,'v 3o-6ll Daw
t\1,ncc,-clir,e 20 mg/1:g l'I 2 7 dc\'S
~.1:nccY'ime 10 mg/kg IV 6 2-J davs .. •
O~·(le.iatvc,me 22mgf<g IVarlM 5 Daily
Oxvtetracvcirne llAl 20mg11t9 l~l 2-'1 7 aao,,s
0X)1e:rac1tli~~ llAJ 20 mg/kg IM 4 3 days

N = 1n1ravenous. JM = Intramuscular. LA= _org,atilnQ

Ow,ded scnedule of tlM> fwe-ila1• :teatmen: ,:e,,ods l'hth an mteNen,ng perilXI o: ·o da~s
Negat.~e status determmee bv serolag1ca; procedures
Oivrc!ed vmhm 12 da\'S

or elimination of carrier infections has been accomplished elimination of 1l1cir infections are only obtained after 12
include haematological and serological examinacion or months have elapsed. 20 s Serological U?Sting to determine
treated animals. the subinocularion of blood from treated the success or elimination should cltus onl~ be conducled
animals into susceptible splenectomized calves. and splen· after sufficicmt time has elapsed to allow-animals to revert co
ectomy of rhe treated animals.20• sero-negativity. Any animals serologically positive after this
However, treatment to climinacc .4J1t1µlnomn infections in period should be regarded as treatment failures and either
most endemic areas is contra·indicawd because or 1he high retreated or separated from the rest of the herd.' u
risk of reinfection and the relative ~usceptibility of the cattle Because of 1he possibility of transplacencaJ infection.
population in the absence of any long-cerm immunicy which calves should also be screened serological])', and positive
accompanies low-level persistent infections (premunity}. 111 cases treared or eliminated in order to prevcm them from
With the advent of sensitive serological 1e~1s. a method becoming a future source or infection to adult canJe that
has been developed whereby 1he source of infection have been deared or infection. ft must. nevcrll1clcss. be
(chronic carrier cat lie} can be eliminated from a herd. It in· borne in mind that the serum of uninfected calves born of
volves tesring the herd In order 10 identify the carriers which infected dams may also react positively in serological tests
may then be slaughtered,!!5. 105 segregated and kept isolated for periods ofup to three months after birth. due 10 the pres·
from che rest of the herd. 190 or sterilized of lhe infection by ence of cole>stral ancibodies. ia;
chemocherapy.a-; 190 • l!lo This approach ha~ successfully Sterilization regimens or A11ap/11s11w infections with che
been used in parts of the US/\ and Canada. but is usually tCtracycline clmgs are summarized in Table 47. l and include
only feasible in areas with a low or moderate preva.lence of the parenteral adminiscraiion of tctracycline.30 oxyterracy·
anaplasmosis. 11 • 1116 dine (S:\J,ez. 134 r7Q. 207 21a O>-')'letracycline (L-\J,1n 20,
The disappearance ·of antibodies co A1u1plaw111 spp. minocycline1~0 and chlortetracycllne.Z1q as well as the oral
from the semm of treated cattle, detem1ined serologically, is administrauon of chlonetracyclioe.r.s 191• 226· 2•11 It would ap·
most often used under field condition~ to indicate M1ccess- pear that the du rat ion or an effective le\·el of antibiotic in ani-
ful sterilization of infection.' 96• rn; However. serum from mals is an important factor in the successful eliminatiQn of
animals in which the infection has been cleared may still infection. Treaunem failures frequently occur at tl1e 101,·er
react posith·ely in these tests for period;, va~~ng between dosage le,·els and when divided treamients are gin•n.1 ,o. i-o
. nvo :m<l ~ix months fnllOlving rr<>atml'nt, 13°· ?06 nr t'VPn Funhennnre. r~pnrtP<ily ~urc-Ps~f11l crr:nml'.'111 rc,gimen, h avl'
longer. panicularly when using che CF tesc.GS. Ir.!. 170• 20• Both not always yielded conslstent results. For example. although
che CT and Co. test titres subside more rapidly than do CF cwo administrations of O>-")'ICtracycline Lo\ (at ~O mg/ kg body
test titres, 82 · 19; indicating that the former tests may be more weight) at an !menial of seven days reponcdlr eliminared car-
reliable indicators of successful eliminaiion. 134 On the other rier infections in non-splenectomized caule,20' three admin·
hand. it has been found experimentally that consistently is1ratlons of OX)1etracycline L-\ (at 20 mg/ kg body weight) at
negaciVe CF and C'..A iest reactions il1 animals following the seven-dar inten,als during the incubation period iailed to
 Bo,ine anaplnsmosls 609

Table 47.2 ,•eatment 'l!gimens for RATE OF ROUTE NUMBER OF INTERVAl

DRUG
the stenlizauon oi An5plasme ADMINISTRATION TREATMENTS
,nfecuons. 1Mooi!iad irom Kuttler.
1979''l'") Qxvte1racycline 11 mg/kg IV 3 I or2dats
-Gloxazone 5-10 mg/kg IV 3 1or 2 o-a-.-s
Q,e,.ctroc•i-hnc 20 mg/kg lM 4 7 dars
lmido..-.•b Smg/kg IMorSC 2 14 da}'S
OX';tettacyehne {LAl 20mg/kg IM 4 7 days
~

lrnidccari, 5 mg/kg IMorSC 2 7, 1.; 0,21 davs

lrrmlCM:arb 4-6m91kg IMerSC 3 D~
m,.:ocaro
1
--- 2 mg/kg ,Mo,SC 3 Daily

Gloxazone 5 mg/kg IV 3 Oailv

fmidocarb 4-5 mg/kg fM 01 S:: 2 i'<.1avs

IV= l~travenous; IM: lnuamuscolar, SC" Subcuta~eous. LA= Long-actmg

eliminate A11aplasma infections. 5i· 1~; The observed dil'f.N- High. but sa!'t?. dosage levels of amicarbalide fail to elimi-
encc between the efficacr of the dn1g i.11 eliminating infec- nate A mnrginall! infcc1ion. 52 and it would appear that the
tions in the Cllrricr smge and that in the incubation period, repeated high doses probably required for successful elimi-
and the finding that four intramuscular administrations of nation result in such ~everc hepato- and nephrotoxicit) that
oxrtetracycline (L.\, at a rate of 20 mg/kg. or S:\. a1 a rate of the use of this drug for chemosterilizntlon of ,.\nap/asma is
20 mg/kg) at se,·en,day intervals. also faih:d to eliminate precluded. The high dosage rates or glo~azonewhich appar·
carrier infections in splenectomized caule?"4 seem to sug- enll~• eliminate ti. marginale infections1tlll· IO!I also resulted
gest that the host immune response probably plays an im- in fatal toxicos1s in all treated animals In one trial. 1-lo,..,.ever,
pQnant contrihuwry role in the success of treatment. combined treaunems with glc)xawne Hnd oxyte1raq•cline.
Oral administration or chlor1erracyclinc for the elimina- or gloxazone and imie\ocarb. arc more efficiem in eliminat·
tion of Anaplasma infections may have advantages in that ing carrier infections than is treatment ,d1h either drug
tre,ument on a herd basis (in the fom1 of feed premil<es in- alone.'lll!-110• 11 ~ Attempts 10 eliminate inl'ec1ions v.ith a
corporating the drug) Is possible.1\lO but the consistent combination of oxyteiracycline and imldocarb ha,·e appar·
elimination of carrier infections by this method is often dif- entl)' been unsucce.s.sful. 112 On che other hand. the combi-
ficult 10 auain because of differences in amounts of medi- nation or four O>-'ytetraC'yclinc lreatmems (LA or SA at a rate
cated feed or salt-mineral mixes consumed b}' individual or 20 mg/kg body weight at seven-da)' intervals) and 11,'o
animals. t iz. ,95• i 9 , imidocarb treatments (5 mg/kg bod~• weight a1 I-I-day imer-
Susrnjned-release o1'iytetracycline bolu,e,- administered \'als) (either oi which treatments have indMdualh· been re-
ro A marginale carrier cattle re$ult in a reduction or elimi- ported to eliminate carrier infections), appears to eliminate
mnion of CF test titres in most animals Inn fail co eliminate infections more consistent!~· than does either drug used
1he infections.33· 3 ' alone. 220
Alternative rreatment regimens 10 eliminate Anaplamw Callie in which infecrions have been eliminated shortly
infections in caule invol\'e the use ofimidocarb and/or glox- after primary info<'tion exhibit no residual inlmunicy upon
a,:one. as well as combinations of these drugs with ox·ytetra- challenge \,ich A mnrgi11nle.'· 0 • 195 in contrast, cattle
cycline (Table 47.2). As with the treatment regimens that cleared of carrier infections are susceptible to reinfection
invol\·e rhe use of tetracycline drugs alone [ [able 47.1), ,·ari- but show resistance 10 clinical anaplasmosis for periods
nble degrees of success have been achie\'ed.a. 201 • 20<;, 22 ; ranging from a i'ew 10 30 months follo\\~ng parasite elimina-
There appears to be considerable individual rnriation in tion.~,s. 1a3. !9<H9z. 20~
tolerance to imidocarb 112 and severe mxicit}' and e\·en mor- Furthermore. re,exposure of carrier animals to .·\11a-
tality may result at the dosages recommended for chemo- f)lasma infection shortly before or during the period of treat-
51eril~tion. For example. although it has been reported mt>m may result In n•inft>C'tion ancl fRilttrP 10 P.lim lnarP thP
that the administration of imidocarb at the rate of 4 ro infections. even when 1rea1ed wirh parcmerally adminis-
6 mg/kg body weight on three consecutive day~ is a sate tered drugs at dosage rates in excess of those required 10
treatment for the elimination of A. mnrginnle carrier infec- eliminate infections under labora1or)• conditions. 116• 1m
tions in catdc.109 mortality mar occur in calve~ treated with TI1is augments the recommendation that programmes
three doi.es ofimidocarb ata rate of5 mg/kg hody\,·eight at aimed at the elimination of the carrier state arc best con-
24-hour inrer\'als. 225 ducted during the period of lowest vector acti\ity and
610 ,u.,10~ mni-.r: Rickeusial and chlamydia! d,seases

should be accompanied by the imphtmentation or mainte· has been proposed as being a possibly more suitable
nance of veclor control measures. 1 ; 115 Long.term oral ad- me1hod for the elimination of carrier infections in herds
ministra1ion of chlorretraC)'cline for 45 to 60 days or longer where re-exposure cannot be prevented. 110

References
.\11,\_\J:<1, t.(; .. UJIUUEI\. D,l.. ¥\\'JUJAMS, 1,D.. 198<>. A ~lUd~· of th.I.' U,l.\it:ft}'Of nm::opl;ir.c clltyme•!ml:tod tmmunoi()rbi.!.nt :S.S:i«Y for mt:.l~uring
lmldocarb d!propiona,c in catde.
t;2-17'i.
fl""°"'" i11 ~~1~ri11ar,· SciM<•'. 28. an1lbn:i) lO \,u,p/n1mn marginhlr in t:1ulc--Serum. Ar,~:rulltm t't1cirftta1'•
Juumnl. 63, "6-79.
2 ADA.,1-s. Lt..4 tOOORO\'IC. A.•., •• ,r,.J, I hech~mo1hor:ipeutic.rffictcynf 1q &ru~,~G. G ,t 1.y-~8. Immune rt•.sponsC'~ of c<wt.'- m ;:i. ch~·:nfc3Jly

lmJc~h dlh,·drocl\ioriile on cnnarrrcm bo,inc ,n..pla,mo;lsMd tnodifltd Nmp.'mmn anrlgen,Amuti<tr,1 }011,nnl of \~tcrlm,,y Rt:garr:,.
bob~,IO>li I The Cffl'<:~ or a =glc trc,tn•m. Trop,ml -~~;ma/ Het1/t/1 39. !f:l.>930.
nod Produerlo11. G, 11-1'8.
:ro B)G,\tXF.. R.O., lg.80. I..abor.J.U)r)'Ond 11cld observatjon..,; on the u:scof
3 ADA.\tS. 1.G. &Tooono,,c. k,.\,, 19-;4 Th\? chcmothcropcutK: efficacy of AJwplit.Yno c·c•urmli as a \.'acc:ine ;qt~fnsc an:1plumo~i...,/.iinlx,b1'\"'
im1docarb diliydrodlloride on concurrent bt,vine-o.nnpl~,mosis and \ ·;,11..•rir:.aryJo,,rnnl. 11, 21 ..22.
babcl>!osis. II, Th• ,•lfcctsofmultiple :.re,um"'1ts. rmµi<ol,\n/11101 ::1 The('on1rol of~ome
81(°"..-\.I.XE, l{.0•• DE \'O~ • •,.,.~ K.\MKQ\\ MAS, PR.. J,970.
/1,rt,///i Ulld PnJd11(//bll. 6. ;9--11-1.
uck·bomedi5(>3i<'< m South ,\irk... 8111/mn ,1, /'Offiu !111.-n:a:10110/ d~
~ ,\J.\\1, 5.A. \'\'11.:.QS, q. &-C.AMPD1.U, R.U., l9i'8t ('tpl.-nmrnta.l bO\ine £p1:0011t1- 81>. 89-1()(),
nnoplasmosi•· Cllnlco•pAtho!Qgu;"I :md nutrinonttl ,tudle>, /lcS'l(lrt/t 111 22 e:r.G"!-. It,(" u~:c;r~ttO\-T...~. f,\,'., 1$84 1-\n n.nap!3.(m~t:r.~erocon\·m!on
Vttertm,ry· Sdflnce. 25, i'fj,..81,
,1udrofc•hti ,ubje<:tcd 10 different lt,vel, u! tkkconnol en~ fatm In
s .\J.J.8Rn'TON• .\..R. .t.-SEGCR. r ..a.., iY62. ·1 he 1raJ\!r.port :and c:u:,~sron of bll\" th~ Whdhook disJnct of South 11·os1i\!rlc• :O:amlbi~ Pmc"-d/11gs of1h,•
plgmenli in •naplru.mosis. ,\nwrico11 /011'lm/ o/Vt:,•rlnnr,•flt•sea,rlr, 23. n,/,r,;n,/r l\i,rW Congr.,,, <J11 {)twru,s of<:",nule. i,-21 s,pt,mhor 198..
1011-1018 Durbu.,. Rcpubht o{l><tuth Africa
6 AU.L~ P.C.• .&. li.'tl1T1.EJl, ~L. 1g81 E!rfft of ,tnnp!n.sma 11uugi,wlt>lnfoctlon ;t:j lllRD. /J •• 1~73. '1<:onot.1l •n,lpllhmo,,f, In;\ <alf, four1111 1of 1/to• S0111/r
upon blood t,;~!JH 3nd t!-h~roc:,.1~ m s:pt~~11'>mi1.t•d ~l\'l•;;. Juurnt,I OJ" Ajrlra,i \ ~,wrfr,af) \SS(J(lmlort, ·M, ~iO.
Parasitology. 61. 9}1-956,
24 1uou" 1.t,r. si JriQr_,~, ).,:\t .. 1t,t98. \lorphology and d<-1\t?lopmt-m ol
1 "'vrru R. ~.L k;\MPAAULT T~t-., 1981. CJlnlcaJ C'hcmi.JU)"Of
\W:.'-', P.C.. .-11111p/1<1tt11J 11111,glnnl, lflickctt~iolt~: \n~i>l•~mat~<eae ,n cuhurcd
anoplusmosls: Blood chl.,,..k~I chQnge, to infected mature cuw.. lrodllS~npulmi< I \c•n: lxodld•cl cell,. fm,rnnl o/:,1,>ika/ ft:tnmofogy.
Ami'ft('(Jn Jrmmai of~ C/l.lri11t1ry Rt":$tYJf'C'il.42~ 3:?.3---3:!5. 3:i,, ;~;g;

a AUJ~.P.r 1r,;·uT1'\J)I ,u.. ~ \:\HR.,mr T • i9'11. r1inir;,1chc•m1(1t)'nf :::; 80\JL\~GT.Jt. 1•.. nuu;.uut.\U'CR, C.,M .. li!\SNl~Tit\, (M... M.\Ct:.\\, JUI. & •·m·n.
anapla\.O'lo~,>-- Com~»r4'th'c i,crum prott..ll chung~ elki1ed b)· !\.H .. 19:1. Am,,pl.,~mn.5il'l: C'.t,ntrol c,(1~ tirs.toutbre:1k l:-1 c.,n;ida bv
auenuiJU-d amt ,·1rulcm Auuµ/11,msa J11"f"$tt:a{4• i,atau.•s. _;\m,,rt,m, ~crologic~.1 fdcmitk.)1ion 3nd ,1a11ihtcr. Ctmnditm /autr:n/ of
Journal nfVrrermary Rf!:!t'Mf<:lt. ·I:?, :i1~3:?S. Compn.ra.m-c ,\ktli.titt<' 35. ?49-2Si
9 r\."WliJt\.OS. f.L.1 IO:>:tl>, 1.1,W., ~1()HHJ~:-.. H.D., IIQt.HliJIT, I), l,t LU <iJ: .. 1!};!:
..~n"phumn- mt1rgmttlr. H~t-moglobln pattt-ms: 1n (')Cp1.·rin1mtally
2U AROC~, \\ .f l(Uli"'LA. 1.0. • rf-'l«o"· cc., 196:;. (or A'"''''""
a.nap!.\$mO.\b. fm1tt1nl ofthtAmericnn \tlN41llllJ ,\{Cfi1atl.U~1x-tariou,
tnfected }'UUTil; c;n.l\'er,.. E.r/Ntrlmrmal J>nraslrok>J:zy. 32. 2~2; J J.r;, 948-SSI.
10 A~'iHO~""\'. r,,\\ ,, 1968, ~rm11~~111r,t nnrlrruml Stilt'> J.., :s \·1..-ctor o(bovim.• :r. OTIOCK; \\,£.- ,on;\l\S, n.s .. nat,,rx. 1.0." J~!\i:.s. t.\\'•. 1s'65. \uto ..
~n•pla,mo;K Irr. l'r0<:<•«ll11g, of th,• F/f1/r \'ett/011al 111npuu111osfs antibody ...,,udil~ 111 bo,inc nnnplasmu-.i;. Am1r'rie<il1 Jou oral"/
Coufer,rrl'f 1968. Stillwater, Okl:ihoma. \'t:ti'nl'ary Rt'.f•Ym:lt. 26. 250-2S3.
u .\~Zli\XI, o:s. H.R..Ull.\, H.n.. FtlltU, L '- ,. G.uu:t'O, c .. tyJ';'".
\'nr-dnatfon :z:11 IROCJ.. \\ I. ••T.\f'~U:\', (".C" .A: WI Wf'.K. t.0;, 1953. H!gh•l("\'t! Jtm.!Uln)cin
\\1th .-\11apfflJ111ll n•n1rt1/.., Rtspr>n<r aft~r an "'}>tnmcntnl thnlkngo (chlortetracycllnei do,;.3ge in anaplasmo~1~. ,,;mcrtrm1 ioumat of
\\ith.~11apl1r<mn mllf!1l11111<'. rro111enl -\nl!ntrl f11!'111/rn11d P11J'111ctl0n. 19. Ve1~n,,nry·~11rrc/r , 1.: 510-51:l.
83-8i.
~ 1moc•. ,, f .. N:..,lh-Os c.c." ,1,aU·\\ t-H, 1.0.. 1!l~- An expcrim\.-nt 1n the
1.e ,\kUSt. R.l.~ 11- M,\MULI, 1.. 19za. L.iboratory S.tudiL•sc of an3plasmo~is hl rrc.attncnl o(nrut1.• ;uittpl~mf>~i, with u:1meyclini: h~·d~ochlondt.", \'ortfl
ennlc tri.:awt! whh oxytctrn.c)'clinL·. Jautu:11 ofthtJ Amt"rfctm Vrrerlnt1rv Am~rlcan \ 't:,1rln<1rum. 36. s.;;-.5SO.
,\fedfml ..UftX/ation 13.1. 317-519.
JO ll!A~\:. (_('., ).UE'-\\'E.~ U). • l!>'f_j,,, f.\\'.. 19.;9. Th'-' tda1ton
)lttC)t.~. W.L.
13 ,\UGusrvx. , .,. &. RIC..\LX£, R.u.. 10;.r. 1nap/a$.ln(l 1n1cct1cm u, t1 giraiic. of trc.s1mt•m 10 hem,u.olngial ch;uag.c,...1n Ano1pl.t~mo~1,-., Prnut.-dh11>'> nf
011ders1,poor1 Jq1m111/ of Vi·ter/r,ury Rt".f<'/Jfdt. 39, 229. 1/t~ U11J1,d Stl1/'1 /.11•,-srAd: $<mi11tt)'A<S1x:fmlo11, l?;-18 O<Ct·mlwr 1959.
t4 u...uam "·' , c:,nnoui, J.w. &c111t1srr.~~r~. J.J·., 1961 Eryt.hrUC)'tC" sunivill s~n Francl•<O. California.
fn experimcmnl a.napJas.mo~J~. r\mork.a11 /():tmal oft '•"1i!nn<11)' kes1..""'0rd1, 31 11-ROclaJi.sR\ u,,, & ,·rol.tlll u.o... 196'1. Haurnatotoa found m 1.,i.ld
?2.. S90-S~6. mt"mbcr~ o( thl.!' OtdN Anfodnct\·Jo ln £.u.t Afric-a Rulftthi t>l Epi!ootfc
15 L\R.aET, A.r 11.lC!\nJ~C.:nt. lti-. \'l, 1.. LUXl)f,JIL~•.\ "·· Dt.0m, I .fl.4, 1-'0CA.,. /J/.w.r,sofNric,,. l~.~2!19,
t->:. ,999. Compamon of ,urfacc protein, of,\n,1p/1umo maig,1111/e J.2 Bit(')\\;'\. c.r..D.. \\1UU, J.>..U. f,.'fl(R<.1.11~J,.1968. Cht.tmotht.>ra~· or
g."Q1>n in rick cell culrute. lick srulv:iry Kl•ncu. and c.ill~. /11/,:a/Qrr mrd it-X~rirncnul A,wptlr.mtt1 mt1rg}unlc lnli'rli(nls. Tri11ls with ~n
lmm11rrlt)-. fi7, 102-t 07. experlrucntlll dlthl<l,omlcarba:tone (a-u11to,1·eth)·l!Jl)"O\al
«6 B.\RSn', .\.t., V,t,l ..'lfR, G.U., ~1\l..Ot. 1•.J.4. \IOOUlRf r .C'.. , lid:'. dUJ'Uoirmlrntb:i.t.oncl. u,m,h \ ·ctirlttofJ' /oumttl. f 2:.;, S;,t,j.-:U4
Charnctctiution of ;1n tmm11nopr0ttc:1~·~ protcin comp!('~ ~r 3J B\'.FORO. a.t....).."'OCA~. ).,~l. lt'lL\JH, , .., .• i9s1.SU\Glintd·l'l1.t~)t :ume-n
,'.nap!mm11 mnrgi1tr1!t b)' cloning ,!lid ~J)rt>slon of the gene roding for indu-.10ns~ .:.. rev1e,, ~ current appli(1lt10n !or an:tpl4'1Smosb ther:tpy
pol~'J'CPtld,•A111 !OSI- /t1fi'CII011tm1/ /111n11mlt)·. 55. 24:?8-2J3S. (>rtxtt-J:ngA of111.t S.-1'tnt/J .\'mi~uwl .:\11aplu.uno1t1 Ca,1,'ef\'.nt:(•. Zl-:?3
1- a..i<R.En. P.• 8E\'lRll.>v~. l.. OMOt.L'V, P.l.. 8RC)W:<, t.O,D- 811SllflV, S.R.M,. Octn~r 1981 Si.nk,iUc. l\,1i$.!'1is...1l1>pi.
GL\.IIU \J.t..... Xl'.AL. a .., .. SMI LU, H, I.. Wltbt fJ..lf .. 196~ Btotogtcal actNlty 3.1 urroao.11.1 rust n. JJ.... ~<.le.-\, K.~1. ~ nAm, J,A .. ~sa2 l!.fi«ts or :a
of some a-dlth1os.cmfen.rba:1.c,n~. •Vawr1. 206, 1J.it.\.1l.:a l. ~-u..:taintd·rd4:a1.c o).jitrracychne bolu!o for Jnapbl~moHs.corne~:'}, 1ht
18 !lo\AAt. o.x,. r.\R.J:lr l\. ft.J., Ot VO~• .'\.I., tx1,,-itt1. P. ~ 'ftOn~,'l'lL H,r., 1986. A Bo1•in1Prcr1t1io.,11u l7, 123-1:?S.
 Bovine anaplnsmosls 611

J;i un~lm. 8 1 , lUSa:Jt J,: •. ~Q<,.\!\', ~.M • S,T'fl\"°"· t.G.~ JL\I.Q J.1.. :9&1. ..,,. lll\l(Jf>O.IIU()" , ... 1 4, fJfDU..! 0.M. 19t'2. Sh,.1d1l'>Ofbuv\m: e~1hr•'.1<}1C''- in
ChcmorrovhJ t~" of ,'C<'tOr· bum.: 1m:ip!u..maH11. w;.th sust3.U1'11 ~iea..-.i· >:»pl•,mo,;, Ill. Roi• qi cllcmjcal nod phrs1e11I cht111ge; In ,,,ythrot,<cs
bolu,~ •\m,•m·~m Jo11m11l<1J \'twr/,1111'' J..'1.5<"'cl,. 1:2, ;!~.2089. ln tht' ,n4.·1.:h2lnlqn nf an.t.tnu:i in 1plen1.'ClO~d ca!\'~S_,;,,u•,i<,w
3Q C.\LLV\\. L 1_ 198,.~Protcm~ttl :mlf ri(kettsln.l dl<t~a~ Animal H-tulth J,i Journal af \ -.•:.,•rittan R•·~~rrh. 23. 813.-.820.
AU#fttl{(l \'ul. J. C:anbl"tr,1· AuSlr.slion (;Q\~mmcnt Publlshmi; ~ni,c. ss 011'1•111 u. o.o.. \);1~ru.>.\t1£. ,,.•, , .\UCTU'111, 1\,, 1q8l. Tun"-"1'1Lt.i,10n of
3':' f'\k,('J'\:, (' \,. .,, U ... n M, -"f0h11C, M,. 19--(, tUta.nl'f>U~ hypCrst'IHili\H)' .-\uap!n.m:n nmr,:f11al. !tnm .'.! n:11ur.11tr~Jnfc..-c1t-d \\'ild African g:i:.ant rr11
nnd 1\Qantll>od~ prt,t.Jucuun'" t ,111lv 1t1)L\(:tf:'d with u,·t or tn.tr11\•au:~d tCrt,1.•to111.~~ptmhtmw..:.. W.:.itc.•rht1u~e-, 1u o cnlfln 1'1g.cna. \~t?r/na,,,
·\m1pla:,1H" m111Jlmrlr, in ho,·hw .mtJ 01,.'inc erythrocytes A.mtrlctm Ptru1.1(1oto10•.8. 337-339
/01,,nalt>{\,.c114•rl1mr,· Rfs,•in,:h, :ti. 1059··1063, .:i,6 OOSO\ \\, •M\'lt.Mi:O. J, IL\RI. L.T LI~. C OIJJfl8(',, C. 6-S[Of.A. C., t984.,·\
38 ,~,1'-""n:-.:. , .. , .• ,tu...:_ n.M ~ 01,11c. .\1., HJi':'- c,•U·m"''d!attd lttunUJ1lt}' t:tpfd ..14tn!ni: procl'dttt,· far Am,pltwut, mdQJ,lnfllc• !n ho\ine
rcl.Jt1.·d w chnllt•n1;,... ~k'.),:urc of crutli' fr'locuLiti.-cl \\ hh v1ru1,mt and ~I') thwq·tc~A.ml'f'ittm Jrmmnl IJJ l ~1rrlm1ry• RtSt'tlrri:, 'f:i, 2.1-~:?1.J·l.
numuated ~.uru, o! J\11aph.J,11w murgwn/e.A.1)1c,•rlMtt/our,Ull of
ii IC K1UAU, w.r.• U\'.C'tJt:-.;
-..n. It \l,,\·(.Osmt.f,., R-\•• 19;q. EffiCAC}" of
l~wri,uuy R1.~·1)r.:'/J. la l tG;'c 1172.
corramic1n: L,\-200 adminl'1Ct«l during •h• pwpa1m1 pcn<Xl of
~i9 c.\.~C),. C..\ .• ~n t.$. 1">.~1. ~ 1-ti-.:11c, ,t .. lf\7i. Cell- n1tdt:.t1..:-d 1mmuJW ~n;iplu,mn,I, Pr,xv,,11,,R< of t/1</JJrtl ,111111111/ MN•tlttg of;Ji, U111t,J
tt'$poni.e to rirul~nt :.lnd OH\'nuawd, \m1P1,w11a ,1;my11uJi'1: at?mlni&tctcd
to cnulc in Ii\·~ .uul h\1'1t"h\71U"l1 rorm~ . •,\ttii'ricn,1/m,mnl of\ 't'Ji?ri,WI)·
'itm,J; .·lmnml fl•"''"" 1U.u1d,11lm1, 28 (~tober...2 '\"o\'tmtbct 19:9. San
Dkll(>, t::.lifomt.
Rom>t(II. 3ll )73-1-:S
t.a l·LJi:~IAO. \\ P . .., ,t>.~o,rf.itt, A., .• 1'183 Ac.quirud (t'llul.u r~pun)l\'(•r.c~
-10 C\MO~. C\., Wt.1~:,·.ut. "· 'L "'ltilC ,\t.. fllUM.)lO~. 1. . .;. ~USO'.'-, o.n., in c.aul+:clu..u1.-d t•f _.\,u,µ(wmn mn~ln111~28 inl'.lnths bue:. ~ ·r;tm:tr,;.·
:974 \ppend•~c-rel3ted 4nlil,en J1r<>ductl<on hi P,mmnplnsmn /nlllTUno/~. an(I ln:mrmt.1/Klllt;,,/r,~·· -~. 6.Sg..G6.l.
caudowm m dt'(!f t1)1.h:oq1.l"•.:; .t1m1~m:11n Jour,Jul ()J ,-..CLn1tm)'
ll"1Mrcl1.3S. 1m-L;31. stt u "~L.\O, ,\ , .• ~T1uttit, o woon.\N>. L.r." Jr:UTl'LER. J.:..t .. 198.i. tmmune
ro,p,,r.,6 of ut"~ ~nllgenk•II" >1llt1uloted ond ch•ll~ngc <"l'O>ed
.n t;Oi\Stu. ,,.L-.
197.1 Transmts.~ion ot .-t,:avlrumn ma~nalt>t,ytheamle-
with ..1,10p!tunu1 mari1n11t.,durin,:: uc.1-. Jn((',1.,uon or m.'1ltmtm with
1lck Booplulus r,UctQp1m. (Jm.:tttdn11d JounmJ <{.-tgrirultumf emit dt:~.1me1hut.n~. \11t,'r((',,,1 Jm,rnal of V1•rcm1tt'lr;• RPtMrch. 15.
.~nimn/~l,wce,. ll. JBS· 1!13. ·131-;Jli.
12 t-o~~n.L. ,1,L .. ,v.u w,1.i,.,.19i2 Tran,mi.>JJonoL,t,u2p!r.,,mo n~rzol"
f;c> Mn,~ .• HU ,n1, n, ,:... lU,n(;llt~. \.,.,.. ft,\tUILT .•\.I'., MCrl.\\" \L,. T,f. ¥
by lhL" Qttle lid,. &xJ11hifii,\ min,1111,u. ·lWttrtfi.:111 \!~.;,,fi1Uzr))OUrtlfll..
P.\t'.\lflt. ,;,01, 1y~.
E>.p«"'!-lon nf m:ifor ~Jr(;tct" pro1efn:? antigenic
48, ,i;;
Wirinm... -during .1cm~ An11r1t.usu11111111rgi1111lr ric:kcust..•mfa inf;,c1i,;u amt
l j COKHM. W,M .. C0'1Rf.l. r.s.M #,{.(J'rr\t H\l,tr-. \.I , t9i8 Bo\"in"abc>nion lmm1,11fr,'i t>ol. &.l&-84 t
a~~x.-intrd wi1h A,u,ptasmt1 r1w,,:inal,.•. c:.t,mulfnn /oumnl tt{
6 1 HU.t,;.?to. I.~, P,t\UM t.,H .. \1(..l..Ulkl T,\;,. ALLHLO, t>.fL1'-UMlBET. A..f •• 1989,
Com,x,rntin• ,\1rdlr1",•, -t.2 227-228.
l>ett'Clion ,m.J qu:mtttMfm'I nr Am111lmum1 m,,rglnult' fn c1ur!ttcanle
.w c0Juuu1, ,u:H. \·Mt...Usfi. o,. C'.t\Rli.:Q~, C...\., m,nc.. t.1 .• ~unuut• .:.L. u,lng ,1 nuclctc Qc1d probt-. J<>:mwl o/C:1111/<111 Microl//fllog;\'. r 2()6.-212.
rRn"ll\O, r.5 ~ u r. A.t .. 1!l311. C:.cm,p:\ri<un or &hl'\-C.: m~lhod( nf
iminuninulon ag~hl~l bu...1nt1- ;tr1r11>1'1"'mu:1oh: .\n txruninAtlon or u~ uw. •,.• • ~-\HH ... H, 11 IYiU, Chang\.·~ ht M'IUIU \.Om.;ttnmnlun u!

po-st\.'Ur.cln~ ~1Tr-~--i,. tm,,,/am Joumul lJ/\ldt'tl•tnr,• Rf'U'1t1d1, .tl, phyg<lt.')1()'i11·'\lunul,11ing r:u.·cor 111 e>.pt1rlml't1t.JU)· Induced bo\im.•
Jnnpln,mo,I, l't<'ftmlnAr)' hnding, ·\m,r/c,,r, Jo,rma/ of\ ~terlnar,•
l~!i1-10fill.
ffl'U'tltdr . l7".f»07-609,
~5 tO!OUtn. D.f.. \\".V,'.\:l'SI. t..(~ ",\IU\1,. 1.~, 198:t ltl'CTUd~ttm(t: of
;t,:apt11sma nrn,w,u,t... lmJuc'--d b) lrnmwtO'-upprt~lon ,,ith 63 1non a.., . f ·"•Lt'\' ,1. ,. G\I l.\t°:fltM. 11 .. 195L T1tc, u1te or aureumldn fn
C\'clopho... phamtd~, W:1•ricnuJQu111al of\·,,u,ri,w.ry R~ktln'h, .iz. !9-2:J. an;::,phi..mn.,, .. ·\'nrrh ·\m1.·n·crm Vt1rr1narfn11.J2. 547 ...:,,;':J.

...:6 cox. s-.R. -6- 01,101,ouu.os. o.T.. t,12. 01!munstrnton 0 1 an :iuto.mtlhody '.'14 Fm, IJ;R, u. t. t;\\ 1Ft. H,l- 1973. ,\hnnlnn in cow',; tnu.cula.tcd \,ich
as~octatcd \\ilh •1no1p!u....rn6sis. ,1111,.•ri~(m Journnl o/\'f!t,-,rt,uuy Rt:1,t:"h,c-lt, ~1:t1pln.sm,1 nm,Y.:,~..al~ n,~rrog.-,wttJ~. "· 59-67.
3.l -3-:-1;. 6.S t->tA..:( 1i.. o,tt., ).;1M1cs, o ..\ "nut.st-w,~<..\t..19','q than1ctcri1..uJonf>fthr
.;- cnt~Tf.S''-l'" l,f, a tMJ•AOLO. J,B- 19;16. lnh1bititm of Annl}losm.a ma~inal? lnclw,lc,n II mi Un.,: mt•mbrune or ·\r112p/11~mn mn,gi11nt..~ b;'
inlC'C:1.ion :n cau.tt" with ox,-1t-1rarychne h\·drochforidc, Pror~,':dlngs oftlu lmm\rnoforrht'n labt.•mn~. ....n:,·rimn Jqur1111/ (Jfl 'rtt'ri1u1ry• R-.">tOrth. 40.
6()1/: .n,munl .\fc"c'IllllfO/tlw UnitIt(/ Sllllr.).'\mn:dl H&."11ltl1 .~5..<0t111titm. 7i7-782.
?8,-.,30 ~O\l!fflbllr 1956. C.1t1C:11to, Illinois. fitl HlA~X.ll:\ , 1,,-.. 10-n: •·, At HUJIJ·, 1.w., 196.L A tf"VIC'\,· nhht-tr4?Jtment or

,a riAnr.s. u.F., m:-.tOf'\luu.o,. <•• r. ... Ron, . 1.0 .. 1970 ,\napla"mO!'-li .tul'lplMffl",;i.~ 11,o,.·._,.,,c1111,;s oj (ht> ,.~mtrth Vmfnrml Ana,,!n~1110:1J1
tronsnu~1on: tntN.-uJruion hy 1hc.• oculnr ruutl'. ~iettrch m \ iurl,:ary ru,,ft•r,,11r..·t 2£-2; ,\pril. 19b:? Reno. 'lct--ad.a.
S<.·irm:,.•. 11. 594-.5'9.5. 67 rtR,\.,n.1,. r.r ., um+. J.,, •• 1~;. A propo',.Cd method of prctmmunhdng
..;') Ut FCUJ\IU} . .... T., );::\:(.}\'\1T'I. U.1 1
., ~,<·,a1rn1-. T,C P,\1,'ilHt. G.U .. S.U.\RfJ C,f.. ,~4t1let wfch nifnlmum Jnc>eu1:i of .o\m1pJ11,m11 r,mtgJnnlC". Rt'lt't1ft/J tu
"' ,1c:u WA1s. T f., 1~a3, l'>c.1-tcclftm nf c1utJ,, mmmtll}" fntecled with \ ~IPt'illllf')1 <..dtta"I!, 8, .i t,i,,-,i 18.
,,1mplasnu1 marg,,wk 1n n rnwo11 ofcndem1c1,} h)· nbl"-d PCR and o GO fRA~);u,, r_i. nu1-t·1 J.\"- • c.ku"t•tr~. t.c., 1uofi, C.:hlorrerracytU~ for
com~uu,,,. cnzymt."·tlnkc.-d 1nu11unu..arbcm n~y u1i,mg n.-,;ombmani clir11ht3tion c,f .1n;1p1a..,mu..i..., In .;.11rrl~t cuttlt•. /utlrJUJI of tht .-4,tu"rittm
nus1ursmf;1cc pm1.c111 S. Jourrwl of Cllni<al \1/rtoiuotogi·, .J6. :i;-;sz \ '(fft'f'i1t11f\· \l,+Jlwl,L<:mcitllltm, 147. 35:t-J.56
su ur kl''t;~. C",," QUl:\L.\'\. 1 ~zb. SplcnectOrn)" in dnmt•<1;tftutcd :i.nlmals 6g HU'loc:'H, U.\t.., \U f.l\'1. \J\. I I \J(;GUIIOi. 1.C. t.J>.AL\lEIU.. q.K., 1~~8.
and It.., M-:quclli.c. \\1lh ;;,pecl..il reference w nnaplasmosis jn "'h...ep. E.-<prc-~1on o! .,\uupk,s.,,tu 11u,~,,1Jf(' maJor sutf~c protl'ln 2 \-arla.na
/;'/r:wllb and Tuvlfth R••r,orts. uf the 1Jln-ct1Jr f1/tc.-rntmuy F.du({lflOn rmd during pt.•f<i,q,mt ~-die ru:k<'n<,•1nia.J11f,..,._~,l()t11111tl /mm,mlt). 66.
/11>:<rnctll, Pnn I. ~6HRO. 120(1-120~.
;t nr,,x,,, "-"·· o·u ... AA. 1•.J , \(HP-.<., ,i:_r.,... ,)t)JUH' ... u .. H)70, ~,ic;n:md 70 c;""Vt· 1.11 .. :~61. U1tm<1n,11·auon ut \Jt11Jtltm,u, m11rguu1l1"
~mmunohcinol}1lc an1.•m1tt :uuJ incn1.i nr cal'.'1.':).}our;wf u/:he.-tmfflM11 witJ> {luor~n-111 d)·tc, .:icrfdlnt- orangt-; comp.am.on with CF L~t .md
I ~i,rirw,y .l/et/irolA<S1kifrt/n,i. 11>5. 1861-1869. Wrfo,ht·'I. ~rain. Amt·rJrmi /uumul nf\'t!liriua,y ~ r d : 2:l,
52 uc ,·o~ ,.1 .• ti-,nnow,u..~. P.~ .• ro1 rnn, J.'-\\', "· ).l.J.Ua.)1.\,, rs. ::r.8. ft.82..JUl•.-
A1nll"'1J'b;ilidc: ·\ lh<.'mpct111c 3j\CIII fot anapli1<mo,k. Ondmt,p(JOrt ;1 ton.\\ ,L.. lto\hlltT. \..f STlUnt. 0 .• Jl'.u..,tER. ~.H., l,.!..\.C\\'U~. D.P.. J.:Ot.\.~ .
/1,mmnl of\'c•t,'rinnry lle,rmd,, 45, 203·2QA ~.M •• f;ORHA \f. ,. ;& 1918. DNt!C"tion of.htapltlJJ)JQ
\(('.(,U1Jll. T.c..
~, un,M.\:,,;\. G ... 15f:S.O. lntnMniM10,1 of ont1p1.~,m6~h.. \11writtuJ jl)11t1:ttl "f m11,si1111lt Jn:~..-ccM tkl \ l-("U>h b~ u11in1:t t"Juned l)~A prnh~.
\ C,rrrimuy Rtu11nrth. 11. 5-ib. Ptu..-«~ll11g10/1/1,• Sn:lomrl. l<ml,·mrofS<l,•Jt~. 1//ul, 85. 91~923.
612 SfCTlO\ n0<u: Rickeusial and chlnrnydiat di;c,a~es

7! GOJ·'l-. \\',L. lOH~WS, \\ .c. S KUTrU:K. ~-L 1.984. Uc.,\'tlopme:u ot :m 91 J,,. r.,1.,Mttt. G,11,. 19'.)Cl, Crrtlr nckcn..~ml.: dur1nt:,
J.U:'SfR.s.r UtH..'-.
indirect fluore.sct>m vn1ibod1· 1es1. ,bin,g mtcrcu'luorometry a, a pcrs~mm Annµ/n,,,m murgmal~ inft.·chcm of c;autc- fn[,r;fon muf
dlagr.osuc u:,,1 for h<ui11c (lr,apJnsmnsl~. A11u!rltt111 {ou,1111! of\'1111:rhran· lmmwlit)·. :;a. 111 ~-1! lll.
Rc.<N1~1t. ,-;G, l080-·20SU. ~ t,. ~0\\ I l.:,... IL Di ((lL\JOt, ._,1·,. 11!\L\1f,lt, <.,J., \1(.(.Ulill 'L, ~JIUJ R, p. II
;;t <.Orr. \\'.L. m t•~MJS. Ll, 1
."' .:-UTit.tR. "·'- .19M. AntJp!Jt~mn marglnaf,.. ,~cuw.o, 1., 1996. -\Jmbod)· ng31ns, ;,;n llmtp!n1,m11 mnrgm,,lr ;\ISP5
£p,.·l)·th10:JX1u UL'Jt)'on/; U.-ttin tC"a<tions ~,·Hh bo,lnc c~ihtOC1,1C~. cpiior,c c,1mmuo to 11d. nnd ~·throcy1c s1agr~ 1dontih(S p-:r,.1!-hmtl~·
F.x1wnmtmUJl Pa.ril.JIUJ/ogy, tu. tOJ-1 lJ. tnfcetcd c-ouk. /OC4rt1tr}QfC/lt1/wf ,\fit1,1b1olo~., "'.l-1. 222:i-22.JO.

7,1 no,7...u.rtt.. F i,.•• tQMUO\"IC:. ft.\..~ n10M~' ,-:,c.., 1«.~6. Jm.tnuni:>:.Jtion 93 KOCA~. "',,l 1981). l>C'Ldopment of, \1tapf11#1Ul mnrgina:~ m 1xodid ilck~
ng:,1nst annplo<mosis and babl-<io<h. P~n I. Cv:ilua1l0r> of Coordina1~ d\.',·~l(1pm~nt ot rickt1L,1fl <1rw1m..m and 1ta hv,., lit'~, In:
lmntunfaation U$fng mlnlnmm mfoctJ\CC.k"'-"' umkrlabor.11.011 s.,m .fl5. tl\.lR.. ,. , .. fed.,, .\lor11h1Jlos...~·. l'h.rs1of11g\'nt11J fhthm·1u1trfll

condition<. rro1M1m,dW111111d Pum#ro/ogir. 2i. 127--:a;. F.rol~ a/Tirk.< F.ngl•nd: tllls I lorw<1od I 1~.

;s tMP.11 .• 1933. l"hem!cnt h!Qod studie~. V Comporttth~ <ntdie«>flnl.,~t 94 Jo:.M a,RHU~. >--I, li\\1X<i, <i.o\ ,._ U\ltc 1 J.A.. 198..5. TrWl.smi~icm
);QC.\~, or
and unlal:rd blood fiJ1r111"' of bo,in« ln he,lth o1nd during Anopl11Smn mnrglnalr bi adult D1m11t11:e11mraffdmo111 d\1rlng feeding
anapJn,smost~ LA. mn~lnflle inf""Cti¢n1 :i:nd ptro5,la.\mosb 'P. big.?n,mum on c-uh·c, -ll11,·rz.tn11 Journul uf\',•1,·rfna,y HC"1t•m,·Jt, i6, l5fi5-1l6i,
lnfcctlon). Oml-.•tstd/lOl>tl Joumnl ()j r,,,....,,,mr:r ~,:it•nunnd.-\mnml 15 J.OC',.,,. "'~~t thi, , ....\. .. U.\lh, J.\.."' UAtUIU~.,1 .• 198~ Dtffl~ln'.'1lt.\tUmo1
l11d,mr;·, I. 371--101 the indu>1on oppcnd•~• ofAnnp/"smn mn'f(lt1n/,• m mmphill
76 I-L\R1', C..ll, 110\'•S~U ru, f., BLHf.LH, 1,, '\;\:PS,\)\, ll,M, 1., ).\{H,\IU)\ , , 19(1, (Nn,:r,c;;wror 11m/1·rt1m1 Am11rl17an /numnl of\ "ttC'Tlnn-,, R,~,Ylrrh. is,
lmldocarb for 1hc c•ontrol ofbo~io,is a11d JnapJa,mo,ls.. Prot:r,·d/1111, 180().181);.
uftit, 19th World Vtt.trlmtrJ' C:cmg"'-SS. 96 W.(JCA\, to..M 1 \\'"'·· ~.A • um an m. u. r., tum. J•.\ .. 19~ Mt.u-phutog,1c,U

;;- HE.XS"ISG, lot.\\'.• 19l,ll :\ulm,,1 Dll8ll.se.1 fo South A/rim. 3rd L-dn. Prctarfa: chara:ctt'ri"-ti<"> uf 1."0l11me-.. of.-\m1p/~1>111 mtttgl11~lt" Thcller in mitl~ut
r.1m1rnl NcwsAg<"11cy Ltd coloni1.~-of O.•rm,1ctnlQr nnclrrw11fStllt).. Anh•ti<on Journa: of
h~1mn1J1:r Ri!tt•ar<J,. ;J. S1SG-.;i1J3
-s ll['.:k\' t.:r•. ;,.o~,u~. D.D•• fLt. ll,L w:('U)L\.SS IL\"lt. 4 \Oft.l. ~ ».. 1.98..,.
er; >:OC--\~. K.S.S., )I\Ut, I-"·~ t.WlM#, <..,\,. lltlk>, Uhn1:(.tr\JC(U/t"Othw11!mmu
Effl"C'ls and USl' of J modified Ike ,lnapJns,mJ margfnale \'l1.Cdnc J:"t b<'o!
ht.ifers in Califomfa,Juumtll oftlu~.·\Jm•rltmk t't1Nri11an· _\f~tilCIII mnrgfna!tiTht-Ucr tn IJ.·mmw111or andt•rso11I 'I tilt"'- ,md Di•muu·,·mor
l,'(lt/ahJU,v 1S.'\y1 \mtirlttm Joumu I c.,JVc•r,•rl1111ry Rt:tSt'llfdt,.; J 191i6-19/(i
,1$$()1'.{(ltimt. lffJ, fllH>9.
;g, um..u.co.. R.f .. l973. 1lu:. cuhun.! of..\Jw11/asmu morp.l11(1(t, - currt'11t
ga KOC\\, ~:'11 M,\JR, ,.,\. t.\'111S:(',
1i..\. ",:nC.\'rtO\, 1.(,.. 1981 Tr-an-.mf-s~lon ot
.~zaµlnmm ,11a'l{1,wlt'Th.l~ar b) l),,rm11n•t1tor rmd,•rsnni.Slilb and
~talus. .Pr'tK,wfmgsofl/JtJ Si.ttll Nnrio,:nl A1m11IUJmOJis CI.Ju[~t\•nc.-.
(krmll(,•ntor: <1tt"£1i/1i ,S;irl. A.tnfl'i<'OII /our,u,{ uf \,..;ttrinn,:i. R4.'>t!tllC11.
1~20~1.trch !97J. I.,, V"t,~. US'\.
~:?. 15-;R
8o tt10.'\1.GO, a.1•• 1o"c, t.,, .. ,.r...
o~o\,·~. fi.! ..'""''n>JO"U ..,.r. 1989. .-t.,mpla,>ma
99 ,:oc,.l"o. ~. ,: .. nl!..L ~.r,. "u \JR, 1~,.. t ~ . °'1uon,,ra1ionof .\tU'1J1kuma
1,wrg:nnl~ in ticlt tl'U cuhur-.·. lmctriea11 Jcumru' uf V....ttlTi-'1.ai,· R,'UJ..•tlr(}t.
111aTlJlrN1/, Thell,,, In tltk• by tick 1r.m,m,...,lt111, onlmal moculatl~n .md
50. W2ll-:?(ll2.
fluf'f"'C;)CCI\I cuirUmdl' i.tud1I."\, 1,n;,rfcnt1 Joumat uf\"t-h!rlnt1rv R1'"J.t'ftt,•I,,
81 UO\\'ELL. C:.J .• WAlJ:LR, J.11. &- Nt'\IILL t'.~l.. JJ)j'8. Trek,. mit!S nnd in.\t'.Ch 141.183--186,
infe-s-cingdnmc.".St1c .'lnimob in Soulh Afi'ic-,1. P3.rt I. l)e.,.;c:fiption~ and
Biology. Department nf ,\gticullural r,-chnJul Sor\in'>. Rcpublk oi
te':.a J:C>C\\, ...,, .. \ J,;\,\IHI. l,11. l,·DOO():, \\".f·,, .,,u. Ullro15trul"tUn.1 n!
o1.n3p\a.smiil tndu,inn-.tPilwhusli..d 1~li:u,,) .;ind &h("fr appNtd~ 1n
south Afrlca. l>clonc,• IIUllctln ~o. 393.
ln1.1c:1 omd hcmoliud ttl)throc..i-it") omd in compWmem :U.ation antif'.t"'lt.
82 r,n.:\.R. P.R. KHflllR, J.P•• .\~IN. LL & TH\R?. \',.1966..C:omparor{v,: -~.mtri,a11 Jaumal of\ 4't«in11r;• RcUl"¥4tr:.J,, 31J. 1l?.-l-1130.
P"Y'Si><enceofc.pm.,"· rubl.'·agglutiaoiiurt end ro:npleJttent r,~tlon
10, ):OC\X. .:,M- n~AIU!. f.H,, t1SU, t,:.f'. & lUl()(.t. \\' .I., t978. u:1r.1 srrurtutal
ttl:.l.ted to dcsUO)' th~ c.uriN ,1au~ of
tc.·~t r~at't!ons in cattle
!ucnliz,,1Jm1 of an:aplo~11I nntigc,11 !P,1whu<k;i l,olott' ,,ith
anaplas:mo"is.. Am<vlc1111 Jout1tt1l of L'<'Wri1wr;• /WJ4."m·/f, 27, 372....JN.
ftrritln•conjugAtcd .mtib<.>d>1,Ar1wrlctm /01mml uft~urinm,· Rt>,.~1,&"11.
83 JOlt~S"TOS l,..,\.V .• TR"UE\tAN.t.:.F•• Ll.AlCH. I: 11\\'U.SO~. \,J, 1980.. \ 39. ! 1'.ll-, 11;.
comp,rtson ofdlr0<t nuurc...:cn1antlbod) ond G1tnua st~g for the
Jt'l2 ~OCA~ ~'-1 •• \\'t("J..'\\'tl\f. >:.11., £\\'J'\:C, ~.,\., 11.\IR, 1.,\, 4- {lAtti«)'.\, .: J., t~8.
~t·mQncrn dlninosi, or !i1\apla"U'ltQ.\h. i\1urmlum \ "nc!rln.nl) /ounml.
Prcltml~ol\· <tudi,,ar 1h,• devcloprncnt nr 111111plntma mnTlJtni>~ In
.J. 222-226.
,alh·n~ g!Md< ot ;1duh r~c.-rllng1'trrmmt•mc,rnmf1.•rst1m ttcla ,hu~o((m
8.i JO~.t.s. E,\\ ... Sll'OC\... 1.\' ~ 1966. BO\ine .1J'U1pl.bmo,h,: ltsd.it1j.,trJOl'o.i11,. /011n111i uf \ "ru1111111)· /1,-s~nrc/1, ·19. 1010-1013.
«rCatment :md con1tol Jounwl oftlnJ Amer/um \t't1crl1lhr}' ,\!t:dkol
tOl ~Rlltat. J.P." AINfl( M.. 1q63. Ana1>I~!'loJl10?oj!'lo. \torpholog.:c dt3!3Ctt,rlsde<
A<JO<'fruioll. IIS. 1624-1693.
of ~,c yar.1>!1< ln the blooo or ,-a!ws lrlfl-cted with the Cn,gon strrun of
8& JOSts. r_v... ~UFWt:ll, 1.0 •• !\"Ok.MA~·. ll.lt. & Nt0t).;.,. \\'.I •• t9('8.Allnp{njm,, AnnpliHmtt matl(mah• 1,m•riMt1 /uur,ud tJfl'r11.•rl>mry P., .Jc"tJtd1. 24. 6

mnr;:i,?nf~inrec,ion in young and .:agc.d c:mlt Amtrir,111 Jrmr,wl 11/ 676-61:li'.

V,r~rina')' Res,:orc/1. 29. 535-54.:.
UM 1rr.1u:1[1t. J.P•• N<Tlt: ,\t..... ,r.HflOP.)U't1 w.. 1~6-J. Arll!pl,L«m~,.
~YI. ll~e
86 10:0:r.s..E.,\\ &-~vP.~L\S, n,o., 1962~ B,J\in('~n:apJ.Jsmo,.i,: 11tc<me-a~. II.!>' p~thogon,._l, of Mi.'tnfa pcodu«'<l b)' lnfl•ction with .•.,:11plnsmll
dinfcal di-.ignCK!~ ~nd pro,wlcbii. Prrtet'l~,tmlf"nfrirt J,011r1/J \'nti<>nal ·\mtritfm /ow 1ml of Vt'/t•1mary Rt1Jc.·«1t'11. 2$. 3:;3-35:?.
l1:npflt.sn11J$iJ Crmft"h.'ttC:4!, ZG-2i 1\prU J962. Rl~O. ~c."\'3.da.
105 Kn'r"'Sn'~.o,. 1L 1u86. A bt.'nefit•to~t nnal)''.'h of th~ progn.rn tu ct,'!dic:m•
87 IORC.t~SON". W,J,. 01! VOS. A,J, &. bAt.GU~l-f. k.J., t9M lnh~<1l\i~· of nJi outbreak of ,1n;1pl:1,.i;mo'l.i1; in Cti,n:td.-i Ptt>f,'t"fllngsoftllt! ,.Ounlt
cryop.rescr.~J HaiHsf11 /,011Js. Tll,b~ll, bij:onlJ10 and A1wpl11.m:o r'.i'lllfillr: J,u41.nrntt()1to/ Spnvoslum 011 Vd,:ri,u,ry Epltlt•miol<lg)" wul Et'Ut1omk1.
fnr caitle-aftf.'r thowhtg, t.lllmton :ind i:~c:ub~tion nt 30 •c. \'t'tt•rtttt1ry 18-2/ ~owmbtor l~)&S.. Singaµot~.
P,uasi:dlogy. 31 :M3-?5 t
to6 ~t.R. ;..1... 1~)66. C:hn:cal and ht'mnmlt1g:iC.U i:omp--3li-sor: 01
88 K[unt\lo\.~. Ts.. c-otr,w. 1.-\.\\'. & N,\UD('. i. W. 19'}11.. l'inm Pt)/S(lllfttgl Ana1,1a1nu, nut.tJ..t/1:at~ iJrtd Altttl)fn1ma ,<'mtt,t.:. .1\1'11:ttum ftJUflM.: of
find i.\(rt'OlOXltO>.'t:SO[ Un.'Stt~k fn So:ahct•tAfru:a. Q}pt' Town: Oxf<)rcl \ ctetrimi.ry· Rt!jc_'llrt:iz 2i.9-11-~)-16.
Unl,,."'~it)' Pr~~ SOulhi'rn Aft1C.t.
10:- i,;:unw. a,;.t..1967, .i\ ,.nidyoflht' immm1ologcl."'J.I rc.lruionship or
89 t;;>'.LLV.. w..a.. 197~. v~w,rbt(Uj' Cl/1u'ca.l DilliJllO-"il, 3rd edn. London: Ar:aplfl~·uu, ma(Jlmolt and l\1u1pl11$ma ,w11r,,L~. ~m:IJ !n l'n.~r,t:111)·
Ballli~re Tlndall Scltilt'P, 8 . .;tr.-i;1.
90 n'liStt:k. R.H, .i., JUS.itC. M .• 19;9, In i'itro culm·Jt1on ufA.ttaplasma 108 »r.:UTTLCft. s:.t.., 19;1.F.mc-o,q,· of oxytttr(lc..)'clint: nm.1:,
,,u2r/(inoi,. tn\~:on of rotd dc,·clopme.m m non•i!1ft:'Cll"d tU')"IhroeytctS. d.ilhiost.miC'arb.l.l.orte in ,M m."oum~m of bm·int!' nnaprasmo~is.
i.mtrtrnn Journal c>fVt'Ittrlruuy R~«lt. 40. 1774-1 ~6. .~n:ttkm: Jo:,mnl ofl'itt'tllWt'}' Rts,i'luh. :i2. I 3~9-lJ;i2
 Bovine anopla.mosis 613

m~ ,unurt. J.t.. , 1,,1. P1omh,tug thL'~i">t\lllc 1:11;tn1~ for the- climi11:1Hon or 129 u), 111n. li..r.1c i;t:\t H, u.. t~,;J. G.Jmc atlnpl'1-\lllU.. l'- lhc lsol:ul\ln nt
•.\t1nplo.rma mnrgimtll' in rhc curri~ anim~l. Prnc..:edlnp;s of rhr i31h \naµlns,,w orga.nl.. nh from .mttlopc. Zf!f1sth1/ft/urTrn1~11mttlt:it: umf
Annual .\11..,..mngnj th,· l/11/tc~d S1mn Ammo/ J/tc1(1lz ,¼i«ta1:Q11, ,Z-4.. 29 l'umsitoli.gw.24 19~-19:'
Oc:nber 1971 Oklah<>m• City, Oklalwana.
130 l0'111R, )i,,j •• OTU'\O, P. ,. 6- U\CA.''tA· w •• 19';'5, Su,;ccptEbUil}· of8Qf~l1
no ~,m··u.a. ~. 19':7 \.nmbtned trr.1tmcnt with .t<thh!o\l.!mic.1rbAZunc c;mfo to C\'J)t'nm~ntal mf1.'Ct10f\,, with .\uaplflsma mnrgioale a.'ld
.and 0~1cuacy-clin, 10 cllmin:nt! A11aplaJt1ut m11tginnt...• in(c<uon, tn &JIJt,:.I,~ t,tgr,r.foa. Z.·mratbtott fi1r \lt1t•1m11rmrtli:i,r. 21, g..!-44,,
~pknt!Ctor'ni:lcd cah,"'- Re'$.t•rrn·J, in lr"t11rfoary· St1rnl'",. 1:l. ;11->-!il'l
1;0 1v~o~ c;.1., ~986. lofc•:n<m,\ '1 rQpu:wl n1se11~rps off>omrsric ~~nlm:,I.~.
1,1 lriUTiU:fJ, K..1- i9~. Coinpamti\'C tc:Spon,w 10 pretmmunh!ati<m U\ir,g i l>rlm,., U.K.: l.onRmon ~,orn1t6c and Technical
uuc.•1iu;1tCd ,1.>wpla.<ma mtttgJ110/tj \ lrulcmt ,.\ nwrgt,w/tand :\. ctmrnlt
1

fn difrcrc:nt "f:C group.. rroplenl,\nwwl lltWltlJ and Pwt.lt;ruon. -1. 132 1or1.c.. J.«.:. 19-f;. Blood mr~tu~.htnsin bo,inc unapfasmo,is. Amrritar:
19~-20:l. J,mrntd oflt1~,,1,u1r,· Rt.wlf'(l1, 6. 2&1-2'8.b
mt ).U1'1'1..UC. ,:.L. J&7.J RC\ k\\~ Current ~,atu~ or t'Onaol and uc::mttl'JU \\ilh 13,:1 :.1\<.,o~u·iu. R.\. & Vt',\"tl' 1,, .. 198.i. R6pmh~ ufc-Mth: upon re•cxpo..;ure
druw,. PrMN,luig,. o[tl:..• Sf.t:lh Natlmwt ,wapftwnm,s Con/l·t...·m·,,, l9-20 U.).>\naplmmanmrgtnoftaftr: climinauon ot chl"Qn.1.eaurtcrmk-ccions.
Morch 19;3_ La., \·c~3,. N,wnd:t. 1'1n,•1ltwt loumal of\ 't:t,•rftuuy l(e5i,·1Jrch. ,15, 695-697
113 1,:1r111..tR,t..:.... t9;".'" f>i.1gnu"'1soi ;,nopla~mw-,-.: Are,.·11.·w. Pm,·,·-·d,n~}; 1.:µ '.\11")0,.IOU. M.4 R[S~IV.\\'. u.w.• \',\IJfoli~. 11.w .. s-r.,uueR, 1:.11. ~ fR.\.\;~.
\\'orb.hop on 11af'mop<utt.tf:es (..\11a1:1!a..~mosts ,md ilam's:oi.Ui, J;-?2 1>,w .. 1973. Efftct offt'ti! daUrintr:.i,~nouitreatmrnts with
M01ch 1975. CL~T Coloml)i<i. 0'.<)1cU'Uc:ydlne hyd.rnd\lnrideon rht.r c:trrier sram" ofOO\inc
11,1 tu-rru1t l..L.. 1919, Currc.m 0napla5mosh c,mtn>I technfq1.u.,..,, in Lhi.! an:ip1n1mo',.1-> Ju11m11t of 1i:e.-lmt.rttt1t1 \.'t'tt•n·m,ry· .\frdlcal.-1.S)·otitlllott.
Jnltcd State,, /o,m,al ofrliq 5<,mh Afrt,nn Vflcrfnar, ,U.,<><"tatlon. 50. 167, IOII0-10$.
:114-320. t.% M ,\l.<l,1c.w, n ..\., ~I\U'°!.QS r.r "HtA•.-.:~ r.w .• 1q78. Effid:tcy of :i nl,"\\'
11; r,..-m.£11, 1:.1.. 1~80. Ph'1nnaco1hcl'apeu11cs o! drug< u<cd in tren1ancn1 nf ()~1~ttt1.cydin~ formul~Uon 4;,~hb1 clinical 1m~pl4\mo,lt.. ~t~riran
Jn.1pla,mo,!ll1> ,1nd babc1.Ja,!, /our,u,/ oftlw ~m,ricn11 Vrr.•rlnn') tm,rual of \'e;trinnry• R~vruJ1, 39. l 10i-1 ·I l0,
\ltdlcal •...,,odntil>I,, 1'G. 110,H 1011.
13t> ,1 ,u1utu.L w.n,.. 19,:i,. Samii:' nbsen!':ltron11 cm annplt1:,m1.1ih,.. Conidl
us );VJTU.R, ,\Jwp/wmae-xp0c.)url· on th~
:i..L, t933- Influence o! 3 ~~nd \ t•111ri1:,1r/r,11, ». -1---
''Ucccs~ artrearmcnt to ~tjmin~,e Am111fh.mu, ca;-rltrlnft.-n1on .. ln caulc.
137 MA~l'll-l.RUJA1, r ... Y~l M~'..C,\ll>f.XTrY, M.• 1U8.t, 'l ht' rffctt or.v,,:p/n.Jnta
:411wricnn JQ11mnt ofV~ttrhlllf')' /ll'$tVtff'h, -M. B8~-88J.
mtt.f8iJU,li'0Zi the~rrnt~ lie- pathwu~· n\ brwini• cryihrocytes
u- i..vrru-a. );:.L. 19&.t ••'11ropfa.li11U1 infe,tkms in ,,1ld ~d Com"'suc (:tJm1N1mrir, 3/M/i;mi.,rr;·nntf Ph,w/o/Q/f.!,'. 78; 851-1!;;4.
"'UfflinQnt£ A W\i\.'\\. /011f1Wl of l\"lldlifi• l)Jst!d,Ji:t. :!O. 12-20.
tl& ~!AX~. o.,-:., Rtitl~ ,s 19\.\3,. \n.!plasmo.i.£~Xl11. S1udi~ concrt'ning thC'
116 urrn.mt A..L. ~ \DAM>* 1.,r;•• 19:. Lnl1m•m,:o of dt'xanH:lhll7.untt tH1 th~ muur~ of a:utelmmunttr Am,-,franJour,w{ n/\'1tqrf1U1')' Rl'St,t;rth. :N,
rcrtlldcscence 01 Anap!asma marglnnlt ,n ,sp1cnC't'tomu:ed c.;1.k"~ 70:.i-i08.
1mcrf<011 fo:mrnl of\'ut~ri11ary RAA·11rc/1, 33, 132,-1330,
13q \L\SON. c ..,., \.'OtWAl. 11. ,.,.. 1981. flu.• tr.to.!!ur of BouµJlllm mlrrop/ti$
tl'l 'lUMU.fl. to.I... 1011~~Q~ L\\' 6 t,.1\H'SOS, j.L. 1't6Q. Chcmt.Hllt(npy to \t"Mintl! b.oc!1dac·1 fro:n i.nfc.. tN'l to u:'lhtfo~tcdcottJ<, ondtr tkJd
\'lhnirw~ AnnfJ/tt,Wtn m,ugi,ml~undt•r field und laboratory cnt1dition1o. t'r,nditioni. \ 'ett,r:,wr.•Pttrt:J.jttt,log;·. 8. 1sa-188...
Pr<Kt:l·dlngs of:-hr<J.ith .-1,munl Mr;;eting of tire Unired Srat<'1 •.\nlmal
HooltJt .1.S1ocumo11, 2 ..; ~on:mh<ir 1qttu, l.out~,'l,I~. ~en:uclo.}·. IJ(I \ W.7.01..\. , .. i• >HJTTltR. i..l. t~t. Th~ ~if~t'.I of ntinO(ycHnl' 011 :.lu~ ~lCUh!
.J1uJ CJrrit-t pt UM..., of..\Jr<,µlf1.Smn m11,nlmtl(' in cxp(.'.rimcnta11~' lnfl!t:rl-d.
uu 1;u·rrtxn, i-;.t...."' ... ,,1N1~. 1.r:... 1978. {t('Jatl\'v ~ffic.1q"of cwu o~t\.•lntt')c1inl'
~p!~ncctomi7.00 ~ttfo Proc~-dingr oflhc5'.·, 1rmlt .\'atlomll
!cmnulatioru. ;ind doxy()"tJlne in 1ht.· lri":ntmcm at nl-uu: ~l.t.~rno,h In Jl11n11/usmo1;JCo111,,r~ne,·. 21 .23 October I981. Srnrk\illtt. \Uills.~lppi.
sphmce1omi100 <tth'Cs...rlmt'ti!'m1 tourunl oft 4'.'/rrmn~· Jt4\"itlft'I:, 3~.
l-17-~9. WI \t(:0ut1U:. T,C., .P,\L,,tR (..J(., U.UU:0. U.R., 0A\"1S. \\'.(;. Al 8.\RBr:T A.I. \~ftg.
ChJr.1<-terf:r.ai:011 of an hnn1unoprot.,.; ti,·,, su1 face prot~ln compk, of
w nrrruA. ~ t...4' roooRD\'IC. fl..\., 1973. lcchniq~ofpNmuni.L'ltton for
AutJp!uJn:n ,,~,q:,,inh• b} 0\:Ar!oninR ond Ltii:prcs.sioiL Prvn."t<illngto;·
1hc control of an~prn,m<1sh PfllM'liing$ of rho SL~r/1 \'nt/0110/
rlrl! F.ig!,rli Xntionn! ~ct,•ritu,,y JJemcjx,m.'{it,• l>ist~ Conft•r.•IJ(I-. 10-1:!
,\m1JJ/nsmosfsCtJ1Jfi'h't1rt•. 19~() Mnn'lh 1973, !it"' ,·egas. °"\\-vada.
As>ril 1989. $1. Loul<. ,\lfnn~•ota,
1~ CUTTUJt. i..:.L. \'t\O,r. ,1..r. • 'l\1t"M)~. f,C.. tlJ;tl. U:')t" nf :u, ~pcrirtu.mrnJ
~J., IAH1'JfflfJrltr.R. D, .& J.,\..\tb I,\., J980.
Ill~ \tCllAf.CM. ~ •. 8F..RCUt, f. f\'tlOR.
long,~ctlns oxytorrocydlno tretram~'<lnll.,\1 ln :he U('OllJU't>t of acute
Can,parhon ot g.hn.;;ir,n~. -in ~ll'ecth',• hut let:\k :inapla-srnarlt!~. \,ith
anapbsmos:ii \'r<u,.mn,.•.\l,:dlrlwJ/SmallAnima! Clinie:fln. 73. ur:-182.
lmrdornrh h}-drochtnrldt". R,,-,..ttrth /11 l'',w,t,mry• Se(i'.J1C('. 29. 198-202
12:1 lUl'TUlt. ~-L &?.Auro I L.. 19118. Ch(IJ'Qctcnstlcs of an ~ttcnumcd
l43 :-.iut.,run·. 'i: • .a. {!'\Jl'...•O(\., tt. "" iq;,:, lmlch.><:,,rb diproplon:uc 1herJp\' In
·lntn,la.mw ma,ginnl,• 4lf d.eer origm a, :>n .1n:<pla1mosis v.acane
~t•nyJJ1 .inopla!>mo,i:o;a1ld b.1be5]osb rmpl«tl Au;mnl llt111tl11w,t
r,ov1c111.v,1m111 ITM/tllflllil Pf()(/11cJio11, 20. s,-91
Pt'l)(/t1t1lo11. 8. Bl-i'O,
12.i wrn t-"R. ~ L Mu-c..:c-. 1.1 .. &Jo1r.,;~os. L,w.. 193.. :,ero1ogrcand dink'al
r~ponse.,, of pa:-~unmuni;,~. , 1:sccmiucJ, .ind pnniou,ly u1h:c.·t~d i:ntdc 1.,1.; ,),1lU,OU)'! '\. 'S!':O.IPS.OS. "· \L 19:-.;. R:tpC"att.'d dru.in~ In the Lri:-i'ltm~nt of
,o cha!tengc.• c....'.\'pt>"'Urc by 1wo dift"cnmt .+v,apfasmn i,:ol.1tu.., •.'111h•m;an Kell) Ml onopl~smo,i, u,ln, lmldoco,h dlh)drochlorlde. Trui;/(111
!01,r1111l t,j\ iliritr(lfj' ffl~~1rcil, ,s. ma-.~·>'>6 A,umat IJ,'iiltlt rmd J110t(uaton. i. 139-1.18.

1w5 l.\\,Nl~f.t, 1 ...~, l~';";' J'h,: m~hnnlt::ll 1mn,m1!i.,lon 0(1\naplmmctund~r 14~ ~1NJ.OftS. Lf., 1Utt',UE$H, n.; .. fill MS. P .. ROU\\.'l-11. R.J.i. C:..\UU\V. LI._

::thodc.s!t1n rondluc>n, Rhodr.$fon Veti:rinmy/rJf•mnl. S, ;4.•:;o tc~8j Prcparn.:fon ond J:1bott11.ory testiOl:l, uf .t ftOLN\ ,..,_t'cln~ C"OnlalnlnJ.!
Bnbf'Jitt bot•i.t. Bal'i'S!o big,•,ninn antl Am111/tW1111 ui,volt•, Rtstarth in
1~6 L\\\'RJ,,u. r .\., :~1,. t:,M .. ~o~" \t.. ,..,,.1., 1uao. The esrcc, nf\\-a.ron
V,11,,mwr,. St:4.•fJrt. 3:? l~'"'-197.
th~ ~onual or d,,,.,.,,.,,
oru~estock in Rbodc<la 7.hnl>abwel. n,.,
Vq:qrirwn· H«ord. 10,.112-11;. u.6 ,111 ult. 1.r... t,.c», lndk~tlort" :,nd UmltatiOrt\ ot anlibiotk 1b~mp, In
un-1plasmosh.. Proct.'t!d11ijl.'f of tltt!S:'th :lJJ11t1td 1\Tt-~·1/ngof th~ Un11dl
c:; u'.\'co1. 'i:. ,u>.• lC1...ttun. \,.r. & )lAC'.C)Nlt~I L tt.,.'\ .• 11}8::. 6o\1nc
Statt•s ·l1Uuwl /lro.ftlt 1hscxJorio11.
J.ni,pla.smosis: CllnknJ. ht!mti.;ologk. J.l"ld $~rologic manffe.i.ta.tiOu$ ln
cows gf\.~n a Joug•acting o~'}'Le lrnC)'Cllnt fommlation in tht\ prcpateru 1,.7 \.111.U,R. 1.c.. 1S1s6.. Th~ prc,·tnUonand trc,arn\t'!nt or anaptasmOSis•• im:ul.s
,~riod .•-tfnitrlra11 Juur,u,I of \lt't,,rbwry R('YtJtch. a.. lJ(..0-1302. aft/ti' .\'1111' )Ork A1·1t1f,.•111y o[Srl,mcl!, 64 . .J~-4.~.
t28 usc:tu...-..:. s.u t. )lAG0~1c.u. K.A.., 1ge1 The proph~·lattic efficarv of 1"8 .\JIIUR. J.C',., U\"\"4 IU •• TUA8[\:J, S:.J. ltQGU ~U\. W.T. t9;a_ ·\ mtth()d o:
tong•.tC"tlngo>..\tt"tr.1ryrllnt.• (Tctmrn~'t'ln/J .. \ ..?001 in t~-p1..•tfml•n1:d ,,:re1:n1tti d~ co bt' u~c.-d ,n ,hr 1r~1umc:n1 or :an.'\pJ~mos~- fC-)-ult~
O'f>\int 2n-apl~s1110.,{'\. ProM.Yli11p;i· oj' 11J1.~ .Xtt:,•ntl: Xirtlmwl .Aunpl1b11101i! ofw.~tinit \\irh Attrt."fUl\)'Cin and Tcrn1myt.in. P,'0Cn."dingtofrl~89tfl
':o,if,·r.~:c•. 21-23 OorobN 1981. SlOtl:\'lllo. Ml<Slsslppl. Attmu,t ,\lmi•,go[tht• Am'1Tltrm \'t1Ni11nry 1bfodhtion~
61 ~ ,,c11<>, 111,ux, Rlckcusial and chlamydinl diseases

n,~ tn.'alm~nt
11$ \llUJ.:fl f.C-,,, U-\"'r, H.f.., tOHP.b~'"T tJ.I, ... OGU~b\, \\, I,, '-~i3 168 1'\JI.J.:r,. a.-,. 1,n, \ 'Tlhun.)tud), uf l1m,nc.-,w.apl.",mow... \,i1h,p1:c1otl
of CJ1,,p1osmo\h h\ l.oui1tl.;n.:1 wi1h .iurcomytia ~nd twr~um·dn JoumaJ r.:!c-rcnc\! ;o the t:hcmpthtropy, O,uf1.:mt:J)QQ'1 Jnr,nml Qf 1.~:1.•1·11tnry
0[1'1,····'m1"n1Ym \ 'r-h·nnmy ,\frthral A~1imo11 12.2. -~.:)...J~l.:! !)(11tnn•anrt J..mmal hulmrry, -I .?6~1-180.
ISO ,11,ttl\\ , ...:."' ·•H..\kMA, ,.~ ., 1~r;9. C:omp,u7ttiw 1..•1lk.l\.")' ,of drug, ln Epidtmlulo~
,,~, l'\Ul. :r,.;,1.. l',\Mlrrl•·J• R.J,, \\11...\0:\, ,q, l(f ·\\ft\tLt. H, .. iJ.. ,~u.~.
bh11nc ;nnpla,mo,l>. '/it1J1ic,1/ .~n/11111/ Ht!C1/t/1111vl P•0</11<1/011, 11 ur lw,\~no JJ1a11la!iomo,h ht b~eJ cuh 1.·~ in nonhetn r1ut."t.';1 ..1and
222-226 Atulralinn ·"t.•rumry)aurm,I. :lb :!ITT-:?71
l;>t \SOIJ.U\ J.H. 80\\'tJ;.~. I' \1,. ..:XO\\LL<.. 0.1•,. '.tlJU..\\',\J'\;, t.~.• tl<K.1' ft.[., 1;0 J•f.\R,o,. c,c., 111«,c..1. ,,.. L .i. ,.tirwrn. 1.0.. 19,;;.A ~t\t~h· 01 terru~vclu11-•
,,.-,sr.,-ro~. i,c;,, t1u<.1n. , ..,\. • u,\u,uuu '4,f.. 199~. Comp.m ..nn of ;i dr,~i,:c L,-,, ~·unk ,,hlch ~u·.: 1rnuplMJ110\!1r1 turd.el'\ JmmwT of rl:.-
1."Dmpc111(,.c- t.nhth1tlun F.l.lS.A an.d lht c.ud 3ffl:'tlutm:uJ¢n tttl tor dH..· ··\nwricm, ,,c1,·r,,tm:, \lwl/~11 .o\~r>tlllflun. 130, .?.CtO-~?.
dca•etltm or arulboUfeto 1uA,111ptu,rnn ,m,,1::1trut.,:u\rJ.-lnt11,tu~,,m ,;1 PrnfQC,.,. 1 .11.u.fiou. n.1. snrow>. n.~4, Gorn 01:\:0 11..:...1~:-- l.10\'fnl•
t,•ntrulti In cauh.•. A11>1tnlln11 Vt'tif'lttnry· /1.1.Jt1111!, :; 1.i;(;-:?.;9, iUl.aplCL<mo!tiS uan-,n1L~ion ~tudft~ l:or,l.JuaN undcr.:or,uolfod n,uut.i.1
1~ ,:oxrt..,lGtw. ~.• J.\.\1U. "''-\... U-\"· \1.0•• Mtl.'ttO'\, n.o.. Hl'.\I\J:\. H-'"" e.'(J)O!-W1.• tu .i l),•mw.t:t111t1.Jt mut,nuni ~ l\'tnt"'l,,tu~\ 1nd~1,.-~nl1w. otrc.J 01
m-,.nt·, ,1 .. 1981. UrnJr4tfon ot cultut~·drri·.\-..d -..olubl(" anti~"'" m tht lowi t>.l\h:m O:c:-gun •1,u~Tim,1 /Olu1111/ ujl'"'"rhuuy Rr•,·,,reir. "38. J.il-~5-t.
;1._!;g}unnruinn h.-"M for hew-inc h;ih~io.,:i-. ;ind nnupl.u:ino,1~ l t·t,•rln(lry P.t.... 11>.::,6. Tn)n4iml$,:on ,,r11napl:tsn,rn;,is.•1mu1Uo/drt• .v,•w ~Ork
lj:? 11 11 ko·.
Pt1ttUIIO(f1i!.1'. 8. ZlJl-297
.-kudL•n.yof ~t·Ji.·n,:t"..'O;. 64 .;o-u:t.
1S3 .MU~tlt,RI ('Ht, U,G •• BlOIU('o. [.f,, 1,.,"()CA",;. t.', \1.. <,(. ,.t r 1\\\ 'f'l'i"-. \\' .l 4,
1:".l l•Jr\'<O,.L. ).fdGfl. \ I ll\.,i;.:, )I., ~M~OVH,$• .\. !c )U."1 a.. r .. 1985, fu,1cn
s:uAm. 1·,1. l996. ~tab1h,hnu.. m ot the ttc,. '1:.1ri~ b..odttbc ~bomt!
.~1wp!mm,1 r,·11ttt1J,• ,·ucdm.• dR.~•11111,t an.,p!Jsmo~i~ m c.ttttl', Rrlrf....lJ
c-~ude pathot~n •.\nqpf1u,mt umtginak 1Ri:kl'a.,..falt"<! .\n3pla"m;.i1ucc~c,) ,t.,..._.,..,,,/Jf'}' ],mmnl. l-:2. 5~3-!1!1H.
in tkk civll cuhure. Jcmrnul 0J'i\ktlir11l f. nuw111lo~ ll, 65f",,..hE>.I
11.1 J'IJTGJtu.H.,. r 1'l':'q Epizoouolo.J;)' and coinrni ot :tn.at,Jld..~mo,;.ts in
!';S \1UAA\', P,l,:.., 1q89. ~·l<>h."CUJnt \'ti(dllC!" ~,~.U:l..\l ;11'\lffl.JI par.l~ill'-& ~ (4t(ft1r.
!')OU th ,'iirk.1. Jm,mflf ()j tht St.>111h ,V,frrm \"ct1.'nmu:i· .-b..~ fttt{(m. SO,
I, 291 ...299..
3-57 ..3,?.
1ss ~ur'l. w.o .. 1935 Bonne :mapla.,.mo-.is~ rlC 1r.ut,1ms:-ion a! -tnuv/(l,mUl
1;-:; 1'0TC.11.1tr,a. 1 1 1~1. r1e~ 1rnn,.:m>t,i011 ot J.n:1pt.a..,mo>i-.m South .-\Jm'iL
margm(l/rtto ;i bl:J~.c.-wa!\ltlwcst (Comu,c:l:at't~ gm, Q11d,·r~r,•1)-0()rt
Pmc.-:rt1.lngs ofthi' lmtrnati<11111I (.'rmf1.•rr11ci ott r,rk Btt1'1>:gJ·mul r:01111rJI,
Jouruflf O/l'tnt11tt1U)' -~llm<'t' a,ul ·\mn111t ilU1r4$tf) S. 9-20.
27~J9 JQnutu'\' lq~JI Cr.:ahamstm,n.
1s, sE!Ti!.. w.o,. 193:9. O\'tne onap~m10tjs: TI1t 1r.msm1~!on of :lnnp!osmn
171,l ro1G1rn R. r 1. 1q8":' l>l'mand tor lht: tJruJ~r,,icpoon ,.u:..·ine~ ~,RQm,{
Ot# and l;prtr,-1l,1oz.clQ11 etms to du,• hh..''°b1,.;(\i. rOt,maim:us 0Jl1ifrcms1.
ur~·bornt.! Lf1,c~c)o! dom\.·,uc- '<itod;. v\b~lr,i,i: \rmamtl c:,mj('h''I• ••
(),u/1.·ntfJXKJrl Journuf r,/Vr1erl11ary $rin:rl't1fld.'Wimnl J,ut,1s1ry·. 13,
1m 1hv lqn~~ram f>mn ::,erif$ Hvlnrmi:, 10 So111Jtc·m .\Jr.ai ·,. R,•1:t'u11/Jh•
9-lb.
\'Nwrni' R11S011r(ti, Pn.·tc.,;'tA.. 12--11 C)Ctobcr 1g37 l~ttt<.on!erencc
t.,"i ~·liff't., \\,(.).. (o .,u:x.,:-.m·K. n.., .. 19,13-- lmmum7...ation ~l ca1tfc- a,gatn,t Cttmrt.
hCitr1'\3tcr ;md the control ttf tkk·bo~ c.;ls-.·J~. redw:u<'r. g:,Ustc-'1-nt."...,:,,
1:: V1.Jtl'i1t'1J1. t. r • t~l89, \•t>lt•nmuy !t<'.SC.i.rch lmucute~ Ondl":".t1.•poort
and ht>.un,~ter~ CJntl,·rsr.~f)(}Ofl Jrmrnnl of\ ·n.,. 111wf)· Sc:w11n·amt :lmmf11
ti npubti<h<'ll d,1a.
lru!ust')· :!O. 138-158.
1it.t f"(lfr.11:l£n,, ..., .....,u" f.U.. 1~1ij,, Freit.'1...tH1~1ng or o1:\m1p!,11m11
tS8 ,1:J r1 \\ .u. A ou 1011. 1•,t.. 193:. Ba\·fne a:nap?a...mu~i..,= ,\ mt1hud uf
ma,i,fn,1/1•. Omf,•r1tr1xmt Jcwnml of \ 't•t;.,r1m1f) R~tMrrl'.• .S8. 179-180.
r,r
(lbitUnfng pure ~mu.int- A,m11ln:t.mlf m,1rfi,wli.and .Am11,!1w1m N'11tmlt
by 1ra11>ml,,Jun through Jmtclurc, lifRltt«mh lfeprm o/ rlt~ Owrmrof lj'lj f'OTl'Ul!lll\.. • · ~O· \,, \;.,'\I \~t,&\\', 11.\\, ,_ f\\'l'°G, '"'\ • J48J
\'t•11:nl1(1~ · .~·,i•{«J. and .r1n;mfll ludc,.'itry. Unfq11 qf ,io:t1h ,\jrlm, .~;?() U,•mun"tir.. uon ,11 ,-\uaµ!asmu ,111u1im1!.t• lnffi.~ mitlgut of HJ:.1pictJtlmlus
.t1m1u...\r,rfT'lt'a11 /1J1111u,J uf \ i~1.,•rftu1r,· Re1,wNir, .;,1. :?l;t-226!.
159 ")RIU-U.>. M 4,,\'51;.R'.\·(;Al.t>f~'n'\ . ,:•• 1982. ·\bt:1nmai bo\1Jlt"t'"n1h~1C
m~mht,i;lnt• prot~Jn:1- .,nd Kl)'\.'OprOltin'II during and after lOJL"Ciial'l \\Uh t8n ,,o,r,tl!'lf-R,J.l,i,, \{.\111-IU.O. 19H!l \'t.•tt!rin.an,•ft\•~.;ir('hln\Ht\114.'.
.\nnp/11.<111111111,rg/1111/e /JifH:h,•mfral n11d Hi1>11h,1·s/(a/ R"""""II c)nder.:opuurt. Unpubll<hcd data
Qummwlnwo,t.s HM, 66-1-672 181 POt<.n,,1,~. F r. & .,.,otr-,...t, ,,, .H•. 199L Vt.'t\.,rino1~ R~·.m:h tn..tirut~.
J60 ~OfUO!\, '·"·· j• ,ru:£11. R,J,,., H)KMf.'.$•f.\UL.K,1-.11.1.c .• :q83. '.\'t,.-onat31 Ond,•t"~rt Unpublbh<'I dot,1
•n~pla<mo<!, tn ~ coif ,\u.<1rt1/11m h~o'rlflll')' /bun:,,/. SC. 3 l8 18:t lit>J<.1:nu, t-. r .. !\un1rnL,:-..o, n ... cu<;c,. 11.e 1981 \m.?rt'.pL.; to trJn\rrtil
161 ourm1 11.1>., s,;,:oc.:.\.,. ~,1•• HMM. J..\.11 tw1~1,:,, , \. J.9lh. Suuni.ng Am,,,!am:i, mmgh~lll'\\1th lll111wl,o((tt r:tfipt•s.a.nd \Wrt:OX)'1 rnk1tr1.ws.
ch11r.1c1t,rlstic..; o( colouie,- ur .·~utplttsma Nfl.f)luUlleThvlll't Jh. Onder!!-1~,x,ort /01u1wl of l ~·li!rbtwy Ut•,.t.w1d1, -t8. 1 l9-122.
D,1r11ui~·,,,~1Qr nnd,1rsrmi$tih.·!'lo. lmr.rlcun J.,1,w1tl of\ '6mm,n· /l,.,,"',r/r, 18:. J'Qt<tn:n "· r. t .. \A-'\ 11.r,,uu1<<J, 1.-t,, 1!1{11~. l.'«llcition of ,\"11111kJn:11
.;2 .1006-1009.
nmr;.ln11/t frum Rli.µt<t'Plk:luJ.. ,\""'"" m~tlt!-. Ond(.r.H1•poorrJourm,I ~if
16'2 ~',O, •,{~tL. S()J..'-.~.... 1•.M .. PJ;JU 1 J-'1~~ wv• I. T.n.• l~:"5, Smdy OJ a.n \ ·c1t·rl1100· rt,·Ji.._·,m:h. ;r. 283-:.!86.
;i,uenu~m'd .-im,pfti.wm 11turgmulc..•\'aceinc in M.nieu- nn1ur31
16, rono1n1R. ra ..,,. ,. \'\ fu:,snt,m., t ,•• inn3. Jnt.:cu,·o). \'uuhmcc Anet
chnllcn~1..· o(immunily in mt tn2001rc ;He:- •\m,·rlt.·tw Jaumnlo(
immunog~n1dl\' 11f 1\1u1pl<1.-=11u1 n•111ral,• li\t' blood r.1~one:
\ "';.•t...•riuary· Rv,,1.virch. 36, 6'.lt-633
o,ul,•ts«f)OOrt Jm11n11: ojl .:tdtum)' R,-s,~rcli• .50 . ..!9 31,
163 onM, t wri.-;1)!\ . .\.;... t~\:\unt I rts.:r.• 1980 \ comparnth ..- ~tud\' of
Hl;i l"lll(dl~. I, I 4 \ \ '\ Rf :'\$ilUJ<{t. I , •• 191:J.I ObM-r\",ttlffO\ on thl' timt
e>,,J"u:nmental Jrmplu~mosi:,,.. in &'1J inditliJ1.1nd Hos rmu:,.< r,1tl1l"
rc."quh·~ br H1nm«tJIUfius &mw,. n~ m11hnc and aduh, :a tfi1n!tmn
.Ji.r;xrmfiwt '• 'i11t•riJU1t)· Jmmml, fi.fi. 2S2-26f
ibwpltmuo nrtrf'R:,u,t..· P(l}('ttf(j11gs,1/ ''"" Etghrlt l\ .1J1ltl Cmw.is, ()JI
16..t. 1\\1,, t..o. & )HU.fJL ~ .. 1971, Anaplnsmosb in a m·wbo:n calf. rih· T/is,.,,s.:,o/Orrt/f I~ 21 St-p:0111bor 1984. Ourb,sn. lwpclllfcof>outh
\ ,..~,wary Rt<YJrd, 100. 58. ..\frlt'il..

l\>3 P.\U1Ul. (i..11 , Jf.\lUSCT..\.)'., IM\'l'i, W,C.. & '\fC(;LtlRE, T,C.., 1986. 18b P(nr.lFTfft.. r.r., ,.\, rtf~S&Ullr. 1 1 , 1901. TickUJ.n'im1,~1on or
1mmu.ruz..1tle>n \'\ilh an ts(llare·l'Qmmon M1ri'i1.:c Jlro:e:n protC'<'t'I- enulc 11mftltJ;;n:,n l·~r:rml,· (),u/i..rrsr,pu,,tt /01,mat of\ twrumr.· Rcsn,l(h, .>;
., .1g.u11,., 1tnupla,.mv.:-.i,. S<-ft·m t', 231 l:?9.9-:3-0Z. rt-7.
~.,, .. 11,,wn. .,.,... c;rJn ,, ,t u .,n:,,.., w.c... &r ~1(:'f;Ulftf,
1ti6 tt.U..'\l~. G,tt .• oauu.li 1ffj rttl rc.1"fl£1\. r.T,,,. ,.,.,~ Jt£'.\~1lUR(,,, , .• 1t,ttt7 The pc~;:-.umcc of coto,,r,ll
Tc 111&1S. tmmunizntion uf card('t,·Hh J 36·ktfod.ihon ,,.rf:icc.' ptmetn .Ana,,lmma 'lntlbodit'""' .wd lucidcnr.: uf in llll'Tn mU1.,mi...~1m, Ql
induct.·~ pi:OlcCU\'n •tain)I hmnulo~o\.b .1:1d lu:wm10Ro0.>.~1m11ta-imt1 A1111p"umtt lntttuon, m cal,·L':). undl"r Jnborauny i'.'Ondi:1on.:-
ma.rgmak challenge J11fi.1,:w,m mut Jmmuml)·. Sb 1526-1531. ()ndrmcy,oor1 Journal uf \'<1;:tm11')1U.·~rcJ:. $~ • .5.ii-:itlO.
lb'"';"' $"AL\1Ut. (i,IL ~UAA,M;J.k\\'t,, f .rt., J..C>C'.A~. K,M Jt DROW!\, ,,.•c, 1S9'l, 1a8 RAI.\, fi.J., Jo,rr•H, ,,.\ kL\nTIJ\, r \I ,, lglib, b.\pcriffl\'nl:\l fnlr.j,nll"-'\00 Of
.'.\lolec:utor basl> for vaccin,• cfC>\'elopmcm ,1g,.1in>t 1.h~ 1.:hrlichinl .N11111/m111n mmgm,11,· tn ,:rO:\Slm.•d t:•1lv0 by th1: Ufl.!,1.,~U uc-k
pal.hogeon.'4,u1µ!tumc, marghwlt•, Pt,,m11uft1Jr,· fodaJ·, l~.1J31-.261l Orn11hWoM$. "'"'tJ!.IZJ'I, f:hriron, 1S,, •m-~.
 Bovine.- :mapfasmosi~ 615

189 fd \''-".\·8£iU.I:). \. noa:cs;.,210. A..., \f\YCM~O. :,:., (U)~'hTTI, '.\L\,, ,\'!,,(), 1•,,1.• :.t()ft m'l<.1.JP.,, ft.J.fJ-!-ltlf ,.~. 1. , •• 1g.;9. Ep1Mm!olog~,
und c:omrol of
DU"""' G. '- tYC'1U~l, '1.<.. 1998. fi\'lllU~llrm of :,n '"">'m••hnk.td Jnnpl,1..<mo,1...,, /011rnt;I oftlw South ~\fri«Jn Vi11ntlnmyA,~<Xf111,,m. 50.
tmmun0$0rbtmt ."i:)..ioa)' U>ing rt."tumhfnanl m;,Jf>r~urface protl,'m 5 for 363-.!66.
~rotog:<"ol dltigno~1~0! bQvin<> :mnpl,'smo:tis ln \"<'nen:~a.. Chmral ond !.!'l'9 1tcx:tM r.1. 6. •,,·Au..\t:r, \,-.n.• 196'6. A rapJd ~tJ.lalng technique for
Dit1gr10J.1ic l11QQrmn()' lmrmmology, 5. z:.,-262. .:\JJnpln.rn1". Amrtltµn Jmmtul aj\ 1'1t11i1tt1.1J· Rt'$Mrrh.17. 112;... 112.&
J91l MtCllt\, f.f,, 1961, Man~cin<'nt ofanaplS!imOii~ in beefca.u1c. 210 SC-111!\0U-:R~ vu, 1t,, ftbTtc. ,,. k\\'OKA1SCH. n,, 1966. Veq?.tric~n!lt:
1'rtx.'«{ffnJr;:1cj1hr ~,ientl, .\'ario,wl :1it(1p(a,nuni1 C(Jr,fo~nr~.:? l-:?.! Un1cn,uchun~1 mH ·\uup1'.a11tu m((rxim1/41 uml .o\, ,..._uuuk L••lf,nl,rift
Oc1obcr 1931. ~1nrk\'IU0. ~hssissl1>1>i. Jnr ·rr;>pt•ttm(•dti,i und PtttusUofngiu. 11. ::u7....:ffi0.
The ctfttl
)91 IUC'.ftEY l ,, • .IIK(J('Jr;. W+l • );Ui·\\'I.Jt1 1.0~' IU'\t..), L.W., \9':'fi. t965,\., \n1111fasmosis X\'11. Th" relation or
211 ~cunorurR, W.F. & JU:otnC' .,1.
01 (cC!dJnil low l<"'\'l,of chlono1nin·e!ln~ for e.indt!<I ~riod.< on :Jutcnmmune pm~--sts (O .:rn.:u,,..mia.Anwrl«m Joumol of\ tm•r;,1ar.·
t!lo- cnme.r srnrn oi an:1plASmosls. nu, Bm.•ioi· Prncrtnonl•r, ! 1 R1.)fr.'t"(h. .?6. 239~:!43.
73-75, ~1:i: <r.CURQtT>FR, \\.f. I. Rr>TtC ''·~ tgU:;n••\Jrnpta..mosr~ X'\111...\n .in.a.l;.:,f-. Qf
192 k1C:IU!\', C,J .. 8KO<.:lo:'- w.r,.. i,;.utwEn. I,{), "l(')StS. E:\\,, 19';'':" R(':$JS1Jnt:e auHMmtigens u\ 1nfoc1ed ~tnd oomln1 bovine Cl')'lhrocytes. Anlfri«m
to "l'\Jp!a.sm(l:;i~ af1rr cl!minru.ion or IOLl"fll .•\m1p!n1ma ,,wr,zi1u1J,• lurmml uf v~1~n11my n,~st'<lrch. 26. 679-~.

:93 si1cun·. t.t."" w~,, ~

.
t1tf1...-aion~. \n~rlc:,m Juumo.J o/Vt11l'.rimuy Rdt'arrlI. 38, 169-r;o.
1.0.. t93t Fffic:u:y o'f chtortetracyclin1.' aRJ,1.rtr.1
::13 S(Hl\OJTUl'R, w,r• lU>HC ,1 .. t~&a-. Btood ,l!-n:m fa<.tor5 ai.~ec:f •.-.11h
crythrophagoC)10-,.t1io in calves wnh o.napl~,1nu~h;. Am~ricau /<r.inml uf
b<>vull.' anopla..ntosi, when .1dntlnfs1cr.<l fw,- thole~ to <:>ale ln a ,·~mrl,wryR~Rarth.1~.19~U-IS9S.
n1c-dieau,d fted t>lud,...,d ~,fl.mincrol mis. Procn:dlng,of111,, S.11w11h 2M )t Hll~i!\"tll c:..,..,
UATUI <,, 1988, R;,dloin11nunoM... ;syfor .41Mplnsm,,•
Vmior:11/ ,11111p/fUmo,1s Cr,11fer,'11u, 21-2.1 tlcu,ber 1981, ~,.,,,,lit, 11mrf.:{1,r,lt· amlh<ld!C'!> in cault.Amerlt:mt Jounml r,j'\ '.?tcri,mr:1 Rt$4'ltrtl:,
Mils!islppi. 49,SQ.l-;o;
191 .,,ru:. ~, .. 1~,. Sn,41e, on ,napla,mo,;i,. Ill. ·\n au103J11ibod1 and :!Jj ,r.c.t"· r..1.. ,. \\llfT[, n .. l!,l'S:.?. The hlMUIMlht,lnl>)' or:u,o.pln!,,fflC,).n,.
,,.mptom.1ttr mac«.11ytfc anatmfa..J\11wricr111 Juurual of\ 'i':rtir.ntj• Pmr,YY11n)?- oftill' f'o!lr/1, ,\'nt1om1J .,l,u,(llasmwl.,r.1mfi"1t.·u« 2~2-
&J<>,m:li. Si Hli6. Apnl 196~ R<'nn \~\'ll<l.,

193 N~IC, M.. 1968. Annpla;moS,S. In;\\ U-'M/\N, o. km~T\r.. "· ~('d">J, :.:i6 ~u,1,10,1,,. s,p ,. TRIP,\OII. 11.r, 1~85-0h~"rv:ulon~on crythrocytic
/11/«trous Bloo,f D1~1ast'sof\4an cmd J\nlmal~. \'1)1. 1L Z\\'W Yor~, acctylchoJll'le~tcra!Ct' In rxpurimt:Huill)' indu(t:d u.naplasmo:,fcin c'n.,lq~$.
1
\ t•1.,•rl,u1r,• PnmJl:olOl!J.·. 17. 231-2.S·t
London; ~c4dtrnic Pr,~,.
180 Fi~TIC. 'l,, 198(). \napJa..-("nU)SS$. /n: A.\tSTUl.l, tt~l-, {ed .• Bot'llli' \ftvlici,:,.
~11 suo.s-t. o.~.. \\'~tLS..G..t. :&. \,·.,uut. F.1.,,•• 1961. Th.er ~cuvir:y o!am-lcurbaHde
a~tt.\l &r~ih iJ1iz~mtnn. rt:r ~"t:t!rutary· Htc.'Utd. 73. ;35-;.;o,
111111 Sul7f('1y. ,·01. I Samu ~rb•ra: American \'t1<rin~i, l'ubltcauor»
216 ..:M1n1, T.&.KIUQUR,L, l·.t... S89J ln\',estigm1onsinm lhc namrc.
l9': Rts-11r.. M,. 1981. An:.plnsmosis# In: JU~'TIC )1. & MCSn'kP.. 1-
1.--dsJ. Dmrasrs
rnus.1uon and pcevtntl<>n uCTe.,;as or $0urht·rn C3ttJc fever. Unltrd sm,,&
1t{Cb.tt~i11 tlw 1,o,,1N. Th~ flni;uc. ao..on •nil l.ondon: Maninus
/}e/J(lrtmem of.¼(r/culrurr. Btlfl.'l1U of,\111111<1/ lntfrmr:,·. Bulluti11 :So. 1
'vljhoff
1-lri6. \\a.Jhlt'4tort C.o\·t:,rruncnt PrinrinSt Ofticc.•.
19.8 JITSTJC. ,~. &. ~,u_u K. J.P.. 19B4. F:rtnil) A.n:tpl:i,m~l,1Ct'J:f.'. In: tklf,Q. ,.ft. it :n.9 ,r, rrrt.R, LJ • ,utUR. .f.. 19s:3. Th<' ap~rcnt <'Til.dicatfon o( Lhe
1101 r ,.c leds). lkf'KD.' 's .\!a11ufll of5y::m'l11fe Bt11:ttrioro10. \'nl l ~n-3pl-3'.'>rr.os~ CJ.Ul{'r illHC \,•hh antibiQtlc-~. V,w·nnm,· .\ftrlior.t•,$mn,I/
ffaltimor~ oncl London: \\1Ulo.m< l\- Wtlklns. .l\m11wl Clink!tm, ~s. 486--iHa.
199 rt;~Ttc. ~! •. SnUK0\1( <:. 1; \\'fil.'fUI, c.1 .. l968, J\J\ ilct~nUc"\h:d AIU'IJ)lmmlt .!?~ fi.TAAI\, M · J:0.::.\',:, ,':.~I .. HAIH. J.A, ~ fi\\'ll\'(':, S,,\,. 198:?.
margtnnlt! ,~ecii?L" PuH,·,•tttfiuJ?$ of tltit St-t11!JII}' s,.~-ond ..\nt1ual .\ltN!tmg ()/ or
lmn1un(>t)iochcm1c•I MwUing ,\t(t1f/l1u11111 m111l(fnafr11u,ller In
tin• U1thc>d :Stmt•, l.itl!5tock SaniwryA<s«fmion. t968. ~ew Otlton,, U.wnal·cmor tmd"'·""m Stile... ,,ith pcroxid,,ic•antlptrox!d.a1e
US.\. tc-chm,qt.:e. .-tm.1 rirarl Jaunwl of\ 'trerinm)' Hcst:a.rrlt. 43, ~;!)...'.l,U,
~ RJ~"TIC'". \L Q \\.'Ant.\C!-t. .,. "'~ 1963. :\J1ttplD!1on\O~);\,, \ l. StUd!e• ,tnd ,\ Xii .!,iU'\\'Aftt. C.G. IMMI U\.\~ \,, (,f(l~USH", ,._ ,1.-(;ltlll,n.. li-::9, rh~ 11$-\. of"
hypot.h-t-sis contcming the rydC' or duveJopnu:n1 of th~ i:aus:aih 3G«?nt.. 1
~
-:.hurt tmd IQng ..\Cling o-x)1eir.it.')1:llne for the trt•mmttm of :\napll'J$J1ttt
A.mr.ican /ounw.J of\'.!t1.'rlna1)1Rl-.Sl•,1rd1. 24 _2'ii'...2:r:'. marglnnl~ in~1tncrwml1l'd calves. /01111111! ()flhr Somh .,v;unu
:!01 r:ulll~. n.H. & to,·1. P,'.. 1.9r.. lnfecti\'it}' ufA11a;,Jr..mm mdrgtnflfr-aitat VL"ttrmmy /ISS!)C11Ul0t1, 50. a3-8s.
lngdtion by poumlioJ Insect vt-ctc:>r.ii. At1h.>tie1t.n Jounznl of\ t•11•r.",wr,· 21.2 "111 l,Ut, D•• rsu:nirw,.. W,'.1,1 ,t.t.TCH. G. A ,.;urn,F.R. K.-l ., ,~ai,,
RC'.,~11iJ1. 38 !t\2~ttu0. f rani.mluton 01 equtne oabti;:n')s1, -and t>ov'lnc :snaplbm<nh b}
::o: F-OltFRT:S.. ILH•• 1•usn. w \., ,1(:CJ\f>R\, H.J.,, ~uu. f.\\' 6- c:()u.1,•,, y.c_ 1!169. f)••mu,trntora/l,/pkru.<. /or,mnJ oJtl,-• .v,,w l'llli' Iimomologfrt.1 Soclr-1)',
Culltidat ,m<I Tob:midnc as por~nual voc:10~ of anupl,,.,mo,is "' 8$. iS-76.
~. lis.....,i<.<.ippi ••-tnnnls of tlJI! Emo11wlogict1I Socimy,,fA.na•rfttt. 62. RGl...068. X.C..1 ~ftUI-NI, "" ~oc,:..:. ~-~t .. llJ\\',\RDS-. \\',. l,\\'1~r.. L\., lt,\IR, ,_,+ & a.u:mos. !t.J.,
:!03 F.()H\-, T..0.. 197:! Th(' inhlbitOf)' eftec., ofimicJoc..a.rb on cixpc-:imrntal 1989. Oc1~1Jon o( ,-olonlt'$ol ,lnupln.,mn 1111111:inn/,• ln,allv.uv iiJ~nd,af
,n:,plasmosb 1n ,plrnf'aom}zed cal\'~~ R1.-sMrcn 1,, tt~u·ri1u.>1;>·S<'i(•nt:e. thte(" D1•muri·e,:;n,-:ipp. infected a~ n~inph .. <>r aduhs ..'4n1ttr!c1J1: J1J11mal
13, 5H1-:t:?2 o/Vtltriunry R1'1~1,v:;1, :tU, l lftl-l.38S.

20: iwtn. T.O.. J97J '[h~thcr..1p}· of bovine anop1a~l(J:j,1S- ~\ith 1m1dm~arb. 224 .,,Olt!-.1.. ,,·.u., ,qaa Ondcr.'itt:pOQn \·~1crinary ln\thutt. Unpub!lshl'<I
i'rtH:~dingJ of th<• s1,.·t1, .VmionnlJ\m11,tu..m:n,1,co,1fl'n·nt\', 19-20 <lain
~larch 19;3, 1.a,wi;,i.. :-.c,·•d>. 221 b"!Otflsit w.u .• 1990. Ondc:ir.t~poort Vl.!11.,rin~ry lns~lfut~- Unp-ubl~hed
to-S 1.0B'f. T.O.. .UltAAULT. l,tu \L\ZZOIJ\, \'_ r<O:,.)_. l,E." IUMOB.\01' A., 197,i, d:tt.1.

lmmunlt),' tn bo,in~ n.naplnsmosl!- :tiler elimination of :t,u,µtnsmn :126 ,wrn. ,._.,_., !TAOUtR. c tt.. 19:'8..·\n~plosn1osls:,\ rtglOnal wologlc:11
,nargfnah• {nf('(.rion~ wtth imidocarb. Amttfcan !tmn:(l/ ofh1uri,u,n- ~ur\'n :,md nnl :mtlh!o1Jr th~rap\' in infot'"Ccd h~rds~Journul t'>l tJ:~
Rl!Jt1«rch. 33. 993-993. Am,.ncnn Veurtm11;o· ,\Jt.Wfrol .~$Jocl«tlo11, I 72. 1310-1312.
2u6 f.Omt. T,O, & M;,.i.t.ou, ,· .• 1..q,~ Hllmumwm ohhri;amer-.;tah~ oibovmc ~:n :,.'\\'}Ff. O..L... >tt;JU)()(.11, W.J
~OA>IW~. JJ..n.., 1901.Antmlia~~oc-.a,,-d \\1th
;,i.mspJ~smost!<. \\ilh imidQttlrb. Ami"rirrm /():tmol o/Vet(!J/11ary Rt.<,·orrls. anl?strus in b«-(lui•U-rrs !nnru.lored \,i\hA1urpl,uma m,u-guu,(.
33. 1931-1933. Endocirwlcg. and O\~n•n cruu1ses, n..
ril,gan1>1Dto·. 22. &13-<,.lO
207 tlOJn\ "T.O., Sl)IP...DS. f.f ,i ,\MERAUlT. T.f.. 19:'8, lJaminauon 0(1hc,camer .228 nu ltf~. , .. 19~0 A1raplt1>Tr1a murginnf..• 1 Cit•n. ,p~c. So\"'* Thi! :nar~1izl
,I'll~ o( bo11ne nnupl~S111osfs with n long-actl~o,1·1e1rncycilnc. point< rn th~ blood oi <•nit .;u1lcrtng ir~rn n •P«ffic di>e..,,., lr,•wn of
,\mrrie,m /011r11al o/1 'oreri11t11y Re;enr,;lr. 39. I 1IS-l 116. tlw Ci,ov,.,•mmem b.'t11rinnry .rJ.n.rteriol(1g1~tl of 1/lc Tra,u1.raaf. 1906.. 9: i..fi.1
616 <1. nos TIIRtl; Rickcttsiru and chlamydia! di5e11se.s

~ nu,1u M, .\., t910. AnnJJhtsm(1 mfltgi11a!,· ·.rl'tn i'Jnd ~J>t'C n<»\. ~ ,\ enc\ld~ a 19,kilodahon protein conwn'Cd m nJf recot,.1111-=d Ana,,!o.sma
pro101.o.u1 o( cuult· .\ Ca'U$C of Lhe so•c:dJNi ¥3ll,idmcss. Tnum'<l,nl '1>«1~ /t:fi\tcuu muJ lmmm111)•, 00. $139-5! 1-1.
.\f,•dlcal lmmra/. 5. 110-11 l.
.: 1;, \11.C.\l:\:Q, O , C \RIO!\. r., .. In, A,1, * RISTlf.'. ,1.. 19i'3. EttitJ'1 of
23011iEltrk. , .... 191u. ·Ane11,Jwmm mutttiiwt..-. An1,•\'. ~.:nu ... ::1nd siwcfc.•-. of1he ijltenuawd i\nnpfmonn mnr.gi.,urlt' vtccrine under 1.-iburntnrv and fw1d
Ptotc>zoo. 1"i'l111,i1rrio11• 0/1/11• lloynl So<·ttl) •/•0111/t,lfriro. 11. t,9-;2. condi1ion_.; m Cofombin. Amartean /0111 rial ()fV"t~·riNnQ ll,....~,Ych, l~,.
"-11 nouJ·a.. .\ .. 1911. FunJ,cr im "'-"1igotJous in10 .inapla.&rno~h or South :?29-233.
Afr!<':m c.it1lt!. First R~/X>tt of tltr L,Jr,,c1or of\ ·,.,,.,rnnl) /l\~i!r:tf't'lt, Unton
2a6 w,~~e,. o.ci.. 1,;81. lmmunoi;M1uhn rospor.;es of caulcossociate<I "11h
o/S0111/1 Nriu,. 7--16.
rrcrud~·nt -1nn,,1n..,mn m11tgfnalcinftc1lan, Proo¥d:ngs1,1f1ftc
~2 rn~tWl. \...1912.. Galb:iQ:k11~~~ u(1mpon~dtanlt" and the prourc:,rve ~4.·ml: \'mfottnl ..t,w,,1,ism,Hfj Ctut}1.'f'r'lt<"I!, 21-:?3 Oetot,tr 1981.
inoculauon agains1 thlsdls""*.Agrim/111m/ Jm,mnl oJ 1/rr 11,1i<>11 q/ Swr~,iUo. ,U,sls&ippl
,orrrll .'l]r/(11, 3. l ~
2.i'"" \\.\t.L\CE, ,, JL. i'Jti:-.1.o~Of t 111·1htt"M.."\!t1c .1ce~khollnt•\tt1r.1~e ~cuvit~
Z.l3 nco~~,~.!oC .• \,·,t...o"' u,r ,. 'l·'-'°°' r.L ~~:: &lbdi<,. Tfo-l/r-rt17:ind .ind ib reiati..onShip to o.smotJc fr;i~Uity o[ c::~1.hl"'OC)il.'.,. in bo1.'int-
,..,,.,.,,/«$>1:n <pp. lnfoci!n~ ~obh: .,n«IOJ)<' fll11wtm~,s ni!;"t ,Ham,. .tn,1pLbmo.)i\.Amtnt,m /()urnal of l'l!WrJ,u,1:• R,•kflf\'h, 24. 55--61
1838). In s,ou,hem Afrlcu. Onde.rstt'JHJOrt Jo11rnttl of\/1!11..rr·maf)· H.t·. . l"arch.
J\I, loJ-16<,, :.!..a8 w.\~Ont..\,, c... • ,,u~\\IA, \\.1r..., i~,1 Scv-,re anapla~mo~ti- ln .t 4•dt3\'•0Jd
culf. 81d/,•t/11 ofFpW,,,1/r Dis"""''"' ,lfrir«. 19. 21i·11 I
~ llfO\lf"W.~. ~,.. :9r. A rev1l·wor the- kno,,fcdg4: ot'\'ecmr, Qf bQdttti
anapl~moib., /11~,,'nL-... r..>..-. l-d.,. Workihop on hCfflQp.ata'tt1~" .:.1_1! \\rt(:-.."Wli\t lt:.h.. tr,;()( " · ~ \I tUIUU)!\, ~J., E'Wl,\:(j. , ..\ 1ro,,n·,t IU).'" h \111,
(onapJo;m0>hand ba1"',io,lsJ, Cl-\T. C.1ll, Co!Qmh1~. 19;;, JJl-118. L\, 198:-. Jnftt·tl\it~· of th.rec ,1\1tnpli~mamargt.nllk btib.u,..., fo:
J>erma.umor w:drrsoJt/, Amt,rlttm /oumat nf \~vwrlllnl)· Ri'fl'arc/J . ..S.
~ 'rOOO=tOVtC. A.A... GOSl..u.JU'.., t.r .... G.-\.RClA, o.• l!t~ L,.ilunt!on of J. nl!w
96-99
lnng,-oetin.R ox~1.etrocvdlm.1 fonnutnion ag.il1:,t 31~p1Mm~JS- 1r1
Colom.blnn cattle. Truf)e,unttfi::in wtcl P,trruiwlosi·t•. 30. 236-238, .;.t,,;o wtt::St\:UCJnt..n. c.. 19;-o_ K.'\:pt•rienc~ ,vuh n.'\·~ri11 In the t/ratment of
236 TOOOnO\I(." M..t\., lOXG. ft.ti.$ M.(,'CAU(), ..... 2977. Comp.,rbon or r,1pid .ina.rt:t5.m0"\ii. in d,tln. c-anle. Hr" ll/u,l Book/or rl1e• \ fnt•ii,:nr.: l'rnff.t.ii<m.
1; 9-1!.
c:,.rd aggluunauon le~ with lhe romp!tn1cm fixmion t~t !o: dia~no>i~
of .-lttaplll$mn m.nrgi11all' infection In c:m.te In Colombia... \ 'IJu:rmlil)' 23i "IES£.'i:U0Tl'f.lt, • i. 197j. Resc-ntth huo lht' rl'lath'l' hnponance of
.\1/aob,ofngi.•. 2. 167-177. T,1banldae r1np1c.•r.. In mt't'h:um~:d di~c-a~c trnu-..,ni,'S!o~. Ill. The
23';'" l'(>l>OROVIC, A.A,. WPr;.Z., t,f\., 1 OP£Z, .u; • (,()'.'o'.1J\U7., r; f .. :9;3- Bo,ine •pid,•ml~ioit'' oi ann~ln.mo..J< in• Our•c'<-Sclttom dru.l\ form. T.'tlpln1f
bahniosi" and aiuspl~t1:0IO'l....; Cu111rol br prt<munillOR and J\JJlmal llt!t1l1lu,ml l'tot(cu-tion. 7. 15-22
chttnop;oph~1axs.s. Wt1f'rm,~11Uil Parrosm,l{)s::,1, 3; '-'2-10 i 2.52 \\111.IA\~S. t.1. ~ ius,~. t.w .. 19GR. HlocWI tratbfwHon, rluriO!!,p:'nt!nt
23,8 fODOitO\ IC;. R.A,. V14'G\tt-.(), O.,G., (.:()~I.At.ti.. E..h 1. AO:\M>. LG .. Ht73, hminc:l.t13plol....1T10""i~ .~1m..-rlco11 /cmmal of\f1..•1t!ri11t11'}· R'N-1;."t'l1th, 29.
Chemoprophrta.,·h :!j::;lin.."'1 flalm/<1 big,•,1111m .,nd H<lh~trr argt1'1thlt1 703-,10,
infec-cion, ...,m,•riam Jmmwl of\tmrlnm,-Rf'K(Tf(lt. 3-t. l l:i:t-l 161 . 253- \\1tSOS .\.J. P,>11,;u11:, 11. • nu1.,1,s, ,..r.. 1si80. li'!perim<!nto1l
:.(59 TRUI.fSU>OD• .M..s.. sw1n.1u... c, ni:,uc. r.n .. 19:,. Ut1,int' !eL1) r«t'!pOM41 u, imn1unll:.1do11 o( cal\1.."S aga-Jt1"ltAlw1,tn.m:~ ma11:fm>ft" Infection:
• 1,wptasnw nu,rgfnt,lt•, ArtJ11rlc111, Juuwnl ?/ Vt11t•,J,:t,r)· Rm111d1. ;tt Oh,.i!n•um,n on thr- use of lhrin~A t'(•ntmh• and.-\. ,111u~'11111l11. \~c11,r111nn
1089-1090, Pam,lrMo;iy, 7. 305-JJ I.
240 rnuu., ,,, J,..f, '-' ,, ~~>:i-:, ,.J.. 19;9-. Obs(?r.o.tion. on th<:' p:nholnJll· ,,r '2'.1.1 h'ILSCl~ ,1,,J. 'lHUOl.\'"'· j,.1l .. 19;8. Some CIT1.•ClS o: tl"Cfuctd l~t:~· {ni3.i.l1
Anaplmma r,wrgmalc.• inft'tfions In tnufl'., \ustralicm. ·tdC'ltllCtt$ ltl on the de,·dopmc,nt or 3.n3pla1omo.!loil. 1n Hoi lndi,u~tto5-S itte~
\·f;l'{('rtl:llf'.\' Mit•IJC'-'· ~. 7.5. Aus1r,,ltm1 \'N.:n·miry Jmlf1111J. 5-;. 121 I ?•I
u, f\\1JltAu~ ••,r.t .. 1.962 Control o1 :mtipl;srno,,, l)y tctdmg~n .\n11h1rnic ::,as \\ll~n, ,.,.. Tf:Ul;;.\~.\.S. J;.F•• t-Pl ... lri:!,., G. *- \'JC.SOIU.J."11 .. \,r... 1918. J\
(Aur,mmyc1nl. l'ronwfh1gsr,f rlw rourth ,\t1flo11a/ ·\rmp!as1m)SJ$ t'Ompari.Son of four ~:otogjca11c-sb in lhe d('tl-ctlnn or hutnorol
Con.f;.1r,•ncv. 26-27 April 1962, Heno. i\c,;id;.i ,trUlhDdltS 1n Jn11f'JCL..,"1t'>J.i:S in C;1Ult.•. J\1151rnlit111 V,•1i.•ri1J?f')' Jour111H. ;-t
242 uu.t-.~:tilkG,, r... 19;0, .N°OIC)On babl:slu1o!'> and an.u.p!;i.,;mo~l"> of C'4-stfe m 383-38$
:i.·1adaga.)t,U \ .A. Immunity und premuuUitm, B. lipi:r.ooilo1o~·. Rt'11 t1~ 256 \£nu11,t\1. 1.1,, ttK.\\'tsuus. Y,, 1981. nu.• 1ron,mi\.~lon o: •.:..ndpT,mm1
dEtmwgi't't tf~ \IL'cl,"t:fn,· \ ,!t,.!rimurc-d,•.t A7)':. Tmpl('Gu:t, 23, 439-434 um~m,1,, 10 c:mll· bv blood ,ut'ldnR nnhmpnd.,. R1..·.futti1 t~uritu1rtth.
24:S ,,,'>ER, 1 "- ~ A'-!DA()NtO, U 1,. ll:)87. Oi\'A prol.H: ... for 1hl"dtte-t1icm 1,r 33. 37-14
.-l,u111fmn,o ,1•ntTt1h1 rmd A1w11/u.<ma nu11,.in,1I~. ()1111,:N~,·µoorr JQtmud ::5":" ,.\l)Q(i f L, ~ I ILl f.Jl, t>.• CID,\.."\, ,\U·. ... USCOI-.. ' .:-.n, t,OV. Tta.n'tn'l!~JOn (>)
<>/Vf't{t1{11ar_v Rr.,w1.1rh, 5-1, n2J...tt.l7
,110('1'~.tl mufJ!ma/t" Thcll~r b)' nm.le:~ of Ut.•mmr.·mormt-dir$<Jnfs111e...
2,U \~~Ut. L.~. \ICG\JJKf.. T.C., llht,\ffll, C...H O..\\'h. W.t•• !o)U:•.\P. \,. P-tl•\~C'j, C. It'd on in ld:i.ho Hrld·snfecu.•d. chronic (";lrnt:r tm\'. .-lmtr1co,, Jaumt>J of
"- );.,·owt.r~"· u.. v•• tR•• 1992, TI\C 1\nt1p!hsmo margilr/lll' m:>p5 gem.~ \11•urina1)' Ri?:o:,wuh. 47 226§,-1271 .

•
 48
Ovine and caprine anaplasmosis
W H STOLTSZ

Introduction J\Jl(i]J/asm11 memetemm. the name proposed for a hithcno

unrecognized anaplasm of sheep and goatS.ro IS closely re·
Anaplasmosis of sheep and goats is an anhropod-bome dis- laced to A. 011/$ and may also cause clinical disease in sheep
ease caused by the intra·el)1hrocytic rickcusial organism, and goats. bUt. unlike A. ovis, it appears ro be less patho-
Anaplasmct011i.~. It is usually a subclinical or mild condicion. genic for goats than for sheep and is not infecch·c for cairle. 6J
but moderate to severe clinical disease is generally charac- A11aplam1a margi11ale. a 5pecies pathogenic for caule, may
terized b}' fe\'er and a "ariable degree of a11aemia and lcterus also. under cenain circumstances, cause latent infection in
thac may occasionally lead to death. Recovered animals re- sheep and goals.~ 16·~~- 29· l<. 36
main carriers of the organism. resulting in premunicy. 1,·hich ln Giemsa-~tained blood smears A 011is is morphologi·
persists for life. callr indistinguishable from A nlflrgitwle,sr.· ;;; and appears
Anaplasmosis in sheep was first reported in 1912 in Zim- as irregularly shaped, almost spherical. imra-crythrocytfc
babwe..; Although a number of repons or ,'i/lnp/asmn infec- granules s1aining a deep purple colour. The organism~ (in·
tions in sheep and goats in Africa soon followed.,. ~ .,z the clusion bodies) vary in diameter from approximately 0.-1 to
repon by Di Domizio in 1919u is considered to be the first 0.8 µm,?:!, 35· 37 1\i1h an average of0.5 µm .35 Within erythro-
authentic description of anaplasmosis in 11on-bovine hO}I~. cytes. 60 to iO per cent of the inclusion bodies are located
In 1924 Sergent et al.~9 reponcd an ..i11apkis11u~ lnfecliiin in marginally and 30 10 .io per cent submarginally or
sheep that was not transmissible to cattle. and Lestoquard22 cemrally.12 3·1• 56· ;, ln comparison. up to 90 per cent of the
differentiated the catt$8tive organism from the A11nplasma inclusion bodie:.~ of.-\. margi11ale arc situated marginally.34
spp. of cattle. describing and naming it A ol'is. Differences hecween .4. t)llis and A. 111argi11a/1>with regard 10
Ovi11eand caprine anaplasmosis occu rs in many parts of the intra-erythrocytic locarion of organisms have been
Africa. ~orth and South t\merica. Asia (including the Middle found to be s1atis1ically ~ignificanr.51
and Far East). the southern and cemral pans of eastern and The uhrasrn1cmre of the margin.ti bodies of A. Ol'is in
western Europe. and the former USSR. 2· 13· 19· ~8 - 32· 3;. ~3 • :;3. sheep er)'lhrocytes is very similar 10 that of A marginale in
5·1· 5~· ®· 1>9 However. the disease may be more widely distrib· bovine erythrocytes. 1s, :w. H Development of A. 011is in sheep
uted than present data indicate. and gone erythro~1cs. and in the tick vecmr. is presumablr
The condition in southern Africa is or relati\'ely little sim ilar to tl1e cycle of de\'elopmem proposed for ·l. 111argi-
economic importance. Because of its often insidious na- 11t1/e. m. 1- .,s. s; The inclusion bodies of ii. me$nlltl!mm are
ture. the deleterious effects caused by A. ovis infection. morphologically indistinguishable l>>· both light and elec-
such as weight loss and unthrifiiness, are probably errone- tron microscopy from A. 011i.•, but a distinctive feature is tl1a1
ously ascribed to other causes. Although mortality due to fewer 1han 30 per cent of the inclusion bodies are simared
anaplasmosis i11 sheep and goats is seldom observed in marginal!~· in e11'lhroc~1e.s.
southern Africa. it would appear that under certain cir- J\pan from the morphological similarities. complete se·
cumstances. overt disease. abonion and mortality may rological cross-reactl\'ity has been demonstrated bi:n,een A.
C><'Cur. I 3. 35 ovis and ti. margi1111leY· They are nevcnheless immunologi-
call)' dis1inc1 as-infection \\ith one does not pro\'lde immu-
nity against the 01her.l<i A11ap/11s11wo11/s and/\. nwsr1eterr1111.
Aetiology and life cycle
on the olher hand. appear 10 be immunologically more
Anaplasmn 011is is the main cause of O\ine and caprine closely related. but cross-imnnmity is neverthele~~ incom-
anaplasmosis. [t is more pathogenic for goats than for plete. a situation somewhat reminiscent of the relationship
sheep06• 63• 69 and onlrvery rarely is it infective ro cattle.2<l· -in between A cemra/e and A. marginale in cartle.'>l
617
618 ,lcr.,~, mm; Wdtct1Sial amlchlam~dial d1scast',

Epidemiology determined. Ba~ed on the observation~ ebell'here that .4. 01 ·is

and A. marginal~ may share common ticl: vecrors.48· ;a one or
Anaplasmosis to sheep and goa£, ha5 bl'en reported from more of the tick vectors ofA. ma rginale or A. ce111mle rcponed
most tropical and subtropical pan~ of the \,·oriel i. 1~ 28• 3';, 13• from Souch ,\frica:is. 39 may• also prove to be ,·ec1ors of A. mils.
~. ~i. 5n, titi. ,;s With the exception oL\amibia.A. m•is has been llowev~. during experimental studies on the lick transmis·
reported from all the countries in ,outhcrn Africa and i~ be· sion of Theilerill .,,•paraw.5,\1 nhipiccµhnlus ,•1,errsi t:l'err~i and
lieved £0 be pre\-alent in most sheep- and goat-farming areas. R11i11icep/rnl11~ ererisi mimericus transmitted only T. s11pnmra
In these areas. small numbers of pa~asltcs may frequently he from splenectomizt.><l sheep infected \\~th bo:h T. sept1ratt1
detected during rourine microscopic examination of blood and ·I. ods.
smears from sheep and goats. Considering that microscopic F-xperimemal studies have indicated transs1adial trans-
examination of blood smears and serological techniques rail mission of •\. ovis by Or11i1l1ot/oros lahor,•11sis II Rhiµicepll-
to distinguish bet,, ecn A 01iis and .•1 marq:i11ale.20• ; 6 thefoct alus bursn•0 and R/1/piceplialu.< 111m11ic11s.'" Transovarial
that sheep and goats exposed 10 A. 11wrginale may yield sero- 1ransmL<sion in Dermnce11 ror ,i/11arw11 has also been rr-
logically positive results without being immw,e to ,I. ovis may ponecf.1? :\dul!S of D1mn<1ce11wr andersoni have been found
be or epidemiologic significance. Similarly. calves infected 10 be na1urally infected:18 and imras1adial 1ransmission b)'
,,ith ,l 011/s ha\'c been noted to react positively to A. margi. aduhs of this tick species has been demons1raced."8 Other
na!e anugens m the complement lixauon and rapid card uck species that have been mcriminated as vectors include
agglutination tests. and yet were fully susceptible 10 Hyalo111.i11a p/11111be11m/; Haemaphysa/is orophi/a.37 fJer•
A. mt1rgi11nlt! infccrion.20 thus rar 4. mesaererwn ha~ onl)• mace111or albipic111s. D. on:idemalis and D. t•ariabilis.'.S
been recorded on the Dutch island, Arneland. but circum- I laemaphysalis pw,cuua and Ixodt!s rfci1111sare suspected of
stantial evidence seems to indicate a possible wider occur- being vectors of ·l. mesneterum.63
renC'e In nortl1ern and nonh-eas1ern Europe.63 Considering 1he ease with which A. 011is can be transmit·
The pre\'alence rates of A. ovis infection in sheep or goats tctl by the subinocula£ion of blood. tra11smission by iarro-
\'t1ry ~-onsiderably within endemic areas, and ma>' be influ- genic means or mechanical transmission by biting Oles
enced by management factors such a, the comrol measi.1res would appear w be distinct possibilities. I lowever. t!Xpcri-
employed and the movement of li,·estock from non- mcmal auemp!s co transmit the parasite bi means of bites
endemic to endemic areas.35 Although no ex1ensive surveys of the sheep ked . .\.Jeloph11g11s 01•i11us, a likely candidate as a
ha\'e been conducted 10 determine the prevalence of A ov/$ methamcal vector ofA 01,is in sheep, ha, e failed. even when
in South Africa. limllcd microscopic observations in sheep sheep with high parasimemias were used as the source of
have revealed a prevalence ranging from 33 10 90 per cem in the infection.68
endemic areas. ;9 . .,.. Similarly, in '.\101.ambique. its preva- Intra-uterine transmission ha, been reponed in sheep10
lence mnges from O10 85 per cem in sheep and from O10 90 and in goats:1 In sheep. rransplacemal iniec,ion has been
per cent in goats.2 In India it has been shown that approx- found to occur during the se<:ond and third trimester of
imately 60 per C<!nt or sheep and 70 per cem of goats are in- pregnancy. but no lesion~ have been observ..-d in foetu,es or
fect«!d.27 while in Kenya the prevalence oi infection in goats neonatal lambs.''7 In contrast. a high percentage of anifi·
from diffcrem localities ranges from 22 10 87" per c-ent.s.1 dally infected female goats aborted or resorbed their foe!·
Although she<.•p and goars of all age;. arc suscep1iblc> 10 A. uses following acute infection.:1
m•is Infection. 1• S·I. at, 110 older animaJs suffer from a weater Sheep and goats that have reco\fered trom A. ot•is infec-
reduction in haema1ocri1 values. St>."~ tion remain carriers or the parasite. and in areas where both
o, en clinical dise:i.~e and monality are rare in non- sheep and goat5 occur. sheep may be panicu!arly imponam
splenec1omized animals. 11· l, but sp!enectomy drastically as rescr:oir hosts because of the lower pa1hogenicity of
Increases the susceptibility of animals and usually result~ A. cwfs in this host.;;;
In a recrudescence of parasllacmia •.relapsel in carrier \'1'11i1e-1ailed deer (Odot:oile11.,; 11irginimws1. eland (Ta11-
animals. 6· -. ,~.2D. 2ff,J~. ~7• sr, le may thus be assumed that any ro1ragu.;oryx) ancl blesbok rDanwlisms dorcas pl1ilfipsl) are
condition thac results in suppression or immunity or de- su,ceptit1!e 10 expt.>rimemal infection wilh ,1. or•1s. 12• ta. 34 • J 6
creased resistance 10 disease in animals, such as debilitating and 1hese hosts could ~l as potential reservoir hosts or In-
diseas~. concurrent infections (particularly ofo1her arthro- fection for sheep and goats.11 a: Although wild ruminants
pod-borne rickeusial and protozoal illfectlon, ~uch as ltnbe- usually develop onlr mild or subclinical lnieclion. severe
Jio spp.. Tliel/erin spp.. 1"ryµnnMoma spp. and disease may occur after splencc1om.y. 18, .l 1
Eperyrl1ro;:0011 Ol'is. 2· 5 · ~ 1· ! 7 • .;9 , <;<, he-cl\'}' helmimh or tick A \'3.riery of :-onh American. Asian and ,\frican wild ru-
parasitism.31 malnutrhion,13 and other stress fac£or~ (such minnnt species has been found to be suscc.r1ible to ..\. mar-
as pregnancy). as well ns variations in the palhogenicily of ginnle fnfectlrm (see Chapter 47: Bovine anaplasmosis).
different parasite strains and indh;dual susceptibili1y. could ;'liarnrall)' acquired ..\11aplasma infec£ions isolated from blue
lead to a higher prevalence of clinical disease and monallcy. wildebeest {Co1111orhae1es wuri1111s), Coke's hartebeest
The cick vectors of.-1. Ot'is in southern Africa have not been (Afc<Jlapi111s lmse/ap/ms coki1) and Thompson's gw.elle
 o,1neand caprine nnaplasmo,i, 619

(Gazella 1/wmpscmlil resulted in mild lnfeclions in calves adult animals usually de,·elop only a mild form of the
and provided liule or no immunit}' to subsequent A. 111<1rgi- disease.g,, 56 Various stress factors and splenectomy in-
11ule challenge.15 Positi\•e serological reactions to A. margi- crease indi\·idual su,cepubilit1 and predis:pMe 10 acute
. s., 3t ~, Su -
r1a/e antigens ha\'li also been recorded In impala Uepyn•ros d 1scase.
melampus), wmerbuck (Kobus e/lipsiprym1111s) Gram's Splenecrotn}' of chronically infected (carrier) animals is
gazelle (Gm:e//a granrii) and eland,2~· 26 but anempts 10 re- frequently followed ll}" :i severe r<>t,rudescence of parru.l-
Co\:er A11aplll$1tW spp. from fmpala and Grant'$ gazelle b}' taemla and mnrtifestation of clinlc:11 disease. indicating the
inoculation of their blood into c.attlc have failed. These importance of the ,pleen in controlling the infcc-tion and
n:~ult, ~eem to Indicate the po~~ible existence of hitherto mainrnining immunity." · 1"· 20 , ' ' "'· :,9
unrecognir.ed .·\m1plasma spp. in wild ruminants, som1t of In cxperlmemal transmission studies, in which A. 011is-
which may be more closely related to:\. ot-i.~.11 as has been i11fec1<xl blood is u~ed as the inoculum. the prnpatent period
demonstrated in the case of a sable ancelope Hippotragus may \'ary from 5 to 40 day,. \\itl1 an average or abom two 10
11igerl in South Africa. 61 Circumstantial evidence would three weeks,i o. !::. :15 5" · \; Generally. the duration oi the pre-
similarly seem to indicate lhe susceptibi(iry of wild rumi- pa1e111 period 1s inversely proport ional to the number of
nant hosb font mesaetemm.1/3 parasites in the infective inoculum.2.1 Similarly. following
Katurally occurring mfcctions \\'ilh A marglna/e in wmstnis<ion by a single pair of aduh Dermace111ar a11der-
sheep and goab ha,·e not been reported alrhough these spe- so11f. a prepatent period of sllghlly more t11an ,l\ week, h~
cies may become latently infected with ,1. 111mgi11ale after been recl>rded in a splenec;1omi1.ed sheep,-111 whereas the
experimental lnfection.30 Prolongad rC>pea1ed passage by prepatenr periods are shorter when there are larger num-
subinoculatlon of blood In sheep ma) lead to the gradual bers or ticks. ·,a
adaptation of A. marginale 10 th is hos,, which results in a The parasitaemia gradually i11creases 10 a maximum
higher parasitaemia during infection and an attenua1!on of level one or t\\'o weeks after th<' pnrashe.s ha\'C (irs1 become
the organism forcaitle.9 · 14 lG,Jo. 2•· 36··'"·"'' microscopicall~ deteciable. 3 · 19· 56 · 57 In non-,plenecto·
Sheep and goats cannot be conslstcntly infected exper!- mi;:ed animals, the maximum parn~itaemia may range from
mentally"ith field i~olates of A. nwrgi11ale.- u;. :?S lndlc111ing approximatelyO,l to-l percent in sheep, and approximately
1ha1 t11cre are po~siblc s1ralr1 difference, of A. 111argi1wle l 10 12 per cent In goats, bul in spleneclomizcd animals the
,,~th regard to it> abllit-y 10 infect these hos1>. Furthermore. maximum para~itaemia is more \'ariable and may range
the carrier state ofA margiiu1/e in sheep and goats may be of from 2 to 90 per cent. 19· 20· w The course oLl 01,is intecrion
~hon duration 11 29 In splenectomlzed goats this tran~ient in spleneccomized sheep is identical to that of ..\. mnrgimtl<'
carrier siate has been found not 10 exceed 208 day-s.:!'• in caule, with the number of parasites doubli11g approx-
A11ap/asmt1 Ol'is has only rarely been shown to be i111,'lrely every 24 hours.:.;
infective for cattle. :\'umcrous attempts to transmit it 10 During the period of rising parasitaemia, parasites are
non-splenectoml;.:ed 48 and splenectomLled canle ha1·e cemo,·ed from the circulation by phagoc)•tosh of infected
failed,-· 9 . 3Q. 14 · ' 7 '>band this characceristk of A OL•is is often er}<tl,rocyres. This results in a gradual decline in the num-
used 10 differentiate it from A. margi11ttle. This difference ber of parasites and a concomitant development or a pro-
would ,eem to indicate a greatt>r host specificit} In A. 01,is gressil'e anaemia The haema1ocri1 1'11lue gradually
than in ..\. 11u1rp;i11ale. However. short-term suni\'al ofA. Ol'is declines and rMches its lowest level a few dnys after the
in splcnectomized cah·es has been dcmonstrated.• 8 bu1 peak of parasitaem!a when parasite numbers are already
t!t t,._.
confirmation of infection req uires the subinocula1ion of reduced.
blood into splenectomizerl sheep. In one study. 20 A. 01tis The severity of the allaemia is 11suallrgrearer than can be
could not be reco,ered from a splenectomi.ied calf at I 7 accounted for by simple removal of parasitized erythrocyte-.
days pos1-infcc1ion. but rhe animal was ~,1bsequem!r round alone. Severe anaemia in the absence of a high para-
10 have retained the infection for up 10 262 days. Xo loss of sitaemiaJI. 63 suggest> :m autoimmune response which re-
1•irulence of A. 01•is for sheep ha.s been ob~et\ed foilO\\ing sult:.5 in thl' phagoc}10sis of infectt'd ns well a, uninfected
mainumance in splenect0mizcd calves.10 · 1o erythro,·)ws by the rNiculoendothclial sr.s1cm."' In non-
splenectomized sheep and goats. with average ma.\irnum
parasitaemias of 0.8 and 2 per c;ent respectively. the comt·
Pathogenesis
sponding average reductions in haematocrlt values were ap-
The pathogenesis of 01ine and caprine anaplasmosis ap- proximately 40 and 50 per ceot.s.. However. in individual
pi:ars to be Mmilarto that of A. 111nrgi1u1/<>infec1ion in cattle animals. the reduction in hacma1ocri1 values may be con-
(see Chapter 47: Bovine anaplasmosis) and, as in bovine siderably greater. Similar reductions in haematocrit ,-atues
anaplasmosls, anaemia results from e1: 1hrophagocyro- were obsen·ed in A mesnererum infection. despite the pres-
sis. i ; Unlike rhe situation in bO\·ine anaplasmosis. there ence of a low parasitaemia,63 In srlenec1omi1.ed allimals.
appears 10 be no marked variation In ag~ )usceptlbilily 10 haemn1ocrit \'alues may be reduced 10 well bclo\\ 10 per
A. 01•is infection in sheep and goats, and both young and cenr:10
620 <tc110~ nwn lhrkensial nnd chlamydia! d,~~ases

rhe corresponding drop in lhe haemoglobin concen- of anaemia. such as pale mucous membrane,. thin \\'3tel)•
48
.
crauon. J.l l .56. 69 1'f se, ere. w1·111ead to h ypoxic d amage to blood. and increased heart and respira,ion rates are
various organs. which may ultima1ely resulL in the death of evidenr.5 · 22 • $9
the animal. lcrerus is not consistently observed. even in cases which
Sheep and goats that rcco,·er Crom A. 01,is infection re· end fatallr:'5· ll9 Submandibular oedema of 1.lle jowl and
main carriers of the parasite.22. J.1, l:i ventral part of the neck may be notlced.J~ In milch goats. n
marked drop in milk yield ma>• occur. and this ma~· persist
Clinical sjgns for several wceks.H
The :emrn of normal appetite and an impro,·emem in
The clinical or patent phase of tlte disease la:.1s approx· the general condition of the animal may serve to indicate
imately one lo rwo weeks. 19 • 30 In animals that survive this recovery.31 However. recovery from clinical disease Is
phase, eonvalescence begins wilh mcrcased erythropoiesis. usually slow,07 and anaemia and emaciation may per·
which b characterized by reticulocytosis, anisocylosis, sist for several weeks wh,:,o 1h11 nutrition of the animals is
polychromatophilia. basophilic 'stippling of immature inadequate.);
erythroc~·tes and 1he presence of Howell-Jolly bodies. Con· Abortion and resorption of embryos have been reported
valescence i~ usually slow and mar last from se,·eraJ weeks in domestic goats infected experimemalli• 1,~th A. 01,fs, 3
10 a Jew months.<lll During this time. 1he 11aematological and 111esc phenomena occurred panicutartr In those animals
clinical signs gradually return to normal. which had developed a high fever and severe anaemia. Ana·
Anaplasma 01-is infection in sheep and goats usually re- 1>lasmao11i,was transmitted tmnsplacemall) and frequently
sults in subclinical or mild disease. but occasionally severe caused anaemia nnd death of the foems iu 111ero. rn foetuses
clinical disease and monality may occur. ,\Then mortality and live or stillborn kids. the A 01•is-parasitaemia ranged
does occur. its rare is seldom very high and may be due, in from 1 to 12 per cent.
pan, 10 concurrent infections with other haemoparasites. 11
Because of the greater pnthogenicity of A ouis for goats.
Pathology
clinical signs of infect ion are more fr~quelltly ob$erved in
them than in sheep.;' ~i.sa. 6'j The severity of the lesions vary .according 10 the duration
Fe\'cr is not always presem and temperature rcanions and the severity of the disease.35 Numerous reports include
are quite variable. A sliglll rise in body temperature may descriptions of necrops)' findings, 1. 3 • :?Z. lll. 3S. $ 6 . 60. Gs
occur tH the onset of ptltent parasimemia. followed h>• a re- The gross lesions to a large extent resemble those found
mm 10 normal for a few days before it again becomes C!l· in cattle that die of anaplasmosis. The carcass gcmernlly
evaled.3 The maximum rise in body temperamre usually appears pale because of the anaemia. and mar be slightly or
coincides with the peak of parasirncmia.22 · 56 and may range moderatC'ly icteric. The blood is thln and watery. and ema-
from •10 to 42 0C. In some animals the temperamre may Ouc· ciation, serous atrophy of ralry tissues. and anasarca or de·
tuate during the clinical phase of the disease.21 while in oth- hydrarion mar also be apparent. Increased amounts of
ers it may remain elevated for more than a \\'CCk, before straw-coloured fluid may be present in bod)' ca\ities. TI1e
returning to nonnal. 3 • :tr. lungs are pale or congested, and oedematous. The liver is
The clinical signs nre usually most pronounced at the enlarged and yellowish-brown and [he gall bladder may be
height of anaemia, which occurs a few davs after the parasi- enlarged and filled with viscous yellow-green bile. '.\'t>phro·
taemic: peak and approximately one to two weeks after para· sis and enlargement of lrmph nodes and adrenals may ;ilso
sites first become microscopically detectable. TI1e :;everity be c,idem.
of clinical disease is to S()me exremdependent on the degree Microscopicall)'in blood smears. evidence of a regencra·
of the rna.'(imum parasitaemia,2 2 but severe anaemia may tive anaemia. characterized by the presence of reticulocyto·
occur In the ab$ence or a high parasilaemia: 11• .c. However, sis. anisocrtosis. polychromasia. basophilic stippling and
e,·en in animals with low haematocrit values. no ob,ious lloweJJ.Jolly bodies, is found. ln animals swfering from
clinic,tl signs of disease may be observed. 1• 3 • 56 but signs or se\'!lre anaemia a monocytosis \\~th erythrophagocyrosis (as
rapid fatigue and Se\lore respiratory distrt.>ss, indicative of evidenced by the presence of 1>arasi1ized erythrocytes. or
lung oedema. may become evidem when they arc herded or their remnams. i.Jl macrophages) and eosinophilia (after a
forcefully exercised. 3 · 56 Occasionally, animals may die number of remissions of i;evere anaemia! may be obserYcd.5
,acutely upon cxertion. 1 Changes in parenchymal organs characteristic of
Generally. only slight lisLlessne.ss and weakness,22• 56· 69 anaemic and hypoxic states may be seen in various organs
or a loss of condi tion5 are noticeable. The appetite usually (e.g. centrilobular degeneration and necrosis of varyiug in·
remains unaffected. but reduced appetite or anorexia. ren$ity in the li\'er. and degeneration especially in cells of the
rumen stasis and constipation may be observed In some Se· convoluted lUbules of the kidneys). The canaliculi in the
verely affccled an im als.:<. ss. 6o. 69 lf\,er may be distended as a result of the bile stasis. ·111e red
On closer clinical examination of affected animals signs hone marrow is u~ually hypcrplastic, but in chronic cases
 O,ine and caprino nnaplasm1>si, 621

may show e\idence or depletion. Haemosiderin aceumula- The CT test has been round 10 be a reliable technique for
rlon In reticuloendothelial cells is us11ally most extensive in the detection of infocllon in goats: 11
the spleen. Similarly, a competith•e inhibition enzyme-linked im-
munosorbent assay (CI-EUSA) ba~ed on a major surface
protein 5 (MSP 5) 8-ccll epitope which is consernid nrnoog
Diagnosis
1\11aplasma spp.wwas shown to reliably clcte<:I diffNent iso
The possibility of anaplasmosis should be considered in all lares of A. 011isin both acute and chronicallr infected goats. 33
cases of anaemia in sheep and goats in the uopics and sub- Results suggested that the sensitivitr and specificit) of the
tropics. test was high.
During the acute stage of the disease. a diagnosis in the Serological techniques fail 10 distinguish between A. 011is
individual animal is usually made on the basis of clinical and A. m11rgi11ale and such differentiation may be 01 epide-
signs. the presence of the organism in blood smears and miok>gic significance. D~A hybridization technique$, using
haematologic-c1I evidence of infecrion. However. when clin- either radio-labelled or biotin-labelled spocies-,pec!fic
ical signs are most pronounced the degree of parasiraemin probes,• 1• &I can differentiate between these species and ap·
mar have already declined c:onsiderably and confirmation pear to h(' more sl·n~itive than Lhe examination oi blood
of the diagnosis may be difficult.56 ln addition. the pre<ence smears bi' light micro~copy for !lie detection of A oris tn[ec-
ofa regeoerauve anaern1a during this stage may complicate t!on In sheep and goats.
the microscopic detection of parasites lA/1(IJJ/a,ma lnclu-
sions at rimes being difficult 10 differentiate from basophilic
Differential diagnosis
stippling in reticulocytes and Howell-Jolly bodies or other
stained particulate maner in erythrocyte,:. :-.ticroscopic de- Th<' clinical signs of anaplasmosis in sheep and goats are
teetlon ofinfccrion during the chronic or carrier stage is also non-specific and 1he disl.'ase, therefore. should be differen-
orten extreme!}' difficult because or thl? exceedingly low tiated from other infectious and non -infoccious causes of
parasitaemia. Jn doubtfol clinical cases and during epide- such signs as fe..-er. anaemia, icterus. listlessness. inappet-
miological suneys. serological test~. subinocularion or ance. los~ of condition. constipation, and submandibular
blood inro susceptible animals, mrd 0:-.IA hybridiza1lon oedema of the vemral region ofch" neck.
techniques may provide additional confirmation. Fever, listlessness. lnapperance. and loss or condition
The subinoculation of blood from suspected carrier an- may l>e caused by a \'ariety of infc:ctiou~ and non-infectious
imals into susceptible splenectomized sheep or goats is an diseases including l>abesiosis. eperythrozoonosis. ehrli-
effective war of de1ecungt1. 011is infection. bu cthe technique chiosis [Ehrlicllia Ol'i11a t1nd Ehrliclri« (CyroeceresJ phagocy-
is too expensive and time-consuming for roullne applica- rophilr, infections!, theileriosis, trypanosomosis and severe
tion ..,, \·arious serological techniques including the internal or external parasitism. Of these. cperythrozoonosis
complement fixation (CF) test .30 · Sll. 6 ' capillary tube agglu1l- and vcrminosis (e.g. haemonchosis) are probably the mo,t
na1ion (CT) tesi.2 ' 3 1 direct- and ind irect fluorescent anti- likely 10 be confused with anaplasmosis in southern .\frica,
body (FA and IFA) tests 1~ and the rapid card agglutination and either may occur concurrently in the same host. Ep-
(CA) 1esi3°· 6 ' hO\'C been used. Due to 1he high degree of er:,,rhrc,zoo11 01•is frequently occurs as pan of a mixed infec·
serological cross-reactivity between ,t ol'is and A. tion with A. 011i,. 12. Iii, J.1 ;; but the former Is distinguishable
margi11ale.'Jl:J· >ll either of these may be used as antigen. but by its generally smaller size and eimacellular location.
A. m11rgi11a/e antigen is most frequently employed because Malnutrition, plant (e.g. Bm.~sica $i>p.) or mineral (e.g.
of l ts greater 8\'ailability. copper) intoxications. or deficiencies (e.g. cobalt and cop-
Sheep and goats. on a\'er~e. develop positi\'e reacuons per) may also cause anaemia. lc1erus may to a grcacer or
in the CF test from a few days before the onset of patent lesser cxtenc also occur in certain plam and mineral (e.g.
parasitaemia. 18• 56 t\mibody titres are highest during patent copper) intoxication~. In addition 10 kierus. inappetance
parasit.aemia and lowest during the carrier phase but, gener- and wc,1kness. chroni.c copper poisoulng Is also characrer-
ally. sheep and goats develo1> lower antibody titres than do lzed by haemoglobinuria.~ 7 which is not a sign associated
caule infected with A. marginale. The majority of sheep and wiih anaplasmos-i~.
goats remain serologically positive after one year. buc some lt should also be borne io mind that the usual!)' mild
known carrier~ have been obsen·ed to r1:11ert 10 negativity clinical sign~ of anaplasmosis may he exacerbated b) con·
against A 111nrgi11n/1> :rnrl , \. n,,i,: nniigf'n.'18· .'m ThP r:A tf'SI cnrrf'n! lnf1><"1inns or non-infpcrinus condition~.
similarly seems 10 lack die sensitivity to detect all carrier in-
fections in sheep andgoats.30, ">!l
Control
Uninfected lambs born of infected e\,ieS may reac1 pos-
itively in che CA test a.n d the CF test for variable periods. but Both specific and supponh·e therapy should be initiated as
amibod} levels diminish or dlsappear at the age of approx- soon as possible, preferably before the onset of severe
imniely three or four momhs.67 anaemia. in order to avoid morrality.31
622 "en"'" 11,.,,. IUc~~u,lal .u1d chlamydia! dlsea~c•

Chlortetracycline and oxytNrac:,cline arc effective for the Judicious comrol of' external and imemal parasices
treatment of A. Ol'iS infection,.35 Anapla,,mosis can be should be practised. 35 Since intercurrem viral. bacterial,
succe$$fully treated by the intramuscular administration of rick.ett.sial and protozoa! Infections may precipitate
oxytetra<.·ycline at a rate of 8 m 10 mg/kg body weighl daily. disease.35• ,f these should also be comrolled wherever
adminisrered in t.'(luallydlvideddosesat 12-hourintervals for possible.
four to fi\'e days.~• The adminbtration of dMded doses al 12· ="ellher oxyiecracycline (12 dally Intravenous adminfs·
hourly inte1Yals is desirable in order ,o maintain higher blood trations at a rure of l O mg/kg body weight ) nor a combinu·
levels throughout the period of treatment. Howe\·c.r, two t<> tion of ox11erracycline (four intromuscular administralion~
three daily Intravenous or Intramuscular adr11inistrariom; of of a long-:1cting formulation :,t a rate of 20 mg/kg bCldy
oxytetracycline at a raw of 10 mg/kg have also proved to be weight at se,en-day intervals} and imldocarb (two intra-
effecrive for the tre{1m1ent of acute anaplasmosis in sheep.;~ mu~cular adminis1rations at a nne ()( 5 mg/kg body weight
A single intramuscular administration of a long-acting for- a1 an inten·al of l.J days} eliminates A 011i;carrier infections
mulation of OX'}'letracycline. at a rate of 20 mg/kg body in splenect0mizt.>d sheep. although these treatment regi-
weight. Is abo effecLh·e in preveming mortality in sheep.67 mens do result in a transient suppression of parasilaemin to
Symptomatic t reaLment and rest arc bencficial. 35 Sup- a level where parasites are no longer microscopically detect·
ponive therapy may consist of the admitustration of hae- able in blood £mears. 59
matinics, tonics and fluid. 11 Gastrointestinal disturbances In practice. sheep and goat, art• not usual!~ vaccinated
such as inappetence. constiparion and rumenal stasis may against anaplasmosis. Animals could be immunli.E:d by ani-
complicate recovery a nd, where indicated, attempts ficially infectini; them wicb A. of/is-infected blood3 ~ possibly
should be made to correct such disorders by the adminis- followed by tetracycline treatm<.'nt to comrol the infection.
tration of appetite stimulants. rumenomrlcs and mild laxa- Care should bt- 1aken whc>n selecting a sui1abl1? donor an-
tives. In valuable animals suffering from severe anaemia. imal to a\"oid th!? accidental transmission of other blood-
blood transfasion may be Indicated. borne diseases. One or two administration, (atan Interval of
Adequate protection against the elemems and easy seven davsJ of a long-acting oxytetracycline formulation,
access to palatable feed and water ~hould be pro,~ded. In administered during the prepatcnL period in experimenrally
addition. anaemic animals should not be subjected to un· induced A. 01•is infecrion in sheep. has been sho\\11 10 pro-
necessary exenion and rams should be reseed before being long the incubmion period and to reduce the sevC>rity of the
used for breeding.35 clinical reaction.;o

References
\..-..:O:o,,,,. 1~0. A.t1apla1attt 01·fs.. .Muiuh1> Re:pt>n of Lhl? Reglunal Vcterin.af') IU OOSAn1.:i-.:. '-. ~ bf()QUAllO, J. & \111.t.llllft•MASC<'J.tJltT. ,\.. 19~ P'.bs.:l.}t<!' :3
L8botntu1 le--- for th~ mon1h 01Junu-:uy. ()cp.;m.mrn1 01 \'~h!"rinnry lra\·c,rs ltt phtc\!nt.a cfo nttbcsli!lt<l (ll'LS c.•t tfl· lttrtp!Afmuo,•IA ,'18'frlt
Sc"ice, Repuhll,· ofSou1h Alric.,. ,\/(1/fr111,. 3J! 320-3?1.
2 \R.'-IJLD. 11,:\l ,: 1MA\".\SSO.'i ~:v:0.> UL'* ,.\., 1983,. lic.t~and urkbomf! u ou JOit. P.J .. l•l)-; Am1.p!:s!lmo'!.j\ (absu:at·ti. 77,~ t t.•r,·rinn?' &•-.·,,~d. l 'f.
haemt.>pazil~hes of sht'Cp and gonts. ~3$onQJ dl5t:ibuur.m in 126&-!?67
~lc>zambique. World .4rilm«I l!~•·I,,... ~5. 2~35.
12 • X-tC.k. l:., 1£,1.::t. D1t' F.mpfflngHrhkci1 t.lt1t EJcnttntUopi• li\r V1nµ/Ju,m11 u,.iJ'
l """'°·· n \t ..,, ,. ' " ~lfl~R~. r H ltlP'll \r,.ri(llAv"ntt~t, In fm11rm t'f1 R(wr und E/lt'f:>'lhm:0011 011/J.. Dttt111dw r,opc1n11uvli:J1tllt'llt X"111chrlft•.u;.
goo,, In S0u1h. \Inca anllklall>· lnfec,.,fo11h ,.\r1<11i/11,w11 ur~. Small 411,..;2
Rumi,wnt Rr-~·md1, 3, 191-197
13 r,tt,JlDT ~.,., l\4:.:6 ..\ pre!!1nimu)' cummunic,uion on ii':r octurtcl'\c~ a!
4 B.l\'.\'S, L.r.\,. 19:1tt. Annpl:tsmo!-t~ t>f Jh(•t.,p. t.'i.ouri,tul)' foumn/, 68,
.;!)()...IOI, .:anapl.llmo~h m ~hn4p and ~o.itfl m J1.1h~:"dn,~. Jonmat ,;f(Atn/Nirttnl'i'
PnritdQK:, 11htl TltnntJitmits. ~. 2.ar~2.rr
s DE Ae>ct., c.,.. 1.930. AM\ort note on chromr t'lna;~ino,t, a.nd ~ndetio.s.lj
m small rumlruuus atcerspltu·u.-ctom~. S!xli?t.1ml: Hl.'J>Oll ofllw /)TM..•toro/ u 11£, \'('\C.. M.\., .. •!lS6, \mmn.l JJi$t1USi-S iu ~ltllt .4(1/c't1. JrJ 1..lfln. Pzertorio:
\ 'tt,:rfl111r,· SH1·ln!S wut..-\nlmul 1,u/mrry. Union u/S-01,tlt .tl[ric.a. 3-10. C~ntra! '="''~,,~en(:' Llml1l'd,
6 ::n: KOC~.<,,.,. QUl'W,,.,s-, J... 192...1 ,\ '-hort prdlluUusr) commumcat.ion an 1.5 J \.T'U.l. v.1t.. J9'4 BC!'1:Uo11 mkro~t>p)'~lu.d~ o(...\rtaµlas,nu, ,,i,J.s..
an.apla;:mo~1,,;: of -.hccp a.~ obsen:~d m St1ulh ..>Jrica.. Bull,•m: <16 In ~u·zt .~mt'r'l(QI! Jc,t,fJU)/ of f,·~•;erJnnl)• llsN•(Utl:c 30. J&il-189.Z.
de Pmlwlogot cxo1iq110, 1i. 651•653, 16 J..ES.Srut rt.u. ~ ~m.1 re.- ,t.. JR7'9· ,,, ,,fun cutrl\Jtion of,1 ~11,1f1>.t11:n
1

; Dt ~on., c,1 4dJUl~LA.'4:, 1•• 1926. Splt':t~torn~· In dc>rn~Mfr;ued -.rnimah.. 11f{1,ymttlt•· hwu-sio.n ofa.nd dl'\"t'Jo1lmt-nt in uonlnfoettid crythrocytr,
-andil'5 ,c-qut"l.at•. whh $pC'ClaJ r~t·rcnc-e 10 :tnapJn1'mo~1~ in sht·~1>. AWl!fJ,(,fVt Jaunml ofl/~1r?ri1U1')' Bt•sl'ttrt/1 '10, l 7i4-17;o.
Elrr;m:J, a11d Trt'i:1frh Rcyx1rlf '1/ t11r DFr«thr()f\C':,·nnmy F.dum:frm an<I 17 ,;oc":-;· i.:.:-,t t!Nlti. o ...,.\•lopm~nc of 1,ut11tnima ,1utff:U1!1k m
Hegorcl,. Umou ofS,,111/r .I/rim, 369-1811. i.xodtd Ude-.: Coordinated dl'1.~lupm~nt ot" J. nck~t..i:11 or~an:..m
a rit t>Q,,:2111. <,.. 1919 oe.n· Aa11,111,.<,1111 mm1111111/, (:o,pl di loll, nrl and tu h,:Xc t1ck. l,t. ~,v1;1t, f.,., 11\tk.J.~. 'cd$J ••\!vrp/Jolu~·.
~ngut •ncmlca, Form~ nnaplosrru,uc.ht di plropla,.'ffloJ. Clml!a Plty;Jq/qgy mu/ &!/1e1dour11/ Eroloi;,•o/Ttcts. England:lill!,
l'iWin,1ri,1 t t1L.<:u-g1111 di fXll/tfll s,1m111ril1 di tglcna. ~2. 203-31 ! . Hon,1>0d t :J.
~ 00~.\n-t:...\, ~. & LtiSTOQUAfm. JI., l9JO, LeSrutaplMmO!'t('.Sdt~rurninnnt.s. 18 l;IUl!i'I, r.P, A SU,""Tlr. \1 • l!163. An1tphtl\mo!\lS. vu l:.'t_perinll'Mll'II
Flo11111· , "114!rin11in~ ct Jrmrnal rh• M.ttfl'rim: r1i1Jri1tttit('lt tftf :,m1«lmil. 82. .-t,,aµllt.tmn ot1I, lnfl-ctlon inn whltt•tail("(! deer Dama r·1tgi1r1lmn:
!2;,-l39 ·\tnl"l1t'Gtt /oJJmal o} \ Cu•rin.ary Rc1n1rcl:, '.!4 •.56';'"-5i2.
 Ovinl'and caprineanuplu~mosi, 623

19 ,um.. n. 6'.' roLRtll., f. 11367. Swdles on \no11losmaarrs 1n(«tion. I. l'' '"'"TtG \IUT. L.t .. l~Ul- l.ur fr.igc dcr Obmri.g(•J <l~r
U>ursc ur !'ipontnncon... 11\ft•cuon~ 111 ,pJcnacton11sc.:J Nlw;orfa.n "heep ""hnfpiropl•5Tlln-en in .v,crbnl<l...,han ,rrans~uk..10n1••1..1r/"fi,r
and gont,, 8111/t1f11 ofbi1/UJ{)tk /)is.,,,;,• 9f,Vriro, lb. o»-;o, ur1ss,·11KlmJit!<-i1,.· uurl pr,,J..·11-...q;iz•· n.,,hcfl~·;md,·. 67. 176-18\1~
20 -.;u n u.R. K,L 1981. lnfocuon uf 11ipk·nt'<'tomi1..ed cul Vt·~ whh .~,:a11la..~,,m ,.u 1~·,;,ru;\lftv. t .•, .. 1935. Un nou\,•nu \CC1cur d:tn, lu 11..i1hm~lon d~

<>1 iJ. Amau~m /ounu1l t>/Vtli.'rlnmy R<'St~nrtll .;2, 209...:.-2096.

1 h~rnopar.i~iu~dC-$ anin1ilU\ (!Qmes,tiques• <'lm111tmlum1, lnl,or~:.,i.•,
~cum:mn 190tl .-,,males d•· t'lnstum P,u11::n·. r~1. 250-:!58
:!l ~vnum. J.A ... t98;; .•~1u111ll1m,a mftttittn~ in w1ld ;lnd domt~l11,'
.J:2: HA.~'.fG.,\Jffl 1·.t" 1ttt;. !"•rmttr.<mor stl:1uum. \'t'f'lchr dc.•.i-
ruminants;.\ rC\'lcw Jm11rr«I nfl\'lldll[t D:toos.'S. :w. 12-?0.
h<'inop.uJ...itt-~du mouu,11.;1innp!n.~mo ovlsc.•1 T1wllt·rit11,1-t.·ond1:a.
22 Lt.:>lQQUAtm, 1-•• 1924. ~u>.knn: n(lt\.' ;,ur 1.es Phopin"trm>)-1.~ du mouton 1Jutlt·1fn tie ta S:J1."l1~~.: de Pt.ulu,logit• ,,.ro11qm,. 3o, -i~ao.
C!n Algcnr t 'anapt.a ..mosc: ,,1:(lp/asm11 oui.J. nov, ;p. Br,llttiu tie /11 SOclt!rt
.:3 rnsnc, M,. 1960, '-napla$mod~...,1t11V11rcr.t :,1 t1e1crinnrySd~nc(r,;.
ti( 11nthdfog;r,wo1ft/t1t·. Ii. 184-787.
112-179.
23 u ·~ffOQU.,RD~ r-.. 1926. 1cs pimJ)l..as:mos.cs du mou,on ct de l.i chC'vrt ..i I HbTU.;, ,,,., 1.9t,a••~t,)p1.-.,mo... 1,
ltr. \\.'EJ.'li\1.\S. I), I, Kl\ft(;, )1,. l.'d,-.•
Art11iL'tS flnsritul Pasrtur di'1gt•n·r. I, 122.-317 hif«tlor,s B/0otl l>1m1.,a rtf.1/m: 0111/ Anfmt1/,,. Vol II ~...,,. York.
:t.:S u<.-..11 RES. 1 t!H9- t~ \'ncd.n:uion d(.)S bo,ic!e1 contt~ 1·,'.lmlpJ:lsmoS£--. l.undon: .~endemic l'reo,s,
L"An,,pfa,s.ma inocu!~ ~u moumn N ~ la chh"Te ~·oth.'nut1 dan$ J1 llh'trc. M.. 1981, A11apl,l,mo1,-. In m,nc, ,1. ~ w 1~1, HI, r.• eds:. l>ls,r.1ts
rorganism~ do cd l.-S.~s M..irru1lt~ Cl Jeur !l'.J.OG est alor, un excellent of<.:attl< m 1/11• Tro/li<s. The H;igu~ Boston, l,indon·. \lamn11< :-:ljhon'
,·,ccin pqur I~< bO\icJo< c,intre l'arophl<m<>,<e I., pl"' gr~,·~. (1111/,tirt d~ l'uhhshcrs
Ir. S.Xi~r/ d,. l'tl!hofogfe l','I011q11,,, 12, 76S-7H 16 ltlb1 IC. M.,. ~l&l~i)\lC ... "\\r'ftnR. C.J .. 1969 An .)I IClllU)t(.'ti '\1tt1pl1u·mn
23 LOHR, t.r.,. :,,tJhn, u., 197.3, Ga.ni~ :m.cpla~rno"1...: 7he l~lnt!nn or ,1u1rginnl,•\11c:cint!. Proen:"tUngr 1J{1hr Unluul S10:t"1 lJ1 l~lc,,el; Samr,:ri
,\,wplmmo DJ1tanim:> frnm antrln~. OitrcJ,r;ft J'm 1ra111•1111:, .1l:.t1111ml AsstxlaliUlt. 7:!. :>6,,,(\9.
P:,rn.sitologi~ ?4, 192.-197 1; R\"H, ,.s. ,(j;IL!lER1'. ,•. ~ " ' IHll.. J.L&. ltRn .'., Jt•• , ~ . -\napla1tm0fi,,1nd
u., 197.\, Dcrnr11on ,n s;,unt> of t1uomcen;
:26 I.OHfl i:..... , MO)...', J.P,J .. A. Mr.1'if', concurtttu coppt-r in10:<1c..uon in ~ht:ep.)011,mll ()ftl:,•.~m.:rtrA'fl
lL'\d .,gg!u1lna1lon anu'bod,e, In lnrracl)1hr~1icorg•nl<nh, 7.rlu,·1,rift l\•1,•r1111uy .Hidical w,,<fnMII. 133, 312,-315
f(lr TrOJJfllmtdt:1111md Pttr<t#ml()git. ?5, 2J7-.22th t.R ,1\H, ,.,.. \\'1.latL ,., .., 1uo,t~.f. ~i•• 196.1 R1;"latiansh1~, ut ,,\int to
bo,inc unapla....;mo:,i~. ("omel/ \ ,ttrrlnnrla11, :i 1. ,J07-; 14
2; wso•. c. ,.. 198~. ,\n:lpl:wno•i>- lnfot·tlvu, Tropic>! 111,eu,~, or nomc~tlc
Animal>- Harlow, UK: l»n!llllan Scientific ond Tl,;:hnical. l'l ,1 H\;t;,·1, l,. uo, ,,nt, .\., P\RR()T, , ... 1.ESU')QUJ\IUl, .... t·l.\\:TUJU:llX. £. &
1n,o<,i-Krn. H ... t~J l , h,oculailtm au momon ~k l',m,lpl:t'.'olti<" du bocuf
28 LU.\\,'!,, LU.W.X,.ztL\,(;,Q.~.• oou.11.P.,\'U,F.. \"l!\',lt JI 7.U.,Hu. -r.,...
Bull~t,n ti,• !n Soc-:r:, tit• Pt,t1.•11!Qgi.,.. exoliqur, a;, 29-5-.:~.
~r", •.G .• JIAS, 1•.:-.. CJ\O, u.r "•,~,. v.1 .. 19,e; Stud} on .axu,ph~mo~i~ 1t1
s••«. ldentlficatlon nr 1he pn1hogcn ~rtd ol),er\'adon ol the clinlc:al so s1un,,A. ~.l".4' .,:\.:'-\I c•. e: . tg8:, C:hcmopmphyl,W~ or \uaplru•ttn m·l.i
~~ns. Chi11("Se Jat1mt1f c,/\'t't~rmuryScitmC1tamf 7edmc,ln?J', J. 5-8~ lnfoction tn ,http ,uth ..t. Jong,acting oxr1ctro1tyclinc• \ f'1,·ri1:nry·
P1m,sitolo;,o·. 1'" :!SS-:!58.
'°
~9 \l \.b, ,. f'Ut ..,,~c,. C,M., 1981. Chnr.,cceri~ation o(A11nplmm11 morgftml,..
,:.1 "4'm 1.nr.\lii..£. t,· 191:1 J-'ln,. Ki,•otmrhu.anuotM:•t rl~t"-\"ml:nmm'.."n \·on
id1,.-ctwn 111 ~plt'm:~1nmi1t·d Jonv,.~tk g03JS.. Al/lt'1N'(W Joumnl oJ
MA.ritina.l-po1nt"S ,·\lu,p/11.<mn n:ttr-tV1wJ,ri ltn 81m \'<HI ,d,afcn in
\ t11•rl,:ary· H~c<m:h, -•2. t42-115.
1

t>c.."1..nch,OM4frii,;.3.. ~tJ;n,,r w:d ,\fantlt~m.•r (l"1'llr1:ll,:I:~ 11'txhc1'lS(IJnj1,

30 M,-\t.,;.<,:"ir.,u. ff,,\,, ECt..OL.\O, \\ 1•. U:<.COL,. S.O, $ ffi\ ... ~ , L\\' 196l. 28,o;ll-~l:!.
A!tnplas.ma oi'is en Jdoho shce;,. Amrrir,111 Journal oft"M~rJ1111ry u st:11tWl\.)t.. \\ ., 191J l:5l"Ob~1chtungc:1 Uher di<" An!lpla,mo~,~ ur.<l
R,~._'(Jrth. J'?, 199-201 Plrup!J.<mo», der '-chafe und /.icgen In Dcu1eh,O.ialrlka. a/:schr/ft fi.r
31 :\l\1.1.Jt:J\. ti.. P., DWJ\'[01• .:,,,Ji\,~ \1"1 H01'MA, ,,.s. is;::. ·\ttap1J.i:mo,i_i,, m l11/el,:11nrrJkr"nl:l1dlt>ll, {>ltf/lJW1Tt AmnkJt~u·n 1111d ll)gltnt dn
go-:us; Rc.•pon c:,n clinie~ cases~ lndlti11 V11wrlmtT)'lour,u,1, SG. 693-694 Jfnusll<"r,.•. 13. ~49--35.?.

12 MOil\'\ R,N. &.-PATl:1.'\>-.'., K.C., 1968. rn:., li.ll!'n.C(I oi tt11115)la..~n1<~'"L, in S3 'H1\l.!SHO:\f, ,.., ·~i,i,An'1rtfa~moi.f~ in 1~1..f:resi:m lamb~ R~fu11lr
rurn1.11a.ru.s m ~Onhl'tn Jndf:i. lndft111 \lt't.:ri11aryfaumal. U. 6&.'i-689. \•f•1rn,1t11'11h. 24.11,-2lB

33 '\l>l)'.\G U. L.W., :t,<,urR"Rl., c.. ftU}lA.~Grn:WA... 1'.tL. \!OU\ \J;i, T.J-.. '\1COUJKt.
0 ~ - ~H0'11'0lL ~ .. Wll\ML" ~ o. Ul!AA~CU\\\'A. J;,11. N '\\('CtflHI r.C:.,. t980
Clonod D:';,\ prob.,. ld,•nury A1111p/mma o,,;., 111 goat< JUtd m O'al a high
I c.. ),;X('J\~·us. O.P. &- P ..U.\11 "· C.11.. t995. Dtltttinn of .~IU'l(Jl1Umt1 tWiJ
ptC\"ttfonce of in(ffliun. /at1wnl o{Qltliti# .\lirr0'1wlo~. 2;-. 2730-27.3$
l!:k<tlon ia goats bf malor lllrfaoc µrou:'.n 5 comimtll\" lnh1b:llon
c1~·mc,lfn.kt.'CI lmmuno,t.1rb1.•m l'\SS~}1, Jaunwf uj ,;unlcul Mfuob!ulogy ,S ~INII,\, 0 ~ f. P.\flL~. n,c... l!,6G, i\nopl0$.mO:.i"I '" t;(l.:Sl'\ a.nd IK'le'Cp, /,ufum
Jl, a:-,;-s;g, \ ',•1i:ri,ttt1J•/our,m/. ;3, 490-,;9,l.

JJ. :\.Ut'l. \\ o .. 1939,. O\'in~anupla....md~i.!O~ni.e tro.n~mh,,lon oC:illnpitismn S6 ·WUTl"L1l. e., ..\,'lHU" tf.LJ, '6-1'\'1!.IUU", M.J .. l95ff AnnplnsmtHWi,~ tn

01'is: .u1d Ept.71•1!,10.:«111 t/lll$ ro th,.. hlt.'3hut:k U)auutl/.1.::,.1 nllitfmn,: 1he Unlt<"d Stau:,,.. bp,t•rlm\"nHtl srndle,. '"hh <http ltnd ~on,.. •-lmt'rfcnn
Jnumnl a/\ 'rkrlutuy R,..._(,•fl«h, 17, ~87'491.
011dt'$lt/lOO'I Jo1,rru1/ (1[V,•1Prlnary·Stf<lt('t ,md ·ln111111! /1:d,urry, 13
9-16. 5? "-l'1.1rr1J1. l .f .. ·r.·:um \IJ, '-1.1 ,. e.v,no. LR•. 1955. An...plt'l..,mu~i<. in 1hc1:p
ln 1he Unhed .;;r.ste!. Jw,r1wl ts} 1h~ A1utrlCY1r1 Vtwri1lt11J .\lwlf(al
li ,.;:!.fr-1,. \\',O., tg68. A.111111lt1.111u, OlflS ln(et1inn. 8ullt•;rn dr I. ·o_ffi,,:
/U$C~·Jmio11. 11; ;?.J.i-24;,
/111,.•mnr/onnl dos Epl,,001/1/'$. ;t), ,1~9-,w,,
5,6 ~TILUJt,o .. c:ow.w.t ...lto\.l~mr.'i..P•• jOUSS(I.S,I,\\ ,<;01'{11\\1.f,.Jl.,J.
36 xnr;,.. \\.O.& nu 1orr P,t,.1t1;t:!, Bovine esn:iplasmo,b· A mt!thod of MCC:.UJRS. 1·.c.. 11189. l.>.rrmuumormuftno11JS1il1.·....; A natur~ \'t."<'tor nf
0;11.a.111.Jng pu~ ,irain> ot .1itttJ)famu, mm'g?)u1lr :ind .'\mt11Jnsmn uuunl,• 1\ttapln:m,tt1u,i.\ C.t!'-toqu.1rd on 'ihct!p ln Idaho. PmtM:dlngJ oftlu:· t·:"Rlu,,
by tr..tn"Sml"-~!on through ;lntcloJ)t"o, F.iglu&.1mh Ri1Jxu1 nf1ii, t>:rcrror of 'Vutlonttl JJt,mnparasin· o,:'l.lvi.~i· O;mftR.nt,•. Sr fm,U. :,Uuouri. 10.-1::
V'r:ermnry Srrrfc,•s and ~11fmt1I JurltlSII')', Untcm d'~mth Afr~a. 3-20.
April 1989.
:r Pt•M~<J. 1' 196&. PitoplMmtnb- A rt!\'it?\\ . Rt'/uaJ1 t~·u·r,ntJr'llh. 23. :59 ,--ra1 rs. w u .. i9ne-1"0. \ttcnnnry RC"<,('.;,rch h1,tilut<". 01utcrsc:,poon.
s2-:-. Uupu1;,li"1cdd3to
:,s VO'fGtrTU1, r.T .. ig:-9. Epi1.omiolc,m•1rnd comrol ofan3:pla~mo.cns in IIO ~"n"'-v""'·, , .. 1111 ""'""' 1\AO, 1989, ~nnplosnl<i~J, In sheep in AndhrJ
South Afrfe:a. {OW't:OI of1ll1• South A)i"lcan Vttt'ri'1fJ?' .4$..(IJ<illti(Jn. 50. 1>md,•,h. lndw11 Vc•r,•rlnnr,• /(Jttrm,I, rN. o72.
367-372.
61 ).S.\..l)X. 1.C•• 1982. Hnl11.•J/Jr. Tltt.•il,rrfaar.d
TIIOMA'I, S1f... ML(OS, D,1. &
J9 1•nt,.1rTtrt. r-. T,.1981. Tirk trnn;.ml.ssron oranap1a~mo~h In South Afrfcn. ..\wlpla!mn spp. inf«ting qhlc :mu:lOpt'. lff111HJ1ro""u ,,(~•·r ll t.irrb;
Pnxeerllng,. t,/ zltt rmrr,muonnl l''o11fer~11r1: t)Jt n:I.: Bforo,ti·o,:d Co,ur.oJ. 18381ln ~outht:n ,\frka.. Oml.t/'$1t'PDOrt /oumol of l ·,,1,•ri1mry: P.ff.t*tlrch.
Crn1Mm..S:CJJl''1, Rt•publft af!i<J11tll Africa. 17-29/amwr,• J9BJ 19, 163-166.
624 ,a,m, nuu:r: RkkeusiaJ and chlamydia! diseases

6~ Til,,\un,.,~, o.. 1913, .·\naplasmo<i< dee S<ho(e in Oeuisch·O-,t-Mr!kn t;(; 'rOV~ff, L.\ On.UttU. St.A. O\\j\"'IJ)I, S.~ ...\IIMf'J), "°·J.. :;iJ.Anfml.t
Ber/Ju Tifrarr:hc,,,, \Vo,:1u.,11schrift. 29. ,;93-39-4 duo 10 anap!iu1no>1<in Iraq, goo,~ I. Cllmc.al ,nd haem~io!ogical
ti:S lHt.(N'B'1C:(i, o .. \'"' \'OR.~TI~N1J(lba-f. (,J,.\,H.\".,. ruut. !I..,,.. 19;~. 81ood features und,•r licld condnlons. /1111/rin Ver,r/11my
J(),ur,ut/, 60, .>i'O..Si8.
pnra-;hes or fflcep In 1he ~eth,:rland.s f: tnnpln,flnCI mcstWu·nun ~p.n
1R1c~et3i1llC$. An•pf3.,m~1ace~eJ. \'et<rmw)' Quarwir, l 1~22. 6".'" 1...,uc. J,L .• 19s- 0\1n(!' anapl:umo.,,i,: /n ,,,.,,o tm~mi5-<fon :i\ h rt•latr, ,a
)tn(!,~o! i~1ction. -\mt•rfrnu Jo1,r1111l of\t~t.erllin.ry Rt•uard1, 18. t()()...
b4 \1S.Uk. ~.~ .. ,,rnHO"tfU, JU. 6o' l>l '''Ml, _D,t- 19,9l .o\n .-t,mp!(IJ11U1 ce11trnk
101
DXA prob.- th"1 dltrc«•n113t~s btt\\ """ 1il:1111I,;1m1> or/, .md ,\11ap/a,m111
tn(Jrgtnn/c Oi'\A. \'rll'rlnury '11~roblo!ol()'.Z8. '31~32.5. .a ,_we. J,L. • co" >1 , • 19,w. res, of ,he sheop kcd ,11riouT1r1g11;01'111111 JLI
a.s .a \.'t'Ctor ot .intmloJmn oi'ia t.e,10<1uard. Arrt.:ri,'"'tw /,x,mol of
h-5 \i:,,tR, t.!>- MC.:G,\.IIRf, J.C.. f'\UU:k, C.H.. nM'I>. \\',C \KMP, \ .. rW,\:,00. r. l'"cttzrb:ar:: Rt•si..v1rclt, .;'i. l 060-1062..
"x:,;-0\,~. D,P, 1"-, ltt9:t. TI1~A11n11tmnu: mnrg1n11lt: m$t15 gene, c,.cud&
• 19·kllodah(m 1>to1cln cun.,erwd In oil n,cognt1<'d A11aplasma ~p,>Clc•.
69 ,,,,.,,:, "·" nu,·•·
1.. 1qfi8. A111111lusm11 ads lnf•(Uon. 11. Parhogcnltlly of a
Nigerian go,11 ,.roln lot Dur<h $hecp and toal$ 11111/tti,: of Ffll::,J<ltl<
/11/ttt(on 1111d /mm,111/ty, 60, S13!h'il44. DiJ<AA'10f.tfr,co. 16 ;3-11(),

,
 Index
No·n!S StroplOl'OC'.Cll.<Spectes Ill: [;84 parawncbral Ul: l 7i2
111 a11yset111unceofsde111ijlc 11ames. a long Ta,vlorello i!t/uigc11/1alis 111: 2085 pec1oral 111: 19!7. 1921. 1922. 192'1
mlc: rcplacPs r/ze gm11,< 1m111,· where //w abortion In pi~ ll: 692. G~G. 777. 809. 822. periorbiml Ill: l 772
lm1er is fd,•micnl 10 the ge1111s name/11th<! 935,979, 1105 Pre,~7· '-'ocard Ill: 1917·25
line nb,w11. abonion in pigs. a~iau:d with spinal.;,;
.V1m1/~rs fn ho/If i11tllcnu: rlw major emf)' on .lrconob<lctl.'r/11111 ~pecil!..< Ill: 1952 ,ubruraneou, ill: 201!l. 210i
the s11iJ}ec1. bntaellools Ill: 1542. 15•13 1'>.!ldi11 species Ill: 2l 20

e
eryslpcla. Ill: l!ll 0 Amcfo 11i/01im578
leptooplmoJ;.111: 14•18 .l(c1111/111mtW!I/I 3 16 . 3 17
nocardiosis Ill: 2012 lc~rfribrl() e1lumolgig,ie11s Ill: 14:?S
S1nphyfoc«c11sspecies Ill: l 750 acet) lcholh1e Ill: I 892
aatd\\Olfll: 1138 SttcptoroccuHpecics Tl!: l i7G. I i78. ..iclwlepinsmn spt!Cies lll: 2076
,\badina \irus If: 12.52 1781. 1782, 178.i .~d,ol~plLmnn a.-wnt/111111 Ill: 2076
alx1masal abordon tn shN1p/g(lats 3>14, 3•17. 386. ~38, - - lnidlffw/1111: 2076
aspcrgiOosis Ill: 21 JG a5 I, 55.;. 568. 620. Ill: I 655 - mndlc11111111, 2076
c,roi,Jon 457,901 . 955. see also cro,lon~ abortion in ,hccp/goat$, associat~d 111th kl1ro11101Jm·rer (Alc11/ig1't1L'I') srlosox1dnns
abomashis Ill: 183:?. l8G9. 1870 Arcn,10/1"creri11m specie~ III: 1952, 1953 Ill: 2084
~bort!on 360,365.545, Ill, 1.;.i 1, M59. 15.15. brucellosis Ill: 1512. 1528. 1530, 153S acld·fast barUll 111: 1!177.1994
1560. 1S70. 1617, 1623. 1904, l!lOS, campylubau~rlosls 111: 145!1. 1479. 1480 ac,d•f.m l)mphnngit.is Ille 1983
19-16. 2()1 l lepto,plro,ls Ill: 1448. l44il ac1do~is, ruminal Ill: li25.1727, IT.IS
abonion .u;sociat,-d with llsteriosls Ill: 1904 ac1d•s1ablo p,comavirus(cs) 11: 1:lH)
babesiosis •ll 3 salmonetlosi!> lfl: 1594. 159-3, 1596 1ci11011y., j"ixirm Ill: Hti5
Chlom)Ylia spenC$ 3.'iO, 55-1 nru,(esl 11: 970. 9,1. ~!JO, 104!1, 10;;0. arnc.hc,rseslll: li5i
myrnsis Ill: 2121-2 1071. 1073, 120; acquired immunodclklcncy S)'ndrome 333.
nco,porosls 382. 383, 386 1·ersi11in species Ill: J(:\20 3•13, ll: 741. 744, 7•17, ,51, Ill: 2015,
Sarcocysris spocil'S 363. 369 3bomon In \\1ldlife Ill~ l 546. 15411 2111. 2137. 2142
toioop!nsmosis 34-1 . 3,17 abscess disea~c Ill: 1923 actinobacillo5ls 11: 712, 1285, Ill: 16-16-70
viru, lnfccrion II: 1029 abscesstesJ .ktinobncillusspecles II: 770.111: 1531,
abonlon inAfriran buffalo 11: 1044 abdOn\Cn HI: 1921. 1922, 192·1 15&1. liS5
:ibonlon in canines II 1205, 1207 cnroio,-a,cutnr system Ill: 1950 Ac1111ollllcilluJSpl'C'1e,, ,nfoetions ll· i i 2.
abonlon in carde307, 382-3, 413. •156, ·11'19. cer,.brum III: 1639. 1948 1285. lll: lil46•70
551, 555, 600 feet II: 1288, Ill: 1727, 1i29, liJ0-33. Act/110/Jncillt1s t1c1i11omyro1~mco111itnns
a l'l f)rt!on in cattle. as~QCl~ted whh I i55. 19·19 Ill: 1531
Arcanolmcteri"m spi:ci~ Ill: 1952 genital sy,tcm Ill: 1952
--r':f{lltlli If: 7(\9.1261.111; 155- !6'.2.
brucellosis UJ· 1510, l~ll. 1514. 1520. hearl Ill: l g.;,J
1652
1521 1535 Intestine Ill: 1505, 1920. 1948
- - n/llltli tnCcctions Ill: 1652 -4
ca.m pylobaaeno,1s Ill: 1-15!1, l4bl. "17!1 ktdll("~ Ill lb:>$. lb~,. IH2
- - 1£i:11ietesif infectiom, 111: 16'18·52. 1756
F,$r/r~richincoll UI, l5fl.l, 1570 !i,·cr lit: 1949
- -11101/ri see Bm!.;ltol<leri« mo/11.'I
lft11t1/l<Jf}lti/t1HOIIIIIII.< !II: I638, 1655 IU11g Ill: 1740, I742. 1950
l~mphnode;.tll:1768.1771, 1781, 1782.
--pleuropneumnnfol' 11: 911,914.
h:tllO~Jliro,ls 111: 1448. 14S2
Ill: 1.\9•1, 1631. 1661. 1670. 1778
l.)·ttw dis~asc Ill: 1442 1949
mytoplasmosls 111: 20iG. 20;;. 20711 mammary Rland Ill: 16-18, )947 --11leurop11eu111011in~ i11foction II: i77.
Ill: 1632. 1661•7, 1950
my,o,is Ul· '.?111. 2121 mutdplcsltes Ill: l ,54. li6ll.1785.1921.
--scminfs rn, 1531. 1655
nocarcliosis lit: 2012 19~6·53
,.n1moncllosis Ul! 151!2, 1586 myocardium Ill: 1653 -mis Ill: !!;ST. lGiO
virus(csJ II: 834. sn. 8i8. 953. 102\J. 1031. pituirniygland 524, Ill: 1·187. 1639. 1946. --$11is infection 111; 16,0
1049.1183.1189.1221.1253.1269 1947 ~c1i110/,ncu/1rm {£11/)nctcri111n} suls ll'.. 1958
Yeninia species JU: 1617. 1620 respiratory iract 111: 1950, 20 15. 2018. - s11/s lnft>ct ion 111: 1!'158-60
abonion in cquids 427, 429.•131, 5S9. 583-9, 20!9 Arrino,nodum m(l(/ttme 111: 210,
II: 829·52, 924 926. 928. !250 skcleutl ,~-stem Ill. 1951 -11cllv1fori Ill: 2l0i
abortion in cqufds. assoclnted with skin Ill: l9-19.210:' - - s1,ec1es Ill: 210~
M1:n11obacwri1m1 species Ill: ! 652, !653 ~pine 477 A1ww111ycesuo1•1s lll: 1942
Cor;..ne[JQfter/1m1 <pecies Ill: !91 i ,;seem Ill: 1505. 1920, 1922, 1948 --1,or•isinfcction Ill: !942·-1
c-n<lomeLrlt!s 111: 1555. 1557. 1564, 1570 abscessnilon Ill: I ;5.1, 1;r,s, 19•16-53, 201:;. - - ro11golemis see D11mu11ophi1r1S
leptospiro$b 111: 1H9 201 i-25. 2107. see also 1111der CT!ll{(Q/eJ/.$/S
mycosi~lll:2121 ab,ccss(e<l - - dermntOll()ITIIIS SCI' f)ermll/OJ)l1i/11,
nocardio~i, 111: 2012 dlsscmina1ed 111: 1754. liGll, 1785. 192!. congo/emis
~olmonollosb II!: lf,08, 1610 1946-53 --hror'flgint,lis II!: 19-12 1962
Slranglei 111: l ii!. mUiary 111: 1919, 1948 - - hyo11ogi11nllsinfection Ill· 1962
- - PJ-OgcJ1es see,4rca1101N1cturlum I0<!-1, 10~6. 1325. 1340, lll 1512, 1546, - - IPpillum to. 481,514
pyogimes 154;". 1547. 197•1. 19,5 - - mnc11ln111m 514
--suis see Ac111wbact1/um (E11/x1c1~rl11m/ ganw ranching 4i0 - - marmore11111 10,514
rnls role in foo1-nnu-mou1h disease Ii: 132;;, --po1ttf)<JS1tm lO, 51-1
Act/11omyret-llke bacteria Ill : 1962 1340 --spanam 10.514
actir.omyce1c(sl II: 849.1 11; 2009·41 Afncan elephant II: 1234 - - 1cs111di11arf11m 538
~ctlnOm)'Cosis ti: i12.951. Ill: 16.:;o, 1942-4 African face fl1es8I - - tilo/1011i 10. 5M
actlnomyco!lc Aincan forcy Ill: 2104· 7 - - raricgntw11 3, 8-l O. .:36. 480. 483. 507
granules Ill: 210. Afncan field rat lll: HIG 514. 53a. n: 1on. 111: 201 1. 2029
mycctoma Ill: 2107 African grey parr01 II; 7S7 Amies· medium Ill: 1481. 1631
acme abdominal syndrome. neonatal calves MTicnn horse sickne!» 93. 98. 227. 276, 4.31. amoebit infcclion5 316-17
111: 1831 fl:685,849,928, 1025, 123J -q3. l24,. ampulllds TU: 1637
acllle cntcrot}l'hlocolltls syndrome. hor~cs 1250 amyloidosis 11: L400, 1413
Ill: 1611 African horse <iclme!J.!. ,'irus t 53, 15-1 II: anaemia 435. 485, 498, 4$9. 500. ;;;G. 581.
:ieute g!andets Ill: 1502 1221. 1227 U:,ll
acute haemolytic crisis 442 African redwatcr see bovine babesio,ls cnnle 265. -113. •156, 480. 483 . .:86. ~89,
acuie imerStirlal pneumonia, hc>rscs II: 682 t\frican relapsing fc\'er~ 500. 577,562, GOO, II!: 1448. 1449. 186;
acme mast!tis. ~hccp Ill: 1829 • Afrlcan 5',ine fever 1. 27, 80. 225. 2ii. 2,8, .:quids 425, 1129. 1024 1772, $ee 11/so
acute renal faiiure, cquids 429. Ill: 1610 II: i'7i,983. 1088·1112.1383.1!1: 1605 equine i11foc1ious anaemia
~cute rhinitis. pigs Ill: 1492 African swine l'evenirus 80, 1~4. II: 975, sheep/goats 277, ,138, 441. 498. 499.
acute Trypa110$omn 1•im.t infeet!on 266, 2'iS 1089 Ill: 1450. 1831. l867, 1919, :?157, 2158
Adelaide ruven'iru~ II: 1184 African uypano:.omosl5 251-87 se, 11/,<o anAemia. haemol)'tlc 413, 414. Ill: 1452.
ad~nocarcinoma ti: i26. 755 ll')'J)anosomnsis 1921
adenoma(s) JI: 75:i African \\1ld c:n 535, II: 1131 Annnindeua 1iru~ 100
adcnomatosil., inte.,rinal, pigi; Ill: 1623 African wild dtli; 378, II: 1130 A11ap/nsma specfes SOI!. 536,537. 58~
.-tdt'rlfJl>iridne II: 668. i98. 817·2•1 , 8i7 :\fr!ka gesigsvllee 81 .4naplasma 1,,,,.Js 524
adeno,irus lntections ngalacrfa 111: 1448. 1557. 1570, li12.seenlso - - cr,udaum t 595
canle II: 819-21 , IJI: 1481!, 1489 mill,. produc1ion decline - - cemmle26. i:>94
bor~es ll: 823·<l ngar,s) see under /111/i1•idut1I 11ames - 11uirgi11ale 11. 15. Ii, 21. 26. 80. 90,508.
pigs I!: 822·3 aggression Tl: 1400 576. 59-1. 617
sheep ti· 820. 821-2. 1260 :\JDS see acquired immunodeficiency - - margi11nle and Eperytlm;::0011 sp~ci('S
adtmo,irus(es, 11: 668. 798. Bl 7•24. a;; syndrome 576
adhesinslll: 1561. l6i9 ,\ino drus 98. 99.11; 1029 - - mt•u1e1crum 617
adhe'Sions. p~ritoneal. pigs lll: 1631 Akabane \itus 98. 137 --oui11t1524
adiaspiromyco>is Ill: 2 11 2 Akat,anc virus disease 98, 11: 1029·3-l -o,,fr24. 442. 578,617
adjuvams 2-14 Afagoa• I type 3) Vlru< II: 119-1 - - phagocytophiln S24
aluminium hydroxide 244 A/bi.fa spoades 417. II: 1417 nnaplasmosi.~ 3, 81, 83, 275, .;17, 459, -ISi ,
Freund ·s complete 244 .o\lbl:in 1a11ga11ylci!11Si.fS24 Ill: 1436
mineral oU-based 24•1 - - 1vmi:o/or524 c:anlc see bo,~nt anaplasmosb
Aed.:s t,kdimorplws} spucles 148-9, II: 98i !!lra/lg4111es see Arhromobatter <neep/go.its 24. 578. 617-22
988, 1007. 1024. 1040 nlcelaphinc herpesvirus t It: 895, 896 anasarca 355
AedL-s Ul'dimorp/111.,) ltt:fO•prl It: I 19S. 123~. Mcelnpht1s b11.wln11/111s 561 AncnlnB7
111: 1798 -cokiiG18 - africana ~
--nllxlpfcwsll: 1008. 1191 A/c,salr:e$ ll: 1227, t3:l9, 1422.111: 1548. ,lll•J'll/$/011/{/ 1:t111i11111 280
- - flrgenti!Of)LII/Ctn/11$ It: l 043 1689 angiopa1hy 111: 15,0
- r;n//(11/us 144. ll: 988. 1041 algal diseases Ill: 2140-43 Angolan bom tick I0
- - cnmp1orlrp1cl111s S75 o.hmemaryiract mycosis Ill: 2120 Angolan brown car1,ck 21
--casplus ll: 1044 Allerton clisea;c see bo\'inc herpcsvirus 2 onisoq'losis 620
- - circumlutefJlus-139. 142-3, II: 988. 1043 infection .411011/relesspccics II: 10·11, 1111-l
-do/zfo/111: 1044 Allenon strain or bovid herpcs\'iru~ 2 A11opilelesl11111crof/1 II: 1184
- - dunraws 11: !043 11: 1268 --s1eplm11si II : 1234
--juppi 139, 144, U: 10•13 Allium species •11 7, 603 nnorexia If: an. Ill: l ii8. 2067
- - linuarope1111/s II: 10,11 nlopecia 356. 953, Ill: 2008 nmelopc II: 834
--mcimoshi 139. l-13-4. 147. 11: 1041 .~lo~... lagopus II: 1127 anthra.x227. ll: 105i. Ill: 1788·181 l, 1856.
- - oclirace//$ II: 1044 alpaca II: 83~ l860
- - ta1miorril.w1cl111s Ill: 1798 alpha co:dnlll: 183(). 1840.1841.18,7, 186-l. ill l'C)llld~2?8
- - 1111id<'mnn,$ 139. 1-13•4, II: 1043 1a,o lmestinal Ill: 1475. 1803
- - l'<'XOIIJ aro/Jit>11.<f1 II: I045 nlphaherpesvirus(es> u: 829. 860. 868. aa:- aswono,islll: li88.1802
- - l'(')/(11/$ ll: 1044 nlpha1irus(es}: 155, ll: 1Ol-1-2 l, I023 Amhraxin Ill: l 807
--zombaensis 139. 147 alvcoll1is JI: 84-1. 1279 Amidorras mal$11p;t1lis 516,553. II: 1045,
,\edlmorph11$ teP Aetfes ,-t,11an~lla cm('l'lfe11S 88 l2i0
aedlne mosquitoe, sec Aede.s (Aedimorp/111,;J amnurosi, 11: 1370. 1640. seenl,w blindn= antigenic variation 259
species ambnrr-madow Ill: 2026-35 aml-fnflammatorles Ill 1612
:lt?pyr~ros meu1111p1,s 353. Tl: 1270.111: 15!2. ;1/11/J~l'Oltl/llO Slll"cies I 0. 480. 483. 498,537. .411ri/ocaprn 11111erirn1m 11: 820. 1205. 1227,
15H 54:? Ill: 1689
Aem1Nic1erspecles Tl: 769 Am/Jlyommll nnwrict11111111514, S83 an1lm!crobial~ Ill: 1612
aerobic spore-bearing bacteria Ill, Ii9 I - - usrrio11 IO. 454, 480, •183. 5 14 apho,phorosls II: 1191
,wrophobia II: 1151 --cajen11e11s,;,6I-I. 538 Aphthoilirus 11: 1307, 1319, 1326
Mrfcan,1nimal uypanosomoses 251-87. see - - coi111erms 10, 45•1, 480, 51'I Apies River ,iru,; 155, ti: 1252
,;/so try-panosomosis --gem11111 10. 4S4, 480. 514 arho,irus(csJ 153·63. 11: 849. ,00~. 1014·25.
African blue tick lJ . •106 - lt/Jbra,mm 3. 4-8, 480. 483. 507. al 4. 1029-3•1, 1090, 1183. 122!, 1252
Afrie311 buffalo 4-18. -.61. 468-i0.11: 820. 876, 542. ITI: 1,31. 1947 ,.\r1:m1obacrer/11m P.l'"gl/lles II: ,25. 729.
 UI: 1452, 163.8. 1720. J;-29, 1738. 1743. ;i1rophic rhinid< II: 920, Ill. 1492, 1·19 1, bodUnry h3cmogloblnurf~ Ill: 186.1, 1865.
1;-1-4, 1919. 19,13, 19-l6, 1962,2152 1707•11.1;'"43 1866•7
·\(<flltOOOCNrlum pyoge111'S Infection a1rophy Btlt'i/111$ nmlimcis90, UI: I 788
!ff: 1946<>-I optic nen·e lf: 953 llaclllu,, Calmcnc-Gu~rin 241. T60
Arciicfo)( II· 1126, Ill: 15'18 te•rlclt 355, Ill: 1530, 1657 lkld/111,f ~peci~~ IU: 1791, I i92
,\relic rabies II: 112; thymu~ ll: 9S5, 97'1 81t<"illt1Hereus1ll: 173ll, 1791 1801, 180:;
AfC)'0/1//ora l,mgimll'ls Ill: 1·1811 turbinate Ill: 170, - - llchcniformi.< Ill: I623
Arc1111l'irufoe II: 1082 ·.nu u, ;92 - - mycoides JII; 1791
Arrnoedsvlnktc Ill: 1886 -\Iy(o/lL~8, - - µili/on111:, see Closrridium viii/om"'
ursen:c poisoning 525, !II: 15811. 1692. see ,lry/om.< 11igro111amla111s 90 - 1h11ri11gie11sis UI: 1791
tllso 1111dcr poisoning ,\11thmeromy/o U: 1098 - - 1h11ri11gfm.<is iscaelc11,is 14;
nnerftis see cquine,irnl anerftfs \ujc;szky's <list>n<c see p,~11d(ni1hie< bac~·snarch re~t II: 1400
,\rt~rlviridne !I: 9-23-H .\ujes1.J..1 ·s disease virus II: 690. 829,909, B.\CTEC method Ill: 2001
anhritis 1442. 178... 178.5, 19~6.1951, see 921. Ill: Hi30. 1778 bacwrnem!a Ill: 1912
11l~o <welling. joints au101y,1,, II: 935. 1or.o bacu.'rial anhriti> JI: 7,14
cnule Lil: 1513. 151... 1511, 1634. 1637, At•(lrirftt 108 bact.erlal icteru~ Ill: 1919, 1921. 1923
1655. 1912, 10-li, 2050, Will ~~imte1101'irt1$ ff: 819 bacterial fnfornons. canlc II: 877
ho= lit 1610. 16! 1. 1r,.,;2, 1768, 1,72. avian chfilm)'diosis 550 bacterial ,epth;uemia. horw, ll: 849
1912,2015.2019 avian tuberculin Ill: 1982 Bnc111rl11m e111erm·olitic11m Ill: 16!.
n,
lnunan., 743, IU: 1440 awns lll: 16-18. 1949. I 950 B11c1,•roitlesspecics infoetion Ill: I i20. 19~6
pig,m: 1631, 1652.1760, 1776. 11a1. /J(lr1eroitles nsacclwrolyticus 111. I;.i..
1785, 1909-12. 20;-1, 2075 - fmgilislll: 174~
sheeptgoat~ II: 736, i41, ;.13, Ill: 1655,
1921
"11dlffe Ill: 1548, 1549
8 - - me/n11il1r,gmlc11s see Jlre1lf)telln

-
metn11i11oga11lca
nodosu~ sue L>lrl1l!IO/N1m·r
nnhritis. aupal/tarsal !II: 2(1..Ji. 2051) B dbea$C II 970·i·I Boc1eroid,·s} 1mdns11.,
septic JII: 17:?.1, :?019 haanno~tler ,pon.~iel.tl• 111. 1869 - orolls Ill: 17·1~
nrthritis-encephnlius. goatS II: iii. 733. Bnbe,; node, fl: 1155 - - ruminia,la Ul: I 74•1
741...J Bttbes/11 ,pc<:i<'s 406·20. 426. 431. 618. If: - - sp/a11clmicus HI: l 7•M
tlrtllrodemm species Ill: 2096 ,51. I ll().1241. li50, Ill: 1436, l4•1fl - - rfu!Tniowomicro11 Ill: 1744
,lrtluvxlerma bOll//amitw lTI: 2096 Babl.'Sia b:g.>111i11a 11. l 5. 22. 24, 406-20. o.6. - - wnformis Ill: l ;.;~
--gypseum Ill: 209G 59-1. 602 - - 1•11/ga111s Ill: 1,129
- - mcun~w,m 111: :?096 - - hlgim1f11a and 11/Wf')'t/m,zo,m species badger Ill: 1g;;i, 1975
- - obwsr UI · 2096 3i'G &1lai!nopt~rnac111onmrnra JIJ· 1546.1.549
- - Ottle Ill: 2()96 - - blg,w1i11a Infection see b<lvirt<' Bnlt//1111/ll/tl 316
- - persicolor Ill: 2096 b3bl'Slo<,s - - mnndril/{lris3 ! 6
anhrog.iypo<i< - - bo11/s J;;. 406-20. 576,602 hnfanqpOSthitb
cattle II: 95..1. 1029. 1033 - - /Jo1•1"$ and Epe,;•thr,,:(11111 specie, 5T6 cattle UI: 20i6, 2078, .wen/so infectious
pigs II: 688. 981 - - 11,n•isinfoctlo,1 $ec bovine babcsio~is bovine rhinouacheitls
~hecp1go3ts ff: 912. 990, 1029, 1049, 1050. - - cnimlli 4, 15. 22. 24, 26. 425-32 ,hecp goat~ Ill: 165i, 178-1. 1934. 2153,
Ill: 2046 - - ca/mlli infeclion sec equine 2154 ..~ce also ulceratl\'c
anhropod-bomc ,irus(csJ seci a rbo"irus(csJ piropla.mo~l, b,danopos1hitis a.nd \'Uh'l>,'.lginids:
anhropods 15;. 52, ll: J095·8, lll:2029 - rn11/sJ/ 280. 426 ulccrati\'t, d~rmato,T,
aniodaccyla If: 688 -cm,.,a,138 balantldfos!~ 376· 7
Arumowot vi= II: 10:i, - - dtl'ergun.< a.;.5 f)(l/1111ridl11m cofi376·,
An·imm/,fs all}~i11ic11$ Ill· l 4•16 - - Cf/Iii 425-32. 11:
1230 Bans method Ill: 19!\6
- - 11iloricu.< lfl: 1,146 - eqtd infocoon see e,1ui ne !laozl ,mis l:17. 155. IJ: !)87, l0Si
(tS\!i\11.S 54 l, Ill: 201\1 p1ropla,mosb Bso11lc cattle 266
A~coll test m, 180, - - /olinw 438 barley 111: 1650, 1950
ascomycetc Ill: 2095 - - mrrroti 126 8annal1 ,,ms I00
A~com,cotinn Ill: 2095 - - mmnsi lnfec1ion ,13s..12 bnrren·i:round carit,ou 353
llsfll!'llirldaell: 1087-l 11\'I - - oc,•1d11111s Ji. 406 Ba.rur ,1rus JOO
Asian blue lick 11. 15,406 - - 0/'{l/ll 90 ba$al cell cp1th~liorna If: 7.57
Asiatic buffalo see water buffalo - - Ol'i> infectio:i .i38-~2 Bruidio/JO/r/S'ij)eCi<'S Ill: 2120
AS!ntic redwaier see bo,ine bal>csiosls - - f)l'lT(JIICftOi 435 Bns1'dlob<JT11s ra11omm lll: 2113
Asiatiese blouuosluis 11. J 5, 406 - fJ!•rro11ci1oilnfection 11. 24, 26. 435-6. ba,ldlomycetc Ill: 209.5
nspergillosi< Ill 211~· 16 5;i; ha.~illi,rc hcmo~lohlnurlt, s~e bnctJlruy
,\sp!''&f/111., ~pecles Ill; l 452, 1~23, 210 I. - - sngenr/ -.3& hacmoglohinurln
2121 - - rra111m,11111i 4, 11. 2.1, 26. ,135. :;7a basophilic ~tippling 601. 620
Asperg(l/11sclmr1111s514, II: l4li, Ill: 1639 - - rrm11mmmi and E1>l'ry1/,rozorm <pedes bastard ~1rar.glc~ Ill: 1768, l ii l. 177?
--/11miga111.< lll: 2114. 2121 5:06 b3t•earcd fox II: 11:10, 1139
a.~plratlon pn~umonla lfl: i;oo - - traucmmmi inr~ctinn 11. 24, 26, ·135-6, hat~ !I: 681. 6112. 1124. 11 26, 1127. l 128,
Mtcroid bodies IU: 2 I 16 576 I 1,15, 1146, Hll,. see(l/sO /11d/11dua/
a.,trocytos1~ II 1400. 1413 babe,,io,b Ill: 1436 11n111es
111,1ro,'iru; ·~ 11: ,03 can le sel' bo,,;ne bnbi~osis be~ded app~arance or ,;epm IIt: 205!
As1yfus atromamlnrus Ill: 1588 dogs 280, JI; 1158 Bearricc Hilhirus ltlO
ataxia II: 13i0. 1413 equid> sui, equine piroplasmo~i> llemwer/11 l>IIS31m111 31 '
cattle UI: 16~7 pigs 11. 24. :!G. 435·6, SiG Bebaru ,irus !I: l 023
equids U: 1009. 1019, Ill: 2019 sheep/goais 438·4~ beei. papuleresiomatitis !I: 1282. 1289·90
pig, IU: 1631 bab~siosis. cerebral 413. lt4. 4 l T. 477. S2,. bello"ing. rnbid c,mle 11: l 153
sheep 11: 998 603, II: 1158 Belmont ,iriLs 99
Atila.r palu,ll11os11.< fl: 11311 baboon Ill: 19i5 B~mb1,x species 68
benign a.-,aplasmosis 3. 26 blood-stained urino Ill: 1958, 1959 bovine anaplasmosis 11 15. 22. 26. -159.
bi>nign b3llcslo,ls 3 blood.<udtng me~ II: 1205 594-610. Ill. 1-152
benign bovine babesiosis l 7 blood)' scours, pli:s Ill: 1810· 4·1 bovine babl':<i<><l< I I. 15. 22. 24. 2,5. 357,
benign bo,1ne thcilcriosls21. 22, 2~ bloubosluis(e) ll. 15 406-20, 459. 481. 11: 951. 1158, Ill· 1-152.
henign enzootic pare~is II: 130;.9 bloutong see b!uetonguc 1639
benign foot rot Ill: 1733, 1735 blo\\1lie~ Ill: t;98, 188i bovine besnoltio~ls 80. 351 ·8 , U· 12i I,
btnzlm,dnzole amhclmimh1t-s II: 1254 blowfly-srrike Ill: 2026 Ill: 2029
llerenU 282 blue duiker rt: 1072 bO\lne bonkers II: MI i
Bernmah ,;rus 11: 118-1 hlue car. pigs I!: 935 bodnebruceUosi:; Ill: 1510·23, JM~
Berr.:roa inca,ra II: 928. 1025 blue eye paramyxovini, II: 690 as zoonosis Ill: l.; I0
besmcclike misgchoone see bovine blue tick(sl 11. 15. 106 bm<i.nc cocC'idiosis see 1111der roccldiosis
brucellosis blue wildebeest 353. 11, 8i'6. 895 bovine coronnvirus II: 791. i95. 803. 80-1
Besnoirin specil!S 351 ·3 blueiongue93. 98,227. 5'!5. ll: 668,960. bovine coronavirus lnfection 328.
&snoitia benn~ni 351 1197. 1201· 15.1225. 1229.1285.1288. II: 795-801.111: 1479
- - bl,moiri 80. 90, 35 I 13Sl, Ill: li36 ho,ine(ie1111at<>phl!Mh 4, 10. 35T. 5-10
--.-opme353 bl11e1ongue ,iru, 153, 154.11: 905, 1!21. bovine chrlichlosis 3. 21. 22, 536, 53841 ,
bosnoitiosii80. 351 -8.11! 12N. Ill: 2029 1247 5-12
beta 1011.in Ill: 183i. 1840. 1857, 18i0 Boergoat kld$ lll: Ii-I I, 1742 hovine en1eric ca!icivlnt<ic,} !I, 703·6, 800
be1aherpeS\irus(esJ II: 919 boksiektc 32& bo\ine ephemeral fever 9ll, IOI. II: I 183·92,
cy1omcgalo"irus-llke II: 860 Bolo J.!beasc Ill: Hl3R-40 1225, 1229
bighomshecp553, II: 1205. 1227, Ill: 1995 bone meat nr: 1886 hmine ephemeral fever ,1ru5 99. IOI. 155.
bilateral corneal opacity 280, 11: 900 bom tickisl 4 · 10,50i.11: 1072 11: 1124
biliary fever 280, $ee also equine bombosluis{e) 4 · 10,507.11: 1072 bo,ine eptz.ootic :ihorilon 551. Ill: !436
piroplasmos!s bom-legged tick(;) 15·17 b0\1ne t!pl1.001ic fowr II: I 183
bilirub!naemia ~36 bontpootboslut. 17 bovine forcy Ill: 2107
bilirub!nuria ~25 Boopl11/11s,1111111/ams Ill· t ~5 l,o,1ne foo1 rot $('(' 1111der foot rot
bil!r goats Ill: J SZ8 --auscralls Ill: I~35 bo\1nc gc-nltal cnmpylobacterios,s
b,ologic:al con1rol of ticks 31 - - ,lecQ/omtu~ 3. 11, ·106. 435, 596. Ill: 1459-r,S
biological weapons JI: IO l!l Ill: 1435 bovine genital tibro<is lll: 1459·65
b!oun deficiency Ill: 1743 - - microplu, 3. 1 I. I5, 406. -126. +28. 596. bo,·!ne 1,:enhal mycoplasmosis Ill: 2076·80
birds as ,irus hosts II: 1oo.i Ill: 1435 bo,ine ltacmobanonell,)sis 581·2
bison 360. II: 1423. Ill: 1546. 1547. 16.14. bordcrdi5e~se 11, 970-74 bovine hep:uk necroha~illo,i, m, 1:-36-41
1689. 19i3. 1975. 1995 horderdiseaSc ,,rus 11,946. 9GO. 981 bo,ine herpes m:1mmaJJi1;s see.b,w!n~
a;son viso11360. II: 1-123. UI: 1546, 15,1,. Bcrd~cella bro11eiliSi!JJCic<1 Ji: 769. 92J, hcrpl?Svints 2 lnfoction
1634, 1689. 1973. 1995 Ill: 1704, 2011-1 bo\'lnc hcrp<is\irus I lnf,:ction se,,
biting nles II: I:?34. Ill: 1798, 2029 - - bro11ellMepc/1Y11nrec1lon 111, 1492-5. infccctm,s bovine rhlnom1ch~fll~
biting midges 93-126, !OJ I. St><' (l/s11 110; ho,'ine herpe,,·viru, 2 inf~ction JI: 887·93,
C11/ico;d11, spcdcs --pnrapcriussis Ill: 169B 1268. 1274, 1292, 1297, 1303
Blris nrte1011s arlecam Ill: 1860 Soma disea,e II; 8·19. 1010. 1368·71 bovin.: herpcs,iru, ,1 rJ: 90-1, Ill: 21.;9
--gabonica l!l: 11160 possib!e·,.oono,is II: 1371 ho,ine herpt~\iru~(~ I and 5 11: 675,829.
81\'en,Arm virus 99, 102 Barna disease ,,;rus II: 1368 868, 875-83. 951, Ill: 1681 2149
black bear U: 1227 Bomasche Kr.mkhett II: 1368 bO\'ine h~rpesviruses l !: i98
black dbease ill: I 863, 1865·6 81Jn1111'1rid<1e II: 131>'7· 7-2 bo\'lne immunodeficiency ,1ni; II: i 41
black fruit bat II: 688 Borre/In spedes 80, Ill: li21 bo,inc leukaemia 90, 275. 11: 708--13. 717
black rhinoceros II: 1227, 1234. ill: 1446, 811rreliha}~..mu1: 1440 hovim, leukacmi;i \'iru, II: 708 718. i,12
2116 --and«mmi Ill: l•MO in sheep II: 709
black spot Ill: l94i - - bissetrii Ill; 14,10 bo,ine leukosis. sporadic II: 708, 713
black \\ildebct>St 353,516, 11: 876. 895. - - lmrgdorferi 90. II: 12SO. Ill: 1-1-10 bo~ine l}111phosarcoma 90. 2-s. II: 708· 13.
111: l-H6 --gari11ii 111: 14•10 ;-17
black-backed Jat·kal 363, Ii· 1131 --juµo11im Ill: 1-140 b1>,ine malignant catarrhal fe,·er sec 111uler
blackbuck JI: 1339 --lusiumiae Ill: 1,1-1() malignam cnt-nrrhal fever
black/lies II: 1019. ll95. Jl l: li5§ - - s11/Jlt1 Ill: 14:1; bo,-!nc. mucosa.I disease see bo\'ine ,1ral
blndtleg LT!: l856·1il --sulllll infeetion Ill: I ~37-8. 17-13 diarrhoea: mucosa! disease
blackquarrer II: I 191. Ill: 1856·61. 186.3, - - l(lllltkii Ill; l 1,10 bo,ine ocular squamous cell cnrcinomu
18i5 - - tlre/ler/ 11. 15. 22. Ill: 1-135. l-137. 2157 11:761
Blru,ford's fox of Asia II: 1130 - - 1/u•iler/infection 3. 4. 11 15, 22. bo,ine pap!lloma,fru<{t.'sl II: ;55
Blaser medium Ill: 1481 Ill: I.J-35·8. 1~37 bovin<' papulars1omathls ii: 1282. 1289·90
8/n.ttomyr:rl den11atltldis Ill: 21 !G - - lrlrda~ Ill: 111-JO aszoonosls IL 1289
blastomycosis m: 21 I6-! 7 - - mlaisim,a l!I; 1440 bo,ine papulor ~tomo1itl~ virus II: 1291
blecksit~,e 575 Borrelia-llke~pirochac1~-s Ill: 1-136 bo,·inl! parainnu~nza ,·Int~ 3 II: 877
blepharospasm Ill. 1438 burrelios!s I I. 15, 22 bo,ine parvovin1~ 11: (1()7
blesbokSIG.11: 10·16. 1205.111: 1479 L)1ne 587. IJI; 14-10·•13 bovine pan•iwlru~ Infection II: 815· !G
blight Ill: 1487 boslulskop 111: 19•18 ho,ine petechia! re"er 225, 22i, 543.,1, 546
bit nd mes Bi bOll")'llmycosi, JJ!: 150·1, I650. Ii;»,. 1756·; ho,ine pleuropncumonia s~e corun~ou,
blindevliee 8, bo1ulinum to~n Ill: 1880. 1887, 1889 bovine pleuropn~umonia
blindness4i3. 4i5. II: 8.~•I. 1019, l!I: 1449. bontlism363. 11:685. MOl, 111: 1885-99 bo,inc pneumonic pastemello,;ls II: 673.
1450. J.l88, 15•19, 1848 canhH!: 1158, 1191, 1'117.111: 1639 Ill: 1677-83
CillllC 111: 1637, 1640 horses II: 849. JOJO bovine respirmory disease complex 11 673.
bllnkbruinbosluis U·G llcJl'icola bozd.1 5i 5 6i9. 797,798
bloednier Ill: 1846-52 bovine aclinomycosis Ill: 1942-4 bovme respiratory syncytial nrm II: G75.
bloedpens UJ; 1836-9 bo,in<' adcno\'irus infection II: 819-21 677, 7911,BQ0.87,
blood-stained mlll<TII: 1448 LIi: 1~88. H89 bo,ine rC$j>lrlllOt\' >1'nC)1!nJ ,;ru, infoction
 l\?)ilX \'

II: 6 77-9. 952. Ill: 1681

bo,ine rhlnO\irus infec1ions II: 1322
bovine salmoneUosis 321\. It; 960.
Ill: 1582-90
as <0(111/lSIS !II: 1$35
Bruc~ltn m~llremis Rev. 1 ,-acchu,, ,u-ain
!II· 1531, 153i. 1538
Bru~·el/11 111!()tomt1e 111: l:;.t6
e
Cachc\'allci ,,irus ll: 102'J
bovir.e spongiform encephalopathy - - od, Ill· IS28, 1537, 15-16
caddisnies 584
JI: l388. 1408 -18. !422-3 --01•i,dnfec1ion Ill: 1528·32
-pmnlpedinelll: 1549, 1550
t'alcitit'mion
bovine irypanosomosls 2 6 7-75, Ill: 2029
--s11is90, l!.810, Ill: 1;;10.1:;1J.1;;30, aonn UJ: 1999
bo,•1nc ruberculm 111: lllll2 cndocardium Ill: I 999
bovir.e 1uberculo,is ru: 19i3·87 1535. 15-12.1523, 1546
- - rnisfntec1ion TII: l542-4. 1548 cnlt'diarrboe,1 II: 71)5. 798
b1wir.1• ulcerative mammillit!~ see bovine calf diphtheria Ill: 1·1 41.2. 1950
herpes,iru, 2 inicc;1ion Bruce/1(1 vaccines .tee 1111tlor st roin
brucellosi~ 227. I!: 9i3, 105,, 125~. !II: 1-1-12, calflymphoma II: i08, 713
bo,ine ulcerative stoma1itl~ l I: ! 282,
1452 1535 calf pneumonia !I: 79,.111: 1635
1289-90
cnulc Ill. 1510 -23. 1542 calkl-like\'ims!esl !I: 703
bovine \'iral d1.1rrhoea 328. 459. I!: 946-62,
hum:ms Ill: 1512-13. 1535, 1542. scc nUo Cnlici11ndne ll: iO\l. 703
lll: 1623
l>m1ne viral dJorrhoea and mu,osal diseasu ~lalta fover i:alichirus{es) see C"licwil'idae
1!: 946-62. 119i, 1225. !229, 12119. 13~ l, pig5 IIL 15"2-~. IS.18 California disease 1!!: 21 l'i' • 18
1489, seen/so bovine ,iral diarrhoea: sh~ep Ill: 1528·32 California enccphalicis 90
mucosal disease wildlife Ill: 1S23. l5·12. 1546·51 Cnllosci11r11s98
bovine viral diarrhoea ,irus !I: 675, 798. tlOO. brucelloiiHkin 1e,1 Ill: 1518, !SHI, camel 382, I!: !O<IG, Iii: 1535. 1549, 197S
8Ti. 946. 970,981. !I!: 15i0. 1583, 1588 1623 cam~I ecthyma I!: l300
bo,111e ,1r.a1 d1arrhoea/hnemorrhag1c brucello~omc: TI!: 1:;13 c11mcl orf II: 1282
:1)11d.rome 11: 953 bruin<mrhoslub,el 18-22 camel 1ick II: 1300
bovine ,iral lcukostS90. 275, It: 708· L3. 717 bruii.ins, fee1 Ill: li25. 1743 camelpox II: 1!!00·1301
bovine viral mucosa! disease see bo,•ine Bryanstnn ,1rus 155. ll: 1247 C"meh,stlramctfnr/11s382, fl· 1046.
,iral diarrhoea: mucosa! disease BS2 ,-acdne Ill: 1522, 153 l, 1544 ill: 1535. 15-19, 1975
Bracltinrin tlec11mb~m Ill: I1-1,1 B11bn/11s 1>1100/is 360. 382, 410. 11: 815. 876, Camp fire J>rogmmme 233
brachygnathia ll: 688. 953, Ill: I i09 1270. Ill: 1620, 1638. !689, 1829. 197,1. Campr·BAP medium ill: 1481
Hmchysµirtl ilyodyumterlfli' II: 67-1, Ill: 1428. 2021!. 2045, 20!)6 tamP,l'iol>actur,pecics H: 647, !II: 1,69.
1571 bubollic plague I!: l 123. Ill: 1617 lli23
bracken iem Ile ;55 Bucyrus strain I!: 92,1 Cm11µyioboc1vr coli 11!: l 129
bracken rem pois<>ning 54-1. see nlso 1111tle r buJfulo disease see Corrid<>r di~cnse --Jetussubsp.fews Ill: !459. 1461. 1623
poisoning bu.Ci'alo fli· 'i'i --/~IIIHub,p. 111merea/ls 306. Ill: 1459.
tmldsot Ill: 1869 buffnh, see African buffnlo orwuter bull'nlo 1461
brnndassic> 93-1 26. see nlso C11/ic()ide-, bul'faln-a•~ociatecl lnfoctlon see Corridor --J1,j1111i347. Ill: 1479, 15,1
•pet'le, di~r:,a~c --/ej1mlln(ectlon Ill: 14~·82
brnndpoot Ill· I:-29 buflalo,()cri~-ed 71,elll'ria pnr,1,1 lnicct!on --sp111or11m subs11. l111bu/t1s !II: 1+59
Brassica species 417, 603. 621 see Corridor di~ease campylobacterlo,is Ill: 1623
brax,· Ill: 1869-i3 bu!Talopo~ II: 1302-~ nttie Ill: 1452. 1459·65
break in wool rollicle ll: 1208 as l()OllO~is It 1302 pig> !I: 91).1
breASt boil. horses m: 1756 buiTalopox,iru~JI: 1302 aszoonosislll: 1479
brc.ithing difficult~ see wutcr respiratory buffel gm~ II: 1185 Campyloba,·1er-like organisms lll: 14.69
dis1ress bulfcl~iek1e see Corridor disc;ise Co11didaspccles 111: 2122, 2L23
Bredavlms JI: 800. Ill: 1588 Bug&')' Cre~k viru• I!: 1017 C,,rulidn nlbimns Ill: 21 Ii
bronchiolitis U; 769. 82.2. 844. 845 bulbar paml}'$i$ Ill: 1885 candidiasisllL21Ii
bronchiolo-alveolar adcnocarci.noma It 726 bulbo-ur~1hr!ris Ill: 1657 canine calicMms II: ,OO
hronchhls II: i69. 1320 bull no:,c Ill: li43 canlnl' corona,iru, I!: 780
bronchopncumonia Ill: I ;54 bullac formation ll: 1190 ~-aninedistempcr227.~{44. II: 696.1158
cattle 11: 955. lll; 15!5, 1587, 1620, 2055 Bw1os10111w11 phle/,oromum JI!: l 588 canine d1siempur vinJs !I: 634, 600, 687
~ulds 11· ,69. 842.a&i. 11!: 1503, 1~2. Bunynmwcrn vin» IS4, II: 1038 canine hcpa1l1ls260. II: 1158
2015,201i.2018,20!9.2121 8u11y1w/rldae 98. 155, ll: 1027 -85 canine orol paplllomnviru~ IJ: 757
pig,, 11:913.111: 1492, l-l9•1. 1604. 1661, Bun~ip Creek ,·irus 100. ll: 1253 canin~ pan·uvim$ II: 807
17i6, 17ill. 1950 buparvttquonc 492 Ct111i1odu,111sl l; 1134
;he~p !I: 821 ll-111,11ng11s species Ill: 2029 - - /nrmm 382. ll: 1128
bro"'" ear tickt$ 18-22. 448 Bur/d1old~ria mnlfoi 111: 1500 - mesomelns 363. !I: 1131
bro,m hrena !I: l 13, --/NL•1uloma/M Ill: 1500. 1503. 1505 --simemis It: I l30
bro\\11 tick 24-6. Ill: 1731 bursaucc Ill: 2 l 13· l ~ C.1ptfo.x I!: 1139
Hruc;,/la specie:, 80,347. 5(;8. Ill: LS.19 bun;ltis Ill: 1513. 1548, 1608.1921.sce also Qiprn ibex Ill: 1546. 1549
and chlamydia 560 fistulous withers; poU evil - - p_1r,•1wlca S53
BmC'i'lla species inieclions Ill: 1S07 ·S2 bush fl\';; tilprine 11rthrit1s·cncephallt1s I!: Tl i, ;-33.
Br11c,,lln al,orru.<130. JJJ: 1.ir,.1, 1510, 1530. bushb~cl. 516. 5-H 741 ·4
1535.1543. 1523. 1546. l:xl5. 1588. bu,hplg ~35, U: 10116, !Ii: 1446 caprine.i.nhritis·cncephalitis ,ints !I: 733
1623. 1623 button ulcers It: 9i9. 1092. 1109. Ill: 1604 caprine besnoiliosi,; 351. see also
Brt1t:<!17a nlx>m1s lnfeclio11 Bunonwlllpw virus 1(>0 b~nuitiusis
horseslll: 1512. 1516 Bu11.lcrml'<lium !11: 1-181 roprinc hi!rpcsvirus(esl !I: 829. LIi: '.!!53
sheep1goa1~ Ill: 1512 8\\'amba ,irus 154 cllpri.ne lmmunodeficicnC)' \irus !I: 710
Brt1c;,l/(I CtllliJ ill: !546 8_\'Q111yn 81 capripox virus 80. I!: !268. 127i
Bructllt1 ceinrne Ill: 15~9. 1550 capybara fl: 13:\9
BrllNIIO meli/P.mit90, Ill: 1510. l!i30. 1531. c:nr:i inchada lll. 1743-4 . 195!
1543. 1546 caracal !I: 1131
Bru~'i/!lt1111el/1e1ms infec.tion Ill: 1535·4 1. carcinoma
15-18·9 ndeno- I!: 726. 755
 ~deno· bronchiqlo-alvcolar II: ,26 cerebral trypano,omosis :?69 chronic pneumonia 111, 1661
comagiou, bronchlolo-alveolar II: 71, cervical lymphadenill< Ill: 17lll ·3 chronic rabies infocuon. non-fatal ll: I UO
humnn bronchiolo-alveQ!ar I!· i26 cer.-ical stenosis 100 chronit was1111g diseasl' 11: 1388. 1422
human genital 11: i5, cervical vencbrnl malformation. horses Chry·$omy1111/bir.eps 111: 1798
o,1ne pulmonary II: 71, ·30. i33. 7.l8. 11:849 --iler::in11t18
1397, H02 cer,•lcitis - 111nrgi1wlis Ill: 1798
~uamou< rell 11: i55·6l caule 307. ;ma. Ill: 163R Cltrysops87. 597
.,qunmou, cell. ocular n: 755, 761 ,heeplgoats Ill: l6f>7 Chrywp$lnnc Iii
squamou, cell. vulvur II: 755, 757 tt,n·ic,i-,'llg!natl< 111: 21-19-5 I Chuzun vinis 98, II: 1()29. 1252
Card 1eq Ill: 1544 Cen•1u ca11ade11.<ls II: 1227 Cl~ agar JIJ 1611!
cardia<: foilurl.' JI: 711 --l'lrtphu.; I!: 79(1. Ill: 1546, llH8 1973. drcling Ill: 16..l,. 1906
cardiac rom1 <>f hors~ ,icl..n~ss JI: 12.15. 19," Clrro1,1ridne II: 1373-Sfi
1236··10 --11111µ011 Ill· 1975 cltrini11 Ill: 1-152
Cardrrll'l,.,,.11: 1so7. 1310. 13m - - 1111(colr,r II: 796 Cltrolxtcterlll: 1560
Carey !~land ,1rus 11: 995 Cr•,tnmr specie, II: 1057. 1158 Ciw1ticli$rlue11a 11: 1137
caribou II: 795. Ill: 1542. 1546. 1548 cetacean dlstemp<,r II: 634 ci\1111: 1137
carrier <tate Cha gas· d~ca.w 2:i I CL.\111: 1917·25
East Coast f~ver 451 chalc1d wa,;ps 31 cls,sic:al swine fever see hog cholera
salmoncllo,is Ill: 1602 Chnlqu,m's agar a nd bro1h Ill: 20,9 cla,;sict,1,111nc fever virus II: 690,806.946.
Tliai1drlnam111lau1 theileriosi,-189 chamois Ill: 1546. 15-17.15-19 950.970
carriers chanC'res. cattle 266 Clm.>iceps fHJSf)(J/i 52,:
Actrnobm:1/Jus eq1111ll 111: 11,~z charl>on ill: I 71!l1·1H l L. seeal.<o anl.llrnx closmdoal mnslltls 111: urn,
:\cri11nbl1c//lt1s plr11rop11ounumilll' chcesv gland Ill. l!ll 7-25 c-Jostridittl 11,yonecmsi, Ill: 1831
Ill: ltl61 cheetah II: 1-122. 1423. Ill: 1975 dostridial ,votm<l infection 111: 1830
Cl1writli11111 pcrfrlngc11st').1>c O Ill: 184' chcmo.teriliz.ation •131 clos1rld!o,i> 58,
contagious bo\'ine plcuropneumonin chicl:en liner Ill: 1887 Closrridlmn species 11: 1261
Ill: 20:55 Chikungunya viru, 137.11: 1015. 1023 Clos1rldi11m lm111/i1111111 JIJ: 1885
Ea.~t coiist re,·er 45;; chinclterinchee 328 - - ,·arnis Infection, 111: 1875
equine pirop!asmO$iS 427 Chmcst' \'accinc Ill· 1531 -r/1mn~>l'i90. ll: 1191.111: 1856
foot-and-mouth dk~asc II: 1336-7 01/n111_1rli11 ~pccii,_, see nl$O 11111ter --,1t11111'tll!i lnfeclion Ill: 1856·61
H11dn1111op/ri/11$ µnm,<11ls Ill: 163.2 Ch/amydap/1lln - tlijJ/rilu Ill: 1612
ff11e111op!iil11ssomm1., Ill: 1635 chlamydia --dlffiei/~ infection Ill: l 875•6
Pt1$1,•11rtli(I m11hodt1,1 Ill: 1690 nnd Bmr..>J/11 ,fll!CiCS 560 --./i1//nxi11lcc1ion Ill: 1876
Sa/mo11e//(l ltl: 1563. 1602 I609 and coxiella StiO - - lme11wly1ic11m Ill· 1663. IS66
Torlorel/r/ 111: 2085 Cir/nm} tllo m11rid11n1111 551 - 1101,1 Ill: 1863·9
Caner procedure Ill: l68l! - - SIii$ 5~ I, ;,:,b - - norJ• mfcc110n Ill: ll!tiJ·li
ca.tarn ,irus 155, 11: I!H - - 1mc/1Qmt11is i>5 l --oedm1nrie11.< see Clostridi:rm no,~·,
c-aseou, l~1nphudeni1i6 Ill: 1505. 1917-25 chlamydia] abonion 550, SS·l, S55 --p.•rfri118<'11>'90, 11:911.l. Ill, 1571. 1588.
Castor bean dck ll: 996 chlarnrdial conjunctivitis 556 ll\9i. 1612. 1819. 1829. 1836. 1840
ca1 flea 58·1 chlamydiosis 52•1. 550·6l. II: 973. m;-, --perfri11se11sinfec!lon, LIi: 1819-53
ca, pox II: 1296 1254, Ill: 1452. 156,1 - - per[rlngens type..\ Infection
cn1aracts IL: 9.53 01/(lm_vdophllt1 spucics 83. 550. 568. 11: 7,1.1. Ill: 1829·33. 18•11
catarrhal pneumonia Ill: 2072 Ill: 1531. 1588 --parfr111ge11s 1ype B Infection Ill: 1836·9
ca1arrhnl strangles Ill: 1i71 Chlnmydoµh/11111/Jorws 3·1'i". 551 , 35.;_ - - p~rfri11ge11s type C infection II[: 1.; ,5
cattle lou<t: ny 59; Ill: 1517. 1520, l.>37 1623 1840-44
eaulc plague see rindcrpest - -cnvim., 551 - - /Nrj'rl11g,ms t)'pl! D inf~ctfon
c:mlt,-d~rh·e(I Tlreflerin pttrl'fl infection srr --/~li.1551 HI: IIHC,-52
Ea.,t Co:ISI fevc,r --p«on,111 ·117, 525,551,555. II: 11.58. --1wr[ri11gem<1yp11 f 111: 1838
rnuda equinn neuri1fs II: 823 111: 1488. 1.;89. 1490. 1639 - pil/Jormr IH: 2089
cnud11 i•qnlna syndrome Ill: 2019 --1meumo11ta~ss 1, 559 - - s.tptlcum lnfccuon lfl: 1111>9·73
caudal ~kin fold Ill 1984 - - f1$/llnci347, 5:iO. 551. II: 1250 - - sMd1•/lii infection Ill: !8i.J.5
caudal vcn,1 cava Ill: 1740. 1742_. 1950 Cl1l1ul'//11 spccit.'S Ill. 2142.21-13 - - re1m1i 111; I8i8
cell-assocl111ed ,ir;iomin in cquld chlorcllosi~ Ill· 2113 coagula1kin c;1scade 4,;5
hcrpc.,irus l and ·I Infection, II: 1!3- chlonn:ued hydl'OC'arboms2.;, II: 1158 coagulop:ath> Ill: 1595. l6IO.see11/so
ceUulili> Ill: 1691, 1912. 2019 chlorfna1e<I naphthnlene~35, imrava.~cular toagula1ion.
Ce11clm,~ cillorl, ll: 1185 chlomphyU-rontaini11g algal' lll: 2142 dis.cinlnnted
(cnrral nervou, ,ys,em disorders chocolate ;ignr Ill: 1629. 1631 coarse bont·leggcd tick Ii
canle If: 120; cholecy,rit,, Ill: 1620 ,obalt defk,ency 62!
horses 11: 849 cholera toxin Ill· 156S, 1603 Cocul ,type 21 virus II: 119-:
pigs 11: 912 choriore1inopath)' coccidloldal gronulomota UI: 2117 • 18
sheep II: 1207 and d}'SPIBsin. ca1 tic 11: 9-:;3 01/:ddioitlos immici.< 111: 2117
Cepho/oplws 1110111/cola II: Ion horses 11: 8-1-t coccidioidomycosi~ Ill: 2112. 2111.1a
Ccratopogonidae 93 humans344. ll: 12,12 c<>cC'idiosis 3 I9-29. 111: 1623
Cen1w1hurl11111 sim11111 II: 1227, 123~ c11,,,srom11tJ//(I glo1sll1Mtl8 carrle 324. 325,326. 11:901.111: 1481 , 1570
cerebellar dysplasla II: 9i'2 chromoblm,tomyct,sis Ill: 2lO!l· lO pigs 324. 325. 327, 11: 782. Ill: 1475
ccrebr"J babesiosis ,113. ·114, 417, -17i. 52~. chronic Mrfcan swine fever II: 1092. 1106, sheep/gum" 323. 325,326. II: 1260.
603. II· 1158 1107 Ill: 1597. 2158
cerebral cysticercosi< 11: 1158 chronic foo1-nnd-mouth di:<ea>e' II: 13.18. cocddlosis 1• crypto<poridio<is 334
cerebral Infarction.\ 71i, 4,ll. see also In fore ts 1353 Cod1/io/){)/11ss11idfim1.< Ill, 210,, 2110
cerebml 1helleri05is ~Ii, 4iS·i, 478,479, chronic glandcn, Ill: 1502 O>d1/iomyfn l10mi11i111mu: 67
480. 52~. 603.11: 1158.111: 1ti39. 16-10 chronic- in1er,1illal nephritis UL 1.J4S coenurm c<.•rcbrolis, ,hcep II: 1158
cerebral thrombo$1!\ '175 chronic organ tuilcrculo,is Ill: 1\180 coiral 11xan1hemn.. cault.' see infectious
 bo,'ine rhlnQtraclleili> contagious equine metrllls II: 872. sec also 1111,for poiso11ing
coital \"esicular examh~mn II: 868-73 Ill: 2084- 7 co11marin rodentkide.., 11: ll lO
Cokeshan~bocst 618 comag,ous pustular dermatitis II: l'.198-9. Councilman bodies 11: 1053. 1081
cold ou10-am!bodles5i6 Ill 1504: see also or( Cowdnn ruminnnr/11111 50i. 336,537
Cole.,·low conj1mc1irae Ill: 1-190 comrolof cowdriosis see hcartwater
colibncillo~is aTbovirus diseases 162-3 cowpox II: 892, 1292. 1296-7
cmtle328. lll: 1;;61. 1563 li\"estock dlseas~ l 71-21 ; a; zoonosls 11: 1296
equlds 111: 15.0 mosquitoes 147 cowpox ,1ru, 239, II: 1297
plg,, Ill: l-131. 1561. 1562, 1623 robic,.carr!er,pccic~II: 1136.1137 coxiclla anti chlam)•dia 560
shcep/goats32i, II: 1260.111: 156•1, 15i0 Tabanidac !II Coxlefln burm•tfl 4, 90. 347,565, 11: 1?50,
colll:tocillosh ,, cryptosporldiosh 33-1 ticks and rick-borne diseases 27-34 417 tu:1517, 1520. 1537
collbacillosi<, emerlc Ill: 156-0. 1563. 15&1. 1rypano:,0mosis 68·9. 280 -8i coyote 382. II. 757, 11 28
15i0. 1571 copper deficiency 621. II: 973 Crassi1taceao II: 1401
colic copper poboning. $/!e 1111tler poisoning CrcL1t>.fcld1-Jacoh disease II: l3B8. 1389.
C,lltle llt; 1934 Coq11//lt11if/la permrb,ws II: 1017 1$91. 1'1()9. 1422
horses.585.11: 1024.111: 1609. 1610. 2019 corntal limbal net1vascularizauon. horses Crimean-Congo h~emotThagic fever-I. 15.
colist-pt!caemia Ill: 1563, 15&1, 1568-9 II: l)Q.3 ti. II: ron- 83
colitis x587 corneal opacity II: 900 ~- zoonosis II: I on
coUti, buffalo II: 640 Crimean-Congo hnemorrhagic ferer ,irus
horscs585.58i, 111: 161 l. 1612 CilHIC 268,456.457, 4 73, II; 640. 953 162
pigs 111: 1~31). 1475. 1604 dogs280 cripples Ill: I iM
sheep goats Ill: 1595, IS96, 1S97 equ1d, :?99. II: 663 Cr/s11sp1ra 111: 1440
eolith. spirochneml Ill: 14i5 giraffe U: 640 Crucid11ro species II: 1145
collmed peccaries ll: 1195 kudu I!: 640 Croc111acrorura 11: I L33, Ill: 1975
Coll<ir/1in11s 11rsi1111s 5!i3 wildlife I!: 640 Crohn 's disease and pnratuberculosis
r.olobus monkey papilloma,'irus II: i5, corne,il ulceration set' 1111der ulceration 111:2004
colonic bloat. pigs Ill: 1475 com< m: 1;2~ Cmr11T11riLI species II: 1057. 1191
colonil'.mion pill 111 1561 Co,·011m•iridau n: 70J. 779-804. 1260. 1281 Cro10/aril1 b11rkem10 II: l!91
Col1fvir11s 156 m: 1,;s1. 1562, 1570. 157 l. lS8l.l crow pic'k Ill: 18iO
combined immunodeficiency \yndrome coronal'lrus lnfoc1 ion crowding1oyndrome cornplel< II: 1289
333. U: 823.111: 2089, 2137 c;mlc 328. fl· 79a-801. UI: 1479 cro1,·$ ~s ,·inJs hosts 11: 1006
competiti\·c cnzyme-lin~cd horse< II: 804 c;ryptOCOC(OSi~ 111: 2118• 19
immunosorbcnt assay Ill: 1519 sheep II: 803. Ill: 1597 Cry1Horo«11~ 11,:nfnrmans ru, 21111
compleml!nt ca.<cndc -155 \\iltl ruminant• II: 795, 798. 79!) crypto•poridiosis 332·5, I!: 1260
complement fixation test Ill: 15li. 1518 corona,1rui Infection.,, cryptosporldiosb cnule 328, 11: 961
c-0ngenhal dei~cis 33,1 sh,-cp327
cattle II: 953. 1029·34, 1254 coron:ll1rus es, s~ Coro11ar,fridoe Ct,'PIOS/JOri(//11111 <lflj'leyi 332
pig.,. paramyxuviru" 1!: 68i·90 coroniti~. cau!e II: 120. 1227 - - mri<>agrides 3:{2
<hrep/goat, II: 1029-34. 121•1 Corridor di.ease 21, 22. 227,449.459, - - m11ris 3.12
congenital in(ectlon. humans 3,14 468• 70. 525. [II: 2055 - - /la.<Onl/11 332
congenital trembles JI: 970· ;.; Corripantl\'irus U: 12,lB --pan•um 332
congenital tremors. pigs 11: 91-l Cory11ebnc1eriu111 specie~ !II: 1938, 2003, - - S£'1)/t'IIIL< 332
congenital cuberculosts lit: 1980 2011 CSIRO Village virus II: 1253
congei;1ion C.ory•1wbnc1cri11111 ,ysliudis Ill: 1931 Cum<>rephnlides fo/L< S84
generalized Ill: 1596 - -diphrlwriae Ill: 1i,11 <'Uffing pncumoni:1. pig5 UI: I i05
oronusal mucous membranes II: 1223 - - ~Ill ~ee Rlu,dor1>c1·u.s,'</lli C11/e.rfF.1mMla11omyiaJr11bi110111s 139, 142
Conidiobo/us species JU: 2113, 2120 - - o,is Infection Ill: 19 1i·25 C11l11.•(.llelmuxo11inn/specfes II: 1017 1019,
conjuncth'iri> - - oL•is sc,tt Corynttbtlcterium 1024
cattle II: i;.;o, 8i7. 955. 12;1. 1322. p.,e11duftllll!rc1,/1,sis C11/r.ac:i,x.> rlwi/vri 138. 139. 141 ·2. JI: 1041
111: 148,. 357, rH: 1449, H88 - - pllosum Ill, 1931 - - uuii,i11111u!l 142
cquids II: 8~:t 92i. Ill : 1771. 2104 - -pswdo111berc11lf1Sis ll: 729.11 1: l5N. --romba~11sls 142, II: I 043
pigs ll: i,6. 979 1531 Cufo.• ann11/iros1rfs575, II: 118-1
,heep/1:na1> II: 1073 - - ptc1uloru/Je,c11/0$/s Infection - - flr/e/lUIIWS II; 10·14
conjuncri\iti~. chlamydial 5.'i6. 557 Ill 1917-25 - - g,•li,lus II: l 023
Co1111ochat1...s 81111 35.l, 5 l6. 11: 8i6, 895. --p;•ogr11,~ see Arr:m10/Jac1eri11111 - - mo1emis 11 · HlOi
IU: 14l6 pyog,•,ws - - peru.tig1111.~ 11: 1044
- - rm1rm11s 3..:;3, 11: 8i6, 895 - - re1111le ru: J9:SI -plpieus 14i, II: 1007, UM-I. 123-l
- - ro11rin11s alboj11bt1111, II: 895 -rMa/einfoction Ill: 19!31·4. 1959 - - poidli/JeS If: I 043
- - rm1rl1111s u111rin11s ll: 895 - - s11is see Ae11110Lmcu/11m (E11/}(lc;leri11m) - - rm~nlis 11: IO li
conservancies 233 Slli$ - - triraeniorrhynclms I!: ID-15
comug,ous aboruon see P0'1ne brucellos!s conon mt ll: 82l cullcmc mo,qultocs see Cule.t
contagious arnc Ill: 1917. 1921. 1922. 1924 coltontaU rabbit p;1pillomavirus ii; 75•1. 75i (Mela11oc()11io11) species
rnmagious agalactiall: 1;!97.111: 2046 cough C11/icuides
~orir\lllloos bu, in~ pleu,opneurnonia I7 I. c.mt~ II. Tilll. 877. lll: 1638, 16-00. zo.w. D.nd ,\fiican buffalo l 21
226. 459. 5·l I. 11: 95i. 1 lt3, Ill: 2045-5 7 2050 African elcpham 11 i -20
contagious bo,ine pyelonephrit,s equids II: 768. 842,864 . 927. lll: 1502. 2rbo,ifll5es 98
UI: 1931··1 2018.2121 associated diseases/infections 98· 102.
contagious bronch!olo-alveolar carcinoma pigs n: G94, ns. i76. 792. Ill: 1-19-1. 1631. II: 103 1
11: il 7 166'1 2066. 2067 as~ociated micl'o·organisms 97 (to.hie)
comagious <:'aprine plcuropneumonia shceplgoais II: 725. i36, 743. 111: 1699. biOS)'Stematie< I03·8
U: 668, UI: 2060-66 2060,:?061 coll~'Cting adults 95·6
comagiou., cc1hyma see orr coumarln rodentici<lc poi~onrng ll: 11 Io. control li4-6
viii """'-
filarlnl nemalodcs911 cyt0adhcrcnce 4J.l ex·uda1iv,;; Ill: 2026. 2008
host prefercn<!es 117-23 Cyuxxwe., su Ehrlichio homis II: 823. Ill: liSi, 2110
life cycle 123..1 ciwmegul<wirus infec1io11. plgs ti: 919-21, humans. pus1ulnr 111: 1586
livcsiock J 2 t .3 ill: 1-194. lil(j !11terdit[it11J tll: 1-142, 1720. 1721, 1723-9,
liw<cock and" ildlifi' 98· l 02 C\10mcgnlovint!>·like bccnhcrpc,wirus1csl 19~9
plain~ zebra 120•:?I fl:660 mycobacwrinl Ill: 1979
1a,,0nomy 103·8 cytomegaly. pigs IT: 920 pig, II: 940. 97'9
vcc1orcapaci1yol 102 1..;~oplasmrc bodies 11: l 048. I053, we also pigs, subcorneal pustular Iii. t i54
vet·cor compe1encc of l 02 Jnclusic,111, pododem,al III: I 727
\-.:C,or ,pccie, 108·9 (11Dplasmic inclw,lon, II: 1123. see 11/so sh~1,p1goats Ill: li29. 17SA. 753. !912,
vim! maintenance nnd tmnsmission l 0? Inclusion~ 193•1. 2<)98. seo also Bolo dlsca.,c: o,ine
viru,c, ,~soclatecl with 98-103, II: l 031 cytotoxlt" necrmizlng faaor 111: 1561. 1568 intcrdlgita! clcrmatilis:$18phyloccc;cal
\1ru,cs Isolated from 99-101 (table) cy1otoxln Ill: 1561 dcm1:nitis in sheep
C11/icoi<fl!5 s.pccie,. as vec:1ors 93-126. .ummer scaso11;t1 recurrent 98
fl: 1019. 1031. 1184. 1195. 1201, 1205, vcrru1·ose Ill: 172·1. 1725
1221. 1223. 122,. 1231, 12:13, 1247, 1252 Derm"riris digirails lit: 172.1
Culicoidl'S (Hojfi11a11ia) insignis 102 Dermml1is imertligitnliJ Ill: l ;23
C11/icoldes 11edfordi 10.L 11 I d"Aguilanirus 99. 100. fl: 12.18 Dermariti.rrodosa Iii: 1983
- -bolirino.(95. 104. 114. 115-17. fl: 123-l. d~f, tomb disea;c II: 9;3 Dem1111Iri.<1!/'rrucos11 UI: 1725
12,ia Danuufama lit: 1689, 1973 dermntophUosis Ill: 2026-35
- - /Jrvv/mrtis II: 1031. 1184, 1205. 1227 Damutiscus albifro11s II: 1205 canle II: 127'1
--fmu:ei 114 --dor.:11Sphillips(Sl6, U: 1046, Ill: 1479 humans Ill: 2033
- - COQTCia1U$ 102. I 14. II: 1184 damllo II: t 041 shecp/gua1s II: 1285. t28a. Ill: I i56
- - t'nd/Jr1Pi11/ 1J.I c1am,c10ie• 5114 Demrato}lltl/us ccmgulensis II: tl-15. 759.
- - ~11gu/x111dei 104. t 1-1 Darmbrlllld in humans rll: ll!-10 1280.111:2026
--full•i1horcix 104. 114 deer 11lt!-87. 89 Ot!n11atoph/11u co11gole11sis infoction
- - glabripem1is 114 deer Ill: 14•16.16\tl. 1622, 1829. l<\73,see II: 1284, 1299
- - gu/lJ.,n/.:iani 104. 114 also i11d/1,fr/11nl 11ames dem1atophytes 11: 759. l:!94. 1299, Iii: 2095.
--imicola95, 104. 109-14. 116- 17, dclidcncy 209G
fl: 1184. 1201. 1205, 1227, 123<1, 1248 biotin Ill· 1743 denna1oph110$ls 357. Ill: 2095-2101
- - i11~11,.,nis 11, 1205 ,·ohalt 621 dcrmawsb, genital ulccrn1iw II: 1287
- - leuco.mcms 104, U4 copper 621. 11 • 9i3 durt'e11g11e II: 1129
- - loxo,/Qnli$ l20 pho~phoms Ill: 18!16. 1889 desen rot ILi: 15•1(l
- - IIUIJ/1111$ 104, 114 selenium II: 738. 1311 <lesert rheumatism Ill : 21 IT-18
- - mi/ne, 101. 11·1 thiamme. caule II: 1158, Ill· 1639 Desmod11.< ror111ui11.< II: 1129
- - llffll'el104. 11 4 th1amme, sheep II: 9i3 l)cu1crornyc01111i.1 Ill: 2095
- - llll'OSllSI 1-1 \itumin A Ill: l •U!ll deviation of nasal septum. pigs 111: 170Y
--oxyswma 101, II: 1031 \1tnmin a, 417 diabetes mellirus. raule It: 950
--pa//idipwwis II: !WI ,,iamln E!selcnlum 11: 738, 1311. diamond skin chseasc Ill: I908· L2
--11ycncmicws10.1.11~ 111. I l-12, l5i'l diarrhoea Ill: 1560. 187•l. 199,l. 1997. 2.003
- - ra1111s 11-1 zlnk 111: 1721 diarrhoea in canlc327. ·189. 520,540.
- - sclwlrzel 101 derormntlon. facial. pigs IT[; 1707. 1743 Ill: J.180. 1481. 1586. 1619. 1875, 2003,
- - simflis 11>4. I I 4 dcgener:nion. 1csiku!ar 111: 1529. ser also se~ also calf diarrh®a
- - troplralis I04 1111t1cra1rophy a<Socia1ed with ,iru~(e.~J 11: &10. 816. 900.
--1•<trilpm11i1 II. 1201. 120S, 1227, 123.; degenerative angiopaihy tu: 1570 946-62, 1228, 1260
- - wadai JI: 1031, 1205 dehorning Ill: 1948 lliarrhoea in horses 585. 586,111, 1608, 1610.
- - :11/rnmsi..<98.10>1. 114 dehydration Ill, !59H 1612. 1&23.2019.2090
curly hair coat, cant~ II: 953 delayed ronception sec lnfenllity assocla1c<l with ,iru~ie•l II: 804,823.927.
Cun.,,/aria ge11/c11la111 Ill: 2H)7 dela) ed hypen.~n~itlvhy, wberculosf,. 1261
cut~neous amhnix lit: u1u2 Ill: l!lHi diarrhoea 111 pigs 7117·!1, Ill: 14JO. Mb~.
cutaneous disease:. c:at:.ldog~ Ill: 2140 delrn 1oxin ill: 1840, 1657, 186-1, 1870 1563. 1569, 1603. 160-1 16"..2, 1829, 1841
cuiant>Ou, glanders m: 1502 De11101ft'.tinrec1ion 11: 12;~. 1293 as,ocla1ed \\ilh ,·iru~tes) II: -s1 787-97.
cuianeons habronemiosis II: 759 dem)'elinating encephalorn)'elitl, II: i36 809.900,9;9. 1260
cutaneoui; horr~ II: 7:,7 dem)'l'lina1ing leukocnccpl1alo111rditis diorrh11cn In ~hccp/goots 325.111: 1480,
curnneous leishmaniosis 379 II; 7•H 159<1. 15%. 1596. 1597. 1620. 183;,
cutaneous nocardiosis 111: 2028 deng\lc or can le II: I 183 1842, 1848. 111;5
cutaneous strcptothncosis Ill: 2026 dengue \inis 11: 969. 1082 associatl>d wilh ,,rus(es II: 900, 10i3
cyanosis Ocnmnrk bat virus II : 11 46 diarrhoea
skin II.I: 1603. 1631. 1664 depression. profound. cattle Ill: 1637, 1639 assocmmd with cocddios,s 325
tongue 11: 1201 deranged osteogen~is. canle 11: 95:l nssociaced with cryptosporidiosis 333
cyathostomosis. horses lit: 1612 Dermacenrorspecies537, 11: 1079 nconntnl cnlfll: 1260. HI: 1569
Cyntmclmm spec,ts4 I 7,524, II: 1000. I 1,,8. Dermace111or (l/bi11ic111s 596. 618 post-weaning Ill: 1563, 1369
1,40]. 1417 --amlerto11i 595. 618 rotavlral II: 1260
CynlcNs p,mtcllfma II: J 133 - - lliWIIS ~26. 421 mw~U1:rs· 333
Cynodrm daery/011 524 - - oceidl'111aTis6!8 D/c~rw/Jimml.tfl: 122i. l:!3·1.lll: 1~~6.
cyst($) Il l. 1;;31 - - ri'liC'lllat11., 42,. 43r; 2116
cys1kercosls. cerebral II: 1158 --sill'llrum 618 Dicltuloooaer f&1cwroides) 11orlos11.i
cys.titlsand p\•clor\cphritis, pig£ UI: 1958-60 - - mri(lbilis SIH. 596. G18 Ill: 1720. 1729. li33
cystl1ls Ill: I :,70, 1931 derma1i~ceou~ fungus flt: 2ll>t. 2! 10 f)i,tyocm11t,1 fii11ria infes1acion 11: ;3a
pi~ II: 938, UI: 1956·60 dermatitis dicumarol polsonlng II: 983, see (I/so under
shecp/goa<S lit: 1933 CUii.fl' Ill: 20 11. 2012, 2098, 2110 poi,011ill&
cr!(l11xZQ011ra11rQ1mg1478 digital Il l· 1720. 1i2·1 Dldeiphi~ ~pede~ 3i l
Dide/phis nlbivemrls 39-l Ougbe viru, II: IUil can le 3, 21. 22, S36. 538·4 I. 542
--pflrnq110)'ellsis S53 du1ker 54-4. 598 dogs 280. 344.11: IIS8
- - 1rirgi11iaun 394 dumb 111bies It: I IS2 hors~ 583-9, Ill: 1612
dlgiin1 dermatnls HI: 1720. l ;2,1. see (I/so dummp111drome Ill: 1848 pigs543
dermatlus dunkop form of horse sickpcs,s ll: 1235, ehrlichiosb. granulocytic 536. :i38. 545
dlkko;> b)' mmme llf: 1863·S 1236·40 Eidolm1 helv11111 II: l HS
dlkkop form of horse \icknes.s II: 12:15. dusr pneumonia. caulc s~e infecrlous £imerlaspccks319·27, Ill: 1562, 1571 1588
1236· 40 bo,ine rhinotrnc.heic1.s F.lmerin 11h<111n 320. 323
dimfdium bromide 281 D11uo11~U11 ~9 - fllabnm~tt$iS322
d!min:izine 2111. 282, 11: i 158 Ou\'enhage,in,sll: 1124, 11 31.114S. ll~ti. - - (J/ije,,i 320
Dipcndlglaurn II: 1158 1163 - - 11pshero11lm 320
diphlherla d)">entery - - mloi11gl 320
calte~ 111: I ;41 . 2. 1950 cattle U: ,95. 798, Ul: 1-179 - - nr1b11m~r,$/$ 322
humnns Ill: 1,41 lnrnbs 327.111: 159i'. J83H-9 - - /,rrk11emi., 320. 323
diphtherltlc p,eudomembrane Ill: 1472 pigs II: 983.111: 1428·33, 14;5, 1605, 1623 - cnprlnn320
diph1herotd 1110amma1lon. pigs II: 979 dysrnyelinauon II: 953, 972 - - ,n11ro1,i110 320
Dlplodia lllflJ'dis ll: 9i3, 1158. I l91 dy,phagia 11, 711 . l223, 1228. Ill: I H2 - - rhri$tl!ns..mi 320
diplodios.is477. ll: ll58. J.117.111: 1639, d)·splnsia, cerebellar II: 972 - - cranr/(11/is 320, 323
1897 dri.pnoca II: ii I 725. 736, ii5, 785.111: 2138
- dei,1/ec~-/ 323
dipping 111; 2028 ca1tle lll: 16/lO. 1691 - ellip.wltftJlis 322
Dirofilari11 rope11s 90 horse1; II: 1236-7
-faurl 320. 32.~
- - roem~r/90 pre< u -920. 131a. 111, 11,1;.i
~heep goall. ll: 999, Ill· 1699. liOl
- -gilmr/1i320
diriycowsyndrome Ill: 16.~8 - - grmmlosa 320. 323
discharge. see under aJJctr;:d .<ite, e.g. nos:u dys1ocia II: 1049
- - l,irci320
cischarge - - inrricma .120. 323
discoloration
fleece Ill: 1938
kidney U: 902. ID: !450. 15i0. 16(14
0 --jolchijevi 320
- - J.uchnrll 3:!U
- - neod,•bli,v:ki ;J23
skin ll: 9i9, 111: 1910 ~l>·-onsc1 muco$al disease. cnulc 11,951
- - 11i1wkohly"l:imo1>ae320
dl$COSpondylitis l!l: 195 i l;a.i African bonr 1ick 10
- - 011/110/dnlit 320,323
disease control in li\'cswok F.aM Coas1 fe\'~r 21. 27, i.s, 225. 275. 106
- pnllldn 323
decision~ I 78-21 i 448-63. ~68. 4i2, 481. ~91, 525.5-M.
- - 11(1n•11320. 323
1>ildlife lmeriace 225-35 It: 712, lll: 1436. 2-055
disscminau:d in1ravascult1.rcoagula1ion 413, ,H) pica! cerebral fom1 475·7 - perminma 323
~28. 4,}2, 455. II: 1048. 1051. 1102, 12()6, ,-;,ctinalion ~60-61 --po/ira323
1210, 1?28, Ill· 1568, 1597, 16.'lO. Hi'.l7. P.aMcm equine 1•ncephalhls II: Io 15· l 7 -purd323
1739 a$ 7.oono,i~ II: 1010 - - p1111c1nra 323
di,1emper En~tem equinu encephnlhis virus 15;, - - .,cabm 323
"8nine227,34,l. ll:696. 1158 II: l014 - - spilUJSfl 323

ce1accan It: 63-l l:asiem equine cncephalomyeli1i.~ 13i' - - sttl.i 32.3

ht·llgehog It: 634 II: 685, 1010, t3i0 - - 1wybritlge11sis 320. 323
phocine II: 634 eastern pipiJmc!lell: 1128 --wemii320
Ol\'A vacCinarion 243 Ebola cc,·er II: 1082 £1,ieophora srlweid~rl 90
di\'eniculitis lll: 1959 ecchymosis Ill: 1596. 1691. H\99 eland 1;s, 516.11: 1423, lfl· 1512. 1546. 1547
Dobsoni(J n11d~rso11i ll: 682 cdamps!a Ill. 1892 clcc1roc1,11ion Ill: 180,
- - molllctl!II.<~ 11: 682 ccowurism 233, 235 ~lepha:m see African elephant or Indian
docking ofta.lJ$ It: 757 l'CLemn i1tShee1,, fllcinl Ill: l 75-1. 1755 clephanr
dng rab!e.• 11: I 124, 1126, 1144·5 Egyptian iever I7. 18, 4116-93 el,•phant skin disease 351 ·8. $ee also
dog llddl: 1235 Elrrlid1111sptcles .;4 I. 536, S3i, Ill. 1440 be~noitiosl~
do~ 2;9 u: 123.J, 1236 Ehrlfl'hfn bo11s:!I, 22,509, 5~~. 5:i6, 543 El,•pltns timl;(ll~11sl> Ill: 1690
dog-,iufng po~ltion Ill: 1951 lnfeclion 3, 21. 22,536, 5.18-~ 1. 542 l!lk II: 798, 799, 1195, 1227, 1.\22. Ill: 15-16.
dolph!n$ ll: ;oo. Ill: 154G. 1549 f!.lrrlichin mnis 280. 508. 536. 53i 1547, 11389. 1973. 1975, 19$5
donkeysJI. 123.J. 1237 12•18 - - cliaffems;'s 508. 53t. 537,538 cmaclared carcass Ill: 1~72
dopklou m: I 733 - - eq11/ 536. 54S cmaciarlon HL 1994. 19')7, 2003
l)ors~1 egg ,n~dium Ill: 2027 - - ewingi 508. 509 canlc II: 953. Ill: 2003. 2050
dosing pneumonia II I. 1698 - - muri~508 equids U: 750
l)ougla, ,·iru, 99, U: 1031 - - 011dir1 544. 54S <hecp gonr, Ill: 2003. 2158
dourine 251. 279. 297-302. n: 928, 102;; --01•/11a '.!~. 537. 538. 5·11. 542. 621 embClllt $Uppura1ive ncphrili> 111: 1652
downer cows II: 14 I 7 IH:2157 embryonic death 344, II: 990. 1459, sre 11/so
OPP6S \'irus II: 1253 - - pi:ag«}'tOphila 441., 536, 537, 544, S.15. focull dearh
draa.isickre 4,$-7. see fll$o ccrcbntl f,21.11:99;, Ill: 1;:,4, 2157 caul"30i. ll:946.9.53, Ill: 1461, 19:;~
theileriosis - p/G/)'$ 509, 536 equids Ill: li84
dmgonOi~ 58-1 - rl$t/cii 90,509,536, SUJ.111: 1612 pi&s !!: 80H
on,:ho1ufll ro,crc,ru 111: 2111 - - rumit111111ium ,i, 8. 507-26. 53i emph) ,~nm II: 016, I l 00, 1191
- $perifura Ill: 2110 --srn1w1.m 509. 536. 583 pulmonary Tl: 1l 89
drie-dnc-sn·wesiek1e SP.I! bovine ephemeral - - suis. 543 subrumneous II: 1189
[ever chrilchia cmpyema Ill: lii2
drodro·bol.'ll Ill· 2026·35 as ,monosls 5-l I vmull: 1020
dro,is 111: 1500-1504. 1505 t,anulo<11ic 539. 545 1mcephali1is 34,1. 360. u· l310. 111: 14-11
Drosophila meln11oga.,1rr 159 Hyalomma-rclated 542 bird~ ll: 1018
dry gangrene lll: 1586. 1910 monoc)~lc539 cantell: 950, i·U6, 14li
duhiwhjie~ II: 987 Ill: 1736 chrlichio,ls 536. 538. 54S. 621 cquid$ 386,396.400, ll: 692. 8'19, 1010,
 1014·21. l02S, 1196, 12-42. 1368, 1371, h<>rses Ill: 1610 epldemiology 194-205. 215- J7
Ill: 14-l:! pig,,11·822 Ill; l-i30.1:;59 epidermlciS Ill: 2026. 2030
equids. Ge1ah II: 849 enteritis II: 90 I. lll: 1994 exudauw Ill: 1759,61. :W26, 203!!
equids. Japanese B 137 caule 556. IT: 820. 878,951. !I!: 14i9, 1582. Jipider111opl1y1011 species !II: 209.5
equids• .\hirr:t)' Valley 13i 1586. 1639, 1840·44. 1875 Bpid1•rn1oph,1•ro11jl0<:lvswn lll: 209,
equid~. mycotic- JU: :!116 cquids if: 927, Ill: 18-10·•1·1. 1989 epidid)'trtiti~ Ill: 15H. 1515, ~28. 1529.
equfds. St Loub 137 pit;s32i, 111: 1·128. 1•169.1604. 1;s1 153.5, 1537. 19S2. 1953
goat, 11:,43 1840-44 ca11ldll: 165,. 2076. 2 l ·19-51
human, If: 692. 694. 997. 1014. f018. ,h~pfgoah 326. 556.11: !l7l. Ill: 1479 sheep/go~~ Ill: 1655. 1656, 1657, 1658
1019. 1020. 103,. 1049, 1051. 1242 1595. 1836. 18'13. 1875 epididymit!s·\''1ginltis Ill· 2149·51
non-equln~~ II: 83•1. 841 enteritis. a~~C>Ciaced wuh coccidio,i,,32b epidid,mo-orchith Ill: 1620, 1656
pig, II: 93- haemorrhagic sec rmder haemorrhagic epilepsy II: 1:170
.;nr~phali1h. Callfomia 90 prolireratl\'I: Ill: 1989 epista~i~ Ill: 1709.17-1:Z. 1950.?1J5.2116
Japanll$e 137, II: (xi~. 1252 e111,•rlris11ecro1ica11s. human Ill: 18•0 cphhe.li.tliwtion. alveoli II: ,29
necrotizlng II: 1019 /!nrt>robacrer Ill: 1560 cpitheliold cells l!J: 19;; 1988. 1997. 1999,
Russian spring·summer II: 995 enu:rocohlts Ill: 18i5 21+1
thromboembolic II: 1416 canle LIi: 1619 cpi1heliornn. ha~al ~ell II: 75;
1i<::k·bome 90. 11: 995 horS<l\\ II: 804. Iii: 2019. 2090. 2116 epiv11g Ill: HG-I. 2.149·51
enccphalomalacia llf: 1571 pig, Ill: 1601. 160~. Ir,()5. 1617. 1622 cpiwotic nhoninn or ewe~ 551. 554
focal srmmenical 111: 11¼6. 18-li. 18~8. sheep/goats Ill: 1597, 161i. 1620. 1622. cyizooc!c bovine abonlon 551. Ill: 1436
1849 18-19 epizoocic haemorrhagic disease of deer !!3.
enccphnlomyelttls If: 1311 enterohncmotrh:lgic E.schvrlcl11t1 ({JI/ ll:9GO. 1185.1196, 1213. 1227·9
catlle-ll,.ff:-l77.556. 11: 1033, 1158, HI: 1561 cpir.ootic hacmQrrhag1c disease of deer
fll: 1639 enwropmhog.•nic J'ischvri<'i1it1 coll 111: 1561. v!rus98, l00,ll:90S, 1220. 12~8
equld~386.396. If: 1006-IOi 1370 15s;.a
epi1.00tic in(enilirv Ill: 21.,9.51
pig< ff:791, 792,793,914, 1307. 13115 en1eropath~· cpizootlc lirmphani:i1l, Ill. 1504. 2104·i
sheep u: 995,999 hyperplasck. sheep 11: 9i3
F.pomoplum,., wnlt/11ergi 11: I 1-15
encephalmnyell1is 11rolifora1i,-c, Jli&:, Ill: 1469-75. 1605
c:p~llon toxin Lil: 1846. IS.Ii. 1850. t8(;.;
demy(•llnatlng l l: 736 entcrnrrhagia 322
Epstdn·flarr hcrp~,,iru., II: MO. 86,; 896
Ea,tern cljuine 137.11: 685, 1010. 1370 ,,nt~rotaxaemrn Ill: 1829·33, 1840, 18-16-52
£pt~irns ~erori11m 11, 1116
.'.l:ear-1:a,tem equine II: 1006 caufe Ill: 1840--14. 18J8
equicl g.tmmaherpe,vlrus Infections
\'cne.\Uelan equine 137, ff: LOI(). 1371) pig, Ill: 1840--M
II; 860·66, Iii: 201i
\\'~tllm t!quinc 90. 137. II: 68..;, 1010,
1370
shecp/11ou1:. Ill: Ul36, 1612
cmennoxaemla, neonatal If!: 1837
equid hcrpes,ints I nnd.; II: ,-o
equid he~,irus l and 4 inrecuons 400.
encephalomyocardius II: t:i09 emcrotoxaemic jaundice Ill: lll..'>!l
encephal<m1yocard1u, ,·1rus U: b~U. !! 14. 43 1. II. 6115. 829· 52. 928. 1010. 1025.
en1eroto~1cosb 11: !ill.l, I Z60
1310 emcrotoxigeruc r:..,rhericl1irl ,·o/1 HI: 1561, I 158, 12.50, 1320. 1310. lll: 14.';0. 2091
encephalomyocard,tis vlrusinfcctiun 1564.-6. 1597 equld lwrpcs,ims 2 and (i infection,
II: 1310· 11 cniermoxin!s, Ill: l:i61. 158i; 1619. 18~ II: 860-66. lll: 2017
cmceJ>halopathy enieromxin equid hcrpcwirus 3 If: 1129, 863-73
1eline II· 1368. L~22 htat-lnbile Ill: 1565. 1566 ~quid hcrpe.\,1rus ; see e11uid herp,•s,irus l
hepatic II: 1020, 1,117 hcat·Stable Ill· 1561. 1565 and ~ infections
sponi;ifcmn II; J 156 cntcrotyphlocolitis Iii: 1608. 1610. 151 l equld hcrpes,irus:; If: 860-66, Ill; 201;
endeml~ insrabllity 27, ,; 11. ,12,. 4>1, 598 cnwro,irustt-s) 11:1313. 1319 equid h~rpes,irus abortion 587.111: 1,68,
endemic stability 27. 409. 411. 42i. '139.·HO. cnucle:uion. f<>111kular lJ I: 14811 see also cquid h~rpcs,, rus 1 and 4
4:H •.isa. so-;-, 525. 546. 596. 11: ion. cnzOQtic bo,1no leuko~I~ 90, 275. II: 708-13, Infections
1075 iii equine ahonion sea abonion in cquid,
cndocardl1is e1\Zootic hepatitis see Rlft Valley (e\·er equine adcnnvirus II: 8-19
can!~ Ill. 17.:0, 2052 e112Qotk n3sal rumnur II: ;23 equine adenovirus infcc1ions U: 82:l·-I
hors~ 111, 1912 onzotnlc nasal wmouf\irus II 721. 723 ~urnc antrltls Viru, II: b!J5. 8~9. ~N. ~33
pig,. Ill: 17i6, 1909·12. 19:iO en1.001ic pneumonia II: 673, 77i, Ill: 1635 equine bahe.iosl~ ;,w eq11in11 piroplasmo.is
;,ndogunous uveitis Ill: 1449 call I<' Ill: lu.7 tquinc bc,noltio~i~ 351. sel! nlso
cndome1rhh en1.nmic stability see endern ic stability bL">noitiosis
cattle 307. 308. Ill: 1461, J.162, 1515. 158,. enzoolic 51eriliry Ill: 1459 equine coital esanthema II: 868-.3
1638, 1~52.2076.2078,2079 en1.~1ne•linked immuno•orbenl as.,ays equine coro1:1avirus 11: 804
horse, Ill: li>55. 1556, 15G4. 1570, 208.'>. Ill: l5l 7. 15111 equine coronavims inrl'Ct!on 11: 80-1
2122 cosinophilic club~ Ill: 2109 equine dcgcncrmive mych,encephalopaihy
pig$ HI: 1543 ,>vsinophilic myl'Sili~ 360, 3f>'I. 36- ff:$-1.9
sheep/goats lll: 1480 cpendyrnitis Ill: 1620 equine ehrtlchial abortion 583·&. lll: 1612
cndomecr!ti~. postporruncm Ill: J9r.2 Ep,uy1/rrozoo11 spcc,cs573. 618.11: 1110 equine ehrfich iakoliri< S83·9 Ill: 1612
tuberculous Ill: 1974 l,jwryrlrro::otm species inrection 436. 573-8. c.quiM cncc·phalitis see wuler encephali1ls
endotoxicshock Ill: 1568, 1610. 1612, 1630 621 11:938 equine enccphalomyefi1i.s see under
eudoto,in ill: 1568. 1585, 1603 1:~0~hrowo11 co,·eoides573 coccphalomyclili•
/:11/tydra lwrf.< 11erel< 395 --ot•IN-1.?. 5·15, 573. 621. Ill: 213, equine enctphalosfs 93. 98. ll: 68J. 84S.
Enra,,ux•b<I histo~nico 316 pan,11m573 12·11. 1247-50, sec nlso enccplmlltis;
emeric calicMru.s(csJ of pigs and cattle - - sui5 436. 573 en,·cphalumycliti•
II: 703-6. 800 - 1ega11{1(fe.~ 573 equine cncephalosis ,•lms II: l:?4,
enteric colibnclllos!s Ill: 1560, 1563. 1564, - - 111omiif,i3 equlnc flu 430.11: 766-72, 928. 1025. 1320.
15i0. 1571 - - 1t>e11yo11i 5,3 Ill: Ji68
einer!c diSea$e eJ>E!t)'thro~.oonosls ~36, 573-8, 621, I!: 938 equine gr;iss sickness Ill: 1829
cattle Ill: 1479. 1480. 1586 ephemeral f('\'(•r II: I L83 equine h~rpcs,irus abortf(ln 58,. Iii: 1768
dog, Ill: 2UO t;plrenwro1•mu II: 1124. I ! 8·1 ~quine herpe:,virus m~·uloencephnioparhr
 see equid hcrpe,vin,s I and 4 cruptke venereal disease II: 868· 73 reline infcc1!ou~ anaemia 58 I
Infection> erysipelas 11· .i7. 983, 1109, Ill: 1605. I031. fc:-lln~ infecliou~ pcrllonllh corona,irus
equint herpc,vim, ocular dlsca.~e see equld 1908·12 II· .SO. i87
herpesviru, I and -I mfcctlons ~1ysipclo1d Ill: 1912 feline par,o,irus II: 807
equine hisroplasmos~ 11.1: 2104·7 Frysil}l!lothri.r insidio.<a infcc1ion feline spongiform encepholopa1hy If: 1388.
c11uino infectious anaemia 90. 279. 431. Ill: 1908·12 1422
IT ,1; 747·51 ,928.1025 --rluuiopall1it1e!IO. 111: Ji36. li85.1908 feline ;ymme1rical enccphahipalh\· II: 1422
equine lnfec1!ou, anac1:,la ,1rus II: 741 - - rhusfuparhtac lnfectJon Ill: l 90IH2 F<'li.< c«rtlClll II: I 131
equine inOucn2a.:lO. II: 766·72, 928, 1025. - - 1011illtid II: 1908 - - ro11col11r II: 1228, 1423
1320, HJ: I;'68 erythcma mlgrnn~ Ill: 1·141. 1•142 - /ybica ~53. II: 11:l I
c,}uinc in0ucn4'! ,irus(es, II; 767. 8:19 e~~hrhol Ill: 1513 - nlgrlpes II: I l39
equine lmeslinal domidJosi> Ill. 1829-33 Esclterichiu coif II: 668. 769, 782. 983. 1260. l'erul pig, Ill: 1542. 15•16, 1548
equine monocyuc ehrlidiio<!~ 583·9, lll: 1428. 1452, 1481. 152ll. 1557. 1560. f,•rtlli1y reduction see infertilil)
Iii: 16i2 )58:1. 1588, 1617. li78, 1785, 18'11. 1959 rt,,cr fonn ofhol"iesicknes,; II: 1235.
equine mClrbllllviru, II· 681 ii.<rherid1/11 coll infec1ions 111: 1560·72, 123i-10
equine mo1or neuron disease ll: 849 1605. 1612. 11\ilJ re,·er high
equine papi1101mwiru~ ii: 755 t",Spundi:1 378 equio:hlll: li73,2121
cqume phycomycosis 111: 2113· H Essen schedule IT: 1163 goats ill: '.!01;(), 2061
equinepiroplasmosls4. IS, 17. 22.24.26. em 1o~!n Ill: 1864 pig, Ill· 1778
278. 425·32. II: 8-19. 123i. 124 l. 1250 Ethidlum 282 fibrinogen s111rhe,ls disru1bancc> II: 979
c-Jrriers 427 F.throplan w11lf II: 1130 lil>rinoid degcnerntion ofbloo<l vessels
equine pleurupncumonlu II l: 1744 £111.t<1cterlu111 mis Ill: 1962 II: b!H
equine pro10z.oal m~'t'!oencephaliris Eubenang<'e ,iru, I 00 tibrinoid \'USCU!iti~ II: 902. I 10,
394-400. I!: S49. 1020, 1370, Ill: t+IZ /:11m,•la11om.1ia ntbinows 139, 142
tibrinopurulcnt pleuriLis lit; 15,0
C(Juine recurrent u,·cilis 111: 1.J.19 F.11musca 81
flbmgen plasma olcvmed 111: t 773
equine rhinirisA virus I!: 1326 flbm:na 11: 75!1
eumyc01ic- mycc1oma lll: 1756. 2107
~qulne rhinopneumonici~ sei, equid F.11phori1111 mm1rita11im 52·1
inrerdiglml Ill: Ii25
herpesvlms J and4 Infections fibcopapillomn II: 755. 758
European brown hare syndrome Ill: I 5-18
llquin~ rhlno,irus infoctic.n II: 1319,20 fibros:ucoma 11, 759
European bcO\\TI hare <yndrome ,iru< ll: mo
equine rhinoviruo(ct< II: 849. 1319. 1326 fi~'Te catarrhnJ,:, du mnu1on see blucmnsuc
European h~JC lfl: 15·12, i;j46. 1548
equine salmoncllosi> 586. 5117. Il l. 1608-13 fiwri~I worms 89. I 53
F.uropcan rl•<h,-a1cr su bovine babe>io,is
equine 5arcoidr sl II: ,S5, 758. :ree also FilOl'irirlae II; l 082
European ~'>ine fet·er see hog choh,ru
11oc!ul~ fimhriae IU: I560
ei«;oriation II: 120 I. 1207, see also erosicm<
et;uine ,·encrcal balnnhb II: 1168· 73 ri~urc
,•xophthalmus fl: 711
equine ve1tereul "iMrls LI: 868-73 hnr!1.ontal Ill· I 72i
equine \1ral 11ncrlli> ll: ;;o. fl.19, 9?4-9, ex01oxin Ill: 1619. 1677, 1801 V<!Oic:ul Ill: Im
ex'Udatlvc cpid~nnlli~ Ill: 1759·61. 2026.
1025, 1241. 1150 lisru1'1
2030
lit/11/rili1101,fr11s II: 1319 rec10,,·aginal m: 1556
exudn1lw pm:umonla II: 726
£t/tt11$b1trcl1i>lli351. II: 1233. 12,18. lll: 1512. ,crolal Ill: IC-.56
eye-frequ~ming 1.f!pid1>1lltra Ill: l·lll8
IS17 fisru!nuswlrherslll: 1516. 1608, li85. 194-4
- - burcltelll bohmf 395
cyele$.< 1ampan z; flabby bag s~'nclromc lll· 1448
tyewonn, Ill: 1489
cradi~u1ion, n,1bcrculosis Ill: 1986 ll~cc,d paralysi~ Ille 1885. 1892. 189,1
l:rlXJv'rus U: 1319 flai:cllar protein anrigens Ill: 1560
F.r,•1ma11odidr~ spotles II: I 0,10, I04 l
crgoti,m Ill· 1.586
F.rosio w1g11/ai> lll: 1,24
0 Flaril'irirlae 155. II: I0$2
rl;l\i\'lru~!es) see Flaui<1fridne
Flal'Obtu:rerium pseudomal/i>i 111: 1617
tr0&ions face !lie, 77. 81 ·3. IT: 1339. IU: 1488 Occce ChSlll(CS II: 9i2. TII: 1938
oboma.sum II: s.u. 955, 1224 "nclal ncn·~ paralysi~ 300,11 1: 1905, 1906 Ole, II!: 2029
conjunc1i,·~ II L 1488 iaci:il pnrnlrsi, Ill: 19,18, 21 16 rloridn horse leech JI!: 2113· 1·1
rnrn~a ll1 1488 Fa.<itJgi.u Jw111iJlt'i 524 lluun.'>l l·uc~ pul«rl@thm as,a} 111 1519
coronal)· ba11d II: 1312S faeces \\ith blood Flury,-acctn~s1rnln II: 1130.1134 113,,
e,remal genitalia II: 86.'I, lll : 2152·5 ca11le Ill: 1480 1160, 1164
rem tn: 1;20, 1;2.; pigs Ill: I .30 nushing, skin, pig• lll: 1603, I!!! I
forustomuch 11. 1210 iacce< whh mucus, pig, Ill: 1430 Oying fo~l'S see frnil ba1s
gums U: 900. 1196, 1223 :iwce, \\ilh necroric m;uerial, pig~ !ll 1430 ny.~1rike 111: 1623
ik-ocaecal ,-alve II: 6&l :al!O\\' deer II: 1339. Ill: 16139, 1;;;, 1973, 1onl•hca1 diarrhoea II: i261
mou1h 11:t>l.J 900,953.955, llllG, 1201, 19,5 foam) ,~ru,(cs) II: 717
120i. 122:t 1313, 1346. 1351 false gl:inder$ Ill: 1500 focal symmc1rlcal tncephalomali~
nan.-s II: 6.0. 955 :a!.esandc:raddll: li43 Ill: 1846. 184i, 1848. 1849
nas2I cavity. can.le 11: 8ii. 955 :alse Stobie fiy i7 80 ioetal de,nh
na.,~I caviry. ~heep 11: 1201. 1207 Fannia m11ic11/aris ,7. 1!0 cnute 30i. II: BIS. 946. 953. 1031. 1252.
oesophagus II: G+l. 95S, 1224 Fanniinae ii, 80 Ill: 1585
Ol113SUlll 11: 1224 iarcy Ill: 1301. 1502. 201 l pigs II: 688, 806. 809
pala~ ll. 6~0. !lOO, 1223 farrell',sele<:tive medium Ill: 1~•3,, lo-13 $heep· goa1S II: 971. 990, 10:;o. 12::;2
penis Ill: 2153 l'asciius Ill: 1691 /ocius mummilica1ion
ruuculum II: 644,955, 12!4 Fll$C(Q/a h~palit:a lll: 1!>83 caule II: 9..53
rurnen 11: 6-14. 955. 122·1 fasciolosis 2,5 pigs II; 688. 806, 809. 935. 919. 13 l I.
skin. horses IIL 1758 iatal familial insomnia II: 1392 111: 1448
tongue LI: 6-10. ~00. I l96. l:Z23 Feild herpc"-irus I II: 829 ~hccp1goa1s 34-1. II: 990
vag:na II: 6-lO iellnc c:illcMrus II: iOO fog fewr IL: 1191
,·es!cl~s II: l I 96 feline lmmunodefklcncyvinis II: 710. 741. follicular pl1aryngi1is. horses ll: 800. 863
l!l0b1\'C $\Omatlri,of C31Lldl: 1282, 1289-90 l:?97 follltutlm Ill: 1754, 1757·8
Xii 1'.1,Ui:.X

horse~ UI: li'S,, 1917. 1921 gammaherpcs,iruscesl Sl'<' - - palliclipe, ·1·1. 50. 252
sheep Ill: 1i 53 Cia11111whrrf)<'SVir/1111e - - pnl1mlis ·13.-lT
Fom3Jla· \lasson siln,r staining procedure ganghoncurilis 11: !l 13. 1156 - - palf)t1/i5 pn/pnlis 50
111:2111 g~ngrene, dry !II: 1586, 1910 - - sthWf!fZi 50
food poisoning. humans Ill: 1586. 1754 gangrenous obomasitis Ill: 1869. 1670 - - $111)'11/WflOIII 68
foot ab,ccss(csl a 111: 1729. 1730·33. 1755. gangrenous mast id•. caule Ill 1829. 1831 - - rabar1iformis 50
1949, 1953 gangrenous myositi& Ill: 18S6 glo~>)' bro,"1 tick 24 ·G
root rot JI: t213. 1285. 128/l, 1351, 111: 17.14. Ganjam ,~ms II: 1()71. 1072 glycosurla 111: 1848, IS.19. 1850
1735, 19-1$ Garl».i ,ims II: 1043 goat di,uasc 32S
caule Ill: 1,20. 1722 gas gangre11e Ill: 1829-33, 1857 1838. 1863, goat plague see pcste des petits ruminants
pig;; HI: 1743 1869-73. 18i4 Gomori's methcnam,nesilver srnin
sheep/goat, 111: 1729.1733·8, 1755. 19U) Gassenarr ganglion II: 1156 !II: 2098, 2139, 21-1 l
fo<)t rot. stable 111: ! i23 ga,<1ric ulcer Ill: 1-175 goose parv0\1rus ii: 815
str.iwbem· !I: 1284.111: 2026-35 ga,tri11s 111: 2120 Gordil ,irus II: I05 i
fOOt·Md-mouth di.ease 80, 83,225, 22i, gastroenteritis II: l07 I GQrd(Jmn IMin111h11, II: 1021
ll:647,668,700.892,96!. 119'1. ll9t\ gnstroemcrocolitis Ill: 1617 gouslc·kte 524. 573
1213, 1225, 1278. 1285, 128!1, 1292. C,n:elltt rllompsr111ii 619 Graafl'-llclnct disease II: 717, 733-a, ;41,
129i, 1313. 1316.1319, 1324-56, gazelle II: 83-1 744, 1-101
111: 15-18 Ceig,•rla pathysc~li$ Ill! 1588 Gram-m,garh•e bac,erla Ill: 1568
foot-and-mouth disease ,·irus 79, 80, geflsicktc Ill: 11)46 Gram-negative in1eetions l!l: 1655-67
Ill: 1947 gemsbok II: 1270. 1423. Ill: 1690 Gram-positive rod< and cocci Ill: li49· 1963
and /\frlcan butl'alo 11: 1325. 13-10 gencrall:ted lymphadt!nopath) II: i4-I Gro.m·s zebra. 395
and nnderpest ti: 1326 gcn~ralized mhcrculosis llf: 1978 granulation tissue 11: 759
forage poisoning Ill: 1887 gencts II: 1131, 1133. 1135
Fon :-.torgan vims II: JOii gr.mu.ks. sulphur JII: 1648. 19-1-1. 2012
(',e11e11n gt•rieun II: 1133, 1135 granuloma associ:ttcd \,ith ArlinomJ'tt'S or
Fonuna disease 21. 22. 45•1. <160. 472-3 genital campylob,1etcriosi~ in canlc
foul-in-the-foot II: 1351.111: 1722 .Y/X'ardill species Ill 1-56, 1943. 1962.
Ill: l-159·65 210;
founder UI: 1i25 genital carcinoma, humans II: ,5;
foxes382. II: 1126,111: 1-146, 1546, seealso granuloma
genital horse pox 11: 868· 73
imfil1/d11fll ,u111u1t genital Ill: 1543
genilal infection,;
Fra11cuelln wlar;,11.ris 90, Ill: 1520 mamn,illnl')' Ill: 1641¼, 1962
caulc If: 8ii
freiu,· U· l-113. sun/so hypcra~sthesla mycobacterinl Ill: 19,G
pigs Ill: 1776
fruit bat~ ti: 681, 682, 6&8-90. 692. 1126. nasal 111:2110.21 II
geniml 1racrdisease Ill: 1784
1145. see al;;o i1ulillft/11nl 11/lml/$ purdnc. ulccra\ivc Ill: 1~3i
genital ukc.r:ativc dcrmatosis II: l 28i
rruitny 159 pulmonary Ill: 1943
Gl'OCIWIOIII! pnrdalis 516
rurno11bln II: 9$4, 1158 spermatic Ill: 1515. 1529, 1530. 1531
geophagia 269
FUNGASSAY !II: 2096, 2100 splrochaetal !II: 1962
Gcml!ston "1.rus 155 tracheal !II: 1943
rung! Ill: 2095·2)24 Gcrs1m;mn-Strtiussler,Schcinkcr$yndromt:
Fun~~ lmperfucti !II: 2095 tuberculous Ill: 1976
II: 1388. 1389. 1392
rungu, poisoning sPe poisoning. plants/ gramdomarous disease Ill: 2011. 2107
geslgs1·1ic1! 81 ·3
moulds granulommous Inflammation, intl'Stlne
Getah vims II: I 023
funicul!tis UI: 1,56 Ill. 1999
Getah ,1rus infection II: 849. 928, 1023-5
iuriou, rabies II: 1152 granulomatous mastitls 111: 1756. 1942
giam forest hog II: 1094
furocoumarins II: 1351 grnnulom:uous pMumonla 111: 2011
Giardla ,peclcs Ill: 16 L2
furunculosis 111: 175-t. 1757-ll grapes $e/J TB grape,
giasin-gi>hu Ill: 2026-35
horscslll: 1917, 1921 grass mou~c Ill: 14-16
Clrajfr1t'flm~loptmlt1/is II: 12;0
sheep Ill: 1755 grease II: 1298,9, 111: 150-1
giraffe, II: 1270
Fu.S11ri11m species m: 210 I greas}' heel. horses !II: 1757
ginh heh Ill· 2098
F11sar/11m mo11illforme II: 685. 93,1. 1O1o. grea,r pig disease II I: 1759 -61. 2026. 2030
glanifon. !II: 1500· 150~. 1505
1020. 1158. 1370 Great l.irnpopo Transfroruler Purl. 233
as zoon~ls Ill: 1500
- - 11«ropl1orwn 90.111: 1429, 1720. 1729. Greater Yellowstone Ill· 15,1;
glandular re,er. human, II: 86'1
1733. 1738. J;41, I74:1. 1744. 1946, 19~9 (i!a,,scr's dlsea.,~ 111: 1629-32 grt-en Oy Ill: 188S
fuzz\'•lnmb disease ll: 970-7-1 green lrmph nudes ill: 21~3
glomcrulitis II: 750
greyfos II: 1128
glomerulonephrltis
grey mongoose II: I 12~

0 canle II: 950

horse; lit: 1653
pigs II: I 107
grey-headed fruit bat II: 688
grinding or tee1h II: 1400. 1-113
groo1 bontpootbosluis I,
G:lbek Forest ,1n,s II: 1057 Glossiflll specius 43-68
ground squirwl II: 1135
Gaboon adder 111: IIIGO identification 49-52
gntactan Ill: 2047. 2054
gro"1h rcmrdation
key 51 ·52
gafactophoritis. tuberculous JI I: 1980 Glo.,s,n(I a11sum/ 252
cattle 11: 953. 955
pig,;- Ill: 1707, 1708. 1709. ~066
Ga/i,r~llt1 pun:erofoma II: 113.9 - - lll/Stulli (IUS(Mi 50
sheep Ill: 2157. 2158
--;w1g11iru•a II: 1135 - - ,w.~umi 111oss1,rize11$is 50 Guillain-Barrr <rnrlrnmP JI· 1161. 111\.',
galkoors ,-an perde see equine - - lmwqmlp1$ 50. 252
piroplasmn~i,
guttural 1101,ch mi<o:o~i• Ill: 2115
--jiurlpesftm-if}l'S 50
Gwenl\irus96, 155, II: 1252.1253
gallsickncss sp,e bo,•ine ~napla~mo;is --/11.<cip1-s 1111mi11i so
galslekte see bovine anaplasmosis --Ji1sclpt>s q11n11u11sL~ 50
game ranching, buffalo 469. •170 - - morsf1011s •14
Gamil virus 155 - - morsi1a11.1 a11trt1/is 49. 50,252
gamma imerforon test Ill: l003 - - 111t,rsi,a11s morsi1<111s 50. 252
gamma 10:..in m: 1857, 1864, 1870 - - 1111$/ii so H antigen, Ill: 1580
Ga111mal1erpes11iri11oe II: 860. 864. 895,896 - - pulllc~rn IIUW$1Mdl 50 liAOE:'1 virus infcetion 11: 81S- 16
hacm3gglur.lnndng enceph~lomyeHds ,1l'us intrapulmonary Ill; 1742 hcat•lal:iile ento1ro1o~ln rt!: 1565, 1566
Ill: !Sil mucosa 111, 1449 hea1-smhle cmero1oxln Ul: 1561. 1665
l-lne111aphysalis ~pecies 438 subcutaneous 111: l-149 licddlestonc procedure Ill: 16!19
J/111m:apl1y$a/is acimlifcr 500 haemorrhage in pigs he<lgehog<listcmpcr II: G3,I
- - /Jancrofri 500 cnc.clnn lll: 1428 hedgehogs 11 1339
- - bispiuosn '138 colon Ill: 1428 he~l ahscessccsl, sheep/goats Ill: t 730-32
- - C()11cin11n 500 imes1in.-lll: 1475 heel erosion IH: l no. l 724
- - /111merosn 500 skin rt: l 10.1 heel wans Ill· 1725
- - i111crmedia 438 haemorrhage In 5hCcplgoatt. l!elnt. bodie.., 111: 2158
-]11Jxmiea500 gall blnddc.r II: 1073 helm~ti.'d guinea fowl 516
- - longlromls 500 lymph node~ lit: Jl.;99 helminthosis
- o.rop/:i/n 438,618 pulmonal} anery II: 1210 canle 275. II: 951,961. Ill: 1588, 1623
--pWICl<l/(l•l3a, .\98, 500,618 subcutJncous Ill: 1702 ~he~plgoau.2i7, 328.578.111: 1623
//(1(11mtrobia species iT -9 1hroa1 region III: l 699 hemiplegia. lal}ngeal 111: 21 Ill
HaemQio/JiCI irriums e.tigua 77 haemorrhuge, abomas.11 mucos.~ Ill; 159G Hcndra \'ims 692
- - irri,ans irrium.$ -;7 multiple organs 11: 1102 Hcndra virus infection II: 681 -5
- - meridiann 79 • pre.pure If!: 2153 a, zoonosts II: 681. 681
- - spf11igeffl 79 retina lll: 163i. 1639 Hcndra l'irus respirmory s~11drome II: 8-19
- - rli/ro1Lti poumsi9 h~emorrhagic diarrhoea Ill: 1842 l lendra \'ims
I laemarobosr,, species 7r-9 haemorrhagic diathesis ca1s II: 682
Haemarol)osrCI n11g11srifrons 79 canlc II: 1228 frwt hats II: 681
- - latifro11s 79 sheeplgoa1.s U: IOi3.1206. 1210 lie11if)IW/n,, 11: 692
- - UlliSi'titlltl79 haemorrhagic disease H,mif)<WiTII$, 3.S zoOn OSi§ II: 692
- -=ulu<'llsis 79 canlc 26.1. 266,268. 27$.11: 953 hejm1lc mccphnlopalhy
J111ema1opi1111ss11fs575, II: 701, 1098: 1293 humans II: 103i, 1051. l077, 10132 ca11lt' II: 1,117
Hae111atopota Bi pigs IJ: 9i9. 1088. I 102. 1103, l lOi, horse, II: I020
1-laemmopom albi/1irra 90 111: l4i5 htpmlc necrobaclllosis
Nt1emaroxen11s separams 498 sheeplgosLS II: 1073, ru: 1829 ca11letll: 1736·4 l
- 1-elifer11s 483 haemcmhagic disease, vir.il II: t<M8. 11)82 $hCCf)I ~031> Ill: 17•11
haematurill Ill: 1934. 1959 haemorrhagic emeriti~ Ill: 1829-33. 1836, hepatitis In ,·a11le Ill: 1586
Uacmobanu11e/la spt>cie~ 581 1838. 1874 heparills
flocmobarto11~1/a bul'/s581 ha~m,mhagic tmcriu~ rwc:roric 11: 1048, 10i7, Ill· 1595. 18S5·6.
--l;or·ls infection .581-2 e111tlc. Ill: 1840-44. 1875 2089
--feUs5i6 she~plgoaLS 111: 1843, 1a15 pyaemic Ill: 1617
hnemoglobinoemia 111: I 4-48. 14-19. 1867 haemo,rhn.i;ic enterotoxacmia in young f/l.'pmocysris98
hacmoglobmac1ni,, nephros,,. 428 lamb~ Ill; I11-10·4-I hepatocncephalopathr syndrome 5:,9
hacmoglobinuria haemorrhagic fae«>s Ill: 1430, 1~80 hepatomegaly Ill: 2090
caule ,i 13. 414. ·117, 428. Ill: M48, 1832. haemorrhagic fever vims 11: I080. 1082 cattle Ill: l·H9. 1587
1866.186i haemorrhagic gasuoentc.ritis, sheep ti: I071 pigs Ill: 1450
equids -125. 429 haemorrhagic iniectious cmerilis. pigs hepa1omegaly, ~heeplgoais II: 990.111: 1596
pig~ 435 111: 18-10-·14 Hrpn1011inis ti: l 30i
,hceplgoats 438. 442,499. 5i7. [II: 1448, hacmorrhnglc Inflammation, abomn~al Her,,es1es 1wrop1111craws II: 1129
1,i,i9, 1,150, 1831. 1919. 2158 Ill: 1702 l/11rpe;wirldne a21.921. II: 1285
hncmoglobinuria. hadllary Ill : 18(,3. t6(i5. haemorrhagic nccrotlzing cntcrltis, cquids IHrrp~s,irus d.iscase of nc.w·bom fo:ils see
1866-7 111: 18-10-4,1 cquid herpcs.~rus I and./ !nfoc1ions
postparturient 428. 603 haemorrhagic septicaemia 275,544, hcrp1.<~\·lru.<;{.-s) s"e l1rrpes1•/rldn~
hacmc,globlnuric nephrosis 429. 111: Ul67 ur, 1689-92, 1so; I Jerrold's egg-yolk mt>dlum Ill: 2()01
hncmolyslns Ill: 1755 haemorrhagic r.racheltl~, pig,; Ill: 1781, ns2 /lerJ/iin $pecle;; 68
haemolysis 4()6. 428,499. Ill: 1449. 1917, hairy coat in f1>ot-.ind-mouth dlsens,' Il~rr/11 pallf11S ll; I057
l~I~ 11, 134!1 H,,,.,rorlwMl//s 31
l111ra,11~cular4 l3 • .:8 I. Ill: 186i !lairy lambs II: 972. 973 hexachlorophene 11: 1417
ha,•mol~'Si~ and ic1en.ts >}~ldrome In sheep ilair)' pamer:, II: 1:1n. 1344 llighlands J viru, II: I 017. I0!8
Ill: 1450. 191,. 19.23 ,air)' vetch 357 l1iJl/K!bosro mjipe.~ 59,
hoemol}11c1maemia413. .;t4. Ill: 1452. 192.1 ~airy·,hakcr lamb disease II: 970-74 11ip/lOJJOl(ll>lll~<lflt/ihibi11.< Ill; 1512. 154;
haemonchosis 27i, l,21 ,alofuglnone 492 hippcn>oramu~ Ill: 1512, 1547
/laemo111;/11u comortl/$11: 1396 Hammo11d1a hammondi 382 h1ppo1ragine herpesvinis I Ii: 89;
hacmophilos,s Ill: 1634·4 I - - h~;rlomi382 lllpp<JTmgus 11iger4 I0. 598
/lnemoph/111sspec1cs lll: 1531, 166! Hamaan•like ,irus II: 1082 his1ofatcin slm1 wst Ill: 2105
ffal'mophilusspecies iofeet,ons Ill: 1628··l l Hama1•irus 15S. U: 10112 Hisrophih,soi•i.<111: 1531.1634.1655
llnemophi/11sng11illl: 1.531. 1634 Hama,irus Pulmonary Syndrome ti: I082 IIUroplasmn l:Qp.<ulawm Ill: 2119
- - paras11i1 II: i85, 91 I. 983. Ill: 1629. harsslall spon,-siektc Ill: 1859 Jfismplnsmtl ca11.<u/(1111m var. [arc/11w1os11111
1670. l 778 ~arsS13gsie!ae 111: 1921 Ill: 2104
--paramis infcc1ion Ill: 1629·32 ilanwtttcr see henrtwntcr //isropla.•mnft,rc/111i11osum Ill : lS(),I
--sonmu.; 41; u, 1158.111: !531. 163~. ban·est mites UJ: 1759 hlsioplnsmln skin test HI: 21 :!0
1655. !906 ::ta) nuu.~ 11: 1396 hiStopla.smusis farctmlno,i Ill: 21o.;.·
--s,imm.sdi~cnse complc~ In canle Ha)tlic~·~ agiir and broth Ill: 20i9 l1im>plasmos1s Ill : 2119-20
JU: 1634-41 Hazar~ ,in.is II: 1078 hlsto1o~ic cto,1ridia Ill: 186ll. 1870. 1872.
Jlaemoprore,u mercl111iko1•t' 89. 90 head-pressing II: 1019, Ill: 1941l rn;.;
haemopiysl.,; Ill: 1742, 1950 head-tilt Ill· 19-:8 hoary alyssum II: 928, I025
haemorrhage in c::ittle heartw,tter3, 4. 8. 10. m,35i, 417. 477. Hoffmrmia i11.sig11is 102
cndo1hel!u m II f: 1$37 507-28,541.5N, ll: 1000, 1074.1158. hog cholera !10. 11: 777,806.810. gi.i.921.
C)'C Ill: 1637 1213.1417.111: 1639 940. 975-84. LOOS. l09:?. 1105, 1108,
 1109. 1309, 1311. 1383, lll: 1601, 1605 hydronephrosis Ill: 1657. 1959 lso-crylhrolylit neonatal 431
hog cholera ,irus II: 690. 806, 9~6. 960, 9;0 hydropericardhan fi2 l, 541. ll: 1249. lc1011y.~ strl/ttl/S 11: 1138
hog Ou II: 775-7. 9H. 938 111: 1849 idiopathic braln stem neuronal necrosis
Nonwrin spede,328. 525. 11: l 158, Ill: lii98 hydrophobia II: 1123, 1151 a,1d hlp))(lc-dtnp1II sclcro, ~ II: 1-1 Hi
homldium bromide 282 ilydroprnnmil II: 990. 1049. 10:;a idiopathic mouth ulcer II: 1351
hondsdolheid soe rabies hydro'1llp~1lX 111: 2150 !leal symbiont intraccllulari• Ill· 1-169
honey badger II: 1137, Ill; 1975 Hydrornea ~pecies 7i, 81 llci11s Ill: 1469, 1587. 1596. l5!'17
hoot 'break' II: 1208 Hydrm<1m irr/11111,, !ll ill·thrirt 573,576, Ill: 1620. li55. 2157·8, see
hoof slough I!: I I96. 1228 hrdmthora~52t. 511.11: 1237 also poorthrlft
Hord,mm mlgare 111: 1950 hyena 111: 1975 lmicola complex 95
horingvlicc ,, -9 hygroma.carpal 111 1442, 1514. 1516. 15:?I immunit) t0
hori1.omal flssure lll: 1727 l~rlocflQ<•n,s meiuen:duige,,I 11: 1094 African S1\,1W fever II: 1103-1
horn flies 77-9 h,p~raemia bo,ine herpesvirus l II: 8;11·80
Homer's ,;}ndrome UI: 2116 cattl~. coronary band~ II: 122ft lbo1-and,mouth disease 11 1343-4
hot:.t• fli115 87. Lil: 1798 pig,. skin II: 979 imnn1ni1....1ti<.>n
horse nu 430, 11: 766-72, 928. 1025, 1320, ,he~p, c;(,n:inal) b,mds n, 120i bluetonguc II: 1213-15
lll: l 768 she~p. mouth/nose 11: 120, bovin~ anupla~mr>,i• 603-5
horsepox rt: 1298-9.111: 150~ h}pcrn~lhesia II: 1310, 1-l l:'I. Ill: 1631, nuclei. acid 242-3
as zoonos!s II: 1298 163i. 1880 passlv~244
hOl'$l'>hOe hare II: 757 hyp1.«glyc:1emi11 111: I 8-18 rabil/; II: 1159.1;.1
hosepipe g1.n Ill: 14i2 bypNk~rruosis JI: 9S3 immunodcliclency Ill· 197,, 2016. 2ll 1.
hou,e me. n. 80, 81-3, II: 1195.111: 1487, ilyperp/asln lntertligimlls Ill: 17:!~ 2114.2119.2137 21~0
1,98 hyperplasia immunodelkien~ virus iaic~tion human
hounongJII: 16-18-52, (;56 interd!gital LIi: 172 1.1724. 1;25
:1.1. 3-1,1.111: :?016
Howell-Jolly bod le, 601. 620 Kupff,·r cells 11: 750 immunoglobin hindmi; protein, Ill: 1636
huisvliec 81-3 lmmunosupprcsslon :L12, II: ,5,r
lymph 11ode 456. 4:\7. -178, 480. -185. 489.
human adenO\o!ru, II: 821 canle II: 951
-199. 500. a-11. II: 712. 955. 9;9, lll: 1699
human bronchiolo,alveolor carcmoma enxootic bo,·inc lcuko,rs 11:, 10
lymphoid 544. Ill: 1596
11: 728 lyn,phoid. kidncy ,157, , 90, .W9. ll: 90?
sheer II: 725
human carcinogens fl: 757 immunoto!ero.nce II: 950
lymphoid.spll'Cn II: 712
human corona,,rus OC43 U: 791. 795 irnpala353. ll:i'5R. 12,0. 1339.111: 1:;12.
muco,al Ill: 1596
human endogenous rctrovinist1:sl II: 721 15-t7
hyphaema 111: 1637
human granulocytic ehrllchiosls 5'16, 536, impetigo W: li5-t
hyphomycetcs lll:2110
a45 impOlcOl'}' lll: 2149
hypllomy·cosis dt'~tru~lb equi Ill: 2113-1-1
human herpes1<irus(es: lf:88i i111poum1l11 coe1111dl lll: I i,13
hypocnk11cmi:t II: l 185. 1191
human Immunodeficient')' \'iru< II: 710. '33. inclusion borlie<, set! 1111der lnclu,ions
hypomagnesaemia 11: 1-101.1417
; -17. 751. l29i inclusion hody rhihitl•. pigs II: 919-21,
hyp•>myelogcnesis 11: ~81
human meas!~< vims 11: 687 lll: 1494, 17 JO
hypophy,;cal at>sccs,s1rndromc UI: 194'.",
human mump~ 1irus 11: 68i indu~ions
see also 1111dt1r ab,ces,;(l'Sl
human papilloma,ims(es) 11, iS7 hepatlr.11: 11-16
hypoplasia. cerebellar
human pan'D,im~ B19 U: 807 intracytopl~smlc 11: 646, 6i7. 690, i77,
caulc II: 953. l:?52. 1253
human T-ccll leukocmia vims II: 708 l 123, 115-1, 1269. 1293. 13i0
pig, 11: 981
Humpt)' Ooo1 iru~ 100 intra()topla~mii;. eoslnophlllr II: 664.
shllt:plgoais II: 9i2. 990
/iy(lc11a bnmne(I n, 1137 6i8. 1071. 1273, 12i8. l2i9. 1294
hypopt~ia, pulmonarJ
hyalinc fungi nt: 2110 lna1111uclcar II: 6'16, 664. 6'."4, 690. Ti,,
cattle II: 953
hyalohyphomycosis Ill: 2107. 2111 80i.809,819·23.844•7 861,872.887,
pigs II: 690. 981
Hyolomma species 452. 41'16, s:i1. Ila lOiS 1192.920,990. 1055. 12,4
hypopla,10
Hyalommn. cumel 17-18, II: 1300 in1r.1n11clear. acidophilic IJ:Gi'.;. 913
thyrnic, caule II: 953
/iyalumma. shiny Ii intranuclear. basophilic II: 819. 919
mrhino1e. pigr, Ill: 1-192, 1491
Hyalomma-rclated ehrllchi;ol infection 5-1-2 intranuc!car. ccmnophilic II: 8·16·7 868.
h)vopyon II: 900. 903, Ill: 1488. I63i
Hyaiomma "'-'!O'Pli11111 436 891. 1os:-1. 13;0
hypo1hrichosis 11: 953
- - anamllr:11111 45-l, 487, II: 996 mcoordination see,ua,'da
hyra.~ 3i8, II: 11:ii
- - (marol/cum ,maroiicum 498. 540 Indian elephant II: 757. ITI: 1690
- - m,arolicwn 11.t'c<watwu •126 indil'l'Cl enzyme-link~d immuno,orbem
- -1/Nrlrum 3. 17. ~26, 487
- - dromedarii 3. 17- 18. 45•1. ·187 II: 1235,
1300
0 assa)' 111: 1519
in(arcUs) in cattle Ill: 1637
e<>ntml nervous ,!y<tL'tn Ill. 1638
- - e.,·cmtt111m1487. 540 lbnrakl disease II: 1185, 1191, 121:l, 1221·5 hcpntlc lll: 1639, 2052
- - impresmm 454 1229 myocnrdinJ Ill: 16.'19
- - margina111m /s(l(1ci 540. 5'12 lbaraki \llru~ Tl: 1227 pulmonary 1TI: 20SI
- - marginawm n1jipN3, 17,407,540, ib11x .;:;3, Ill: JMG, 1549 infarct(s) in equlds, mu~cular Ill: 1,72
596. II: l 079 lrtenb II: 1057. Ill: 14•18, 1829 inforct(sl In pigs. splcnlc 11: .979
- - nwrginat11111 1trra11it11111 II: 1079 ictcru~ infarcl(s). kldnC?)"l5i. -199. Ill: 1639. 2052
--p/11111!1e1m1618 caulc .; 13. ~56, 480. -189. 577. 600. 11: l050. infect!on(sl
- rnmca111m 3, 15· t6. 407,427.542, Tli: 1448. 1449, 1861>. 186i amoebic31G·li
II: 1043, 1078 equlds 425. 429.11: 1024. !'.NII. m: 1449. feet Ill: li20·29. 19~6
Hy/Jumirm 87 2090. 2091 gnsiroh11<,.~tl11al rract 111: 2120
hydranencephaly pigs-135 genital Ill: 17iG. 1946
cattle II: 953. 1029. 1252. 1253 shccpigoau 436. ·1-12, 576. 620, 11: 990, guttural pouch Ill: 1769. 1772
pigs II: 690 Ill: 1449. 1450, !NO, 1831. 1832. l86i'. i111ra-111ennc 42i,429. 439
sheep/goats II: 9i2, 990, IOW. 1049. 1050 1917. 1919, 1921. 2158 resp,ramry tract Ill: 1319. 1777. 2067.
Hydroch<H!rus hydrocl,oerls II: 1339 rctews. enzootlc ll: 987 2116
 slon 111: 210i 1723-9. 1949. Se<!1tlsodcrmalh1s Jenner 11: 1296
strc;micoccal. hc>rses II: 928. Ill: 161 2 imerdigital necrobacillosis Ill: 1720. 1,22 Joest-Oegcn bodies II: 1370
subcuum~ous Ill: 2107. 21 JO intcrdigital phlcgn1or1 rn· li22 Johnc <e sickle TI!: 1994-2004
udder Tll: 1!1;9 lnterferon-gamm.i as,;.,>· Ill: 1985 Johne·$ d£se.ase Ill: 1994-200-1
umbilical m· 1;41 in1crmediate foot rn1 Ill: I i34. 1733 joint damage. pigs !fl: 1909
urinary tract 111: 1931 ..J. internal mycosls Ill: 2114·24 joint ill Ill: 1612, li5fi
\'Cnerenl. opponunis1ic Ill: 1556 int,•rstltial emph~s<.'ma II: 678 joim,. ,woll~n 11: i36, 1105. 1189.111: l·Ml,
wound~ 111: 1874. 19-16 lmer,tltlal nephritis Ill: 1449. 1-150, 1570, I r78, 1951. 2071
infec1io11$ bo,ine cenico•\'aginiti~ ~nd 1959 jowl abs-ces, Ill: 1781-3
cp!did)<Jnitis Ill· 2149·51 int<!rsthial pneumonia JII: 21:17 ju,•enllel)mphoma II: i08. 713
infornou• bo,·ine k~ramconjunc1h-11ls imemilial pneurnoma Juvenile rheurnot0id ,,nhritis, huntar.
111: 1-187•90 c:mk'/shecp 556. II: f,t\4, 674. 725. 729. UI: 1-140
infcc1ious bo,1ne rhinolracheitis 11: 647, ;33_ 738. 799. 822. Ill: 1S95. 1639. !681
875-83. 90-1, 951. 960, 1225. 1351.

0
goat5 II:;41. , .13
Ill: 1488. 1-189. l:>83. 1639. 2055 horses 11: 683, 679. Ill\~
lnfec1:ous cnninl.' hepatitis 280. II: 11 58 pig, II: i85. 937, 1106. 1107
infoctJous lnfcnlllly 1~6•1. 2J 49-5 L hnestinal anthra)( Ill: 1475. tao~ K antigens m: 1560
infectious kem1oconjuncrivilis 83 Intestinal aspergiUo,ls Ill: 2116
Infectious mononuclcosi~. humans II: 86<1 Kaalplaas vlrus 155
intestinal clostridlosis 587 kaa,agtigl" lfmfadenitl, Ill: 191i·25
infecuous 01:crotlc hcpacills Ill: lll6S-6 1ntc,tinal haemorrhagic S)11dromc
infectious ophthalmia Kaeishi \irus II: 12.21
Pi!!S Ill: 1-175 Kago,hima ,;m, II; l2ii2
ca11le Ill: 1~87 sheep 111: 182!1
sheep/goats Ill: 148i. 1489 kairomolles 44
intestinal torsion m: J.175 !\ala a1.ar 379
infeciious periodonthls of calve~ Ill: 195 l imracumncous te,t Ill: 1807
infectious pustular balanopos1hhls/ kalwerdifterie Ill: l i4 I
lntrn~1Qpla,mic inclusion bodi(•< sl/e Knmo p:tral)"sis 4. 111
\'\ll\'ovaginhis, ca1de see lnfec1iou&
mzderinclusion~ Karoo paralysis tick 18
bo,ine rhinotrachciri, intrndermal wberculin test Ill: 1974, 1982.
infectious thromboembolic Karon-~erl11n1111ingsbo~luis 18
19Bl:l Kar,hi <'iru~ JI: 995
meningoencephalitis, cattle Ill: 1634
tntranas~l tumours II: i2 I karyorrhe~is. l)<Jnphc>id 1Jssue II: l 107, 1110
infec1i\'e bulbar necrasis Ill: 1i30·32
imranuclear Inclusion bodies see under
infcnilit) lit: l~-18. 1-159. 1-161. 20i8, seen/so f.:11sba \1n,s 98, JOO, IOI. II: 1029. 125?
inclu;!ons
s1erili1y k,na Stt p..stc! de, petits nuninunlS
1ntr3· uterine ,nrecuon
caulc U: 816. 1183. l !SS. Ill: 14•18, 1,152, Kelen.icdne st min II: 1160
equid~ 4-27. -129
1459. 1462. 1510, 1636.1640.2076. keloids 111: 1725
shee1>/gQatS 439
2076.:?149 l.cnnel cough, dogs Ill: 1492
mtr.wascular co•gulation. disseminMed s,e
cquids Ill: I 5:;5, 208..J kcra~nis 31,, II: 903. Ill: 2ll.l
di~emmated lntr~,·a:scular congu!atlon
pig, II: iii, 809. 14•18. 15-13, see also cattle 11:953, 111: 1487, 1488
.ntr:wascular h3emolysis 413. •181, 111: 1867
porcine reprnduccive nnd rcspir:uoi, equids 299. Ill: J.1~9. 21 Ol
,odism In: 2104
syndrome purulent Ill: H88
ridocycll:is ll: 903. Ill: 1449, 1-188
shctp/goats II: 970, Ill: 1530, I537 small rurnin~nts Ill: 2046
1ridol'irido.e 11, 1089
lnfcriilit;·. epi1.001k Ill: 2149-51 kcratoconjunctM1ls Ill: 1904
,rritabilir:y. cattle Ill: Jf,37
lnferilllty, lnf<'ctiou, JII: 146-1. 214!1·51 canlc II: 820. Ill: 1487, 1905
.~o-ery1hrol\1ic neonatal icterus ·131
lnflnmmation, see also oedema: swelling hor.aes 559. II: 86Cl, llf,~. Ill: H87
sc,.Jmmune erythrol)'i.is 431
abomasum Ill: li02 infcttiOllS83, Ill: J.16;"•90
:som~tamidlum 281, 282
accesso11 sex glands Ill: 1510. 1529 ,sonfcot!nic·hyclrwJde Ill: 1986
$l1C1cp Ill: 1487
colonicmuco.n m, 2019 ker:uoconus IU; 1486
»oquinolinealkaloids II: 1058
diphtherofd II: 979 1-eratolym Iii: 2033
/sospo,a~pecl~,3 19-27. 111: 1571
gr;inulomatous 111: 1999. 20)9 kera1oma plantar,:, ~ukatum LIi: 2033
rxodes pacifirns Ill: J.14 l
hendlmouth rnft 1iss11!?l> Ill: 1648 keratomrco,b Ill: 2101
- - fX!/'111/c:nm., Ill· "141
lntesune Ill: 1999 !,,cternh virus l!IO
-riclm,s644,618.ll:996. Ill : 1440.1754
joints Ill: 1543 kctosb II: 1.; Ii
- nibic1111d11.•3. 18
l~mph node, Ill: 2019 Khasan virus II: l 078
-scapul<ufsSl2. 596.111: I-MO
l)<Jnphalicsm: 1502 kidney
i~adid ticks 3. II: 996. 1253 abscessat!on 111, 16S3. L65i. 1742
mandible Ill: 1(150
lxodiplmg«s hOQi·eri 31
penis Ill: 2153 ulnckish lit: Jil50
- thei/erae 3 I gl'e)-i~h-whi1e fot·1 II: 902
pvogranulomatous Ill: 1754
re..piratory u·nc1 Ill: 1768 ,niarcts 45i. 499. 111: 1639
IOnl(Ue Ill: I 648
lnOanunatOJ'}'Oedema Ill: 187S
in0ucnza90
0 petechlation Ill: 160~
white-sponcd Ill: 1370
kJkU)11 Jll: 1734
lnllucnz,a A/equine- I/ Pragu<'/56 II: 766 ina~iektcll: 717-30. 733,738, 1397, 1402 kimata Ill: 1788· J 811, sere also :unhru
inOuen.ta Alequine-2/).liaml/63 ii: 700 jaag~ickre reiro,·lru< ii: il7. T.W Kfrnberle)' virus 99, II: 118-1. 1191
Influenza ,-\rus(e~l II: ,Sa iad,111, 111: 14-,6. ~enl.~o illdil4dt1lll 11llt11//$ Kindia \iru< II: 1253
lm~rspectes rea,soriment 11· T.'6 1ackrabb1t II: 757 klrch11ll:l026-3S
lnOucnr.a lnmcsiown Canyon sero1ype 90 Kle//$lelln ~peci~>S lit: I 555, 1.560, 1612
equine 130. 11· iGG·rZ, 928. 1025, 1320, lnnuai,• disease 21, 22,454.468. 472.3 /..7ebsicl/a species infection, 111: 1555-8.
UI: 1768 Japanese encephalitis 137.11: (ili5. 1252 1612
porcine JJ: 7,:;.,.
914. 938 lapan~e enc-ephalftis ,irus 154, 162. II: 690. a< 1.oono~is 11 t I555
lnlluenm-like Ulntss. humans 11: 1056, 1077 810. 1004 /..1ebsi~lla o:ryt<X"tl Ill: 1555. 173g
lngwal'uma \'lrus 155, fl: 1031 jl'junitis.111: I S87 -p/11111/co/11 Ill: 1555
intercellul.ir organisms Ill: 146\1 lcmbrnna disease II: G-li --1me11mo11lc1 II: 1250, 111, 155j
lntcrdigi1al dermatitis Ill: 1442. 1720, l i21. hma ,irus II: 703 - - t~rrlg,ma Ill: 15,;5
ldem-in-die-kaakll: 685. 1010. 1158, Lassa fever II: 1082 hepatic II: 1051. IH: 1866, 2015, 20:?0
m: 18,8-83 Intent:) ltistoJ>ath<>logical II: 11 O,
klontwol 111: 2026-35 \IH\ • 1 infection II: 897 intestinal mucosa rll: 1596
knopkaak Ill: 1942-4 bo,~ne hcrpe,;vini, I infection U: 878 lal)'ll' II: 1223
Jmop,·elslolae see lurnpy skin disease bo,ine hcrpesvin,s 2 infection II: 889 lipll: 1194. 1289.12911
koala 551.553 equi<l herpesvirus I ;ind-I infeciions lip. shcc11Ill: 16-!8
Kobus deft1.<Stt 453 lf: 835 l~'ll1ph nodes Ill: 1988. 2015
- - Plllp.<lpt')?IIIIU$ defttssa 169 equid herp~~,irus 3 lnl'ection II: 869 mandible Ill: 1943
- - ellipsip,;•,111111$ II: 798. Ill: I S.lli pseudernble, II: 91 I ma:.:illn 111: l!J,13
--/ec/J~ Ill: 1975 suid hcrpes"lrus 2 infocuon II: 919 mnu1h II: 1194. 1289. 1298, :3-1.5, Ill: 17•12.
Koch's body 4SO late-onset mucosal disease. cattle II: 951 194.1, 202!!
Koch~ po,aulatos HI: 171!8 Lnwsoni,; mmue/111/nris Ill: I-IG9, 160S mulciple si1es Ill: 2032
koksidio,;,c 319-29, see also coccidio<i• lead poisoning see 1111der poison in~ myocardial, horses II: 1235
krimpsiek1c Ill: 1897 leafhopper II: 1195 necrngronuma1ulous Ill: 1923
kriptosporldio,c 332-5, see 11/so Leanycr ,irus t 00 nostrils II: 1289, 1298
cryptclsporidiosis l.ebombo virus 1f,5 ocular. dogs 2i9
kudu II: 6-10. I 132, 1339. 1423, Ill: 19;.; le,:h\\'C Ill: 19i5 ocular, humnns If: 1048. 1051
Kunjln ,·!ru, II: I00,1 Leishmm1i11 species 378-81 oesophagus II: 1223
kunkers UI: 21 H u,ish111t111/tt i11fa11111m 378 oronnsal ti: 1289. 1298.111: 1,.;2
Kupffor cells IU: 1596 - - 11111jor 378 o,iduct< Ill: 20i6
kuru II: 1388, 1389, 1391 - - lrt//JiC/1378 p1•nis Ill: 2153
~·wni$! Ill: I ,83· 1811, sec also amhra.~ lci~hmaniosJs 378-8 I pharynx II: 1223, Ill: t ,42
Kyalami \'irus 155 L~ly,tad ,·iru~ s~e porcine rcproducih e and placcnm Ill: 1906
Kyasanur Forest disellse U: 995 respir.nory >)'lldromc ,·int$ pneum:uic 111: I923
kyphos,s. pig$ II: 6S8 lt>nmrs II: 1-123 pneumonic Ill: 2067
lcng.ua a:t.ul see bluet.onguu IJTCJ)UCC II: 1194

G lcntie11hircnuclea1ion Ill: 1488

1.J.,11tl1•/rimw 11: iii, 733
lemiv,ru~ II: 717. i18, 733, i41
puh110113ri• Ill: 1503, 2049. 2050
p~1,granunuuulous Ill: !923
renal Ill: 1959
la bouhitc 11: ,17. ;33.9, 741, iH. 1-lOl lemi,iru, infections ti: m. ;29, i-13. ;4; respiratory 1mc1 Ill: l 712
la cowdriosc see heanwmcr Looffier"~ egg medium Ill: 2027 skin 490, Ill: 2034. 2098. 2100. 2104. 2105
la pcstc bovine sec rinderpcsl leopard 111: 1975 teats II: 119•1, 1291. 1:!48. 1348
la Pied:id :l.llchoacan pnramyxovlrus It· 690 lcopaJ:d tortoise 516 teat~. " ·mer buffalo II: 1302
fa 1rcmblante see scrapie Lepidoptero, cye-frcquentitt)I lit: 1488 teats/udder, cimle It: 889. ! 302
fncerai:ions. rec10-,'8grnol UI: !556 leprosy 111: 1996. 1997 tongue Tl: 1194, 1223
fochl')mation, increased II: 820. 900, 1207. lep1omc11insf1is. sheep Ill: 16'lo tongue. cattle Ill: 1648
1228.111: 1449. 1-187, 1488, 1680 l.epros1,im bac1elin Ill: 1464 turbinuu,. pigs Ill: 1709
lacrosse soro1ypc go Lcp1ospirt1 scrovars II: 810. Ill: 14-lii 1-1-16. udder It: 12!! 1
lnc1ic debydroitenase-elevati ng virus II: 92~. 1588 upper body Ill: 2113
933 t.,,,pzospira ~peci~s 11: 849. 921. III: 1445 utenis Ill: 2076
lnevt'ldse brwnoorbosluis 2 I-? up1ospirn borgp~ierstmii 111: 1516 ,'llcuolar. in rabies It: 1156
IAgopus species 11: 997 - Ct'Ul/coftt ill: 1-1-17 ,-~~lcular II: 1194. lll: 21'19
I.Ages bat \'ifll• II: 1124, t 130. 11:11. 1145. - - cope11/znge11illl: 1•146 le<scr house 0)' 77, 80
1151 -gripf>0rypho.,n 111: 14,16 lethal t0xin Ilic 1801
1.t1go11irus II: ;oo. i03 - - lznrdjo Ill: 14~5. 1416, 144, l.ersltele vin1s 99
lambdy<entery327. Ill: 1597, J8S6-9 - - /cremlzarmorrltagiae Ill: 1445. 1446 Le11<oc:,·to::A>011 c<111//eryf 98
Lambrecht ,;ru, II: 103 t - - lnzl!rrogrms Ill: IJ,15 l~ul.nernla ti: 711
lamellarsuppuration 111: 1733 - - 111/11/ Il l: 1446. 1447 leukoc\'tC•nssoci3ted viraemla II: 838
lamenes,ll!: 1441. U42'. 1951 - - /JOIII0/111 Iii; 1445. 14-16. 144; !cukociicephalomalacia II: 68S. 1Dl0, 1020,
!amen~ - - /(lrtlS.<Ol'i Ill: 1446 1158. 1:1;0, 111: 18-19
cattle II: ll9G.lll: I-Hl.1637.1720.1723 Lcp1o~pirn~ea1• see /..ep/05pfl'tt specie,, lcukoencephalomyditi, II· -4l. 744
hor~es II: 1196. Ill: 1441 !ep10,piroslt< 41 i, 436. 603, II: 914. S38. 97'3. leukopenla. pig,, II: 9i,
plgi Ill: 1543. 161)1, 20il , 207,J, 2075 105;, 1254.111: l•M2. 1445-53 lcuknsis. ,pomdic 1>0,1ne II· 708, 713
shccp/goms II: 743.111: 1505, l72ll. 1731. I.opus amcrica(ll1S 553 lcukotoxin Ill: 1677
li33. 1;35, 1736. 175,1, 1912 --mpe11sis !11: 15'12. 1546 Levimhal ngar Ill: 1629. 1r,.,1. 1661
laminlus ,1~8.585.111: 1441. l-112 - - $(U:iti/i~~ 516 libido loss. stallions ll: 843
laminiris lesions see a/.~o 1lecrosls: erosions licei>:'5.11: 12ll3. Ill: t,58
caulell:954.1191. IH: li':?l. 172-1.1725 lesions 1ighmlngs1rikc Ill: 1807
horses Ill: 1610 cattle II: 933. 955, 1271. Ill: 2030 U11og11nrlw.t ,,1111/i575
lnmslekte Ill: 1885-99 check. sheep Ill: 1&18 lion 11; H23. Ill: 1975
lamstreptoclde Ill: 2_03.; coronru)· band II: 1194 lip and leg ulc<ermion II: 1285. 1287-8
I.angst II: 995 erosh·c II: J 194. 12M lipofuscino..sl~ II: 1413
I.Ingeberg ,m1<155 ~xm:mttlc, Ill: 2107 lipooligosocchurldc. hca1-s1abh, tn: 1GJ6
Langerhans' giant cells Ill: 19•1:i. 1977, 1997. i<?<?1 II: I 194. 1:w;, 1346 l1popol)-S.1cchllrldc Ill: 1568, 1585, 159,,
1999,.21-11 genimlia. raule II: 877. Ill: 2070-80. 2149. 1603. 1679
Lam1111n ,amara 603 2151 L,iteria f111111ovii Ill: 1904
l3J111gi1i, ~nltalia, horw< 11: 869-73 - - mo110..·yrogmc;90, 241. 2~2. 3,-.
canle 111; 1638, 1639 genitalia. 11ig, Ill: 1543 II: Ml7, Ill: 1489. 1513, !f?l. 191)-1
necrotk 111: I i-1 l genitalia. sheep Ill: 1529, 2153 listerin~ls4;7, 11: I.JOI . M16, J:I: 1639,
ulcerative Ill: 174 l gr.anulomatou> 3 17 1904.-6
Lm/onrct~ris 11rx:1lllagans JI: 1128 gum. 11: I Hl4 little red fn,!t ba\ 11: 688
l.nsiosp,mn11m bipi111w11tm II: 10.~7 ht:ad Ill: 2096 lh-er ~b~<:<'SS complex. caule JU: I;'"36-41
li\'cr Ouke!s) Ill: 1865, 186G. 1867 myco1ic Ill; 2120 mallgnom 01.>dcmo Ill: 1829-33. 1869·73.
llvcs1ook piss 111: 1 ,Bl. 19a;. 2020 1874. 1876
comrolofdlsenses 171,217 pyogr:inulomatous Ill: 2012 eanlc Ill: 1863. 1875
mc~suring Impacts 01 diseasc>s 189·206 sheep/go;ii; Ill: 1595, 1917 maJJgnant pustule Ill: 1788-1811, seenlso
roleinAirfcnn societies 182·3 l)'mphad~nop:11hy II: i4·1 anihrax
1,ildlife imerfacc 225-35 lymphadenopaihr maligna111 thcilcrio~ls 493;9_se/1 also 1111der
li\'CMCCk dl~eastlS call!.: II· 12;3 thelleriosis
clas,ilkrulon 183-4 ~'quid, II: 750 malignant transtormatfon !I: ;59
conrrol I 71-:!i; pig.I fl: 1383 mallein te.<1 Ill: 1500. 1503. 150-I
estimating cos,~ 211 ~mphangiectasr,. caulc Ill: 2051 I lnlleom_vces ma/lei see 811rl:/11Jltler/n ma/M
geowaphical infonnation 198-9 ~mphangilis --p:<l!udoma/1,!l see 1Jurkl11Jlderia
impact 18.~-9 caltle II· 1271 Ul: 2120 µs,mdomfl/11•/
impaet on cropse1c. 186 cpizootic Ill: 1504. 2104·7 Malia fever Ill: 1535
impact on cnvironmem 186, 205·6 equlds Ill: 191 i. 2015. 20 19, 2103. 2104 mamml!Uris. cn11lc see bo,~nc herpes,~rus 2
i111p;,c1 on household income HIS pigs Ill: 2120 infection
1mpac1 on human welfare· 187. 202-S ~·mpho Ill: 1917-:!5 mangc35i. ll: 1280.111: lioS. 1,59
Impact on lives1ock producti\ity 185 l)'mphocyu,• 11ansformation test Ill: :?003 .\lannheimia !Past,mrtlla} species 11: i99.
impact on market ncc,-ss 1e; l>'TllPh~ IO>U, II: ;os 800
lmpacc on !he poor 188 l)'Tllphold mmouns II: 708 .,rn,m/1~/min (P1wc11rel/nJ species
Indirect impac1 198 lymphoma JI: i 12 infections 111: l 672-17 13
measuring Impacts 189-206 ~mphoma .\It1m1ltt'imin /Ptis1e11rulln) l,nemofrri.:a 241
ri~k a,,,essmcm 202 calf JI: 708. 713 ll:6i5,821.877,951.952. Ill: i635.
risk managcmcm 135 ju,·emle II: ,08. 713 1a,s, 1639, uMo. 1&,,. 1s9s.
sur,eillance 233·5 ~kin II: ;oil. 713 1697-1702, 17().l
livestock production S)'$lC111s 179-82 thymic 11: ;os, 713 - - glu<"otid11/ Ill: 16!15
commerd:il 179 ~mpho<arcoma 11: iOl\. ;12. ;13 .~1a11to11in nfricam1 II: 10•13
lradltional 1i9·82 dogs3~-1 ~l.tnlO\L~ '.ikin le<il Ill: 2 106
llama II: 834 marbling Ill. 2051
/.y(J(!rOSi/1 25-l
LQf1 /0,1 90 \ larburg disease II: 1082
~·ssa bodies II: 1155 marine mammnl brucello,i~ Ill: 1549
loblollprec II: l 020
lys;a\inJ~ genotypes II: l 12-1. 1146
lockjaw II: 685. 1010. 1158. Ill: 1878-83 mtlrinc mammal, 111: 15·16. 1549
Lyssm1m.., II: 1124
locomotion d1s1urbanccs. pig, II: 979 ~ larondera \irus 155, II; I253
1.oefjlcrolln pseudoma/lci s~e Burk/10/dma Marro.kal virus l 00, JI: 1253
µ!t!11domn//e, mnscules, shecptgoais II: 1284
f.,o/i11m species U: 1000 ~la.son-Ptizcr monkey virus JI: il8, i22
longsiektc 111· 2043·5, .\/ns1a1le1101•iras II; 8 19
louping ill U: 995-1000. 139; .\f(JC(Jcn mulatra 39:i mas11us Ill: 1560, 1570. 1756, 190... 1~6.
as zoonosis 11: 995. 999 macaque monkey ti: 1-123 1!147, 2123
louping ill vaccine IT: 1391 MacConkeragar UI: 1560. 1671. 1617. 16,8, closuidial Ill: 1830
louping i11,in1s 15-1 11: 995 li.5.9 gangrenous Ill: 1829. 1831
Lounbgcll: 1006 macrophages. ;trc.iming Ill: I;05 granulomawus Ill: 1756. 1942
lousiness35; macroschizoms -150 myc01ic Ill: 2123
low conception rn1c$ se,. inf~r1ili1y maculcq necrognmuhimntous Ill: 1962
U)wcnstcin-Jenscn medium UI: 2109 pig,11; 1293 ~taphylococc.il Ill ; I ,56
Lowveld bro,,,, ear lick21·2 shrcp1goa1s II: 1278 tubcrculou, Ill: 1974, 1980. 1985
L.oxodomn africmw I!: 123•1 tear. caule IT: 1296 ma<rit:, in t"alt.l e JI: 951. 1273. 13--18.
l.udli1tet1esnr lll: 1885 mad cow du;ease II: 1408 Ill: l.J.J8. 1479. M80. 1515, 1555.1556.
lumpy ja\\ Ill: 1650. 1942-4 madurn ioor 111, 2107 1557. 15;(1, 1587, l634. 1(;38. 1744.
lumpy skin disease 80. 83. 35;, II: 891. 892, \Jn(lure/1(1 myretomnt/s Ill: 210; 17:;4. 1762, 1781.1923, 194i. 1985,
1160. 1268· i5. 127,. 1292, 1351. maduromyco,1s Ill: i 107 201 !. Wli. 21 Ul, 21-IU. 2141. :!142
HI: 2029 ma.edi-\'lsna II: ;1;, 733-8. i4l. ;.14, 1,101 ~ocinred wilh nocardio~is Ill: 2012
lump)' skin disease ,iru, II: 12138. Ill: 2030 mattdl-vlsnn 1irns II: il8. 734. i~ I. i47 pr,1tothecoric Ill; 2140. 2141, 2142
lumpi wool 111: 2026-35 \laln Drain ,•Jrus 100 purulenr Ill: 1'180. 1917
lung oedema see oedema. puhnonal) ma! de Caderas 2S--I. II· l 129 mastltis in equlds 111: 15Si. 1558. 1768.
Lungenseucht derTlere Ill: 2045·57 ma! rubrn II I: 1908-12 1917. 1921
lungcrs Ill· 204 7 mol31se and poor p~rformance II; 842. 860, purulent !11: 1769
lungs 863. 864 masrius in pigs Ill : 15'12. 1555. l.5l7. l.570,
intemiual emph)'sema 11: 6711 rnaloisc. horses II: IM2. UGO. BG3. 86<1 1942
syncytia U: 67-1 malaria 153 mastiliS in sheep/goats 11: 73!l. ;3;, 741.
lungsicknes, Ill. 2045.5; ~1alforma1ion ;.;3, 128-1.111: 16SS. 1657. 1702. 1829.
lungworms 11: 701. 729 pig, II: 688. 9i9 1831 192 I. 20112046
l.unyo 1inis II: 1039 sht•ep JI; 1207 .\[(1$/0ITl.fSSpeCics Iii: 1446
J,11ttomyia spccles•H8. II : 1195 mnlignam carbunclt: Ill: 1788-11111, seen/so .\/mr1t·1U'i1t 11lgcllifoli1t poisoning, see uniter
Lyrno,(ptCIIIS 378. II: I 130 anthrax poisoning
Lrme horrelio~!~ 587. Il l: 1440-43 malignuni catarrhal fo,er 8-'I. 225. 227. 524. ~lnyaro ,int, II : 1023
L)•mcdisea1¼'587. IU; 1440·43 II: 712. 895-905, 1351 ma)11ies 58•1
lymph node cnlurgcmem Ill: I 773. 2 f.13. see cartle-159. ii: 647, 9GO. I 19;, 1225. 1229, ml.'3sles virus II: 634,660
nl$o w,du ocdema: swelling Ill: 1469 meat and bone meal 11: 1409
lymphadenitis (aU$Cd by AJHV-1 II, 89:'·8 med!0,1inal onthrn.x Ill: 1803
c~us.,heeplgoats Ill: 1505, 1917-25 cacsed by o,·H\'-2 II: 898-9 ~lodi!erran~an r11,1>1 fc"er I 7, 18. ·186-93
,·anle Ill: 2(ll2. ?020 deer II· 899 medium(s) sec, 1111dcr i11tli11idual 11nmes
horses Ill: 1768, 192I.2015,2018, 2019 pigs II: 899 Medlar bodi~s 111: 2109
melaena 11: 711 minute virus of mice II: 806 murine hepatitis virus II: W5. 804
Mvlanoco11ion species IT: 1017. 1019 miosis Ill: 1449 ;111urray Valley encephalitis 137
Me/es melts Ill : 19;3 Mitchell Rlver,irus II: 1248 ~Jurrny Valle)' cncephallris ,•ir~s II: 100,1
•\1el/Cf1 tlecumt,e,,s 11: 1417 mLxcd fonu of horse sickness 11: 1235• Murrinn 25•1
mclioidosis Ill: 1500. 1505 !237·40 ,\fu,..._.a sp,-cie~ 77, 8 1-3, II: 13a9
Mellil'Ora captnsls II: 1137, HI: 1975 :Slokol3 ,iru, 11: 1124. 1130, I H5, 1151.116-l fet.>ding habits 82
Mdophngu$ ov/11us6l8, II: 1205 mollicme JI!: 2046. 2076, 2152 ,\(risen t1e1ltiops 81
\lenangle ,,rus II: 687. 810 .Wollum,m co111agios11m 11: J:!98 --1w111m11alfs n. 111: l~88.
asioonos!s II· 690 ,\/01/11.rcum conrnglo$UIII of horses - - contl11ce11s 82
meninglllsl!I: !5,0. 17S.l, 194i 11: 1298-9, 111: 1504 --co11fisca1a8I, 11: 888. 1270
cattle UI: 1639 mongooses 11: ll31, l 133, 1135, ll!li. 1138. - - crasslrostris 82
catlle, nbrinopurulem 111: 1639 JJ39, 1446, ~ea/ISO i11di1•id1u,/ 11011/l!S --domesticn ;;, 81.111: 1480. !~8,. 1798
horses IU: 1610. l611 ~lononegav!rales 11: 13f,8 - - domestica cnllet·a 62
humans 11: 1038 ~lontana sheep disease 11: Tl7. 733·8, 741, - - t/omesrica curuiforc!!ps 82
pigs 111: 1631. 1n6. 1ntt. 11s1 741. 1401 --/re1.>dnw11i82
pigs. bacterial II: 91-l, !11: 1629 moose II: 1227. l339.111: 1548 - - ln.<IOfJltlltal1>1tl 82
sheep/goats Ill: 1655, 1657. 1702 Mopcia rl: 1082 - - lusoria 81. 597
tubetculou~ Ill: 1979 ,\/ort1en specie$ 328, :;2,5, 11, 1158, 11(: 1586 - - 1111111roi 1)1
meningoecephalomyeHtis, pigs II; 9i9 .\lnra.tolln species Infection llJ: 1•18':-90 - - 11e11i//i8l, 597
meningoe11ccphalitis386, 1l: 684. 82.2, 1008, ,\fora.telln bo1,is 83. Ill· 1·187 - - sorbe,,s 81
1031, 1311. 111: 1505. l5S7, 1568, !586, --bol'is inf~ctlon 111: H8,•9 - - tt'mpesramm 82
l.;&i, li84. 190~, 1905 -cnprMlll: 1487 - - 11,m,o/i$Silllfl i1
canle 111: 163-1. 1637, 1655. 1657, 1906 --osloe11sis Ile 1,187 --xnmho11wl1u8I. 597
non-suppurotlve IT: 903,913 --01,i.flll: 1489 ~luscldae. vector:; 7,-85
pigs m: 1601, 1604 - - 01,is infection 111: 1489,90 M1udn111i
sheep 111: 163,1. 165i - - ph~1t)•lpyr1111iett Ill: 1487 --swlm/(11tS 77. 80
thromboembolic4ii. lJI: 1634, 1657. 1906 .\lorbil/i1,ims n: 634. 660. 681. 63: \lusclnae 77. 80-85
thrombolic IT: 1158, 111: 1634. 1637 Morel'~ dbcasc 111: 1923 d1~ease uansmisslon potential 83
Mephiris mtphitis 395 .\/orcllinspccles 77, RI musk ox IJ; 798
mercury poisoning 357, 524. 525. seQ also :.1organ·~ ag;irand broth Ill: 2079 muskrat553
under poisoning .\lor1icrellu Ill: 2120 ,W11s11•/r1 1•1.1011 395
.\oft'$Qlll)'iO 6; -wolftlll· 1623 mycetoma ill: 1756. 2107-9
mcmldehydc poisoning 11: 1158. see nlso mosquitoes mycobactc,rin! dermatitis 111: !979
1111tler po1soning as vectors 137.49, 157, 597.11:988. 100,1, mycobacrerial gran11lomas Ill: 1976
Motnrlrl=ium anisopli.ae 31 1014. 1023-4. 1031. 118,.1, 1195. 1234. mycobacrcrial infection II: 71:?.
mecritisllJ: 1442. 15-16, 1570 l 2$3, Ill: 1798. 2029 Ill: 1965-2008
l.'nttlc II : 951. nr, 1638 control 1•17 M.veolM1°1eri11m species !11: 2011
equids Ill: 1769. 1952, 208'1 ~-cology ofl~ctor species H 1-1. l 48-9 M:,~·olMCWri1(11J ,w/11111 Ill: 1980. 1983, 1987,
pigs Ill: 15-13, 155;, 1784. 1952 epidemics 144·7, 149 1989
sheep/goats Ill: 1597 ~outht'rnAfri~n 137..li --(ll'ium sub$)>. pararub,•rcu/l)Su
~ letronidazolc 317 ~i•b-Saharnn Africa 14?-9 Ill: 1994. 2004
,\llcrocys1is nerogi1WS(I 6-03, fl: I 057 ,·<lt1or<:ompe1ence ofl39·40. 14,·8 - - <11,i1,m-l11rrt1,·t!l/11lnre compl<'x
microcnccphal) viral maintenance l·M·7, 149 111: 197-1, 199,1
canle 11: 953 ,irus isolation.~ from 138,9. 147-8 -/JQ1,f.<24l. 111: IS13, 1516.1973, 198,,
sheep/gonts II: 1049. 1050 :Slossuril ,1ms 155 1989
micrognathfn. pigs fl: 981 moth.> Ill: 1488 - - /x,1•is infection 111: 1973-8,
mkrophthalmia. cattle 11· 953 mountaln gazelle II· 1;139 --Jarci11og1P11'$ 111: 210;
mfcroschlzoms 450, 486 muuntaln hare 11: 998 --forwirum 111: 1983
Micrqswrum mnis Ill: 2096 mouse hepatiti, ,1rus 11: 791 --lmercellulnn: Ill: 1983
- - ~qui1wm Ill: 2096 mouse mammary tumour viru, 11: 718 - - l:011,aii Ill: 1983
-g.)'p$ettm 11, 1299, 111: 2096 movement incoordinatiun si,e 1111der ntaxla - - /)(lr/111tberculosisll1: 19-Lli. 1983
- - nmwm Ill: 2096 mu toxin 111: 1847 - l)lrlai II!: 2003
- - porsfcolor 111: 2096 mucocutaneous k•,shmaniosis :1111 --$111(,g71Wtis II!: 1983
- - specie, Ill: 2095 ;\/1,ror UI: 2120 --rui1errnlosis:?41.111; 197•1, 1980. 1983.
.\hcroms agres1is 11: 998 muc-ormyco:,,is m: 2120 1987, 1989
M1ddclburg ,irus 155. 11: 1015 mucosa! disease. caule II: 64i, 9().l, 946-62. - - ruberc11losis ~,,sp. caprn~ m: 1989
mild foot r<:>t Ill: li34 1196 mycoides cluster Ill; 2046. 205-1, 2060
mild theileriose:s 3 mud re,•er 111: L-H5. 2032 ,\lycoplnsm(I species 8.3. 11: 668, ;29, 7.j,j.
mfliary tuberculosis Ul: 1971, 1980 ~ludjinb:my \~rus 100 849,111: 1488. 1489, 1490, 1698. li8.;,
milk fever II: 1185, 1191 Mugugacocktail461, ,170 2076
milk production decline 11; I I~. 1189, 13<1$, mulh"1'ty hc:m disease 111: !Sil .\1yt:oplt1sma type 7 Ill: 2076
1343, ill; 1442.1448. 1•161. 1535, 2012 mule deer II: 798. 1422 .\tyropf<(sm,1 ag11/ac1lae II: 1397, 111; 1490
mllkringte,t Ill: ISli. 15 19 mules 11: 1234 "1ycoplasmn agal<1c1/(le ,'8ct!n~ II: 1398
milk, blood,~tamed 11!: J.146 ~lutes· opero1ion 11: 757 ,\/y.:op/nsmn 11tk11/11.,ce11s Ill: 2076
mil,;,-dropsyndrome Il l: 1448 multimammate mou~e papll!oma,~rus - - nrgi11irti 111: 1490. 2076
milker·~ nodule II: 1285, 1289, 1291, 1302 11: 758 - - bo11i11genirnli11m Ill; 2076
miltsiekte Ill: I iflB· 181 l. seen/so nnthra., muhinucleatc cells/giant cells 11: 66-1. 66i. - - boirirh/11/s Ill! 2076
/lllllzbrand Ul: I 788- 181 1. see also amhra~ 677 - - /µ,i1is Ul: 20i6
Ml11iop1erusschreibersiill: I 1•15 multlsystemic vascuUtis, horses II: 844 - botlOCIIII !11: J.189
mfnk396 multlsystcmlc wasiingsyt1drome 11: 808. - - ,a11ade11se lll: 2076
minl;ewhale 111: 1546. 1549 8i0. 1374•83 - - crqJricolum s11hsp, caprlp11e11111011fac
.Minnesota isolate 11: 90,1 muriiorm bodies/cells Ill: 2109 111: 2060
- - COllJ/lllCliMe Ill: 1490 nagana 25 i-87 seenlso trypanosomosis ncona1.11l death
-fells II· -;o :'\3.irobi bleeding di~case. dog~ 280 cattle Ill· J.152
-jlocculare 111: 2066 2,.
:--alrobi ,hecp disease •I. 21. 24, 15~. horst'S IJ: 8-19, 92.8
- - l1yop11e11mo11it.1e 11: 823. 911,914.1307, II. 6-ti. Gl:i8, 105i 1071 -5 pig, 11: 1023. rrl: 1-148. 15,12
1383, Ill: 1494, 1705. 1778, 2066, 2072 :,.:airobi '>hcep dbea,c ,·in,s 4-11 <hccp347, 111: 1528, 1597, 1620. 1921
--h.rorh/11/s Ill: 16.11. 20;-1 Sairol'irusgenU> 155.11: IOil. 1077. 10i8 neona1al dtarrhoea JU; 1:\f,3
- - ilJQr//i11iJ infet11<1n Ill: 2071 ·2 :-.nmib1an red-legged tkl: 22·4 nconacal cmcrotoxacmia Ill: 1837
- - IIJV>,~'/IOl'UII! Ill· 20:"4 :'\amibiese rO(lipoutbo~lui< 22·4 1wona1aJ lonl <)'Tldrome II: 84~. 845
- - hyos,•1101,fa~ infection Ill: 20,4-5 \"a11/IOIIIOIUI$ 49 nuonacal iso~iyllirolyi.b 604
- - myrnidPs sub,p. c,1pri lll: :!046 .\"(111opl1,1·em, .•al111i11rolt158•1 horse, 11,849
- - mycoide:;subsp, m.1-roides Ill: 1490 nasal discharge II: 900. 1341\ neonatal neo,porosi;, 38(;
- -111.waides subsp. 111ycoid<!S l..irgc C'anle II: i'98, 820,900, 1189. 1322. lle?onatal piroplasmosis 429
Colony fll: 2046. 2 I 52 Iii, 1680, IG9l 2050 neona1al ,ep1icaemla 111, 1555, 1557, 1652.
- - mycoides subsp. mycoidt!S Small cqu,ds It: 842. 864. 1024, 1320.111: 15U2. 1664, li04, 1772. 17114. 1904
Colony Ill; 2045 1771 (;.3,2116 ,\'rorick'l'mw species 5S7
--Ol"ipneumnnia II: 738, Ill: 1698 pigs II: .;5, ,,G.111: 1494. tllG~ .\"eorick~usin uloko111i11ico 584
- - 1-er«1111d11111 Ill: 20,6 , shccpl!lOat> II: 1073. l20i. Ill: 1699, 2061 - - lwlmimho2l'n509. 536. 583
mycopla.,mal anhrill$, pigs Ill: 20i2·5 nasal glanders 111: J;;02 .\'t!O.<pOrtl Cllllillllf/1317, 3,1. 382•90
myeoplasmal pneumonia. pigs Ill: 1705. nasal ml'COSi< Ill: 2115 - - e1mi1111m, life cycle 382
2066-8 Kdumu \1TUS 155. LI· 1015 - -lwglies/ 386
mycoplasmal polrscroshis and nnhritis. Xear-Easrem ~quine cneeph:tlomy<"litis neosporosi, 341. 382-90. 400
pigs m: 20i1·2 II: 1006 ncosporosis
mycoplasmns II: 800, 111: 2043·82 nacrobadllosis ll: il2, Ill: 1741.1949. s cl' c:utle382·6
tl/:;Q 1111der flJ}i'Cletl .~It,• duer 387·11
mycoplasmosis
nccrohaemorrh:tgk emcrocolici&, equicls dOl,"1, 3116,90
in cattle !U: 2076·80
genital Ill : 20,6-80 Ill: 1840·-M goats386·,
necro,ls horse~ 3!16
mycoses Jfl: 2095-2124
brain stem neul'Oi\al ll· 141 i neonatal 381\
myc:o~l~
bulbar. infccti,e Ill: 1730·32 ,hcep38i
eutaneou.s 111: 2095·2101
cerebroconlcal 41 i, 477. II: 1158, \'fo.<1011U1 /fJlitlne Ill: l5·l6
l,'UIIUral pom:h Ill: 2115
Ill: 1639. Il\-10 nephritl~
inccrn~I Ill: 2114-:?~
cerebroconical. th!anunc-dt>pendent 4;; c:iule Ill. 1448, 1449
nasal Ill: 211;;
eurtip, tu: 158i, 1760 horie~ Ill· 1652
suocmaneous Ill: ::!IOZ-24
gall bladder muco,a 111: 1590 mterscitial Ill: 1449, 1'1~0. 1570. 1959
~uperficial m: 2095·2101
hcpauc, C31tlv IH: 158i pigs 11: !l-10, 111: 1959
rnyco1i,· afiortion Ill· 2121-2 nephropaLluca cp,dem,cn 11: 10112
lwpacic. W1nwan·u>ngo haemurrhag1c
111,couc dermat.iti> Ill: 2.0:?6
fo,cr 11: 10111 n<'~1hros1s 428. 429. Ill; 1595
myco1i",ndo111etri11, Ill: 2122·3 hepatic, fol'lll, canle Ill: IT.i9 ncl'\"e dysfunc1ion, equids300. Ill: 2116
mycouc plac:t>nutis Ill: 2122
hepauc. iocal, pigs If!: 1450 nel'\•ous~igns 475. 41l9. 520. 540.11: i36. 738.
nl)'COUC,tomat.iu, II: 119,
heput!c. iocal.shecplgoars Ill: 1702 743. 901. Ill: 1569
mycotoxicosis see poisoning. pl:1msl
hepatic. horses II: 845, Ill: 2090 pig$ Ill: 1631. l 778
moulds
llee1cal!C31 ,·alve, pigs 11: J 107. 1109 'leufeld reaction Ill: 1r,55
m)'cot0xms rn: 1452, 159,,see (lfso
ischaemic 111: 1587, 1637 neuritis. opcic ner,e, t'llttle ll: 953
poisoning. plants/moulds
muscular II. l 190. 1211. 1221 neurologicnl disease Ill: 194,
m)'elitl$ 111: IS05
o~<ophagu~ ll: 12.21 horses II: 843. 84-5
myelocnc<:phalitii, equine prow~,,al
oronasal II: l:?2,1. 1228 neurological signs
394400. ll; ~9. 1020. 1370. Ill: l442
pharynx II: 1221 cnnlc II: 1413
mr~loencephalopathy. hors.."> II: 846
Schweiger·Sl'1del ~hc.>nth II: 1107 horses IIL 1442
rnyt~is. shcepigoats II: 128'1, Ill: 1735
skeletal mu~dl' II: l 190 J>ig, II: 69-1, 909,913, Ill: 1603
my<>eardlti< II; 1310, IR46
lhta:nme-dcpc-ndent ccrcbrocortlcal 4 ';'7 Midlife 111: I:,49
ca..-tle Ill: 1~2. 1634, 1635. 1655. 1691,
th,ornho·i~chaemic. hor~"' 11,841 neurologic:al <)11drome, can le Ill: 16.r
2052
tip5 of eah/tall/lirnb.> 111: 1587 neuromu~cul:,r paralytk syndrom~ Ill: 1896
hor~es Ill: 1652
tongue 11: l22l neuropathy, ope le !WI\ e l 1: 834
pigs 11:811. 93,. 130i. Ill: 1i78
necrotic nnd uh:erativ1; ,wmntit.ii.and neuroto~in Ill: 1878, 1887
myomecritis. hors.,_, ll: 926
l3J:'Tlgitb Ill: 1741 · 2 nemrqphilia II: 1190, Ill: 1,,3
myo:iecrosrs. clostridial m· 1831
necrocic emcricis Ill: 1829·33 New Forest dl~ease Ill: 1-18i
myo;imhr nurrltionnl. sheep Ill: 1736
pii:s ill: 1469 Newbury agcni l II: 703
myosiri$
~hccp Ill: 1840·~·• ;>;cwbury a.gem :? II: 703
bouyomrcotic ut· l 756
cattle m: 1691
necronc ioci, liver spleen 111: 1587. l 588, :-.ewcastle disca,.-e ,irus ti: 687
1905 :--t.•wing·s trypcose medium m· 2063
eoslnophilic 360. 3G-I. 357
necrotic rhini1i~. pigs Ill: l 7•13 =-:gaingan virus 99
gangrenous 11!: 1856
rtccrouz.ing cncephalic!:;. horse, TI: 1019 nibble retlex II: 1400
horses l!I: I i56
nucrolizing hc:paritls ill: 2089 ,·1roria11fl gla11co 11: 1254
\/yotu da.,_1oe11L~11e 11 I l ,16
nccro11.z.u1g lntilctlous cntNltls, pl!:$ .V/t1011Jrnfe!J II: 9l4. 933
--dm,1~mo11if ll: 1146
llJ: 1840--14 '-:lie catfi,h Ill: 1549
O<'c."rotoXigenic ~ht!riehi11 coli Ill: 1562 Xipah \'iru, 239

0 'lieeL111ing~train II: 1268

Negishi II: 995
'-l,grl bodies II: t !23. 1125. 1154, 1154;
Xipah ~ini~ disease ll: 692·6
a.< zoono~is II: 692
;\'oc,m/i",perlcs lll: 151i,2003.20!1 210i
~·oama canle 263, 266, 287 ,\'eomelanh-011/011 II: 1040 \'ocardia nstt>rold<'S 111: 2011. 210i
.,·aeglerf,1 316 neonatal anapl~srnosis 601 - - braslf/umls Ill: 201 l. 2107
.Vnegletit1 /011•/erl 31 G nt>On8lal collbaccillo$i, Ill: 1562 --fercinlcn Ill: 20 l l. 210i
- - 01icidis,:1111inrum Ill: 2011 acu1e pulmon3(1', ca11le IT: 1191 Omftlrodoroscoriaceusll:8;6.1098. 1205.
nocardial dermatitis Ill: 2026 bmin 521 Ill: 1436
noc:irdloform actinomycete(s) ll: 849, dcpemfon1 11: 924 - - ,·r1·111k11s II: 1096
111: 2011 t.>ars IJ: I207 ~lolrore11sisGl8, 11: 1098
.\'ornrdioides Ill: 2011 C}'elid~ II: 935. 1207, 1228 - 11/lll'Ol.'IIIWJ 11: 1096
nocardiosi\ Ill: :?011·13 face !I: 1207 - - 111011!mwcomplcx3, 27, 11: 1096
nodular worm Ill: 1919 head II: I nG, 1239 - - porci1111s tlomestic11s 11. 1096
nodules Inflammatory Ill: 1875 - - 110rci11us t10rci111,s 11: 1093. 1096
internal organs. ,hccpti:on1s lfl; 1505 lann~ Ill: 1950 - -$tWig11yi 596. le 988, I098
nasal mucosa. equids Ill: 1502. 2110 hps 11, 1207. 1228 - - wricara ti: I 098
p·ulmonary, equids 111: 1503 lil"er Ill: 1702 Oml1hognl11m species 328. 525, Ill: 1588.
wspirn1ory tract. cquids 111, 1503. 2113 Jow~r hind l!mbs re 1024 16-10
skin. canle 11: 891. 12il !\mph node< Ill: 1596. 159, Omirhoglo..<s11111 species 328. 525. 111: 1;;ss
skin, equlds 11: 1298, Ill: 1502. 2105 n~ddl: 1236. 1239 omithosis 550
subcutaneous 111: 2103, 2110 perlorbhul. c:aulc II: 1189 Oropouche \'irus I00
~ubmucosal Ill: 2103 prriorbital. horses 11'. 927 Orrhom;.·xotririilne 11, i65-,i
suppuratiw Ill: 1649 peTiphari ngeal, pig, Ill: Ii05 onhom~,xo,iru<(t.">1 s&e Ortll1;1myxo1'irida~
nofcl 3 . 21, 22. S36, 531Hl. 542 pulmonary 457, •169. ,J90, 521 5,11, II: 682, onhopox virus infections 11: 12"8. 1296
non-progressive atrophic rhinitis. J>'ll" 68,l, 1191, 1210. 1237.12-19. IJI: 1,,9. orrhopoxvirus(es\ II: 1268. 129,, 1300
Ill: 1q92 1596. 1702. 1114!1 oryx .ree gemsbok: Arabian OJ')':\~
non-progress!,·e fooi rot 111: 173,1 <no1nl. st~11i1>ns II: ll,13. 10?4 $cimitnr•horned os;-':<
nonstcroidal a111i-inflamma1ory drl1g ,ubcmancousU:9'15, 1189. 1236. 1239 o,y.tdnmmah516
tO~iciry 58i wpraorhlrnl, hors<'-> II: 86.1, 92i. 1236. - gti;;d/(111: 1270. 1423. Ill: 1690
Noro,,inis n: ;oo 1239 - - lwtt>l):Y II, 12i0, 1-123
North American blastomyco5is In: 2116-17 10nguu 11: 120;, 1228 0Me<1genesi,. deranged n: 953
Outk(J11ia Ill: 20ll
OSICOlll)'Clitis Ill• 1754. 1946, 1951. 1953
North American red fox II: 1128
northern fur seal 553 canlell l: 1586. 1587, 1942·•
<kw11/1a,.wmomum ,l)i:des Ill: 1919.19..S
gnmulomatous, <:anle Ill: 1942
:"orwnlk virus II; 703 ocs1ruHyclc!rrcguluri1)' Ill: 1459. M61.
granulomatou,. pigs Ill: 1543
~orwalk-like 1irustesJ LI: 700, i03 20;8
horse~III: l6J0, 1Gll.2015,'.!019
nose-ringing Ill: 19-18 Oasrroso,•islll: 1946
1uberculous lll: 1978
nosocomial Salmonella infections Ill: 1608. OlifnnisvlL'I virus l 5S
os1cope1rosis, caule 11: 9-19
1609. 1613 olifamvelslckie 351-8. see also besnoitlosi< os1cophagll1 Ill: 1887. I889
,01·idium 282 omphalophl~lllris Ill: l5i0. 1653, !78~. os1lich 11: 1423
:rndek acid lmmu nizot!on 242-3. see a/st, 1,85. 19-19 otilis e~1erno
immunix:uion Omsk hacmorrtwglc fc, er II. 995 cn11le Ill: 1638
'v11mida 11w/etigris 516 onchocerdo,!s 357, II: 12;-1 dogs Ill: 1762
nuwcsiek1e U: 769, 770, Ill: 150-1, 1505, 011,ov,ri,me II: i Ii otirls intcrna Ill: 1948
li68-i4 o,utar,a :il1Nlrlcn~1 ociri< m1tdla Ill: 19,18
Xyabira ,irus !lll, 155, JJ: 1252 Ondlfi disease see bm<inc pe1ech1al fever 0109'<111 megalorir II: 1130
ni·ala 11: 1423. JI!: 1446 cmy~homycosis Ill: 211 l 011/Q11 faro II: 1130
\Jyamanlni virus 155 Ooskuskoors see East Coast fe,·cr o,·erentlng discas~ 111: 1846-52
.Vycreris rilebalcn II: 11-16 opale)'e fish II: ,Ol o\'ercxchcment stt h~'Peraes1hesla
op,m-mou1h br<'athing. pigs. II: 694. ,76 011ii,11$ masclrat11l I I: 798
Ci) ophthalmta
nmle Ill: 1487
ho~e, Ill: 1-147, 14.-19
o,ine adeno,1ru~ infecrion~ S<'<' adcmo\'irus
infections of sheep
o,ine and caprinc a.naplasmoslS 2-1. 5i1J.
0 antigens Ill: 1560 sheep/goat, Ill: 1487, 1489 6 17-22
O'ni•ong-nyong \'lrus 11: 1023 oph1halmi1i,. cquids2i8. Ill: 1-149 ovi.ne and caprinc hepatic nccrobac!llos!s
oat cell ti!: 1699, 1702. 1705 oplstho1onus 11: 12;;.1, Ill: 1880 111:17-ll
obex II: 1400. 1415 opossum Ji I, 394, ;;53, 111: 14'16 O\'lne and caJ)rlnc salmoneUos,s 32,. II: 973,
obs1ructio11, imeslinal. pi~JII: lC>31 ovportuuistic mycobactcria Ill: 19i8. 19i9 107•1. 1260.111: 1594-8
ocelot II: 1422, 1423 Op11111in ,pccies Ill: 16,18 O\'ine bab<.'1iiosis 438-42
ochm1o~n A Ill: 1452 oral ntcrobar:illo~is UI: I ;.i 1-2 o,inc carorrhnl fc\'cr see blueiongue
ocular discharge Or/Jir•in,s 156. 11: 1202. 1221. 122,. 11..l!. O\ine corona,irus II: 803
ca11lc· IH: 900. 1488 1232, 12-17. 1248. I 2!32 ovinc coronavlrus infection 11: 803. Ill: 1597
dogs m: 2140 orchitis35S. II: 1273, Ill: 1510. 1511, 1315. o,ine chrl!chiosiS 24,541·3
equlds n: 842, Ill: 1;;1 1521. 1535, 153i, 1542, 1543, 15~8. 011ne enzootiC posthitis HI: 1931·-l
pigs !I: 776, Ill: 1709 1608. 1952, 1953 o,inc fo111 ro1 Ill: 1733-8. S<'ll also u11dnr foot
sheeplgoatS Ill: 1699 caale Ill: 1638, 1943 ro1
ocular disease. horses U: 844, ll63 ht>rses Ill: 1611 <wine herpeS\'in1s 2 II: 895. 896
ocular squamous cell carcinoma II: ;ss. ,61 <he11p/goa1~ Ill: 1657. 1921 o,in~ hcrpesvirus II: iii
Odocolleus lzemlonus 11: 798, 1422 Om1nmosn111erict11111slll: 1995 o\'ine imcrdigital dermatitis 111· I 729
-1t1rgl11i11n1t$ 382. S24. 1;18. II: 795, 1205. orfll: 1278. 1279, 1282-6. 1288, 1351. O\'hW len1Mrus II: 73'1
122i. Ill, 1528, 1973. 1995 In· 1736. l 7S6. 2029 O\ine mou1h and gum obscure d!sea<~
Oedac1,s 11icari1u JI: lOli a;. ,.oonosls II: 1282 II: 1351
oedema disease Ill: 156Q, 1Sil orf,iru, II: i59, 12s;, 1289, Jll: 2030 o,1ne pe$11\'irus diw~se II: 970-i<I
pig$ Ill: 1563, 1566. 1569. 1570 organophO,Jlhate poisoning, see 1111d~r c,vfne pre>i;rcss!ve pneumonia II: 71 i, 733·8,
oedema disr;a.e toxin m: 1567 poboning 741. 744, 1401
oedema toxin Ill: 1801 c,rganophosphate~ 52•l. 525 o,ine pulmonary carcmqma ! I: 71 i -30, 733.
oedema Onema! sore 379 7:ill, 1397. 1402
abomasal 522. 11: G-H Omi1iiodar11.,svecics II: 1088 ovine 1hcilcriosis 22. 24
O\'ln,• vcocronl disease 11: 1285. 1287·8 goa1s 11: 7 43. Ill: 1505 pcricardilis 111: l5 70
D11isca11nde11s&553. u: 1205. 1227, m: 1995 horses 300.11: 843. 1019. Ill: 2019. 2116 caule Ill: 1639
Oxford imradcrmalschcdule II: 1163 K:m)O~. 18 fibrinous UI: 1699. 1705
oxpeckers. red- and rcllow-billcd 31. 34. laryngeal 540 pigs Ill: 1631. 1705
!Tl: 2()29 musde; Ill: 18115, 1894 ~hei:p/gQals 111: 1699
O.\yurfs equl II!: 1758 nef\e. equid, 300 perinatal dea1h Ill : 1946
oc~phagal. in canle Ill: 1893 periodic acW-Schiff reai:tion Ill: 20!18. 2138.
Oesophagnl, in hor~~" n: 1240 :?141

G pig., 11:912.1308, 1311.111: 1543, IG3l

po~1enorm: 1505. 1543. 1951
sheep 4. 22. 24.111: 1505
perlodk ophlhahni.1in homl~ 111: 1445.
1449.
puriodomi1i.<, caul~ Ill: I 7•l4, H/51
Pnd1ys1,g111a ~pcdes 573
paral11ic rabies II: 1129 pcri•oophorihs Ill: 2079. 2150
l'ac/~-srlgma lanfolius 52·1
Pnmmphi,romum mlcrolw1hr/11m 111: 1588 p<!rio,dtis. can le 111: 17-14. 195 t
- - pygmneum 524
Paramrxo1•iridae II: 627-96 pcrlpneumon!c conmgieuse bo'1ne
- - tiu,mnus 524 Parlllll_l'J:(Wirus II: 673. 687. 692 Ill: 2045-57
Pacific harbuur "t?nl 395 paraphfmos~ IL: t:2e;, Ill: 2J53 p;,rl!onitis Ill: 1570
paddling movements. caule nt: 1637 paraplegia Ill: 1951, 1953 catlle ill: 1639. 2076
P11ecilomy"•s species l!I, 211 O parapox,iru$ infcctio11~ II: 1289, 1301 plir.; 111: 1631
pallor disease 575 parnpo~sirus(es) II: t282, 1291 persis1ent foo1-nnd-mo111h disease
Palyam serogroup orbivlrus Infections par.isllie amoeba 316 lnfec1ion 11: 1340. 1353
11: 1252·5 paraufuure bo,ine ~almonellosh pe,,ls1ent infl'<:1ion
Paly.un scrogroup ,1.rustes) 98. 11: 105i, paratubcrculosb ll!· 199·1·2004 cattle 11: 946. 950-51
1248, 1252.111: 1588 and Crohn'sdisease Ill: 2004 pig,, II: 9i9
Pangoniinac 87 parnryphoid 11: 1057.111. 1-181, l5f>-l .s,-ealso ;heep 11: 971
Pm1/r:11111 ma.timu111 Ill: l 74~ bo,1ne salmonc.>Uosis: porchw pes1c des 1>e,l!s n1minantS 226. JI: 634. 64,.
panther II: I 227 salmoneltosb 660-69. 1057, 1074, 1278
Pamlwrn /e() Ul: 1975 par,iv:1,cinlasee pscudocowpox pe,thirus!e~) II: 914 . 946. 975
--pardusl!l: 1975 paravaccln!a ,;ru, II: i:!91 ca1lle/pigs II: 97 i
pan1ropiesc bloubosluls 11. 15 parc,L• ihe~prgoa1, 11: !)70
!>apmicolaou sraining method Ill: 2139 caule/horses Ill: 1637. 2019 ,,;Id ruminant> II: 970. 971
paplllomas. squamous ll: 755. 758: seen/so pigs 11: 13 I I. Ill: i 543. 1631 pciechiation. kidner Ill: 1570. 1604
1111der papillornavin,s infections p,merior475. II: ,1 l. 927. 979. ill: 1951 scro,al surfaces. Callie JI!: 1691
papllloma1osi~ n: 754. 1285 Parker's Farm vin,s I00 shecp/gonis 111: 1596
papilloma1011s 1umours 11: 756 panuriem hypocalcaemia 11: 11!15 skin. pig.~ II: 979
papflloma,·lrus Infections I1, 754·6 I Par1101<frid1,e II: 805-17. 914, 111: 1784 Pc1,•vo virus II: 1Z53
camels 11: 1301 Pasadas'sdisease 111: 2117-18 l'fl'/ffcr~lla ma/lei see Burkholderin ma/lei
ca1llc JI: 755. 1292. 1297 pos,ive immuniznt1011 244. see al$tJ />/111cocl11wnis nethiopicus II: 808. Ill: 1975
dogs 11: 757 immunization - - africmws II: 1088
cquids II: 755. 12,98 pastern dcrmatllis 111: 1757 phaeohyphomrcosis ill: 210;, 211 O·l l
goai.s II: 75i. 1284 Pnsu•me/la Spl'Cie< II: i25. 736. 769. ,;o. pha1~hyphomyco1it sinusitis lU: 2l JO
hcmans 11: 757 138,1. lll: 156-1. 1661. 1677. 1704 pharyngi1is II: 860, 863
pigs II: 7.37 and .1/(//mhvimin sprcics 1nreclion, P//asrolt1rc1w; ci11enms 551. ;;53
rnbbirs II: 757 Ill: 1672-lil:l phcnan1hridine compounds 261
~heep II· 756. 1284 Pns1e11relln /1nemolytlca see Mm111hrim,r, Pl1ilaema1omyill 82
papilloma\iru~(csl II· 754 'Pa.sttllNl/ll/ }ltll'/UO/yt/C/1 Pl1flolfclio 87
Pl/pio 11nl1111s Ill: 1975 Pa.s1t'1tfl!lla 11111/r«idu 90. 2i5, Ii: 823, 911 - - aelhitJpicn 88
Pt1po1•avirltfn~ II: 753-62 914. !120, 937.111: 1492. 1623. 16.38, phimosis
papulor $lomntlti< ll: 647, 95 I. 1351, see 1639. 1664, 1670. 16ii. 1689. 1695. cquld.;;300
nlso bo\ine papular s1ommi1is liO.,. 1707. li78. 206i sh~eptgoon, 11: 1287.111: 2153
papules --pestis Ill: 1617 phlcbo1omus lever II: 10..18
camels II: 1300 - - p1w11motropim Ill: 1704 />li/ebo1om11.s 378
~aide II: 1196. 1291, 1296 --psi:'11do111berc11/otfa Il l: 1617 --mss/3711
h()CSCS II: 1298 --trl'lwlosilll: 1G'J5. 1101·2 phlcbovlnrs{es) 155. 11: 1038. IOi/9. 1057
on e:(temal genitalia. horses II : 8G9 p;meurellosis II: 10.;7, Ill: 1672-1713 phlt!gmonn inr(•rdigicnlfs. caule 111: ii22
pig$ ll: 1293 pasteurellosis Phnom-Penh ba1 virus 11: 995
sheeplgoa1sll: 1278. 1284 canlc Ill: 156~ PilOCll 11ini/i11a ric/11/rdsi 395
Skin. horse< IU: 1758 pigs II: 77i. 1109, Ill: 1605. 170.J-5 phodne distemper II: 634
/>mr.cy11/cris selousill: 1137 sheep/goats II: 729, Ill: 1695·1702 phosphorus deficiency Ill: 1886. 1889
/>arr.filarla bovlcaln 81. 83. 597 pas1cur!1.ariot1 of milk Ill: 1975, 1987 photophobia357, ILi: !+49. 1488
paminfluen1.a cype 3 infection 11: c,;;i.5 pa~ture it>ver see tic'k·hornc ft'\•cr ph01osen,l!Mty see pho1ophoble
pnrainfluenza 1ype 3 \'ln,s II: 668, 6'i3. 798, Paww, liarborli 525 phycomycosis 111: 2120
800,821.951, Ill: 1488. 1489 - - td11111um11itwa 525 physhls. horws 111: 1610
paminfluM211 ,;ru5 pear!ssee TB pearh Pl-3 ,1rus in humans II: 673
bo,~ne II: 687 Pea1on ,1ru~ 99. II: 1029. 1031 pic.i 111: 1886
human 11: 687 pedal ulc~rart,·e dennaiosls II: 1287 Pfroma,,fritlnl! 11: 1305·65
para-Lyme dhea,c, syndrome Ill: 1440 P,•11icilll11111 species Ill: 1452 pig nun: 775· 7, 91~. 938
pnrrtl)'$iS 26.11: 1368. 1370, 1885. 1905 Pennisewm c/a11des1i1111m II: 1254, llli 1,34 p,glouse575, 111701.1098
bulbar Ill: 1885 Peprm,tre/JlOClX'CIIS imlolicus Ill: 1744 pigronfm•er Ill: 1921
canle..:75. ;i.10. II: 1129.111: IG.17 penl~ cnkcfalosc see equine encephalosis plgm) chimpani~e papilloma,iru~ 11: ,S7
fadal Ill: 19-18.2116 p~rdegri~p ·\30, II: 761i-i'2, 928. 1025. !320. pi!i Ill: 1560. 156 i
facial nerve 300. Ill: 1905, 1906 lll:li68 pinkeye 83.111: 1·187
flaccid Ill: 1885. 1892, 189•1 Peregri1111s ma/dis II: I l ~5 plnwom1 Ill: 1758
x.xii l'.\t>fa.

pipcstrcam diarrhoea Ill: I 997 pncurnoLOxin 3-mcihylindolc II: 1191 porcinu entcric noro\'h,s'es. Jt 703-6, 800
Pl11is1ullr1S $11/Jfm•tts II: ll 211 pncum<iv.igina Ill: 15Sli porcine emc,ric sapoviru~:es, U: 703-6, 800
Piropl1w,u1 t"lllmlli -12s Pneumm•irinne ll: 687 porchte cmer0\1ru~ in(o!Ctlon l!: 793. I 307-9
plropla,mosls. neonaial 429 P11~u1111;111irus 11: 677, (!117 porcine epidemic diarrhoea 11:;87·9
piroph1sm$ -150 p()(/odemmmis 11s~ptim 1iiff11sil 111: 1;25 porcine epidemic diarrhoea ,iru$ II: 787
pi>goed Ill: 1931·4 pt)l[,x/vrmauris rircrm1saip1n lll; 1716 porcine haemagglutinuting
Pirhomyu., charrnrum Ill: 1755 pododrrmarlris se1nicn Ill: 1n; encepha.lomyelhu. ,,rus II: 791, il!S.
pined kermolys,, Ill: 2033 Pol!Cilogolealbim1c/10 II: 1138 80-I
pituirn.ry abscess syndrome Ill: 19-17. 19-18 poisoning J>Orcinc haemaggluunaung
pituha11· nbsce!'S, goms lll: 1-llli arsenic :,25 Ill: 1588. 1692 cncephalom>l'litis ,;ru.~ infection
;,huirnrygland II: 950.111: 1917 bracken 1,•m 544 11: 791-3, 914
pizzlc.> rot Ill: 1931 · 1 chemicals 11: 125~. 1,101, I-Iii. Ill: 14i5 porcine intl.,.lin.tl 11dc11oma10<1~ Ill; 1•169
placcnia infectl\ity. scrapie II: 1396 copper 417, 4'12. 578. 603,621. ll: 99 - porcln~ ln1esdna1 camp;·lobacwr, Ill: l-lu9
pln~entltis dicum:irol ll: 983 porcine pamt)1)hOid see porcine
cattle Ill: 1513. 1535, 1638, 2078. 2079 food. human~ Ill: 158Cl. 1754 $lllmoncllosb
c:aule. nccrotizlng Ill: 1480 forag~ Ill: 1887 porcine p:irvoviru, Ii: 690. 806. 921. 93tl.
horses 586. 111: 2121 lend 477. 524. 603. 621.11: 1000. I 158. 983. 130i. 1309
myCOtic Ill: 2122 Ill: 16.19. 1640 porcine parvo,irus infecuon ll: 806· 1I .
shttp II: 971, Ill: 1620. 1952 \/arricmi(l llib>ellifo/ia 477, ll: 1158. Ill: 1452
plantation mes SJ Ill· IG-10 porcme pa~1eurellosi> n: 777, l lO!I.
plasma fibrogen lovul increao,e II: l 185 nwrcury3~7. 524,525 Ill: I GO!i. 1704-5
Pili medium of Knbely lll: 1805 organophospha1e~ !I: 1000, l 158. 1401 pordnt- ;iicornavicus\e<\ II: 810
pleural effusion 111 2045. 2()50 pesticlde.524. 52:i. II: 1110. I IS8, porcine plroplssmo~is I l. 2-,. 26. 435-6, ~76
pleuritis Ill: 1950 HI: 1639 porcine pl,•urop,wumonia 11: iii. II!: 1632,
cttnlelll: 1635, 1639,2050 plant;,/m<iulds-li7, 524. 525,378. 503. 1661-i, 1950
fibrinopurulent Ill: 1570 t;21. If: 685,928, 10()0. 1020. 10;1;, 1057.
(loccine polinenccphalum)>eiili> ll. l 307-9
pigs Ill: 11>31. 166-1 1110, U58, 119 l, 125-1, 1311. 1370,
porcine pol)e>ero,;iii,and anhn11s
sc.>mftbrinou, n 1105, Ill: 2062 1-101. l-117.lll: 1152.1588.159;.1539
Ill· 1629-32
shcc.>p/gonis lit: 1487, 1699. 2062 saliJ1omycin II: 131 I
J>Orcine rcproductivc3nd ,espirnton
pleucopcricardius. caule 111: 16\ll salt II: 1309. Ill: !Sil
pleuropneumonia. cmde seecomagious ,make bit~ Ill: 1807 svndmme II: 933-W. 983. 1307 1383.
bri\'ine pluuropneumcmia Ill: 160~. 1778
polar benr l!L 15.\9
tibrlnous l!J: 16'19. I 70S. 2050 pol!ocm:cphalomaluola porcine wproductlv1: Md rt-spiratory
horses see equine plcuropneumonla cattle JI: 1416,lll: 1639 "Jndromc virus II: 690. i85. 806. 914.
pig, II: "'."i,.111: 16.>2. 1661 -7. 1950 ,h~~p 11: 973 924 93..1. 1309. Ill: !630 1-·7 207 I
seroflbrlnous Ill: 1680. '.!045, 20S5 pol!omtephalomyclilis JI. !3f,8. 1ro porchtt, re,pln11ory comna,1ni- II: 784
shul'ptgoats Ill. 1G99, 2046, 2062 ca11le JI: 1033 porcine r~spiratCH) coronavicu, iniectian
P11e11111acys1iscnr/11/I Ill: 2020. 2131 pl~ltl307•9 11:784-5,938
Pmw1110,,-s1/s c-,uimi infeetlon UI: 213i .g poll evil Ill 1516. 1785. 19-12·4 P<>rc,nc ,atmonello,is 11: -:-.:-. 9-10, 983.
P,,~um(IC)•sri:. jiro1vci 111: 2137 pol) anhri1is ru: 1570. I i.54, 195:l 1109, lfl: 1431. l-li3. 1601-6. 1623
pneumoaymn,!s Ill: 2137-9 cimleS55.55G. Ill: 1505, IS68.1585. 1035. porcine tuberrnlous l,mphad~nius lll: 1987
pneumocnterltls 1639, 1691. 20.52 porclnc ulcerative granuloma Ill· 1437
hors,.., II: 92i equid$ 559. Ill: 1557, 1608. 1652, 1653. porencephaly
,heep II: 822 17i2 catth: II: 953
pneumonia UI: 155, 1S70. 1803. 19-16, 1950, pigslll: IG2S, 163l.1705. 1na pigs II 6!10
2116, 2138 sheepi goats 555. 556. Ill: L505, I568. she,.,.p/goais II: 9i2. 990. 1049, 1050
caule ti 678. 799. 951, Ill: 1586. 1620. 15$G. 1657. 1736, 1754. 1912 porpoi,es Ill: 1546, 15-19
1634. 1638, 1639, 1640. 1655. 167i, p,:,lycluomatophilia 620 po~,um 111, l!J73. l 975
1681. l691, li40. 1742.2011. see also polyps. nasal Ill: 2112 post-dipping lamenl.'ss. sheep HI: 1912
calf pneumoni« polyse11lSi1is, p1i:s Ill: l 629, 1778. 2071 ·2 posterior parcsi~ 475. II: ,I I. 92i. 979,
cuffing lit: 1705 pol}'S}TIO\~lis 111: 1!151
enioo1ic seet>nzootic- pneumonia io.lls Ill: 2019 posterior ,•ena c,M1 syndrome. cant.:
cxudatin, II: -26 ,erotlbrinous II: 1190 Ill: l7-l0
gronulornalOus Ill: 2011 polyuriu Ill: 1959 po,th11(,. Ill: 1931, 215'1. $ttnlso ulttrative
horse< 559. Tl: 769. 823. 865. 927. 111: 1557. Pongola virus 155 dtmm1o~is
1608.1623, 1771.2011.2019 poor c:onc-eption rates SI.'<.' infortllil) canle Ill: 1933. 193,1
inier<ticial stt Interstitial pneumonia poor performance ,yndromo II: 842. 860. po>tpanuriem endometrilis Ill. 1!152
pigs ll, 695, 93!1, ll05. 1105. 1107, 863, 86-1 pos1parturirm gas gnngrnnc Iii 1869. 1870
Ill: 1494. 1601, 1603. lf)0.1. 1661. 166-l poor thrift poqpart11rie111 haemogtohinuria -128. 603
I;0,1, I i05, t;09, 1778. 178 I. 2066-8. caulc ll: 950 post-primary di,sscmina1ion, tubcrrulosls
20:1. 2072 pig, II; 935, 1378. 1382, Ill: 1623, !!J6, Ill; 1976
sh~ep/go.11s I!: 725, 729,736. 971. <heep ste Ill-thrift po>1-v11cdnnl reaction ill: 2056
111: 1-187 !695, 1697-170!, 2062.see porcine adeno,1ru, lnfcc1!on~ II: 822-3 po!il-weaning coUbacllluls. pig, Ill: 1623
, (I/Sf> rnncdi-,·isna porcine hnb~slo~ts 11, :?.4. 26. 435·6, 576 post-weaning diarrhoea Ill: IS63. 1569
pncumonic past~urdlo,is poccimicirco,·irn, 211 808. 810. 938 post-weaning multlsy~1emlc ,,.,1ing
ca1tle II: 673. IU: 1677-83 porcine cytomegalovlrus inf~rhon syndrc,m~ 11: 800. 810, 137-1-83
pig~ Ill: 170-1-S ll:!H9·2l. Ill: 1•19·1, Iii() Pour111c1rl11xm,r spe~ie,, l I: I 088
pneumonic ,igns. grial$ ll· 7-J.'-l porcine dermatitis and nephritis ,vndrnme Potomoc/1oen,s porcus ·135, 111, J.1-16
pneumonitis II: 9·10 1>ot-belhed ~ppearancc lllc:?019. 2138
ho~c, ll: 845 porcin~ .:hrlichiosis :\.13 Pmomac hor5de,cr583-9. Ill: 161:?.
pigs ll: 1307 porcine endemic diarrhoea virus ll: 782 pox ,iral diseos<.-s see names ofi1ul/l'idun/
pneumorrhagia. cattle UI: !i•ll. 1950 porcine entcric calich;rustcs) II· 703·6. 800 tliseoses
Pox,•iridnell: 1089. I lG0. 1265-1303, Pse11domo,1as sp~ctes Infection purulent embolic glonw1"11!onephntis
Ill: 2030 tn: 1~98-1506 Ill: 1653
PPD rubcrculln Ill: 1982 P.{eudomo,ws nemgi11osa II: 1250, Ill: I S57 pus. granul.1r Iii: 1649
pregnancy toxaemia 11: 1401 - - ma/lei see Bur).},olderia nwllei pusrulardcrmntitis, humans Ill: 1586
l'reisi-Kocard ab•ces.•ation Ill: 19li·25 - - mnlrophi/111 Ill: 1520 pustule{sl
premature birth 360,365 - - pseudomallei see 811rkho/d11rfa camels II: 1300
cattle Ill: 14~8. 15!3 pseudomo.lki canle:ll: 1291, 1296
ho;sc, 111: 1449 1>seudomrcetoma Ill: 2098 horses Ill: 1758
pig., nt: l4.i8 pseudorables pigs JI: 1293
premunity.: 12, 44 l cmtlell:9111417 sheeplgont~ 11, t2i8, 1284. 1287
Preston medium Ill: 1481 dogs II: 911 Puul'nala ,irus II: 1082
l'retorln Virus 15;, horsesll:911 Puumnla·like v!rus(cs) II: 1082
Pti/tlOWIIO me/o,1i110gi'llfca Ill: 1720, 17,J.J latent II: 91 l pyaemia, equids Ill: I 7i2
pre-weaning mortali1y, pigs II: 935. 938. see pig$ 243 ll: 777. 793. 810. 909, 15. 938, pynemlc hepalitls UI: 1617
also nronarnl death 983. 1309. Ill: 1632, 1778 Pymomo11(1S l9
prickly pear m: 1&18.1650 reac;tlvaicd II: 911 pvelitis. pigs !II: 1959
primary complex. tuberculosis Ill: 197~ sheeptgoats ll: 1401 pyelonephrltis
to:nplete 111: 1977 ps.:udorab,rs ,iru& see suid hcrpes~lru~ I conle Ill: 1931 ·4
incomplete III: 19;; , 1988 pseudo-nnderpest setpeste des petl!s horses Ill: 1931, 1933
primary fod. ttJ.bcrculosis Ill: 1975 ruminants pig,; JI: 938,111: 1958-60
prion dlWlse(s) 11: 1388· l-124 pseudorubcrculosis sre Yem11i11 species sheep/goat, JU: J65;, 1933. 193,1
prion hypothesiS II: 1388, 1391, 1392 infections p~'Odennn Ill· l 754
prions 11: 1388. 1391 psmaC0$1S 550 canine Ill: 1762
PrOC11via cape11si$ 378, II: 1137 P,·orophora 597 tail, hon-cs lll: 1757
proctiri.s Psoroptf'SOl'lsll: 1401
pyogranulom:,. soft tissues of head Jnd
c;mle326 pmrmlgan JI: 99; mouth !II: 1648
pigs Ill: 1603, 1604 P)'llgranulomatou, lymphat;!enif!s OI: 2012
P1eridiim1 ,pecics !II: 1588
J>rocyot1 loror395, II· 1128 Jl)'Olysin Ill: !946
Pt(>ridium aq11ilin11m fl: 755
progres,1,-c atrophic rhinitis. pigs Ill: 1492, p~'OmC,ra
Prl'ro11i11 p(l//t11s II: 1057
1494. 1707-J I . l 743 cauJe306
f>teropo,lidm• ll: 681, 682.. 692
progressive (001 rot Ill: I 734 equlds Il l: 1557
Pturopus (ldm,ra//1m11m ll: 682
prolapse 0fthird eyelids Ill: 1880 py01hora" Ill: I i54. 1950
--(1/ceco II: 682,688
prollfcrati\'e and necrotizlng pneumonia. P) re:da II: 820
--mpisrrmus II: 682
pigs II: 938 p11hlosis Ill: 21 13-14
- co11splcllfa111s ll: 682. 688 Py1hi11m i11sidios11111 lll:2095, 2113
proliieraiivc enteritis Ill: 1989
proliferauve emeropathy, pigs Ill: 1469·75, - - h.>·tm1cit1111,s 11: 693
- - il.l'J)(l1ne/111111i II: 682

CD
:605
prolifurati\'c haemorrhagic cmeropathy. - - 11eo/Jibemlc11s ll: 682
pigs Ill: I ~69 - - po/iocephalus II: 682, 688, 693
prolonged gesratlon ll: 990, I033, 10•19, - - smpulaws II: r,a2, 688
--1•11111p1r11s II: 693 Q fever.;. 565-9. 11: 973. 1058
'.058
pt:yriasisrosa U: 1294 quancr-evil Ill: L856·6l
l'romicromorrospora I! I: 20 I l
puff adder !II: I860 qumcr-m 111 1856·61
pronghorn antelope II: 820. l 195. I205, qu~mng (:'lt!ufeldl rc~c1lon l!I: 1535
:22,. Ill: 168!') pulmonary abscess.,rion, caule Ill: I740
pulmonnry:inthrax IU: 1803 Quercus spcc:11!$ Ill: I588
prostatltls JI!: 165i
pulmonnry arterial thromboe.mbollsm quinapyrnmine pro,ah 282
Prorelescrlstaws II: I 1:)8
Ill: 1950
quinapyraminusulphate 282
Pro101hl'C(I enrichment medium IIt: 2140
quinozolinone 492
Protothl!Cil ,peck1, Ill: 2 HO. 2142 pulmonary aspergmo~ls 111: 2116
Prototlll'l:a morifonnls Ill· 2140 p11lmonary cmbolic anc~m Ill: 1950
--s1ag11ora Ill: Jl.SO
- - u/11u!tl 111: 2140
- - 111icker/1t1mif !II: 2140
pulmonary embolism, cattle Ill: 1740. 17~2
pulmona,y emphysema LI: t 189
pulmonary form of horse •ickness II: 1235.
0
-::opfii Ill: 2140 1236-40 rabbit caliciv!rus II: ;oo
protorhecos!s lll.: 2140·42 pulmonary glanders Ill: 1502 rabbil haemorrhngic discas~ ,1ru, 11: 700
protOIOKin Ill: lll.J7 pulmonary infarcts Ill: 205 1 rnbbir oral papillom~virus II: 757
protruding 1ongu~ !II: 1895 pulmonary scquesira Ill: 1661. 1664, 20-15. rabies 227. 400. H7, 524,603, ll: 6&5. 914.
proud Oe$h II: 7;,!l 2051. 2055 1123·64 1309, 1370. 1401. l~I;
pruritis II: 1399 pulrnona,y cuben:ulos;s Ill: 1!+74 Ill: 1442
pigs II: 913 pulmonary vasculotropic El IV· I in!ccrion n.zoonosis II: ll2:I. ll51·64
prussic acid poisoning Ill: 18~6. seettlso 11: 844 see equid herpesvin,~ I nnd l can!t Ill: 1639
wider poisoning infecrions horses Lt: 849, 1010. 1020
Pse11do11/Tr$c/11,rta bo,vdil Ill: 210i. 2110, pulpykldncydfscase525. II: 1213, lll: 1841, rabies ,im• 11: 909
11 ll 1846-52 rnbies-r~lawd virus(t>.s1 II: JJ45.;
pseuctocowpox II: 11~2. 1ia2. 12B,1 , 1211~. puma II: 1~23 raccoon 395, II: ll28, Ill; 14•16
1291-2, 1297 puncmred ~le, catrlc 111: 1724. 1727 min roL 111: 2032
aszoonoslsll: 1291 purified protein derivative tuberculin min scald Ill: 2032
pscudoco\\'pox ,•irus II: 1291 m: 1982 Ra11giferr11ro11d11scMiho1t353, II: 795
pseudnfarcy Ill: 2104-7 Purklnjt cell des1ruction II : 999 - - mrm1d11s groe11/amlic11. 35.1. JI!: 15,:2,
pscudoglanders Ill· 210~· i purpura hacmorrhagica Ill: 1808 I5-l6
p~eudo-lumpy ,kin dbea~e see bo\ine equids II: 769,928, 1025. 12.11. m: I Ti2. --1ora11d11.<rar1111d11slll; 15.J:?. 1~6. 1689
herp~"ints 2 in fecilon li73 ra~h. see 1111der~kln rn$h
f'.reudo111011(1sspcdes II: 769 human• Ill: 1772 rat Ill: J.146
xxiv t'\Q::..'\

/lattus 11orvcg1cus Ill: 1146 rhesus monkey papilloma\'irus 11: 757 rimJeq;cs1 virus 11: 68i
llB51 ,"accine Ill: 1518. 1521. 1531. 154·1, rheumatoid anhritis in human< II: i43. 744 ringworm Ill: li55. 2os.;.2101
15,li Rhigioglotw a; Rocky Mow,taio gont Ill: 1995
reactivation rhinitis Iii: 2111 rod~nt-assod:m,d viru,{l'SI IL 1082. 1296
equid herpes,irus2 nnd 5 lnfcctfons caute n, 1322. Ill: IG3!l rodcn1s lU: 1542
II: 861 equids II: i68, 844. 661. 863.111: 17.'l roe deer II: 1339
p,cud.or:ibleb II: 911 ptg,ll:i76. m,9 19.21.93;, 111· 1494. rooipoo1bo,luisle) 22·4
rerrudc><:enc<' in equid hcrpe,wlrus I and l 1707, 1708 rooi\\aler $ee bo1'ine bab~o~ls
ln(ections 11: 836 pigs. atrophic II: 920.111: l492. J.194, Ro,e Bengal tebl 11!: 1517, 1518. 1544
rcc1als1rlcturc~ !II: 1603. 1604 1707•1 l.1 743 Ross River virus II: I02l
recto-vaginal lisrulas Ill: 1536 rhlnosporid!osis !II: ii I l · 13 rotavirus infection II: 12~6·6!
recto-,-aginal lacerations Ill: 15:,6 R/1lllos1X1ridlwn seeberf Ill: 2112 caulc326
,ecumbency rh11101rnchem, sPo infecli(>us bo,•inc shci,p/goats327.111: 159i
caule II: 1189 rhinotracheitis !•Cl)'!)IOSporidlosis 33~
shecprgoais UI: 1596 R/rmouirus II: no. 130i, 1319. 1322 ro;a,irust~s) II. ;0:1. 762. 800. 1256, ill: 1481.
recurrent iridocyc!itis 111: 1-H9 Rlripia.,phalm spedes 26-7. 537-11: 1071 1562. 1570, 1571. 151!8. 1612
red deer 11: 1339.111: 1618. 19;5 10i9. 1096 rougct 11!: 1908· 12
red dwarf b\llfalo 468 lllrlpi('ephalus t1ppe11dimlams 3. 18·21. 406. Ro11.,sem1s lleyJ!J/lacus II: 1145
red grouse II: 99 7 448.468, 472. ·178. 538,540,542 II: 668. Ro~~1I Fann virus II: 995
red hancbeest 516 996. 1071. 1072 Rubul,wirus II: G8i
red nose, cattle see lnfect1ous bo,fae - lmr,a 24. 438. 496. 5'12. 618 as zoonosis 11, 690
rhinotracheiris - - cnpemls 4S.i rumenuis Ill: 1736·41 , 1949
red-le,gged ticlcs22...t - - cami1,omlls 454 ruminul acidosis 111: 1725. 1727. 1739
reduced fcnllhy see infertility - - compos/111$ 454 ruminal aspergitlos1s Ill: 2116
reduviid bug~251 - - drmoni 21 . 454, •168 rund1:rpes sel' rimlcrpesr
red.water see bo,inc b~besiosis - - t11W1si c,wtsi 3. 22., 407. 426, -12; 478.
rulllS II: 979. 983
recd.buck Ill: 1446 ~85. 498, 5,10. 542, 596, 6111. !II: 1 135
Runyon's claS(;ific:mion Ill. H'l63
regional ileitl,. pigs Ill: 1469 - - l!l'l'ftsl mlmerlcus 22-4. 498. 618
Ruplrnwa ml>i,·llpm Ill: l 5·16. 15-19
reindeer n: 1339.111: 1542. 1546, 1548. 1669 -ft>llis Ill: li31
Rus,ian ;prlni;•summer <'llC:ephaliti, II: 995
renal failure. horses Ill: 1610 - - gl11brom1w1t1m 8, ll !: In I
Rus1erholz ulcer see ulceration. sole,;
Reotrlrid(la98·101. 156. 11: 1199· 1264 - - koclri 454
rcpell! breeders fl: 810 --lormslmr)'illl: l~l
reproductive disorders
conle 111: 1634. 1638·-l 1
pigs 11: J 307-9. lfl: lii'fl
- - /111111/a/llS 24
--m11/Js,111mwe26, ll:988
- - 11eumn1111i Ill: n:i 1
0
shcep,goats Ill: 1921 - - pme1e:c1mus 26 sable antelope 410,598
reproductive fuilure in pigs It: 687-90. 806. --prams454 Sabo virus 98, II: 1031
933·•10 - - pulehel/11.t 3. 2'4, •15·1. 4 78, 11: 1072 Sabourautrs dextrose a.gar Ill: 20!l6. 21 ~o
.r espiratory disease S}11drom.,. cattle - - sm1gu/1w11S 3. 26. ·128. 436 ~addle boil m: l 757
111. 16.~i-8. 16-lO - - sm1gr1/11~11ssa11g11111e11s IJ: 1235 saddl~ scab 111: 175,
respfratOT)' disease --sinws3. 24·6.436.5!)6, II: 1072. 1098 saimitlin<' herpesvirus II: 860, 896
cattle JI: i98. 820. 876· 7. 952, 953. - - 1t1ra11ict1.< •126, 428. ~35. 436,618 Saint Flori< vin1s II: 1057
Ill: 1635. 167.. 1678 - - lt'(IT/)/lr/Olli 3,26 salivarlon, Increased 11: 1189, 1196. 120,.
cquids U: 823. 84 1, 8-l4, 849. 860. 661. --Ulmbu::fensis3. ·168. 472,478 1228. 1691
Ill: 1319. 1-192. 210.1 Rhi::op,u Ill 2120 ialrnon fluke 564
pigs II: 696. 822. 909. 912,914.920. 934 Rlr0</ocoeciLt sp~des lfl: 2003, 201 l S11lmt1t1t'lia serovurs 34i. II: 6-li. 668. 951,
shcep,goatsll l: 1695 Rhodocormseq11/ II: 770,849. S&i, 1260. 126(1.111: 1481. 1520, 1594. 1602, 1608,
re,plralOT)' di~trel» Ill: 1612. li85,2015 1623. l,85
canle 111: 2049, 2050 - - et111i infection II!: 2015·22 Salm<m1dla spec!~~ UI: 151H, 157 I
plgSI II: 1630. 16&1, 2067 Rld<'llsft1 ,pecie., 8.3. SOS. 536·46 Salmo11ella ~p1'Cles lnree1io115 m: 15,8-1616
,heep,gouts Ill: 20GO. 2061 Rid:1m,ia ccmjwu:rlw,• .,ee (,()/e.sio1,; S;1/mu11e//,11\bonuscqui Ill: 1608
respiratOT)' S)~l~~ial \'lrus II: 687 .:onj1m,1iua,: - - Bavhmorbllkans Ill: 1582. 1589
horse$ II: 67i -co11orf•l II: 1082 - llrandcnburg Ill· 1<>85. 1586
sheep/goats II: 677. 821 - - iw/11eric11, 5·15 --t:hu!eraesuis !Ii: 1601
respirator)' trnct dlscase se<> under - - m'itut5-12 - Dublin Ill: 15 !3. 1582. 1608
respirator)· disease - - 11rowa:(•kii.509 - Emcri1idis 111: 158-1. !601. 1608
respiracon· tract infoc1ion, hors-Os 11, 1319 - - rlckeusli 509 - - \'1omcvidco Ill: 158.J
restnctlon fragment length polymorphism - - ruminn,uium 501 - - \luenchcn Ill: 1608
technique Ill: 2002 --1yplli509 - - :>lcwpon 111, 1582. 1608
retained placem~ II: 1254 rkkeusia.s83. 508, 536- 46 --Trphimurium Ill: 1513. 1582. 1594,
rell! mirabilis !II: 1948 rid a s,,e scraplo 1601. 1608
reticuloc)~l~ 60 I Rm \'allei' rcver IG2. 22i, 3-17. $<14. II: 98~. --l)-phis1,rs Ill: iflOl
rNkulocyto<,s 620 992. l03i-59, 1074, 1080, 12:H. lll: 1588 salmon~llose hy perde sve eqi.une
reunal desrrucuon. hors~ II: 644 humnns II: IO.l,, 104i. 1050. 1082 salmonellusis
retinitis 344 Rift \ ',lll~)· fcvur\'irus 137. 143. 15·1, II· 103,, sahnonellose vnn varke tee potdnc
retrovira! disease of cattle II: 708 Ill: 1588 ~almunello<i<
/lerro1•iridne II: 708. ,18. 733, i.J l rinderpest 58, 90. 225. 22,. 252. 25-l, 448, glmoncllosis 11: 1057, 125.J. lll, 1623
rerro,~rus(esJ 11: i08. 718,733, 74 l ·159, II: 629·49, 660. 904. 961. 1057, I' cryptC)sporidiosis 33.;
RC\'. I mutam srrain ,,accine ti!: 1531. 153, 10i4. 1123. 121!9. 1351.111: 14119. 2029 as 1.<mno~I~ Ill: 1582, 158.\. 1598. 1601.
rhabdomyoly,ls Ill: 1772 and foo,·and•mottth disease II: 1326 1603. 1604. 1612
Rhabdot•irid"e 101·2. 156.11: 11 2 1-98 small nuninant-adapwd II: 66l! ca\!le see,bo,ine salmonellosl:,
rhe,us monkey 395. 11: 1423 vocclne242 ho=s see equine salmoncllosls
 pig, see porcine salmonellosls scouring Ill: 21S8 Shuni\in,~98, II: 1031
~hccplgOal> see o,ine ~nd captine screpfcUL 11:738,999, 1388. 1391-1404, ;ialoglycoprotcin II: 1368, 1389
salmonellosis J.108 sick habitat syndrome 232
salpingitis 30i, 308. Ill: l4Gl. 175,1, 2076. scrnpie-assocl01ed fibrils II: HO!, 1415 sidc-s1rip1•d jackal 11: 1134
2078, 2079. :?130 scrapie-like a~cnr II: 1409 sigma virus l 59
salpi11goi,critonitis Ill: 2078 scratch reflex JI: 1400, 1413 Sigmotfo11 llispufus LI: 821
salt poisoning II: 1309. Ill: !57!.see also strew-worm fl~· 8, 2 l. 67 slka Ill: 1975
111uter poisoning ,crofula 111: 1973 <Uaie tn: 1904
sambar deer 11: i98.. 799 ,crub hare5l6 sih er dollar plaques. cqulds 299
Samorln 282 ,~a lions 11: 700. 1351 sUver-halred ba1 II: 1128
San ~1i1.,'Uel sea lion ,irus II: 700. 1351 sen.ls 11. ;oo. HI: t 5-16, 1549 Simhu group \'irus(esl 98, 155. u· 1029-34,
sandcrack 111: 1727, I 743 selenium dcfiticnC) II: 738. 1311 105i
,andfiies 1Si, 378, II: 1195 s~llers merhod II: 1157 simian agent I 11: 1256
sandfly fever 11: l057 Sclous' mongoose II: 1137 simian haemorrh~gic rever vims II: 92,l, 933
Sang<l\iruS II: !031 semen, poor quality Ill: 1530 simian immunod.eficicm:y,irus II: i!O
S011o1•inl$ II: 700, 703 •emln;il vesicul11is s)'t'ldrom,e 555 Simpl~x11m1hpecies n: 887, 889
Sapporo-like ,irus(e,) It: 700, 703 scminoma lll: 1531 Simuliidnc II: IO 19. l 1%. Ill: 1755
saprophytic mycobactena Ill: 1983 ~cmliki Forcs1 ,·irus l4i, II: 1023 Slmull11m species see Simulildae
Sarcacys1is species 34 7. 360·62. 366. 3il. Semple•l}l)C\'ilCCll1C II: 1!23 Sin ~ombre,llkc vims(esl II: 1082
JS.I Semdo spec:es ll: l 05i. 1158 Sindbis,irus 137, II: 1058
Sarcacysris aricr:icanis 366 Senkobo chsease 35i, Ill: 2026·35 sinusius Ill: 163!1, 17i2. 2JIO
- - ucnmm,363. 3b6 sepuc anhruts. ca!Uelhorses Ill: 1723, 201~ $1phora591
- - bt,ffa/011is 366 septic 1rawnatit- pododcrmatltis Ill: I i2, sitatungn II: 798
--tanis3il sepricacmia 11:849. Ill: 1'1~9. 1557. 15(',0, sitfasr. cattle II: 1271
--cap,aumis 362. 363. 366 1582, 1586. 1595. 1601, 1603, 1608. ~itsiektc HI~ 1951
- - cru.zi 360, 363,366 I ,84. 1788. 190.1. 1905 skeletal drfects. callle U: 953
- tlubeyi 366 canle 111: 1620. 1634.1635. 1690. 1691 ,kin discolonuion. pigs Ii: 979.111: 1910
--tq11ict111/s 366 cquidslll: 1653, 1670. 1n2. 178-1 skin di~rclers Ill: 2034
--fr1/c11rula 37!. 395 haemorrhagic 275,544, Ill: 1689-92, 1807 skin h~iperaemla, pigs II: 979
-fayeri 366 pigs 111: 1630. 1652. 1670. 1nG. 1,s1. skin l~'ll1phoma 11: 708. 7 l3
--fusiformis 361> 178~.1911 <kin pet<:chiation. pigs II: 979
-g{gtllllefl 362, 363. 366 shcepig03tS Ill: 1620. 1655, !65?. 1699. skin plaques. equlds 21l9
- - hnrd,mgerl362 20,16 ,kin ra,h Ill: 1,14 I, 1,142, see lllso el')1h¢1na
- - i1ircico11is 366 Si?prlcaemic amhrax 111: 1804 mig,·ons
- hfrmta 363. 366 septicae1nic b:icterlnl infections. pig, II: 777 horses If: 927. 1024, Ill: I75i
- - l,am/11($ 363. 366 septicaemic disease Ill: 1570 humans nr: 1513
- - le11/ni,/ 366 horsc,111: 1610, Hitl.1652 skin sen;itlvity 1cst I U: 2003
- - 111ed11,iifom1is 362, 366 pigs/cattle Ill: 1109, 1570. 1637, 1653, Sk.irrow ngar Ill: 1463, 1-181
- - mlescheriar1t1 363, 366 1704. 1910 skunk 395. ITI: 144G
- mflwmsfs 366 shecp/ go:u.,, ITI: 159•1 slangkop 328
- - 111011/e, 366 scpr.i~aemic Esc/rer,c/iia <-oli Ill: 1561 slapsiekte 2.~7 ·302. see nl.$0 dounne
- - lll!'lt(Q/Ul 371. 39,l sepucaemic pas1eureJlosis II: 1109, Ill: I ?04 sleeper si11drome Ill: 1634, 1639
- porcifelis 366 sequestra. lungs Ill: 1661. 16&1, 2045, 205l, sleeping sickness 25!. 25.~, 265, :!GI fl: IOHl
- - mngiferi 362 2055 sleepy foal disease Ill: 1652
-spei'rl395 scrofibrlnou:1 polyS) noviti~ II: 1190 slender mongoose II: 1135, 1138
--sulf1ominis 363, 366 serologic:11 cross-reactions Ill: 1623 Slcnkdnlko<)rs see Rifr Valier f(wcr
- !!!11Pfla362, 363,366 S~fO~itiS, pig, llf: 20;'! slipper fonnarion II: 1208
safcocys1o~is 360· 71 5~rp11/i11a see BraeJ1yspim slit-faced bn1 II: 1146
sarcopric mange 357 sliaker foal syndrome Ill: 1892. 1896 slO\\' (mbcJ agghuinaiion tcs1 Ill: 1517. 1518
Sorrruu>mmn vlm!11ale 4 l 7, 524 Shamond~ ,~ru~ 98, f I: 1031 ,lowly C)-Wpathlc oqu!d herpcs"\iru,
Sathuperl ,1ms n: 1031 ~heath ro1 ITI: 1931-4 Infection.< 11:860·66, tu: 2017
Savi Ill: 2026-35 shedding. ,:>quid herp~>'1rn~ 2 and 5 11: 861 ~lt,wly cytopathlc orphnn hcrpesvirL,
St'allby mouth see or! sheep head Oies8l 11:860
scabby ulcer Ill: 1931 ·4 $hccp ked 618.11: 1205 small &r'-'l' mongoose II: 1139
scab~ Ill: 2030 •heep pulmonary adcnomnto•iS II:; 1,-30. "mall ruminan1-adap1cd rinderpest II: 668
scabs i33. i38. 1397, l-102 small ,moom boot-legged tick 15, 16
cattle II: 129!. 1296 shcep-poi.andgoaipox 11: 1277·80, 1285. small weak lambs II: !)70, seefllso 1111//e r
hQrses nt: 1758 1288 w~ak
scald Ill: 1723. 1729, 1735 sheeppox and goatpoxvirus II: IW9 smallpox 239, II: 1123. 1293, 1296
scarring. nasal mucosa, cquids Ill; 1502. sheepµox cclls II: I 2,9 ~mallpo~ ,'iru~ II: 1300
1503 shelly hoof Ill: l 733 small-sported ca, II: 1139
schlsiosomosis !ti: 2112 shepherd's disease II: 1284 small-sponed gcnct II: 1133
Schn\lffelkrankheit Ill: l 707 Sh!ga10,cin Ill: 1561 SMEOI syndrome If: 806. 810. 920. 1307-9.
Schon1ci11-Hc11och'> µurpum Shigu u»Ju-producing E.cllc, ic/,/11 u,/i 1310. 1311
haemorrhagica, humans 111: 1772 Ill: !561. 1566-7 Smimbum strnin \'accine II: 1050, 1058
S<.'h\\'nrt2man rearcion Ill: 1739 Shiga-like toxin Ill: 1561 smooth brO\\'ll tick Ill: 1,31
scimitar-homed oryx 5J6. II: 1270. l42J Sh/ge(la eq11fru/13 Ill: 1652 snake bile poisonin!\ Ill: 1807. see t<lso
scirrhous cord Tll: 1756 shipping rc,·er In: !677, see nlso bo,ine 11m~r p0Iso11lng
sclerodcnna. canle 355 respiralOI')' disease complex snake venom Ill: 1860
sclerosis, hippocampal II: l~li Shokwc ,1rus 155 <neeilng, pigs II: 776. 792. 920, ill: U&l,
sclcro1ic bodies/cells 111: 2109 shori-talled \'Ole ll: 998 1708
Sclerotlnia sc/erQtiol'um II: 1351 \hre1~ II: 11~5 snoring. caltlc 111: l gss
snotsickte sel! malignant cmarrhal fever S1aphylococc11$ ,pecic, infections - eq,11 infec1ron U· 769. -:--70, Ill: 1504,
snowshoe hare 553 Ill: I750-62 150~. 1768-74
snuffling. pigs LIi: 1708 ,hccptgoais II: 128~ - - rqui ::-000pidomims 11: 8·19
Sofjin virus 154 Smphrlococcusmtl'l!IIS Ill: 1739. JTS.I. 1750. - eq,dsimills 11: 769
So/,11111mspeties II: JOJO 1841 --pord1111slll: 1781
So/111111,n kt1'i'bet1$C II· I •l Ii --auri,uslnfcction Ill: 17M·8 --porci1111$ infcction~ Ill: l i81·3
sol~ ab~ce~s see 1mder ab$c/!Ss(cs) - - llllfl'11$,tS1.00llOOi< Ill: 1754 -Wis 11: 785, 91 I. 914,937.111: 1631,
\O~ ulceration sl!e 1111der ulceration - - ,1rromogm~ 111: 1762 1776, l i81
~omnol~ncc. canle !11: I639 - - ~pld~rmidls Ill: I i6l! --suis infoc1io~ Ill: li76-9
South Afrkan bom tld •I ·8 - ,,_,•kt/$ 111, 1759 - suis infocrions. a~ zoonosl5 m: 177,
South African o,ine maudi-,•isna virus. - - imermedi11.< !11: 1762 - - :ooi:pidl!llllCIIS II: 769, 770, 1250.
goat> II: 734 --simultms 111: 1762 111: 1708. li84. 1785. 2121
Soulh Mrican territories types of S1ei11eml!ma 31 S1rrp1omycessp~ics Ill: 210,
fom-and-nwuth diseas~ \'irus I!: 1325. ;tcllatc scarring. na$3I mucosa, equi<l$ strcptothricosis Ill: 2026·35
132i. 13·10 Ill: 1502. 1503 S1rep1orl1rix see D,•muuophi/Jts
Soulhcrn sea otter 395 s1enosis. trad1cn. cau le II: 1273 ;tress fll: 1995
spasms. muscular 111: 1880 S1eplra,1ofilaria species 7!1 <mule Ill: 16,8. 1680
specific pathogen-free herds Ill: 1630 stcphanofilariasis 79 pigs Ill: 1630
,pecilic pnthogim-rrne pigs Ill: 1629 sterility striped polecat LI: 1138
spectacled fruit bat LI: 688 caulcnt: 1461.2151 >triped weasel II: 1138
spenn.abnonnal 11:937, 1189, Il l 1529. J)ig> Ill: 1542 strongylo larvfil mlgran< !11: 1612
1530,2076 sheep Ill: 1530 Slf(mgyloidi·s pap//los11, 325. El: 1734
spermatocode lfl: IS29. 1530, 1531 stlffn~, Ill: 1-141, 1442, 1880 s1rongylo~is 279
~phingomyelinast>·Specific phospholipa,;e cattle 1637 <truck 111: 1840-.14
D ll!; 1919 pigs lll: 207 I. 2074 "1rychnine poisoning 11; 1158. uoe also
spinal abs<:essatlon 4 77 voluntary muscles Ill: 1880 1111der poi,oning
spinal cord h)'pomyelinogen~sis 11: 95.1 ,tlllhirlh II.I: 1904. 2 121 Sruan's mel.fium Ill: 1481. 1631
Spiroduuu, see Borrtlin canle ll: 953, 1221, Ill: 1«2. 1452 study methods 189-210
spirochaetes Ill: J42 7· 55, 1721, 1743 hor,;es 111: 1449 stunted pig.< 111: 1130
spirochaeto~i, see Borrelia 1/reilerl Infection plg< II: 688. 777, 81)9, 9,35. 979. ill: 14.J8. stupor II: 1400
spirol:etose see Borre/la 1/reileri infeclion 15-12.1543. 1910 Sl\'W~Si~ktc If: 1191
Spiru11/1umu species 111 : 20·16 ,heep/go:us 347, II: 970. l 050.111: 1448, subcomeal p ustular dermutlth. pigs
Splt'ndore,IIOcppll rn.nerial Ill: 17S6. li5i, 1596. 1620 Ill: 175-1
1943 scomatitis ,,1bcucant:0us emphysema II: 1189
Splcndorc-Hoeppll pheno,nenon Ill: 2109, caule II: 1205. 1221 ::ubc1uancous ruberruli n resr 111: 1984
211121)3.2llij.2121 necrouc Ill: 174 l subcutaneous 1.ygomi·cosis fl!: 2113
,plemc fever Ill: 1788·1811. see also amhrax ulcerative Ill: I7•1J sudden death Ill: li88. 180,. 1829-33
splenic infarcts, pi~ II: 9r!I stomalitis-pneumoenterit,s,complex see cattle Ill: 1637. 16-tO. 1691.1,42. 1848.
sph:,nomegal)•521. ll:750. Ill: 1'149. 1450. pest~ des per its rum,nants 1858. 1874. 1875
1587. 191 I. 2158 Smmo.11:rspecics 77-80. 254.11: 1112. 1205. eqwds Ill: 1866, 2090, 2091
horses m: 1611 1270. 1293 plgs H; 696, fll: I704
shcc1)tgoars III; 1,;95 51omoxyscalciJr011s 77. 80. 575, 59,, ll: 888, sheep/goats Ill: 1701. 1842 1848, 1866,
Spondweni ,irus 1,;5 1098.JII: 1488. 1798 18,4
sponglform chang<'<. brain 11: 1388. 1400. --11/ger80 sulcl heriics,•irus 111: 690. 829. 909,921.
1413. 1416 <tonerues 584 Ill: 1630. 1778
~pongifonn cncephalopath)• II: 1156 StOOl~fekte 47, ~uld herpewlrus 2 infection IL 9 19 -21.
sponssickte Ill; 1856-61 Stormont test Ill: 198'1 Ill: 1494, 1710
sporadic ho\'inc encephalomycliri~417, strain 19 ,11ccinc Il l: 1520·21. 1547 lntCllC)' 11: 919
477. ,;ss. 11: 115s, 111: 1(\39 strain 2 vnceiM Ill, 1522, 1531. 15,l-l Suld-Afrlkaan;e hontbo<lu!,4-8
Sporadic bo,,nc lcukosis 11: /Oij, ; 13 $tram 45/W ,'l!cclnc Il l: 15l I sulphur grnnul(ll; HI: l&l8. 19,14. Wl2
sporid6min Ill: 1755 strain RB51 vaccine Ill: 1518, 152!. 1531, sumrnermastltis, canle Ill: 174'1. 19.;;
Spororhrl-r ;(/ienkii 11t: 1504. 2 102 15'1-l, 1547 summer rash. horses ill: I;.;;-
sporotrichosis 111: 1504, 2102-4 sm1in SRB5 l vaccino 111: 1547 S1111(/asd11rus 98
,poncd hyenu II: 1133 strangles II: 769, 7i0, ll!: 1504. 1505 suppuration
spri11g lamb paraly:;1~ ·1, :!.2 I 768•i 4 nasal mucosa 111: 1:;(r.;
,pringbok516. 553, II: 1046. 1270 $!ranguna lfl: 1959 of nodules fll: l 649
Sp11mm•1rlnae II: ii, trauss reaction Ill: l:>O•l supra-allan1al bursitis sea poll evil
squamous cell carcinoma II: 755-61 strawberry footrot 11: 128-!. Ill: 202 6-35 suprnspinous bursitis see fistulous \\1thers
squamous papillomm; II; 755. 758 straw-coloured irulr bat II: I 145 S11rfcara s11rira11a II: 1135
Sr Louil< encephalitis 137 streaming macrophages Ill: l 705 suricnre II: 1135
St Lou!; encephalitis \'iru~ 154. fl: 100-I Street ·\labama Oufferin vaccine strain rurt3 251, 2.5.1. 297
stable Oh)$ 77. 80. 575. IT; 869. 888, I098. 11 1160. 1164 S1/SUf(/[afer11.S II: 1095. Ill: 15~2. 1546
Dr: 1488, 1798 s1reptococcal Infection~. hors('~ II: 9'.!8. --scroft1 Ill: 15'12. 1546
stable foot rot Jll: 1723 Ill: 1612 sw,dlownc~rbug~ll: 1017
sraggcr srndrome Ill: 1848 $lreptococcal lvmph;l<Jcnith of s1<tine ,wullowing difficuli)' SI.';> dysph:1gia
sr~Mic<: see ~table ni<'< 111: 1781·3 swnflo"inJl reflex lo!!SII: 1189
Stamp's modlficatlo,i of the ?Jehl-~eelscn S1rCf)/OC()CCll$Speclcs 11: 8-19. 1250. 111. 1452. swnmp cancer Ill: 2113-1~
Stain Ill: 1510, 1515. 1517, 1530. 1537. !5,7 1784. 1841 ,wan'tp fever - equine Infectious anaemia
1(;57 S1reproco«11sspcc-ies infections Ill: 1763-85 swayback. sheep II: 973
<taphylococcal dtmnntitis iII sheep Ill: 1754. SIN'PIOCOCCUS agnlnctitll• Ill: l i8 I sw~at flie~ 81
1755 -6 - dy~galac1iae lll: 1781. 1784 S\\tuting~ickness •I. 15, 35i
s,a,,1iylocorc11s,pccie~ 11: 769 - equill: iiO. 864. Ill: 1768 >1,e~rklou Il l: 1730-32
~w~et Itch 98 1achypnoea Ill: 2138 - - serg1mri 486, 500
swectsiclaebosluis l 5· 16 Taenin mullf,w,1,, coenurosis II! I I5-8 - - rnurorragi21. 22 . 2,1. 449, 4S8. 459,470.
Sweet"·ater Brunch nrus 99. 102 --40/111111 ii: I I.SIi -H5. 4 78•9, 480
swelling. see also oedema tail pyodemia. horses !II: 1757. 1758 - - 1't!/ifera 454. 459, 483·-1
in horses. fnce II: 1236. 12-19 rail shedding. pigs Ill: 1910 thcilerlosis 3. 498· 9. 54-1, 603. Ill: 1639
in horses. head 11: 1236. 1249 rampan, eyeless 27, II: 1088. 1096 annulnta l7. 18, 486·93
in horses. lymph nodes. ($uhmandibular> 1'auro1mg11sory.,;;16. II: 142:1, Ill: 1512, t ... rcbral see cerebral thcileriosls
111: I~oz. I I il J:,..16 1roprc:,11,. 18. 486-93
Joint< II: ;:iG. 1105. 1189. 111: 14-1 t. ins. l'tt,vlor.t/111 cquigm/r(l/i~ 11: 872, Ill: 1557. 7.lmbabwc 21. 22. 4M. 468. 472·3
1951. 2071 2084 r;,,•la::ia <pccie< Ill: 1489
l)111ph node~. mc,enteric Ill: 159; - - e(l11~11iialis Infect ion 111, 2084· 7 1h1mnally dimorphic fungi 111: 2095
in pig,. lymph node, lll: 1782 TB-grape, lit 1978. 1985 them toxin Ill: 1864
in -sheep goats. he;id HI: 1863. I BHS TB-pearl~ lit, 1978, 1985 thiamine dctidency
in sheep/goats. tongu< II : 1207 tear•marks. pigs Ill: 1709 ~aide II: ll58, 111: 1639
submandibularsubcutaneous Ill: 1691 tendo~-yno\id~ she~pll:973
teats U: 889 cattle Ill: 1691. 2052 thick knee II: 743
swine i.lyienu.·~· 11: 983.111: 1428,33, 1-175. equid, m: I i68. 1,72 thimble 111: 1,27
1605. 1623 tendovagmni, II: 1190.111: 1513, 1608 Thimlri virus 99
S\,ine cmerlc calicilirm(e,;J II: 703-6, l\00 l<'ratogcne5i$ II: 946. 950. 973, 1029. 1033, thin ~we syndrome Ill: 1917
<;\\ine ~nzootlc pneumonia Ill: 2Q61>·8 10.:6, 1030. 1057. 1058 :hin ht'ad form or horst' sickncs, 11: 1235.
swince~·,lpelas Ill: 1908·12 teratology II: I2J:l l2-36· 40
swine Ou 11: ns-7. 91-1, 938 reschen·Tatran tnfecuon n: w:i. ~14. thin sow syndrome IU· 1960
~wine infeni11ty and respiratory syndrome I 307 •9 . 131 l. Ill: I 632 Thogoto ,1rus II: !19i
see pordnc reproducuve nnd ,cs, and immediate sl~ughrcr method fhog0to virus inrcetion II: 1057
resp,mtory srndrome 111: 1986 1hompson's gazelle 618
teSt(sJ see 111rder /111liuitf,ull names
swine h1Uuenza 11: 775-7. 91,1. 938 three days IL: 1163
swine v~rulnr disease U: 701, ll97, 130,. 1cranolysin Ill: 1878 1hr~·dapllff.;ickness 99, 101 If: 1183·92.
tetanospnsmin Ill: 1876
1313-17, 1351 1225. 1229
1c1anu, II: 685, 1010. 1158, Ill : 1878· 83
,wine ,-esicular disease ,,rus It: 1313, 1351 thrombi. lymph111ic.s 111: 2051
1 Cit.' \'fru< 155
swineherd'< disease Ill: 1445 thromb~10pen[n
1etr.u:yclines Ill; 1831
~wincpox II 1293·-1 ca11le If; 949. 953
w1rapnres!s Ill: 1951
~\\in~o:niru, II· 1293 p[gs JI: 977, 1102
Thal Rt,>d Cro,;s !mrudermal ~cht'dule
swollen heud form ofhon<csickncss chmmbocmbollc meningoencephalitls
II· 1163
II: 1235-10 Ill: 1657, 1906
Thayer-~lanin modified ml'dlum Ill: 1537
swollsn head i,1 ram, Ill; 1863·5 thrombo<'mbolism
Tl1cllerln species -1. 441, 6111
sylvar;c l"Jbies II: 112•1, 1127, 114·1 ii inc anc~· ll:10:W
/'helleria sp. r~ larula) 5ll0
Syh-icnpra gr/mm/a 5+1. 598 pulmonai,· artery fll: 1950
n1ei/cr1n 11m111/am 17. 18. 458. 48 l. 486. 5·10
Syncen,s cnffor 4.J8. ,168· 70, 11: 820, 876. rhrombophlebitis
- - am111/n1t1 rhellcrlosls 17, 16, 486-93
104-L 1325, HI: 1512. 1546, 191'1 c~nle Ill: INO, 1742, 1950
- - lxm:em -180
- - mffu 11a1111s 468 horses Iii 1610
- buffeli 412. -154, 458. 500
synt;iia n: 67•1. 678, 681·4, 687,695, 733.
- - b11jJ'tJlitommialls infccuon soo.:;o1 rhrombo,t,
738. 74-l. ,80. 791. 796. 1187, 892. 896. cattle II· I l!/0. Il l: 1588, 1637. 205:!
- - f'(llli 4. 17 22. 24. 26. 425-32
937 ccrebml475
--eq11/infoc1ion SPP equine
•i11ovial mcmbran(!S U: 743 meningeal 475
piroplasmo;is
S)'n(>Vltis
- - ltirrf-198 sheepigoats II: 1279
in raprlne anhriti5·enccphaliris II: 74 l 1hrombo1ic numi11goenceph:ililis. c:i:tlc
- - lllwrenai 468
cnnle 111· 15s;. 1639 Ill: l63-1 1637
- lcs1uquortfl 498
pig~ ru: 11as
- - mmnus-154, 458. ·159. 470. •li5. 4i8. 1hymic Jrmphoma LI: 708. 713
systemic adcno,·irus infection II: 820
483,486,500.603 thvrofd. lmpnlred runctlon II: 9SO
systemic di><last?, dogs m: 2140
- - n1111a11s infection 480·81 T!brogargan viru$ 99. I09
sy,rcmic disease,. ~hoop/goats Il l: 1448.
- - orlmtalis 48-0, 500 tick tontrol 27-34
1695
- - or/rmofls infection ,;00.501 tit'l. pnr;ilysls llf: 1755
systemic mycosls Ill: 21 l4·:?4
- - 01•/s.98, 578, 111: 2157 tick pyaemia 5~5. 5-16, Iii: 175·1
~y;tomic pa.,u,ur.-llo$ls Ill: 1695, JiOI
--pnn-o3. 21. 22, 448-61 .475. 418. -180. tfd. toXiCO$h ,I, l8, 21 . 22. 24, 26. 232
483.4SS.500 rjcJ., typhus -I

0 - Pflfl•a Boleni 4i3

--pn11:a bo1 1i.t472
-pnr,,a bo,•is mfeccion 21, 22. ,154, 468,
ticl-bhe fever ,I, II: 1082
tkk·bomc encephalitis 90, n, 995. 997
rick-borne encephalitis virus(l!SJ lll: 1440
Tl /.J-1 ,;mun lll: 2056 •172·3 tick·bornc fever 544·6. 11: 99~
Tabanldae see taban,ds -parmlJa.mbu-161 1icl:-bornc flavivfrus II: 995
rabanids 87·9l. 156.353.111: 1798. 2029 - - µan•a lawrencel 466. ,17? tid-bome rn,hus IT: 1002
Tobon~e/1(18, - p,m•o /awre11ct•i infection sea Corridor ricks 3·34. 111: 1949
- - tlfrrlli<'hflli.s 90 disease a.nd Caxlu//a /Jur,w1ll J67
Tobnn11sspecies87 254 597, II: 1205 - p11n'll ~lt,guga 461 ars&sid 3. 11: I088. 1095
Toba1111< abac1or9 \ - pfln·n pi1r1·11 lnfec1ion "'"" East Coast as ,-ectors 4- 34. 6-7. Ill: I i'98. 2029
- - blgiuum,s 88 fever Lxod1d 3. n: 996
- - wpema11i 91 - pttn•a S<>rtengeli -161 tiger hean disease II: 13-16
--11lgro11i110111s91 - p11ri,a, cra-nsplacenml infcetion .:;o Tfnaroo \"iru5 ~9. 11: 1029. 1()31
--,111llulus91 - rn:oncfh1• 498 10~ ab~ctss, sheep/goats Ill: I 729, I i33,
- - 1aw1iolt1 88. 597 - - .<~para ta 22. 24. -15-1. 483, 485. 498. 5,8. 1755
rach)'cardfil Ill: 2138 618 1'<1gm,iridt1c 155, II : I OU-I, 1013-26
tok.wplasmose 3;1;•.19, see also trembling - - ec111T11um 25-1
toxoplasmosis piglets ll: 983. 131 l - - e,q11ip,•rdt1m$O. 25 I. 256. 279. 297
mngue protruding UI: IS95, 20~9 sheep ll: 991! --fl'll/lS/q!), 90. 251,254, :?56. 297
tonsillitis II: 913 tremors. muscular ---.<imiae-49. 90. 256. 277
lOOlliWCer Ill: 1733 cattle IL: 1 189. ill: I637 - - $l1is 255, 258. II: 1110
tortlcollls JI: 998 pigs Ill: 16.11 - - ll!Pi/~r/ 69, 90. 25'l. 261. 269
To scana ,1rus II: t 039 T1\'µ0111'11111species Ill: 1440. 1721 - - 1J/1•ax 49, 90. 255. 256
toxaemia - - h_1v1I_V#1111en11I' sl!e Bfl1Ch;~p1m - - 1·/M.<. acute infection~ ln C11Hli:' Zti6.
cattle ll: 877 /1.1·orlyse111erlt1e 275
pregnancy U: 1401 - - thdluri see Borrolla 1/a•i/~ri trypano~omes 49. 251-87. 603. 618
toxicoinfoctious hotuli;m m: 1892 ueponema.Jike orr,,;inbm Ill: 14-JO trypnnosomosis 43. 68-9. 90. 187. :?25. 22i.
toxicoinrectious syndrome 111: 1896 Trin1011w ge~meckcrl II: 1098 251,87.481.5.J0.5'14
toxicosbsee muter poisoning Trlbu/11s INresrris II: 987.111: 1736 cattle 21.17 •75. Ill: 2029
toXin(s) Tncl1011to11ns vagi,wlls3 I() dogs279-80
alpha Ill: 1830. 1840. 18'11, 1857. 1864. rrichomonosis305·12. Ill: J.152 equids 278-9
1870 Tricl1ophy1011 specil,s !11: 2095 humans 251. 255. 261
beta 111: 1837. 1840, 185,. 1870 Tric/10µ'1.v1011 f!</lli1111111 Ill: 2096. 2097 pig,~ 277-8
botulinum Ill: 1880, 1887 • - - me111agro11h.vce; Ill: 20fJ6 ;.hecp/goat, 275·7
cholera Ill: 1565. 1@3 - - 01!/.< Ill: 2097 tryl)anosuscepribillt)· 265
c~~o· m: 1561 - - ntbrum Ill: W96 trypano1olc,rancc 68. 252. 263. 265,266. 287
delta lll: 1840, 1857, 186-1, 1870 - - 10,w,mm Ill: 2097 Tr:,pa11o=oo11 ~9
endo· 111: 1568. 158S. 1603, 1619 - - t~·m11·omm 11: ilO. (II: 2096 tryptophan. dietary I. JI· 1191
cntero• Ill: J561.1565. 1566. L585. 161~. - - uiolaceum Ill: 20!J7 1sct$C mes ,l3-69, 251
1829 Trlchostm1s 1•11/JJ<'c11/t1 Ill: 1973 as vectors 43,69
epsilon Ill: 1846, 1847. 1850, 1864 u-1,mus Ill: 1880 biology 43·9
CUI Ill: 186-1 Trltrichomcmns Joetm 305
control 57-69
l!XO· 111: 1619, 1677, 1801 --fovll,s Infection 146,1 distrlbut ion 52-7
gamma 111; 11>57. 1864. 1870 - - mob/lcnsis305
tsetse ny disl:a,c 25l -8i, see also under
heat-labfle UI: 1565 --s1,u305
tl)1)311050lll()$h
heat-stable 111 , 1561. 1565 tropical bont tkk 8-10
lcuko· Ill: 16,7 uopical canine pancytQpcnta 280 tubercle Ill! 1976. 1977. 1989
mulll: 18H tropical horse tick 42i tuberculin Ill: 19i'll. 1982. 1989
neuro- Ill: 1878. 1887 troptaal theileriosis I7. IS. 486·93 tuberculin wst. subcutaneous rn: 1984
ochra· .-\ lit: H52 uoples~ bontbosluls 8· IO tuberculosis 241. 111: 1650, 197'3·89, 1996
oedema d,sea~c Ill: 1567 Trypaddc prosalt 282 as zooriosis Ill: 1973
Pas1e1trf'lla mu/10<:ldn 111: 1,o7 T!ypam1dium lll2 cattle Ill: l9i'3,87
pnewno·3·methyllndole U: 1191 trypanocides 28(),87 horse;; Ill: 1989
prot0• Ill: 1847 llcrenll 21\2 pigs Ill: 1987•9
Shiga Ill: 1561 dimidium bromide 281 shce1>lgoats 111: 1989
Shign·Uke Ill: 1561 diminazcne281 wildlife Ill: 1973. 19i5
lheta m: 1a&1 di:ninazene nccturate282 tuberculous cndomc1ritis 111 1974
\'cro· 111: 1561 F.lhidium 282 tuberculou$ galactophorfti~ Ill! 1980
7.Cta Ill: 1864 fnilure 283-5 tuberculous gronulomas Ill: 1976
ToxoplMmagondii 337·48, 371,382.397, bistory 280..S I tubecc:ulo11s lymphadcnltls 11 !: 1987
938. 1623, see also toxoplnsmosi; homldium 281 tuberculous mastitis Ill: l9i4 1980
toxoplasmosis 33i-UI. II! 973, 1158 homidium bromide 282 tuberculous mcningills m: 1979
Trnbctkrankheit SPesc:rnpic homldium chloride 2112 tuberculous os1comyclltls m: 1978
rracheitis isometamfdium 28 l. 282 111llcrkulosc see w,du tub~tCJlosis
cattle-Ill: 1638, 1639 No,idium 282 tulp328
plg,111: 1781, 1782 phenamhridlnc compounds 281 tumour(Sl 11: 7 11. 72 l. 723
Trt1g,,1aph11s illlf(llSii Ill: 1446 quinap>·ramin<' prosah 282 Sertoll·ttll lll: 1531
- - scriprn,516.544 guinap}raminesulphate 282 tUrl:e) corona,irus II: 791. 795
- SfNk~ill: 798 Samorin282 turkey rhinotrarhe!t!~ virus 11: 687
- suepslceros II: 1132, 142.~.111: l9i4 strategic use 28~ turning $lrkness '175- 7. see t1lso cerebral
uansfronticr eonservation aroa~ 232 rre:nment 280 theileriosis
uam,lt ft1\'Cr II: 769. 770.111: 150,1. 1505. Trypacide 282 twin lamb liiseasl' 11: 1401
1768•74 Trypacide prosalt 282 type A innuenru virus ll: 166
tran,misslble ga:,trocmerltis II: 780-83, 784, T~'Pamidlum 282 t)1lC B retro~irus 11: il6
787. 1261 usage principles 281 1ype <: oncovirus II: 718
transmissible mink encephalopathy use in catde, goats, sheep 282 rype O retro,1rus 11; 71 II
II: 1388, 1421 U$Cindogs2ll3 t>'Phlilis
transmissible spongifonn enccpholopatn}' use in cqttine infettions 283 horses Ill: 1611. 2019
11: 1388-1424. 1408 use in ,,igs 282.-3 pigs. dlphtherltic 111: 1430
,as wonosis II! 1388· 1~24 Try•pa110.1om11 species sec also pl~. necrotic 111: H30
Lrnnsmission of rabies. oral IT: 1132 trypanosontes ,heepigoats 111: 1595. 159G. 1597
ttanspl3ccntal lnftc:tlon, T11ei/er1a pan•a Trypa11osoma brucci 256. 279. 297 typhlocolltls
470 - - lmm•I brurv/ -19. 90 r::utlr 326
mtuma Ill: 1829 - - bmcei gamblc11!ie(rhodusfensc) ~9. 90. n$<ot:ia1cd with coc:ddlosi~ 326
traumatic lnju~· teats/udder 11: 1292, 255,2SG.261,2r,.2;9 cqulds Ill: 1832, 2015
Ill: 1962 - - bmr~i rl1odesit11sP see Trypa1wso111n pigs 111: 160•1
travellc~· diarrhoea 333 brucei gambiense typhoid nodule!' !Tl: 158:-
crembllng (louping ill/ II: 995. 1000. 1397 - - co11go!e11se -:9. 90. 255. 2SG, 2i!l 1)7.zcr'> disease Ill: 2089·91
 uveiti~ 31 ;, II: 903.111· 14•11 Venczuelen equine cnc:ephalomreliti, 13;,
horses II; 8-14.111: 144!, 1449. 1450 11: 685. 1:i;o
vcnnlno,ls, ,hceplgoats Ill: 1597

0
VCIOlOXU'I Ill: 1561
lJasin Gishu disease ll; 1298·9, Ill: 150.1
verruca vulgarts .II: 756
Ugandn S ,irus 11: 987
verrucose dermatitis Ill: 1724. 1725
ulceration
,nccination 239-44, II: 1050. U59·61. 1242, ,·cnlcal fi5suro JU: 1727
abomasal 457. 490. 499. 11 : 901.111: l702,
1352, !II: 1520·22. l 589-90 vc:,sicles
2120
buuon. pigs II: 979, 1092. 1109. 111: l(i(J.1 vaccines camels II: 1300
adjuvantS 244 cattle II: 889, 1196, 1291. 12!16
caulc 11: 82l
attenuated ll: 1130. 1290 ~'ll!llc. teats/udder II : 1296
conjuncti,-nl. ~'little Ill 1488
convemional 240-11 horse< II: 869
corneal II: 900 mouth II: 1346
corneal. cattle Ill: 1488. 2011. 2012. 2 101 dJploidccllll: tJGt
genenitally engineered 242 pigs II: 70 1, I 196. JSIS
corneal. horses 11: 863 sheep/goats II: 128•1
external genitalia II: 869. Ill: 2152-5 inactivated 2.J l. 111: 1598
live 242. ll: 10-16, 1050 \'eSicular diseases II· 700, 1194-7, 1213
face Ill: 1648 1313-17
gastric. pigs Ill: 1475 live 3llCTIU31Cd 240-41 , II: 1207, Ill: 1520.
1538, 1589-90 ,..,sicular e.,anthema of swine II: 700-702.
iloornecal V',dve, pigs II: 1107, 1109 1197, 1316. 1351
lncestinnl Ill: 1838 mark.er243·4
modem 2.Jl-1 ,·cs1culor~tomatitls90, II: 701. 119;1-. ; , 1225,
lme1tinal, pig$ Ill: 1604 1229. 1316. 1351
lip end lcg,shccp II: 1285. 1287-8 recombinant II: I 160. 1161
Seruple-rype II: 1123 as monosis II: 1HM
mouth. caule II: 953, 1223, 1227, 1228 horst.~ II: 872
&ubunit 241-2
mouth. n: 1351 \'llCC!nia \11USJI: 1293. 1297, 1298. 1302 1·esicula.r s1omatitis Indiana ,irus 15~.
mouth. sheep II: 1351 vaccinia ,iru~ infections. as 1.<,011os1s II: 1194
1.U1Sal mucosa, cqu)ds Ill: 1502 II: 1298. 1300 vesicular stomn1itis N<!w Jerscr virus 99.
penJS Ill: 2153 ,ucuolat!on. brain ll: 1400 JI: 119·1
respimtOI)' tract. equ,tls Ill: 150:-1 \'OgLOal dischari:,,
vestculars1omatitis virus II: J 124. 1194
skin n: 1271 1279, 1287. UJ: 210~ cattle Ill: 20;3, 2l49, 2150. 2151 veskular venereal disease. mule see
SOl~~-Cllttlelll: 1721.1724, 1726 cquids Ill: 1557, 1611, li8•1. 2085 lnfoctious bovine rhincmach<>itis
te,m, c:nttlc II: 1296 pigs Ill: 1784. 195?. 1959 Vt'Siculitis, seminal 5~
tongue, cattle Ill: 1648 vnl\initis catclc Ill: 2076. 2079, 2149-51
see also eroslons; lesion, cattle 308. m: 1638. 2149-51 ,Mcp/gonts Ill: IGSi
ulce:ratl\·e balanopo;thitis and horses tn: 20~ 1·e.,iculol'ims 156, II: 112-1. 119-1
\'lllvovaginitls UT· l 784. 2152·5 ~hee1,1gonts Ill: 1655. 1657, 2152 l'e.swinu II: iOO. 703
ulcerative dermato~ls 11: 1285, 1287·8 Vnlleybumps Ill: 211,- 18 \'land<' folc Goat medium Ill: 20(',3
ulcerativ<: gastritis Ill: 2L20 \lall('y fe,er 111: 211 i · Ill \'/brio cllolan Ill : 1520
11lceratl\'e granuloma ,yndromc, pigs val\'\llar endociuditis. horses Ill: 1652 ,ibriosls II: 1254
11:: 1438 \'umpire brus 11: 1124, 1129 \'lcin spefies 357
ulcerative lymphangitis Ill: 1917 Vt1rictllo,oir11s II: 829 ,illous shortening or ~mall inwstin.i 11: 788
canle lrl: 1917. 1923. 19N mrkpokke 11: 1293-4 ,mcmia
eqmds !II: 150•1. 1924. 2019 \'lJS<rulari1.mion. corneal. l'attle UT: 1488 ccll-ossocintcd II: 1.137
ulcermive cyphlocoliti;, foals Ill: 20 15 \'SSCulitil> ll: Gil3. 685. 839, 841 leukocyre-ussocio1cd. ti: 838
ulcerauve n1Mtb ll: 1285. 1287·8 coulc II: 1271, 12,J. Ill: 1514, IG;Ji, 1639. ,1.ral papular dermatilis II: 1298-9. Ill 150.J
ulcers see ulccnuion: st>c also erosion~: 1952.205-1 ,iral pneumonia
le~lons associated \\ith bovine cphemcrnl re\'cr horses II: 681
11rnbillrnl infection UJ: 1741 II: 1190 humans TI: 681
Umea dlsea~e JL 951 cqulds ll: $,I.; 846,927. Ill: lii2 pigs IL 77, Ill: 2067
lJna viru$ U: 1023 fibrinoid II: 902, 1107 virino hypothesis II: 1388
UIIClJO\'l'ntlunaJ ,in." II: I388. 13!>2 seminal Ill: 1514 ,hlnu IL 1388. 1392
tmdiff',rentiated bo,ine respiratOI)' di~easc sheep/goa,s I{: 1279 \'irulcncc factors
Ill: 1635, 1677, 1678 \'CCLOl'S 3-153. 597 Nnemorihilus .<Qm1111s Ill: 1636
undulant fever. human, Ill: 1512 anhropods 157-6:! pneumonlc pasteurcllosis Ill: 16,11
unthriftln~'Ssee lll-thrlft; poor thrift Culitoides species 93-126. J'S,7 ,ahnoncllosls Ill: 1603
urban rabil'S II: 1124. 1126. 1144·5 mechanical 597 virulent foot rot Il l: 1734, 1735
urea pol~oning n: 1158. see also uuder mosquitoes 137-49, 157.597, II: 1004. tiru> per,is1cncc, foot-and-mouth di<e.isc
poisoning 1014. 1023,4. ]03] Ir: 1340, 1343. 1348. 1353
Ureapinsma ~pccies II I: 2(J76 :V!uscldae 77·85 viruse~seq also f11diuid1111/ r•in1s 11nmes
urateritiS Tabanidae 87-91.597 :inhropo(!-borne 153·63. m1090
cattle Ill: 1931 lick> 3-34 as biologic"! wcupons II: Io 19
pig$ Ill: I 958 1sem.• mes 43-69 .a~~ociated with Culil;oidt>s 98· 103
sheep/goats !II: 165, Velorc 1ims II: 1252 haemorrhagic rr~,-tr II: 1080. 1082
Urgi11ea species 328. 525 venereal Infections. oppormniSlic, equids maintenance mid tran,mission 102. J,i.i. 7
U,oc_yoi1 dm:reo(U'ge,m:11.s U: 1128 Ill: 1556 mosquito-borne. 138. l;i7. II. 100~. JOI~.
urouterus Ill: 1556 v1•nereal ~lntin;. or Klebsi~lln µ1,~11111omn~ 102.'1·4. 1029·34
urovagina Ill: 1556 Ill: 1556 rodem-a,socimcd II : 1082, 1296
urs,,s
nmeriCll/111$ ti: 122, Ven<.'7.Uclan equine encephalitis II: 1018·21 ,irus-likc agents II: 1387-14:?4
- - mnrfrlmus Ill: 1549 asioonosis II: 1018 ,iscernl blackquancr Ill: 1859
urtica.-la, horses ll: 102,J \'ene1.uelan equine encephalitis ,irus ts,1. \'IScl'l'al lolshmoniosis 379
Usutu virus 155 243, II: 1010. 1014 ,isna JI: 741. 74-l
merine 1nfuc1ion ltl: 1560 ,itomin A dcfkicnc)' Ill: 1488
Uul.;uvirus grnus 155 ,1mmin 8 1 deficiency 417
\'llamin E1 sclenium deflcicncr II· i38. t3l l \\'e,;1ern equine cncephlllitl, It: 1017• 18 - - pseudo111brrr:11/o,<is Ill: 1597, 1617.
ru: 1H2. 15,1 Wcs1ern ~uinecncephnli1isvirw, 154. IG18
,iwcnds II; 1131 II: 101·1 yer,1111v$iS II: 90·1
Vici II: 10-11 \\'es1em equine cnrephulomyeli1i< 90. 137.
,·leksiek{e !11; 1908, 12
0
ll: 685. IOIO, 1370
,·omiting and \\'tlSting disease II: 79 l. '92 whi1 ... line di,ease Ill: 1442. lT.?4. 1726. I i,13
,·omi1ing. pills It· 781. 792. 793. 912. 979. white muscle diw,isc II: 738. 1213, 111: 1736.
1105 li55 L.agrcb schedule ti : 1163
\'l'Olp<JOljiC l[f I i?2. 1733·8 white rhlnocero~ II: 1227, 1234 ~ebrn s1riping II: 645, 664
>'U.llb.-!k see orf "hitl'·bcardcd wllclebe~~• II: 89.5 zebra 1ick 2·1
\'11/p~sca11a JI: l 130 white•iailed deer 3!12. 524, G18, II: i95. i!lll. zebras 351, II: 12:l3. 123,1 1237, 1248,
- - t'iwma II: l I'19 79g, 11!15. 1201. 120:;. 1227. 1·122. Ill: 1512, 154i
-[11/1'(1 ll: 1128 Ill: 1321:l. ig;3, IWS, 1!195 Ut,1 lOXin Ill: 186-1
- m/1,es332, 11: l 126. lll: 1546 "ild boar II: 109:,, Ill: HIS. !5'12. 1:,.6. 7.iehl-1':et,lscn modification Ill: 1510, 1515.
\'Ul,-ar squamou< cell carcinoma II: 755, 757 15'18 197;; 1517.1530. 1537, 165i
\'Ulvi1i!,, ~heeplgoaLS 111: 1931 -4. 2152. 2154 \vlldlifo diit'a!,CS Ziehl-1'celsc11 si.1lning 1echnlquc Ill· 1977,
vulvovaginili5 im1ia,1 228<!0 191\1
caule II: 891. Ill: 2(,176, 2078, 2079, see also lln•smd: imcrrac(• lli·35 Zika ,o!rm It: 989
infectious bol'ine rh!noirachc!1!~
sur\'elllance 2:!3·5 /.imhabwe 1h(;lltrl1>sb 21. 22. -15-t 41l8.
she~p/goaLS m: 17114. 2 152. 2153. ::us,1
wildllfo 472-3
vulvularcndocardltis Ill: 1909, 1910. 1950
and Bru~lln species infoc1ion m 1313. Zinga ,'irus II: 1039
1542. 15-16-51 i111k deficiency 111: 1721
conscrva1ion ureas 231 ·3 1,op animals. wilh cowpox II: 1296
fencing 2.30-31 7.oonosis
impac1 Ill disc~se 22S.30 and Caxrella bumerli 565. 568
I\lal!erion degeneration of spinal cord indUSll)' 2.11 •3 anthrax Ill: l788. 1802
ll: l 185. I 190 1iYcs1ork in1crrace 225-35 Boma disease (possible. II: 1371
walrw,es Ti: 700 pa,1otalism nncl 231 bo,i11c brucello$1S tll: l 51 O
"-:ipiti 11, ,as. 111: 1546. 1s,1 role in foo1-and- n1omh disease ll: 1339 bo,ine papulars1omarill, 11: 1289
\\'anarin poisoning lll: l•I 75, se~ also 1111der rick conrrol 33 Br11cel/11 mcliwmis Ill: 1535
poisoning 1ran<lt,t,a1ion 2:13 !lmct•lla mis Ill: IS42
Warrego virus 99. II: 12,18 l\'lllem$ reaction Ill: 20.1; hruccllo~i~ 111, l;.Oi-52
Warlhln-Srar:ry nu,1hod Ill: 1452, 1473, 2091 \\inter d~'\ClllCI)· of cn11le II· 795. 798, bulfalopox II: 1302
w.inhog U:808, 1088, Ill: 1975 !JI: 1479 t>ampylobacicrio,:I, lit: 1-179
warts S(!('a/so papillomuvlrus lnf("cllon< \\'itMlll!tSrand ,1rus 155 cowpox II . 12\JG
heel Ill: 1725 \\Obblcr syndrome II: I020 Crimcan,Ccmgo h~emorrhaglc fo,-e,
imcrdighnl Ill: I ;z5 Wongorr \'lru, I00 II: 107i
1ea1.> II: 892 Wood ButT;,lo :-1:uiqnal P.irL..111: 154i liastem i;qu!ne cncephal!th II: 1015
waJ.ps. chalcld 31 Wo<>d"$ uhravlole1 ltght 13mp !II: 20$8 "hr!ic.hlosi, 541
wastingdi>ease.shecp Ill: 1919 wooden wngue lit: l 648·52. 1756 glanders Ill: ISOO
wa,,dngll:738, 791.1378. llt:2003,201S woodland caribou 353 Hendro ,~rus n: 681. 682
wasting >)'lldrome, post-we.aning woolloss II: 1~00 Nmip,win.ts II: 692
mut1isystemic II: 6011, 810, 1374·83 woolly coa1 in foot-and-mouth dlsea.~t> hor:scpox II: 1298
water bufl'nlo 360, 382, ·110, II: 81 S. 8,6, U: 1348 Klebsie/111 species infecuons 111: 1533
1046, 1270. l!l: 1620, 1638. 1689. 1829, woolsoner'sdiscasc Ill: 1788-181 l. s~e al~o louping ill II: 995. 999
197-l. l975, 2028,20-15. 20<J6 anihrox ~talla fever Iii: 1535
w:ner mongoose JI I 138 \\Ound botulism Ill: 1892 ~!enangl~ ,,jrus II: 690
wmcrbuck 453. 469. II: 798. 111: 1546. 1547 1V11rh,-rerin IJfr11crofti I53 Xi pall \'irus IT: 692
weak orlll: 12a2
cnJ,·es Ji: 8:0.111: 1480, 20i6
0
pseudoco"1'0X ii: !291
lambs IL: 970.111: 1+18 rnblc. II: 1123. I 151-6-1
offspring Ill: 1448. 1543 J?11b11/t11iil /IS JI: !i90
weaner coliti~ Ill: 1623 fkllmont'llo<is Ill: 1582. 1511,1, 1598, 1601,
x~rus innuris II: 1135
weigh1 Jos~ ii: ;43, Ill: 1596, 1997. 2019 1603. 1604. 1612
\\'csselsbron dlsca~e 347. Ii: 987·92, 1057. Srn11il,•locClC'C11S a11re11s infection 111: 1,54
!:l;,4
as zoonosis II: 988
\\'csselsbron ,iru, !37, H2, 143, 155
0 S1r~p1oc0crns mis lnf~c1ions 111: 1,7
1rnnsmisslble s1>0ngiiorm
,·ncephalopa1hy 11 1388-1424
\\'e.s,eJsbronsiek1e34i. II: 987-92, l05i, yellow ie1 er 153 1uberculo,ls Ill: 1973
1254 yeU01, f~ver \'irus 15-1. Tl: 987,989,996.1082 Ve11ezuelru1 cquinQ cncephallt!, IJ· 1018
\Ves1 '.':ilc ,iru, 137. 154. II: 685, 98Q, 1015, ~ellow lamb disc-OSI! Ill: 1829-33 , eskular ,1oma1hl\ ll: 119·1
1057 yellow mongoose II: 1133, 1135 Wes.,elshron ,irus II: 988
\\'est J\ilc ,irus infection II: 1004·10 vellow-b.icked uck 2•1 Yersl11in species infornons Ill! 161T, 1618
•blrds II: 1004-6 r,•rs111i(1 spcc,C!' Ill: l 588, 1617 zoono1k dlscn;os Ii: 692
catsldogs Ii: 1008 YcrJinia spcci1!5 infoctions lll: 161 i-25 iwoegersicktc 11:ili. 733-8. 741. ;.;.;, 1401
horses JI: 1001H 0 ~s zoonosls Ill: 1617, 1611! Zygndmus ;.pecics II: JO 10
humans II: I 004 l'crsi11ia cm,rorollr/c(I Ill: 1519. L~3. 15-H. zygomycosls lll: 209S. 2113, 2120
pigs II: 1008 1550. 1597. 1617. 1618. 1623 7.ygomycoiinn Ill: 2095
Infectious Diseases
of Livestock
VOLUME ONE

ISBN O 19 576169 3

9 780195 761696