PM1ch21 ARTHROPODS OF MILITARY IMPORTANCE

MILITARY PREVENTIVE MEDICINE:
MOBILIZATION AND DEPLOYMENT

Volume 1
Section 3: Preparing for Deployment
Before these Marines arrived in Southwest Asia for the Persian Gulf War, an impressive amount of planning and
coordination had been done by preventive medicine professionals on their behalf. The health threats facing these
Marines had been assessed, plans had been made to counteract those threats, and systems had been put into place to
monitor their health status during and after the deployment. These efforts were focused on keeping disease and
nonbattle injuries to a minimum—and they worked.
Photograph: Courtesy of the Defense Visual Information Center, March Air Reserve Base, California. Image 47 on the CD-ROM “U.S.
Forces in Desert Storm.”
211
212
Arthropods of Military Importance
Chapter 21
ARTHROPODS OF MILITARY IMPORTANCE
RAJ K. GUPTA, PH D; LEON L. ROBERT JR., PHD; AND PHILLIP G. LAWYER, PH D
INTRODUCTION
CLASS INSECTA
Mosquitoes
Flies
Fleas
Lice
Bugs
Stinging Insects
CLASS ARACHNIDA
Ticks and Mites
Spiders and Scorpions
SUMMARY
469
Military Preventive Medicine: Mobilization and Deployment, Volume 1
R. K. Gupta; Colonel, MS, US Army; Research Area Director, Research Plans and Programs, US Army Medical Research and Devel-
opment Command, Fort Detrick MD 21702-5012; formerly, Department of Entomology, Division of Communicable Diseases and
Immunology, Walter Reed Army Institute of Research, Silver Spring, MD 20910-7500
L. L. Robert Jr.; Lieutenant Colonel, MS, US Army; Department of Preventive Medicine and Biometrics, Division of Tropical Public
Health, Uniformed Services University of the Health Sciences, 4301 Jones Bridge Road, Bethesda, MD 20814
P. G. Lawyer; Colonel, MS, US Army, (Retired); Associate Professor, Department of Preventive Medicine and Biometrics, Uni-
formed Services University of the Health Sciences, 4301 Jones Bridge Road, Bethesda, MD 20814; formerly, Entomology Consultant,
Office of US Army Surgeon General
470
INTRODUCTION
Until the latter half of the 20th century, military certainty that a particular arthropod is a vector of
casualties due to disease and nonbattle injuries (DNBI) human disease are:
usually outnumbered those directly related to com-
bat. Historically, arthropods have been a leading cause • Demonstration that members of the sus-
of DNBI. The direct effects of arthropods include tis- pected arthropod population commonly
sue damage due to stings, bites, and exposure to vesi- feed upon vertebrate hosts of the pathogen,
cating fluids, infestation of tissue by the arthropods or otherwise make effective contact with the
themselves, annoyance, and entomophobia (an uncon- hosts under natural conditions,
trolled fear of arthropods). The indirect effects on • Demonstration of a convincing biological
human health include disease transmission and aller- association in time and space between the
gic reactions due to bites and stings and to arthropod suspected vectors and clinical or subclini-
skins or emanations. Arthropods also destroy prop- cal infections in vertebrate hosts,
erty and materiel used by the military, and there is • Repeated demonstration that the suspected
even some concern that arthropods could poten- vectors, collected under natural conditions,
tially be used as biological weapons. The impor- harbor the identifiable, infective stage of the
tance of naturally occurring arthropod-borne dis- pathogen, and
ease is well documented, as is the direct injury, an- • Demonstration of efficient transmission of
noyance and material damage caused by nuisance the identifiable pathogen by the suspected
arthropods. During World War II, it is estimated that vectors under controlled experimental con-
more than 24,000,000 man-days were lost due to ditions.3
arthropod associated disease and injury.1
Arthropods are of greatest importance to mili- To minimize adverse effects caused by arthropods,
tary operations when they act as vectors of disease. military entomologists must be able to identify vec-
A vector is an organism that transmits a pathogen tor and pest species accurately and then prescribe
to a susceptible host. Arthropods serve as vectors appropriate countermeasures. This begins with a ba-
in a number of different ways, from simple me- sic knowledge of arthropod morphology and classifi-
chanical transmission of pathogenic organisms on cation. Members of the phylum Arthropoda have seg-
the arthropod body, as when house flies carry dys- mented bodies with paired, segmented appendages,
entery bacilli from infected feces to food, to the more bilateral symmetry, a dorsal heart, a ventral nerve
complicated process of biological transmission, cord, and an exoskeleton. The arthropods are divided
where the pathogens must spend part of their life into five classes: Insecta (insects), Arachnida (mites,
cycle in the body of the arthropod before humans ticks, spiders and scorpions), Crustacea (crabs, lob-
can be infected. A fundamental activity of military sters, shrimps, water fleas), Chilopoda (centipedes),
medical entomologists is to establish the role that and Diplopoda (millipedes). Two of these classes, the
certain arthropod species or populations play in the Insecta and the Arachnida, are especially important
transmission of a particular infectious disease to to war fighters because of the potential effect they can
service members. There are primary and second- have on military operations (Table 21-1). The other
ary vectors. Primary vectors are those that are three classes are of minor significance. The identifica-
mainly responsible for transmitting a pathogen to tion of the arthropods by class, order, family, genus,
humans or animals; secondary vectors are those that and species is achieved by matching their morpho-
play a supplementary role in transmission but logical characteristics or features to couplets on a
would be unable to maintain disease transmission published dichotomous key. The class Insecta is the
in the absence of the primary vector.2 The criteria largest arthropod class and the one that includes the
that must be satisfied to declare with reasonable majority of the arthropods of military importance.
CLASS INSECTA
Insects can be differentiated from other arthropods have only one pair of antennae.
by the presence of three distinct body regions (head, Insects transmit diseases either mechanically or
thorax and abdomen), and three pairs of legs (Figure biologically. Mechanical vectors include cock-
21-1). Insects are the only arthropods that possess roaches and filth flies. These insects may pick up
wings and most, but not all, have them. Insects also pathogens on their feet or other parts of their body
471
TABLE 21-1
472
SYSTEMATIC LIST OF ARTHROPODS NAMED. ALL FALL WITHIN THE PHYLUM ARTHROPODA.
Scientific name Common name Scientific name Common name

Class Arachnida Arachnids Amblyomma maculatum Gulf Coast tick
Order Araneae Spiders Dermacentor andersoni Rocky Mountain wood tick
Family Theridiidae Comb-footed spiders Dermacentor variabilis American dog tick
Lactrodectus mactans Black widow spider Ixodes dammini Northeastern deer tick
Ixodes scapularis Black-legged tick
Family Loxoscelidae Recluse spiders
Ixodes pacificus Western black-legged tick
Loxosceles reclusa Brown recluse spider
Ixodes persulcatus
Order Acari Mites and Ticks Ixodes ricinus European castor bean tick
Family Trombiculidae Chigger mites
Leptotrombidium deliense Class Insecta Insects
Leptotrombidium fletcheri Order Anoplura Sucking lice
Leptotrombidium pallidum Family Pediculidae Head and body lice
Trombicula alfreddugesi Pediulus humanus capitis Human head louse
Trombicula autumnalis Harvest mite Pediculus humanus humanus Human body louse
Family Pyroglyphidae House dust mites Family Pthiridae Crab lice
Dermatophagoides farinae American house dust mite Pthirus pubis Human crab louse
Dermatophagoides pteronyssinus European house dust mite Order Siphonaptera Fleas
Euroglyophus maynei Family Pulicidae
Family Glycyphagidae Xenopsylla cheopis Oriental rat flea
Glycyphagus destructor
Order Hemiptera True bugs
Family Macronyssidae Fowl mites Family Cimicidae Bed bugs

Ornithonyssus bacoti Tropical rat mite Cimex hemipterus Tropical bed bug
Ornithonyssus bursa Tropical fowl mite Cimex lectularius Common bed bug
Ornithonyssus sylviarum Northern fowl mite Assassin bugs
Family Triatomidae
Family Dermanyssidae Triatoma infestans
Dermanyssus gallinae Chicken mite
Triatoma dimidiata
Liponyssoides sanguineus House mouse mite
Triatoma barberi
Pyemotes tritici Straw itch mite
Triatoma gerstaeckeri
Family Sarcoptidae Scab or itch mites Triatoma proctata
Sarcoptes scabiei Scabies mite Triatoma sanguisuga Blood sucking conenose
Trixacarus sp. Panstrongylus megitus
Family Demodicidae Follicle mites Rhodnius prolixus
Demodex brevis Lesser follicle mite
Order Hymenoptera Bees, wasps, and ants
Demodex folliculorum Follicle mite
Family Apidae Bees
Family Argasidae Soft ticks Apis mellifera mellifera European honey bee
Ornithodoros hermsii Apis mellifera scutellata Africanized honey bee
Ornithodoros parkeri Family Formicidae Fire ants
Ornithodoros turicata Relapsing fever tick Solenopsis invicta Imported fire ant
Family Ixodidae Hard ticks
Amblyomma americanum Lone star tick
TABLE 21-1 continued
Scientific name Common name Scientific name Common name

Order Diptera True flies Culex tritaeniorhynchus
Family Ceratopogonidae Biting midges Culex vishnui
Culicoides spp. Culiseta melanura
Family Simullidae Black flies Deinocerites spp.
Simulium spp. Eremapodites spp.
Haemagogus spp.
Family Psychodidae Sand flies
Mansonia annulata
Brumtomyia spp.
Mansonia annulifera
Chinius spp.
Mansonia bonneae
Lutzomyia spp.
Mansonia dives
Phlebotomus papatasi
Mansonia uniformis
Sergentomyia spp.
Psorophora columbiae Dark rice field mosquito
Werileya spp.
Psorophora confinnis
Family Culicidae Mosquitoes Psorophora discolor
Aedes aegypti Yellow fever mosquito Psorophora ferox
Aedes albopictus Asian tiger mosquito Sabethes chloropterus
Aedes fulvus
Family Glossinadae Tsetse flies
Aedes leucocelaenus
Glossina morsitans
Aedes lineatopennis
Glossina palpalis
Aedes niveus
Glossina swynnertoni
Aedes poicilius
Glossina tachinoides
Aedes polynesiensis
Aedes scutellaris Family Tabanidae Deer and horse flies
Aedes serratus Chrysops dimidiata
Aedes sollicitans Salt marsh mosquito Chrysops distincitipennis
Aedes taeniorhynchus Chrysops silacea
Aedes thelcter Tabanus spp.
Aedes togoi Family Chloropidae Eye gnats
Aedes tongae Hippelates flavipes
Aedes triseriatus Family Muscidae Filth flies
Aedes vexans Inland floodwater mosquito Fannia spp.
Aedes vigilax Musca domesticus Common house fly
Anopheles campestris
Anopheles darlingi Family Calliphoridae Blow flies
Anopheles donaldi Auchmeromyia luteola Congo floor maggot
Anopheles funestus Calliphora spp. also called a filth fly
Anopheles gambiae Cordylobia anthropophaga Human bot fly
Anopheles minimus Lucilia spp.
Anopheles punctulatus group Chrysomyia bessiana
Culex gelidus Cochliomyia hominivorax
Culex neavei Family Oestridae Robust bot flies
Culex pipiens quinquefasciatus Southern house mosquito Dermatobia hominis Torsalo
473
Culex pseudovishnui Family Sarcophagidae Flesh flies
Culex tarsalis Sarcophaga spp.
Piercing-Sucking mouthtype
Mosquito
Labrum-Epipharynx Hypopharynx
Maxillae
Mandibles
Labium
Fig 21-1. The three major divisions of an insect body are Fig 21-2. The sucking-piercing mouth parts of a female
shown on this drawing of a mosquito. mosquito.
Source: Letterman Army Institute of Research, Presidio Source: Letterman Army Institute of Research, Presidio
of San Francisco, Calif. of San Francisco, Calif.
while feeding on fecal material or organic waste, sucking), (2) horse fly subtype (piercing/cutting),
or they may ingest the pathogens and later contami- and (3) house fly subtype (nonpiercing/sponging)
nate food consumed by the humans. Some examples (Figure 21-2). All Diptera are holometabolus, mean-
of important mechanically transmitted diseases that ing they undergo complete metamorphosis, passing
have plagued military operations are typhoid fe- through egg, larva, and pupa stages before becom-
ver, cholera, and dysentery. However, the insects ing adults. The larvae of most medically important
that are of greatest importance to the military are Diptera species require high humidity and are
those that serve as biological vectors of human aquatic, semi-aquatic, endoparasitic, or live in wet
pathogens. Pathogens transmitted in this manner or moist terrestrial habitats. The mosquitoes, sand
must pass through part of their life cycle in the vec- flies, black flies, horse flies, stable flies, midges and
tor, where the pathogens multiply, change form, or filth flies are examples of Diptera that are serious
do both before being passed on to a susceptible host. pests of humans and animals.2
The host is usually infected via the bite of the vec-
tor or by having the excretions or body fluids of Mosquitoes
the vector rubbed into the skin.
Many of the vector insects discussed in this chap- Mosquitoes are the foremost nuisance insects and
ter are members of the order Diptera, the two- disease vectors in most regions of the world and
winged or true flies. Approximately 80,000 to the sole vectors of several pathogens that cause dis-
100,000 species in about 140 families have been de- eases of military importance, including malaria,
scribed, and new ones are being added constantly. yellow fever and dengue. They also play a signifi-
This order contains more insects involved in trans- cant role in transmitting filariasis and forms of vi-
mission of human and animal pathogens than any ral encephalitis.2,3 Mosquitoes belong to one of the
other. The Diptera are relatively small and soft- more primitive families of Diptera, the Culicidae.
bodied insects; all winged members possess only one The blood-sucking behavior of female mosquitoes
pair of wings and a pair of short, knob-like struc- and some other Dipterans predisposes them to ac-
tures called halteres. Conspicuous compound eyes quiring pathogens and parasites from an infected
are usually present, and most species possess an host and depositing them in the skin of a suscep-
additional three simple eyes, called ocelli, that are tible host.
set in a triangle at the top of the head. The Dipteran
mouthparts, while subject to great morphological Life Cycle
variation, are all adapted for sucking fluids, as op-
posed to chewing solids, and are either piercing or The immature stages of all mosquitoes develop
nonpiercing. They can be divided into three sub- only in water, but adaptation to and preference for
types as follows: (1) mosquito subtype (piercing/ particular types of water and water locations vary
474
Anopheles Aedes Culex quito larvae are affected by light and water condi-
tions (eg, temperature, water movement), predators,
Egg
and other living organisms in their habitat. When fully
grown, a larva changes to a pupa.2
Pupa. The pupa, or “tumbler,” is active but does
Larva not feed. This is the resting stage from which the
adult is formed. The pupa rests on the surface of
water unless disturbed, then it quickly moves to-
wards the bottom. It breathes through two respira-
tory trumpets on the thorax. When the adult is ready
to emerge, the pupal skin splits, and the adult pulls
Pupa
itself up and out of the floating skin, on which it
rests until it is ready to fly.2
Adult. Adult mosquitoes are distinguishable
Adult from other Diptera by the long, filamentous anten-
Palp Long Palp Short Palp Short
nae comprising 14 to 15 segments, a long proboscis
adapted for blood sucking, and scales on the fringes
of the wings and wing veins. The rounded head
bears large compound eyes that almost touch in the
Female Female
Female middle. Males can usually be separated from the
females by their bushy antennae. Adult mosquitoes
Male Male display great diversity with regard to resting and
Male
oviposition habits, biting preferences, vector com-
Resting Position petence, dispersion, and flight range. Although
breeding sites vary by species, mosquitoes can be
divided into three major groups based on where
Fig. 21-3. Characteristics of anopheline and culicine they deposit their eggs: permanent water breeders,
mosquitoes. floodwater breeders, and artificial container or
Source: Centers for Disease Control and Prevention. treehole breeders. Because each of these groups con-
Vector-borne Disease Control Self-study Course 3013-G. tains important vector species, breeding sites
Manual G. Mosquitoes of Public Health Importance and Their should always be considered when investigating a
Control. Atlanta: CDC; 1994: 14. mosquito-borne disease outbreak or when imple-
menting surveillance and control programs. Anoph-
eles (including malaria vectors) and Culex mosqui-
widely among species. The larvae are anatomically toes usually select permanent bodies of water, such
different from the adults, live in different habitats, as swamps, lakes, ponds, streams and ditches. On
and eat different types of food. Transformation to the other hand, floodwater mosquitoes lay their
the adult stage takes place during a nonfeeding eggs on the ground in depressions that are subject
pupal stage (Figure 21-3). to flooding. During heavy rains or after snow melt,
Egg. Female mosquitoes may lay several hundred eggs deposited in these sites are inundated with
eggs, depositing them on the water’s surface or on water and hatch within minutes or hours. Included
sites that will be flooded by water later. Each egg is in the floodwater group are Aedes vexans (inland
encased in a protective shell. Depending on the spe- floodwater mosquito), Aedes solicitans (salt marsh
cies, eggs may be laid singly, in clusters, or in rafts.2,4 mosquito), and Psorophora columbiae (dark rice field
Larva. Mosquito larvae, also known as “wig- mosquitoes). Artificial container or treehole breed-
glers,” can be found in permanent ponds and ers include Aedes aegypti (the yellow fever mos-
marshes, temporary flood waters, or in water that quito), Aedes albopictus (the Asian tiger mosquito),
has accumulated in tree holes, leaf axils, or other Aedes triseriatus, and others.2,5
natural and artificial containers. They are very ac- Both male and female mosquitoes feed on plant
tive and feed on minute aquatic animal and veg- sugars, such as floral nectars, to obtain nourishment
etable life and on decaying organic matter, coming for basic metabolism and flight. In addition, most
to the surface to breathe through a respiratory si- females will take a blood meal from birds, reptiles,
phon at the posterior end of the body. The larval amphibians, or mammals for egg development (Fig-
stage includes four developmental instars. Mos- ure 21-4). Biting behavior may be very important
475
Fig. 21-4. A blood-engorged Anopheles stephensi mosquito, Fig. 21-5. Distinctive characteristics of Aedes aegypti
which transmits malaria in Asia. Source: Letterman Army Institute of Research, Presidio
Photograph: Courtesy of Ed Rowton, Walter Reed Army of San Francisco, Calif.
Institute of Research, Silver Spring, Md.
in the epidemiology of disease transmission. Many singly on the surface of the water and are boat-
mosquito species bite humans to obtain their shaped, possessing a pair of lateral floats. Aedes lay
bloodmeals, and some feed on humans preferen- their eggs singly on damp litter or moist areas near
tially. Some will enter houses to bite humans the water’s edge. Culex deposit their eggs grouped
(endophagic), while others bite their human hosts in rafts of 100 or more on the water’s surface. Anoph-
outside (exophagic). Other mosquitoes prefer to eles larvae lack a breathing siphon and lie parallel
feed on non-human hosts, and many species do not to the surface. Culicines all have a prominent res-
bite people at all.2 piratory siphon at the posterior end of the body and
Whereas wind-assisted dispersion records of up are usually found suspended at a 45° angle beneath
to 100 km have been recorded, in control programs the water surface. When at rest and when feeding,
and epidemiological studies it is usually safe to say adult Anopheles hold their bodies at a 45° to 75°
that mosquitoes will not fly more than 2 km from angle to the surface, with the proboscis and abdo-
their emergence site.6 This is an important point to men in a straight line. Aedes and Culex rest and feed
remember when surveying for potential vector with the body held parallel to the surface. The most
breeding sites. reliable way to distinguish between adult Anophelines
More than 3,000 mosquito species have been de- and Culicinae is by examination of their heads. The
scribed. They are found throughout the world ex- Anopheles possesses palpi that are about as long as the
cept in places that are permanently frozen. Three proboscis, male palpi that are paddle-shaped at the
quarters of all mosquito species live in the tropics tip, and an evenly rounded scutellum. The palpi of
and subtropics, where warm, humid weather favors Aedes and Culex are much shorter than the proboscis.6
their rapid development. Mosquitoes may be found
at elevations as high as 4,300 m above sea level, such Parasitic Diseases Transmitted by Mosquitoes
as in Kashmir, India, and 1,160 m below sea level,
as in the gold mines of south India.2 Descriptions Malaria. Malaria is the most important arthro-
of the biology and physiology of mosquitoes can pod-borne disease of humans. Essentially a disease
be found in Bates, 7 Christophers, 8 Clements, 9 of the tropics and subtropics, it is present in 102
Gillett,10,11 and Horsfall.12 countries and infects up to 500 million people and
From a medical perspective, the three most im- causes 2.5 million deaths annually, primarily in the
portant mosquito genera are Anopheles, Aedes and tropics.13,14 In Africa, malaria is one of three infec-
Culex. These are contained in two subfamilies, the tious diseases that contribute most significantly to
Anophelinae (Anopheles) and the Culicinae (Aedes the burden of disease as estimated by DALY (dis-
and Culex). The following characteristics serve to ability adjusted life years).15
separate Anopheles mosquitoes from the other mos- Malaria is caused by four species of protozoan
quito genera (Figure 21-5). Anopheles eggs are laid parasites of the genus Plasmodium: Plasmodium
476
falciparum, P vivax, P ovale, and P malariae. Although condition. A resurgence of malaria in South Korea
P vivax causes the most malaria infections world- in the late 1990s, particularly around the Demilita-
wide, the most serious, and often fatal, form of rized Zone, underscores malaria’s importance to US
malaria is caused by P falciparum. In many parts of military forces today.18
the world, infected people may carry large num- Filariasis. Lymphatic filariasis is another impor-
bers of parasites without showing signs or symp- tant mosquito-borne parasitic disease distributed
toms of the disease, thus serving as reservoirs of throughout tropical and subtropical areas of the
infection for blood-feeding mosquitoes. Malaria world. Currently, an estimated 146 million people
parasites enter the human host through the bite of are afflicted with this debilitating and disfiguring
an infected female Anopheles mosquito. The life disease, which is caused by three species of filarial
cycle of the malaria parasite takes place in two sepa- worms: Wuchereria bancrofti, Brugia malayi, and
rate cycles: one in the human or vertebrate host and Brugia timori. Wuchereria bancrofti is the most widely
the other in the mosquito. (See chapter 36 for de- distributed species, and its infections are the most
tails of the malaria parasite cycle.) prevalent.19,20 An estimated 115 million people are
Malaria heads the list of arthropod-borne dis- infected with the parasite. It occurs in large sections
eases of importance to the military. It has had tre- of sub-Saharan Africa, parts of South and Central
mendous impact on military operations in the past America and the Caribbean region, parts of India,
and continues to threaten the health and effective- China, Bangladesh, Burma, Thailand, Malaysia,
ness of today’s war fighters when they are deployed Laos, Vietnam, Indonesia, the Philippines, Papua
to malaria-endemic areas. In these areas, native New Guinea, and island groups in the south Pa-
human populations often appear healthy, but a large cific Ocean. Brugia malayi has a more limited distri-
portion of the people may be carriers of or semi- bution, occurring in China, India, the Republic of
immune to malaria to which they have been ex- Korea, and Southeast Asia, including Indonesia and
posed repeatedly since birth. The malaria pathogens the Philippines. Brugia timori is confined to certain
are maintained at low levels by the hosts’ immune southern islands of Indonesia in the Savu Sea. The
systems. These inapparent infections can be passed number of people infected with the latter two spe-
quickly to nonimmune newcomers by mosquitoes. cies has not been estimated, although humans in
US forces deployed to malarious areas lack immu- endemic areas have infection rates as high as 30%.2
nity to the pathogen and if they are bitten by in- The worms may infest and block human lym-
fected mosquitoes, they readily contract the disease. phatic channels and cause enormous and debilitat-
The results include loss of manpower, increased bur- ing swelling, resulting in much human suffering.
den on the military health support system, and de- Besides their physical and psychological impact, the
creased unit morale. All of these factors reduce the filariases exact an enormous toll in terms of de-
commander’s ability to execute the directed mission. creased economic production in nations where such
The malaria threat to force readiness that has decreases can be ill afforded. The filariases gener-
confounded commanders in the past still confronts ally cause morbidity but not mortality in infected
our forces today. In 1993, a number of Marines (106) humans and so normally do not have an immedi-
and soldiers (70) in certain units participating in ate effect on military operations. It can, however,
Operation Restore Hope in Somalia developed ma- cause a significant loss of personnel and resources
laria.16,17 The reasons for this, the largest outbreak in units stationed in endemic areas. In World War
of malaria in US military since the Vietnam War, II, a survey of two units stationed in the South Pa-
are a complex mixture of incomplete medical intel- cific showed that they had infection rates of 65%
ligence regarding the malaria threat, lack of com- and 55%. Both units were returned to the United
mand emphasis on and compliance with personal States without having entered combat.21
protection measures, and the complex life cycle of The vector mosquito acquires the microfilariae
the malaria parasite. Medical surveillance revealed from an infected person while obtaining a blood
that one half of all malaria and dengue cases dur- meal. The filariae shed their sac-like sheath and
ing Operation Restore Hope occurred in a single travel quickly through the midgut of the mosquito
Marine battalion located in the Baardera area. A to the thorax where they develop in the large indi-
subsequent investigation of these outbreaks found rect flight muscles. After a number of internal
that the Marine commander did not enforce recom- changes and two molts, infective third-stage larvae
mended countermeasures. Fortunately, the ill Ma- develop. The infective larvae find their way into the
rines recovered and the unit was not involved in hemocoele (body cavity) of the vector and eventu-
any tactical engagements while in this weakened ally to the proboscis, from which they are transmit-
477
ted to a new host during blood feeding.22 an urban cycle. The sylvatic cycle involving forest
Depending on the species, vectors of filarial canopy mosquitoes as the vectors (Haemagogus
pathogens may be active during day or night. Culex species in Central and South America and Aedes
pipiens quinquefasciatus is the main vector respon- species in Africa) and forest primates, mainly
sible for filarial transmission in urban areas; in ru- monkeys, as the hosts. The urban cycle involves Ae
ral areas, the vectors may be Anopheles, Aedes, or aegypti as the vector and humans as the hosts (see
Mansonia species that inhabit the forests. The main Figure 36-10). The urban cycle begins when humans
vectors of Wuchereria bancrofti in the tropics are An become infected in the sylvatic cycle by entering
gambiae, An funestus, An darlingi, An minimus, An forest habitats and being bitten by infected sylvatic
campestris, An punctulatus group, Aedes niveus, Ae vectors. People infected in this way return to their
poicilius, Ae polynesiensis, Ae tongae, Ae vigilax, and villages or cities, thereby initiating urban transmission.
Culex pipiens quinquifasciatus. Brugia malayi occurs Yellow fever originated in Africa and was brought
in rural populations in the Far East between 75° and to the New World during the 1500s by slave trade
140° East longitude in small endemic foci. Specific ships. These ships maintained their own colonies
known vectors of this pathogen are Mansonia of Ae aegypti in fresh-water storage containers. The
annulifera, M dives, M bonneae, M annulata, M virus and urban vector became established in the
uniformis, An campestris, and An donaldi.2 Caribbean region, on the east coast of South and
Central America, and in the southern United States.
Viral Diseases Transmitted by Mosquitoes During 1741, the British lost 20,000 to 27,000 men
to an epidemic of “black vomit” while on an ill-fated
Approximately 100 arthropod-borne viruses expedition to conquer Mexico and Peru. The British
(arboviruses) are known to produce disease in and Spanish forces suffered heavy losses on Cuba
humans. The best-known examples of human in the 1760s, and the French lost 29,000 in 1802 while
arboviral pathogens are in the following genera and in Haiti and the Mississippi Valley.3
families: Flavivirus (Flaviviridae), Bunyavirus and In 1900, a US Army Yellow Fever Commission
Phlebovirus (Bunyaviridae) and Alphavirus (Togaviridae). under the direction of Major Walter Reed was
The flaviviruses are either mosquito- or tick-borne, established in Cuba, where its studies demonstrated
the bunyaviruses and other closely related viruses that a filterable agent present in the blood of acute
in the family Bunyaviridae may be transmitted by phase patients could be transmitted by Ae aegypti
mosquitoes, ticks, sand flies, or midges; the mosquitoes (Figure 21-6).24 Jungle yellow fever in
phleboviruses are transmitted by mosquitoes and the Western Hemisphere is transmitted chiefly
sand flies, and all but one of the alphaviruses are among monkeys, marmosets, and other animals and
mosquito-borne. These pathogens cause infections in
humans and livestock resulting in febrile illnesses
ranging from mild discomfort to severe influenza-
like symptoms and can cause encephalitis, en-
cephalomyelitis, and hemorrhagic fevers. Mortality
rates can be relatively high, especially in infections with
central nervous system involvement or hemorrhagic
symptoms.2 Arboviral diseases are often characterized
by sudden onset and periodic epidemics involving
thousands of cases, which could seriously affect
combat effectiveness in military operations. The
following mosquito-borne viral diseases are
discussed in order of relative medical significance.
Yellow Fever. Historically, yellow fever virus, a
flavivirus, is probably the most important and most
dangerous of the mosquito-borne viruses. It was
first recognized in humans in the 17th century, but
mosquito transmission of yellow fever virus was
not demonstrated until the landmark studies by Fig. 21-6. Aedes aegypti
Major Walter Reed, Dr. Carlos Finlay, and their Photograph: Courtesy of Colonel Raj Gupta, MS, US
colleagues in Cuba in 1900.23 Yellow fever has two Army, and the Letterman Army Institute of Research,
transmission cycles, a sylvatic or jungle cycle and Presidio of San Francisco, Calif.
478
causes fatal infections in these animals. The reported close association with humans, and is distributed
vectors of the forest canopy are Haemagogus species, throughout the tropics.27 Secondary vectors include
Ae leucocelaenus, Sabethes chloropterus, and possibly Ae albopictus (the Asian Tiger mosquito, which was
Ae fulvus.25 Many cases of yellow fever have been introduced into the United States from Asia in the
reported in recent years from South America, the late 1980s), Ae scutellaris, and Ae polynesiensis
vast majority of these were in males over 15 years (Figure 21-7).2
of age who were presumably infected while The first epidemics of dengue fever were
working in the jungle. In the last 30 years, Ae aegypti reported simultaneously in 1779 from Egypt and
has reinfested Central and South America and Indonesia and in 1780 from the United States. The
brought with it the potential threat to transmit first recognized epidemic of dengue hemorrhagic
yellow fever in an urban cycle. Because of a very fever occurred in Manila in 1953 and 1954.28 In the
efficacious vaccine, yellow fever is no longer a ensuing 20 years, dengue hemorrhagic fever spread
threat to the US military. to other parts of southeast Asia, and today it is one
Dengue. Dengue is a viral disease transmitted from of the leading causes of hospitalization and death
person to person by mosquitoes throughout the among children in southeast Asia.29 Dengue fever
tropics and subtropics. It is an acute, nonfatal disease was not considered a public health problem in the
of particular importance to military operations Americas until the latter part of the 20th century.
because of its rapid spread and its ability to in- During the 1970s and 1980s, dengue serotypes 1, 2,
capacitate large numbers of personnel at critical and 4 were introduced into the Americas. 30 The
periods. Dengue fever is caused by any one of four increased incidence of dengue and dengue hemorrhagic
closely related dengue virus serotypes (dengue 1- fever worldwide has resulted in an increased number
4) of the Flaviviaridae family. 26 The virus is of cases being imported into the United States.
transmitted primarily by Aedes mosquitoes, par- Additional details can be found in chapter 36.
ticularly Ae aegypti, an urban species that lives in Japanese Encephalitis. Japanese encephalitis is
Dengue Hemorrhagic Fever

Dengue Fever Outbreaks
Fig. 21-7. The distribution of dengue fever outbreaks and dengue hemorrhagic fever. Map: Courtesy of Colonel
David Vaughn, Walter Reed Army Institute of Research, Silver Spring, Md.
479
one of the major public health problems in Asia.31 paddies flooded with water. Studies have shown
Outbreaks have been reported from Japan, Korea, that the number of Japanese encephalitis cases is
Taiwan, Siberia, China, Okinawa, Thailand, Malay- directly proportional to the density of the vector
sia, and Singapore. They occur in warm weather in when an epizootic in pigs is in progress.32 Transovarial
temperate regions and throughout the year in the transmission has been reported in Ae albopictus and
tropics. Japanese encephalitis is principally a dis- Ae togoi (Figure 21-8). There have been occasional
ease of rural agricultural areas, where vector mos- isolations from Anopheles mosquitoes.2
quitoes breed in close association with pigs, birds, A vaccine affording excellent protection (> 90%)
and ducks. Japanese encephalitis virus is a flavivirus against Japanese encephalitis was licensed by the
and is maintained in mosquitoes and hosts other Food and Drug Administration in 1992 and is avail-
than humans. The principal vector of Japanese en- able for military personnel deploying to endemic
cephalitis is Cx tritaeniorhynchus, which feeds pri- areas. It is widely used in parts of Asia, such as
marily on animals and birds. Cx vishnui, Cx gelidus, South Korea.
Cx pseudovishnui, and related species are the pri- West Nile Fever. West Nile fever (WNF) is prob-
mary vectors in Thailand and India.31 These mos- ably one of the most common arbovirus infections
quitoes breed in ground pools and especially in rice of humans, considering its broad geographical dis-
tribution. It is still considered a disease of unknown
origin, however, because of its infrequent epidem-
ics. WNF virus, a flavivirus, has been implicated in
outbreaks since 1950, but the largest known out-
break occurred in South Africa in 1974. Infection
usually occurs in early childhood and produces only
a mild febrile illness. In 1998, an outbreak of WNF
occurred in New York City with 9 deaths.33 WNF
could have significant impact on military opera-
tions. For example, in an earlier outbreak in Israel,
636 cases of clinical disease were reported in a popu-
lation of approximately 1,000 at a military camp.34
WNF virus has been isolated from humans and
wild birds, but isolates also have been recovered
from animals, including lemurs, chimpanzees,
chickens, ducks, geese, horses, mules, donkeys,
pigs, and cows. WNF virus has been isolated in 17
countries from three different zoo-geographic re-
gions: the Palearctic, the Ethiopian, and the Orien-
tal. The transmission activity of WNF virus indi-
cates a seasonal pattern limited to summer months.
All of the recorded outbreaks in Israel occurred from
July through early September; reported outbreaks
in South Africa occurred from December through
April.35
In nature, WNF virus has been isolated from mos-
quitoes and ticks, but mosquitoes are considered
the major vectors. Studies in Egypt, Israel, and
South Africa have implicated Cx univittatus as the
main vector in these countries.35 Cx neavei has been
identified in South Africa. Cx vishnui complex,
containing Cx tritaeniorhynchus, Cx vishnui, and
Cx pseudovishnui, has been implicated in India
and Pakistan.
Rift Valley Fever. Rift Valley fever (RVF) is an
Fig. 21-8. Distinguishing features of Aedes albopictus
Source: Walter Reed Biosystematics Unit, Department of acute, febrile, arthropod-borne yet contagious
Entomology, Walter Reed Army Institute of Research, zoonotic disease caused by a bunyavirus of the
Silver Spring, Md. Art by K. Miyasaka. genus Phlebovirus36 (Table 21-2). Most arboviruses
480
are associated with either a single species of estimated 200,000 human infections and 598
mosquito or a closely related group of mosquitoes, reported deaths. 40 In May 1993, RVF virus was
except for RVF virus. RVF virus has been associated responsible for an outbreak in Aswan Governorate,
with numerous flood-water mosquito species, Egypt.41 RVF epizootics follow periods of excessive
including members of the genera Aedes, Culex, seasonal rain, which bring an increase in mosquito
Anopheles, and Eremapodites.37 Other incriminated population. Outbreaks of RVF have occurred in
vectors include a biting midge (Culicoides species), areas of Africa as diverse as the dry low-rainfall
a black fly (Simulium species), and a tick (Rhipicephalus climate of Egypt, the wet forested areas of Uganda
species).36 and the Central African Republic, and the relatively
RVF virus has been reported in 24 African dry, high veld areas of South Africa where winter
countries. In 2000, it appeared for the first time temperatures frequently drop below freezing.
outside of Africa when it caused a major epidemic RVF virus may be maintained transovarially in
in Saudi Arabia and Yemen.38,39 A major epidemic flood-water mosquitoes, as demonstrated by the
and epizootic in Egypt from 1977 to 1979 caused an recovery of RVF virus from both male and female
TABLE 21-2
ARTHROPODS FOUND NATURALLY INFECTED WITH RIFT VALLEY FEVER VIRUS
Species Locality Species Locality

Aedes antennatus, simpsoni, and Kenya 1982 Madagascar
aedimorphus vansomereni 1979
cumminsii Burkina Faso 1983 theileri Kenya 1982 South Africa
dalzieli Kenya 1982 Senegal 1975, 1983 1953, 1970, 1975
dentatus Zimbabwe 1969 Zimbabwe 1969
tarsalis Uganda 1955 zombaensis Kenya 1982
durbanensis Kenya 1937 Eumelanomyia
Neomelaniconion rubinotus Kenya 1982
circumluteolus Uganda 1955 South Africa Eretmapodites
lineatopennis Zimbabwe 1969 South Africa quinquevittatus spp. South Africa 1971
1975 Kenya 1982, 1984 Uganda 1948
palpalis Central African Republic 1969 Coquillettidia
Ochlerotatus fuscopennata Uganda 1960
caballus South Africa 1953 grandidieri Madagascar 1979
juppi South Africa 1978 Mansonia
Stegomyia uniformis Madagascar 1979
africanus Uganda 1956 Mansonioides
dendrophilus Uganda 1948 africana Uganda 1959, 1968
furcifer Burkina Faso 1983 Central African Republic
Anopheles 1969
anopheles uniformis Uganda 1960
coustani Zimbabwe 1969
coustani and fusicolor Madagascar 1979 Other Diptera
Cellia Culicoides spp. Nigeria 1967
pauliani and Madagascar 1979 Simulium South Africa 1953
squamosus
christyi Kenya 1982
pharoensis Kenya 1982
Culex
antennatus Kenya 1982 Nigeria 1967, 1970
antennatus, annulioris gp, Madagascar 1979
simpsoni, and vansomereni
Reprinted with permission from: Meegan JM, Bailey CL. Rift Valley fever. In: Monath TP, ed. The Arboviruses: Epidemiology and
Ecology. Vol 4. Boca Raton, Fla: CRC Press; 1989: 51–76.
481
Ae lineatopennis collected from larvae in the field Eastern

and reared to adults in the laboratory.42
The Equine Encephalitides. The Togaviridae
include two genera, only one of which, Alphavirus
(or Group A virus) is arthropod-borne. All but one
of the 15 Alphavirus species are transmitted by
mosquitoes.3 The major Alphaviruses transmitted
by mosquitoes are eastern equine encephalitis (EEE)
virus, Venezuelan equine encephalitis (VEE) virus,
and western equine encephalitis (WEE) virus.
EEE virus has been isolated in equine and human
cases from Canada to Argentina along the eastern
seaboard of the Americas and in the Caribbean
region, southeast Asia, and eastern Europe. The Western
main vectors associated with transmission of EEE
are Culiseta melanura, Ae taeniorhynchus, Ae sollicitans,
and Ae vexans. EEE virus generally causes a severe
and often fatal encephalitis; its mortality ranges
from 30% to 60%. Between 1961 and 1985, however,
only 99 human cases were reported.43
EEE virus was first isolated in 1933 from the
brains of horses and in 1938 was identified as caus-
ing an epidemic in Massachusetts. The viruses iso-
lated from outside the New World have not been
associated with human or equine cases. Culex pipiens
is known to circulate this virus in bird–mosquito
enzootic cycles in Europe.44 The ecology is poorly
understood, and the mechanism of spread to hu- Fig. 21-9. The distribution of eastern and western equine
mans remains speculative. Research investigations encephalitis in the United States.
suggest that EEE virus is maintained in North • Sporadic or focal infection reported
Source: Centers for Disease Control and Prevention.
America in an enzootic bird–mosquito cycle. The
Vector-borne Disease Control Self-study Course 3013-G.
main enzootic vector is a fresh water swamp mos- Manual G. Mosquitoes of Public Health Importance and their
quito, Cs melanura, which rarely bites horses or hu- Control. Atlanta: CDC; 1994: 4.
mans. Epidemics in humans and equines occur
when the amount of virus in hosts living in swamp
environments becomes high and it is carried by virus in Ae aegypti invaded many of the mosquito’s
mosquitoes to birds and mammals outside of the tissues, with the highest levels in the salivary
swamp environment (Figure 21-9). glands.
VEE is caused by a number of serotypes and WEE is found in the United States west of the
causes fatalities in horses, mules, donkeys, and Mississippi River and in Wisconsin, Illinois, and
humans. In 1971, an outbreak of the 1B serotype of Canada. It is also present along the eastern seaboard
VEE virus occurred in Texas in association with a of North and South America and in eastern Europe.
similar outbreak in Mexico. 45 Primary vectors There were many major outbreaks in horses in the
during this outbreak in South Texas were Psorophora 1930s, with thousands of cases and many deaths. The
columbiae, Ps discolor, Ae sollicitans, and Ae thelcter. largest human epidemic, involving approximately
Other species of the genera Culex, Anopheles, 3,000 cases, occurred in the western United States in
Mansonia, and Deinocerites were also found infected 1941. Another large epidemic was reported in 1990
in Texas and may be important vectors in other in the central valley of California.47
countries. Twenty major outbreaks of VEE between Culex and Culiseta mosquitoes are known vectors
1935 and 1971 have occurred in 11 countries but of this virus. In western endemic areas, Cx tarsalis
have mainly been centered in Colombia and is the most important vector of WEE because of its
Venezuela. VEE virus isolations from mosquitoes suitability for propagating and retaining WEE
have mainly involved Ae serratus, Ae taeniorhynchus, virus48 (Figure 21-10). WEE has been isolated from
Ps confinnis, and Ps ferox. A study46 showed that VEE Cs melanura in the eastern United States.49 Isolations
482
Fig. 21-10. Culex tarsalis, a major vector of western equine Fig. 21-11. Phlebotomus papatasi, a phlebotomine sand fly
encephalitis. vector of Old World cutaneous leishmaniasis.
Source: The Ken Gray Image Collection, Oregon State Photograph: Courtesy of Ed Rowton, Walter Reed Army
University Department of Entomology, Corvallis, Ore. Institute of Research, Silver Spring, Md.
of WEE virus have also been made from many other of the female sand fly is typically sharp and painful
species of mosquitoes and from birds.50 and often causes considerable irritation. Extensive
reviews of the biology and vector associations of
Flies New World51 and Old World52 sand flies have been
published. There are over 700 species of sand flies
Sand Flies that are found in a wide range of habitats.
Female sand flies require vertebrate blood for
Sand flies are Diptera of the family Psychodidae. maturation of their eggs. Some species feed once
Nonbiting moth flies in the genus Psychoda are also between ovipositions; others may take multiple
in this family, but only the sand flies are medically blood meals during a single oviposition cycle. Most
important. Sand flies occur mainly in the tropics and human-biters feed at dusk and during the evening,
subtropics, with a few species ranging into temperate but some species will bite during the daytime as
zones of the northern (to 50° N) and southern (to well if they are disturbed in their resting site.
40° S) hemispheres. Distribution is limited to areas Windless or nearly windless, dark conditions may
that have temperatures above 15.6°C for at least suddenly induce great numbers of sand flies to seek
three months of the year. There are no sand flies in hosts. The majority of anthropophagic sand flies are
New Zealand or on Pacific Islands. The genera also exophagic, biting persons outside their houses.
Phlebotomus, Chinius (represented by a single species However, some species are endophagic, readily
in China), and Sergentomyia occur in the Old World entering human dwellings, where they bite the
and the genera Brumtomyia, Warileya, and Lutzomyia occupants and either leave or rest inside.
occur in the New World. Anthropophilic sand flies The type of resting site used by adult sand flies
in the Old World are distributed mostly in the varies according to species, availability of microhabitat,
subtropics, with a few human-biters south of the season, and amount of moisture present. In the New
Sahara and none in Southeast Asia. In the New World, most species prefer the tropical rain forest. The
World, they are limited mainly to the tropics.3 forest has many microhabitats used by resting sand
Characteristics. Sand flies are small (2–4 mm in flies, such as on the underside of leaves; on or under
length), delicate flies with long, thin legs and tree bark, on tree trunks, and in animal burrows; in
narrow, pointed wings with parallel venation hollow trees; and in rock crevices and caves. In the
(Figure 21-11). They are small enough to pass Old World, sand flies tend to breed and rest in drier
through the mesh of a standard mosquito net. At habitats. They can be found associated with termite
rest, the wings are held erect over the abdomen at a mounds, cracks and fissures in the soil, animal
45° angle. The proboscis of the sand fly is short, and burrows, piles of rubble, cracks in stone or brick walls,
the antennae are long. Only the females bite and and caves. In peridomestic situations in both the New
suck blood, thus acting as disease vectors. The bite and Old World they may be found resting on walls
483
inside human or animal dwellings.
Diseases Transmitted by Sand Flies
The Leishmaniases. The public health impact of

the leishmaniases has been grossly underestimated
due mainly to lack of public awareness. Only during
the last 2 decades has it become increasingly apparent
that most forms of leishmaniasis are much more
prevalent than previously expected. 53 The leish-
maniases are endemic on four continents in 22 New
World and 66 Old World countries; 16 are developed
countries, 72 are developing countries, and 13 are
among the least-developed countries. Major epi-
demics of visceral leishmaniasis are in progress in
Fig. 21-13. Verruga peruana lesions on a child in Peru.
Brazil, India, and the Sudan. In Bihar State in eastern This extreme case resolved after treatment.
India, it is believed that 200,000-250,000 people Photograph: Courtesy of Richard Andre, Uniformed Ser-
contracted the disease in 1993. In southern Sudan vices University of the Health Sciences, Bethesda, Md.
15,000 cases were treated and 100,000 deaths occurred
in a 5-year period (1991-1996). Cutaneous leish-
maniasis is on the rise in many countries with objectives is significant (Figure 21-12). Patients seen
epidemics ongoing in newly settled areas of the at Walter Reed Army Medical Center, Washington,
Amazon basin, North Africa and the Middle East.53,54 DC, have included both active duty and reserve
Among the more than 500,000 service members service members from all three branches of the US
deployed to the Persian Gulf region during the Persian military, as well as personnel temporarily or
Gulf War, 28 cases of leishmaniasis were identified in permanently assigned to endemic areas. The number
personnel after their return from the theater of of units represented by this relatively small number
operations. Of these, 11 were identified as viscero- of patients suggests that many more service members
tropic, and 17 presented with cutaneous lesions.55 The were at risk.57
causative agent of the visceral form, the protozoa Bartonellosis. Bartonellosis (Carrion’s disease),
Leishmania tropica, was previously thought to cause also believed to be transmitted by sand flies, is
only cutaneous disease.56 caused by the bacterium Bartonella bacilliformis and
Although not a war stopper, leishmaniasis is a only occurs in rural areas in the Andes mountains
persistent health threat to US military personnel of South America. The disease causes two distinct
because they are deployed or conduct military clinical manifestations in humans: Oroya fever and
exercises in locations where the disease is endemic verruga peruana. The former condition is character-
and its overall potential to compromise mission ized by severe hemolytic anemia, joint pains, pallor,
fever, and jaundice. Mortality rates range as high
as 40% if untreated. Salmonella infections are serious
concomitant complications of Oroya fever.
The other, less life-threatening form of bartonellosis,
verruga peruana, is named for the warty appearance
of the skin of infected persons (Figure 21-13). These
lesions usually develop following recovery from
Oroya fever and may last for up to a year. The
number and appearance of the painless nodules
varies considerably. Lutzomyia sand flies are the
suspected vectors of both forms of this disease.58
Humans are thought to be the only known host.
Sand Fly Fever. Viruses in three families and three
genera are transmitted to humans and other
vertebrates by sand flies. Symptoms of human
Fig. 21-12. The geographic distribution of leishmaniasis. illness usually resemble influenza, with fever, retro-
Courtesy of Colonel Raj Gupta, MS, US Army, and the orbital pain, myalgia, and malaise. Complete
Walter Reed Army Institute of Research, Washington, DC. recovery within a week is the norm, but encephalitis
484
has been reported in at least two patients infected

with vesiculovirus. Sand fly fever (also known as 3-
day fever or papataci fever), the best known example,
occurs in the Mediterranean region, the Middle East,
Pakistan, and northern India where Phlebotomus
vectors exist. Stiff neck and limbs, red face, severe
headache, and painful neck muscles may occur. Long-
lasting immunity is conferred after the first attack. The
disease has been considered of military importance
because up to 75% of nonimmune adults arriving in
an endemic area can be affected.59
In the Mediterranean area, the sand fly Phlebotomus Fig. 21-14. A black fly (Simulium sp).
papatasi is the suspected vector of sand fly fever in Photograph: Courtesy of Colonel Philip Lawyer, MS, US
humans. Although little if anything is known about Army.
transmission cycles, it is suspected that vertebrates,
rather than insects, serve as reservoirs of the viruses.2 40,000 new victims each year. The 1990s, however, saw
a decline in the prevalence of onchocerciasis infection
Black Flies and morbidity due to the remarkable success of vector
control programs in West Africa.
Black flies are blood-sucking Diptera of both public Black flies also act as mechanical vectors of
health and economic importance. Besides being myxomatosis and certain arboviruses, such as Vene-
disease carriers, they are annoying to humans and zuelan equine encephalitis. However, this mode of
domestic animals because of their biting and crawling transmission is normally of minor importance.
habits. Black flies belong to the family Simuliidae,
which contains several genera but only one of which, Tsetse Flies
Simulium, is an important human-biting fly. These are
small, stoutly built, hump-backed flies. Only the There are at least 23 species of tsetse flies belonging
female feeds on blood and is a daytime feeder. These to the genus Glossina.61 These flies are found only on
flies have a worldwide distribution where suitable the continent of Africa south of the Sahara desert and
conditions exist; there must be well-oxygenated water infest an area of approximately 12 million square
for the immature stages to develop. The larvae feed kilometers. They are larger than a house fly and can
by filtering particles from the water and brushing easily be recognized by the wings, which extend
microorganisms into the mouth using a pair of beyond the abdomen when the fly is at rest and lie
prominent mouth brushes. Adults emerge from the one over the other like a crossed pair of scissors. The
water, often attacking in large numbers. In the tropics, wings also have characteristic venation, with a discal
there are multiple generations each year. The medial cell shaped like a hatchet or cleaver (Figure
mouthparts of the female are short and broad. When 21-15). Both sexes of tsetse suck blood and feed
feeding, they tend to stab the tissue and wait for the
ruptured capillaries to ooze blood rather than sucking
neatly like mosquitoes do. Simulium bites cause
considerable pain and irritation and frequently
become infected (Figure 21-14). The punctum heals
by forming a characteristic black scab.
The parasitic round worm, Onchocerca volvulus,
which causes onchocerciasis, is transmitted by
Simulium species. Of the 18 million people infected
with this parasite, more than 95% live in Africa.60 The
clinical manifestations of the disease include dermal
nodules and lymphatic and systemic complications
due to vessel blockage by the microfilariae. This causes
severe itching, papule formation in the skin, and
thickening and loosening of the skin, which sub- Fig 21-15. The tsetse fly (Glossina morsitans) after taking
sequently hangs in folds. The most severe compli- a blood meal.
cations are onchocercal lesions of the eye, which may Photograph: Courtesy of Ed Rowton, Walter Reed Army
lead to blindness (“river blindness”). This affects Institute of Research, Silver Spring, Md.
485
exclusively by day. susceptible host during a subsequent blood meal.

Two groups of Glossina are involved in the Adult worms live under the skin and migrate
transmission of African trypanosomiasis or “sleeping around the body, causing allergic “Calabar”
sickness” to humans. Other species transmit the swellings and abscesses at the site of the dead
trypanosome causing nagana in cattle. The riverine worms. The adult worm becomes visible to the
areas of West Africa, mainly the Niger River and patient if it crosses the anterior chamber of the eye,
Congo River basins, are the habitat of G palpalis, but it does not cause serious ocular damage.
vector of the chronic Gambian form of the disease Occasionally, worms invade the brain, causing
caused by Trypanosoma brucei gambiense. Further meningoencephalitis. Some tabanids, particularly
inland, where the rivers shrink into pools in the dry certain species of Chrysops, are known to serve as
season, the vector is the smaller species G tachinoides.2,3 mechanical vectors of tularemia, anthrax, and
In the dry bush country of East Africa, species of possibly other diseases.62
tsetse occur that do not require the humid conditions
of river bank vegetation. G morsitans is found in Filth Flies
wooded savanna where rocks and trees provide the
shade that protects the fly from desiccation. G The term filth fly is generally applied to flies that
swynnertoni inhabits much more open terrain where live in close association with humans (synanthropes)
the only shade is provided by large animals. Both and which feed readily on human feces and other
species are associated with the Rhodesian form of decomposing organic matter, as well as food intended
the trypanosome found in wild game and infective for human consumption.63 These flies are of medical
to humans, T brucei rhodiesiense. importance because they act as mechanical vectors of
pathogenic organisms, especially those causing enteric
Deer Flies and Horse Flies diseases.
The fly genera Musca, Fannia, and Calliphora all
Although of only minor importance to military contain important filth fly species. These flies are
operations, deer flies and horse flies can be a medium-to-large, burly flies with conspicuous
nuisance in localized areas. Deer flies (genus compound eyes and a well-developed thorax and
Chrysops) and horse flies (genus Tabanus) are the two abdomen (Figure 21-17). They lay eggs on feces or
most common genera in the family Tabanidae. They other decaying organic matter. The maggot-like larvae
are robust flies varying in length from 5 to 30 mm subsequently develop and consume the organic
and have clear or mottled wings. The bite of female matter. The adult fly consumes only liquid food. The
deer flies and horse flies can cause a severe reaction fly regurgitates fluid from its crop onto a food source
in a sensitized host, with considerable edematous to liquefy the material, and then sucks it up.
swelling, pain, and irritation (Figure 21-16). Throughout history, military personnel have
Species of Chrysops known as mango flies, C silacea suspected the association between flies and enteric
and C dimidiata in tropical west Africa (from Sierra
Leone to Ghana and Nigeria) and C distinctipennis in
central Africa (southern Sudan and Uganda), transmit
the filarial worm, Loa loa, which causes loiasis in
humans. The microfilariae are taken up by the female
fly from the peripheral blood of an infected human
host during the day. The parasite is passed to the
a b
Fig. 21-16. (a) a horse fly (Tabanus punctifer) and (b) a

deer fly (Chrysops discalis). Fig. 21-17. A female filth fly (Musca domestica).
Source: The Ken Gray Image Collection, Oregon State Source: The Ken Gray Image Collection, Oregon State
University Department of Entomology, Corvallis, Ore. University Department of Entomology, Corvallis, Ore.
486
diseases. They have observed that a rise in enteric

disease follows a rise in fly populations. Greenburg
discusses evidence that Shigella, Salmonella, infectious
hepatitis virus, and typhoid fever virus are trans-
mitted and spread by a variety of filth fly species.
Effective fly control programs have demonstrated on
many occasions that fly control can reduce the
incidence of enteric disease.63 However, there are
many examples that caution against an exclusively
entomological approach to the epidemiology of
enteric diseases. There are many potential vehicles for
transmission of such diseases, and their importance
depends in part on the habits and lifestyle of the
people involved. Thus, field sanitation and personal
hygiene are as important today as they always have Fig. 21-19. Calliphora vicina, the flesh fly.
been. Source: The Ken Gray Image Collection, Oregon State
University Department of Entomology, Corvallis, Ore.
Eye Gnats
unable to develop through their immature stages
Hippelates eye gnats of the family Chloropidae are except on living tissue. This condition is known as
an important group of synanthropes that frequent the specific or obligatory myiasis and is restricted to
face and eyes, as well as the mucous and sebaceous tropical species. Examples include the Old World
secretions and the wounds, pus, and blood of humans. screwworm (Chrysomyia bessiana), the tumbu fly
Certain species have long been suspected as possible (Cordylobia anthropophaga) and the Congo floor maggot
vectors of conjunctivitis and pinkeye or sore eye of (Auchmeromyia luteola) in Africa, and the New World
humans and animals.63 In Jamaica, Hippelates flavipes, screwworm (Cochliomyia hominivorax) and the human
which feeds avidly on humans, has been implicated bot fly (Dermatobia hominis) in Central and South
in yaws transmission. America. Specific myiasis occurs when fly larvae
Adult eye gnats fly throughout the day. These flies invade the skin. The infection presents as multiple
are nonbiting but may produce minute incisions with painful blind boils in which secondary infection may
their labellar spines. They feed on exudates of sores occur (Figure 21-18). The head of the boil shows the
and cuts and around natural orifices. black spiracles of the fly larvae. Treatment involves
the suffocation of the larvae with an oily ointment,
Blow Flies, Bot Flies, and Flesh Flies after which they can be removed with forceps; care
must be taken to ensure that the larvae are removed
Myiasis is the term given to an infestation of living intact.5 Another method that seems effective and less
tissue by dipterous larvae. Some species of fly are likely to result in infection or other complications is
commonly used in endemic areas of South and Central
a b America. A slab of raw meat is placed over the boil
and held there overnight. When the fly larva surfaces
to breathe, it crawls between the surface host’s skin
and the meat, or it may burrow into the slab of meat
and can then be removed. The presence of larvae can
be psychologically distressing, but they are not usually
life threatening.
Fig. 21-18. A warblelike lesion produced by the larva of Some larvae of Calliphora, Lucilia, and Sarcophaga
the human bot fly, Dermatobia hominis, is shown in pho- (the flesh fly) species that normally develop on
tograph (a). A Dermatobia hominis larva is shown in pho- decaying organic matter will readily feed on wounds
tograph (b). The adult fly lays her eggs on the abdomen
or other live, damaged, or contaminated tissue (Figure
of a blood-sucking insect, such as a mosquito (and some-
times a tick). When the mosquito bites a host, the prox-
21-19). This condition is known as semi-specific or
imity of the Dermatobia eggs to the host causes the eggs facultative myiasis and can sometimes occur in the
to hatch and the bot larvae burrow into the skin. anal and genital regions. Accidental myiasis occurs
Photographs: (a) Courtesy of Colonel Philip Lawyer, MS, US Army. when fly eggs or larvae are swallowed in food, and it
(b) Armed Forces Institute of Pathology, Washington, DC. results in intestinal disturbances.
487
Fleas Lice
Fleas are small, wingless insects that can be Lice are small (1-4 mm long), wingless insects with
important vectors of disease and often are serious elongated or discoidal bodies that are compressed or
pests. Their bodies are flattened laterally, are flattened dorso-ventrally. Three kinds of bloodsucking
usually brown, heavily sclerotized, and generally lice infest humans: body lice, head lice, and crab
possess bristles. The legs are well developed for (pubic) lice. The head louse and body louse are
jumping. The oriental rat flea, Xenopsylla cheopis, somewhat elongated and similar in appearance, but
found on commensal rats in many parts of the the head louse is the smaller of the two. The crab louse
world, is the most important vector of bubonic is much smaller and has a nearly round, hairy body.
plague and murine typhus. The plague bacilli and The life cycles of these three species vary slightly, but
murine typhus rickettsiae are ordinarily acquired all take place on the body or clothing of the host. The
by the flea during feeding on infected rats. Then body louse confines its feeding to the host’s body and
the pathogens may be transmitted to humans in the remains on the clothing next to the skin. The head
absence of the flea’s normal hosts, as when the rats louse lives in the hair of the head and the crab louse
are dying of plague in an epizootic. Plague exists among the hairs of the pubic region. The crab louse
in a sylvatic form in widespread parts of the world, can also be found on the hairs of the legs, the chest,
and many of the 2,000 species of fleas are vectors of the armpits, and occasionally the beard and eyebrows.
this disease.2 Fleas can be very annoying, and their The genera Pediculus and Pthirus contain the lice
bites may produce itching and dermatitis in that normally infest humans (Figure 21-20). Pediculus
sensitive individuals.5 humanus, containing both the head louse (P humanus
There are identification keys for most common capitis) and body louse (P humanus humanus—no
fleas of public health importance. Fleas are longer P humanus corporis) of humans, is one of the
generally discussed by families, which are based on three or four species included in genus Pediculus.
the fleas’ morphological characteristics and their Ferris64 has discussed the nomenclature in detail. The
ability to serve as vectors. The families are genus Pthirus includes the crab louse, Pthirus pubis.
Ceratophyllidae, Leptopsyllidae, Pulicidae, and The most important member of this group is the
Tungidae. Family Pulicidae includes a significant human body louse. Historically, it has had a more
number of species that are pests of humans, profound effect on the history of humanity than has
domestic fowl, and pets; are vectors of the plague any other insect.65 Lice have always been associated
pathogen; and are putative vectors of murine with wars because they thrive when sanitary
typhus to humans. conditions are poor and human populations are
Fleas have piercing and sucking mouthparts and homeless and dislocated.
feed exclusively on blood. They lay their eggs on
or among the hairs or plumage of the host or on Head Lice
debris on the ground. Eggs hatch into yellowish
white, maggot-like larvae. The flea larva usually has The head louse is gray but tends to resemble the
one or two rows of sparse but well-developed color of the hair of the host. Head lice remain mostly
bristles on most of its segments. The larva usually
lives in the nest of the host and feeds on host-
associated organic debris including food particles,
dried skin, dried blood, or excreta. Larvae of some
medically important species undergo three molts
in 2 to 3 weeks. It may take a few months in other
species. The larva spins a silken cocoon around
itself, thus entering the pupal stage. The pupa
develops into an adult in a few days. Adult
emergence from the cocoon may be triggered by
Body Louse & Head Louse Crab Louse
vibrations resulting from host movements. Adults Pediculus humanus Pthirus pubis
may live for weeks or months, sometimes even
without food. If environmental conditions are Fig. 21-20. The body louse, head louse, and crab louse.
unfavorable, or if hosts are not available, de- Source: Centers for Disease Control and Prevention. Vec-
veloping adult fleas may remain inactive within the tor-borne Disease Control Self-study Course 3013-G. Atlanta:
cocoon for extended periods. CDC; 1994: 11.
488
on the scalp. Head lice have been pests of human- Ethiopia; Asia; and the higher altitudes of Mexico,
kind throughout history and pose no health threat to Central America, and South America.2 The causative
infested persons. However, the infestation may lead organism is Rickettsia prowazekii.
to embarrassment and social ostracism. Head lice are Trench fever is a nonfatal disease, and the causative
small (1–3 mm long), elongate, wingless insects that agent is Bartonella quintana. This organism multiplies
occur on the head and around the ears and occiput, freely in the lumen of the louse’s digestive tract and
but heavy infestations may move on to other parts of is not pathogenic to the louse. Epidemic relapsing
the body. The female lays the eggs on the hair. The fever has occurred in many parts of the world. The
eggs are glued to the hair and take about 5 to 10 days pathogen is a spirochete, Borrelia recurrentis. The in-
to hatch. The nymphal stage includes three molts sect can acquire the pathogen by a single feeding on
before changing into the adult stage. The entire life an infected person but cannot directly pass it on to a
cycle takes about 3 weeks. second person. It is transmitted by crushing the louse
and so releasing its infected hemolymph onto the skin.67
Body Lice
Bugs
The body louse looks almost identical to the head
louse, but it usually stays on clothing and makes con- The true bugs may be winged or wingless, but they
tact with the body while feeding. In heavy infesta- always have a proboscis or beak suitable for piercing
tions, some lice may remain on the human body even and sucking that is attached anteriorly and kept flexed
when all clothing has been removed. Eggs are depos- under the head when not in use. The two bugs of
ited in the seams of clothing. The incubation period medical importance belong to the families Cimicidae
varies from 5 to 7 days when eggs are laid near the (bedbugs) and Reduviidae (assassin and kissing bugs).
body; it is longer at lower temperatures. After hatch-
ing, the young lice begin to suck blood at once and Assassin and Kissing Bugs
feed frequently throughout the nymphal stage. Like
the head lice, the body lice nymphs go through three These bugs are commonly called cone nose bugs
molts before becoming adults and the egg-to-egg cycle because of their elongate (cone-shaped) head (Figure
takes about 3 weeks. The optimum temperature for 21-21). Most Reduviids “assassinate” or kill other
development is similar to normal human body tem- insects, but a small group of reduviids belonging to
perature. The louse does not leave the body unless the subfamily Triatominae exclusively feeds on the blood
body becomes too cold (death) or too hot (high fever). of vertebrates.2 They are also called kissing bugs
New louse infestations mainly occur during contact with because occasionally they take the blood meal from
louse-infested persons or their clothing. around the lips of the host. The bites of these bugs are
Crab Lice
Crab lice are only found on humans and die within

24 to 48 hours if forced to leave the host.66 The eggs
are laid mainly on the coarse hairs of the pubic area.
Nymphs pass through three molts before becoming
adults. The total life cycle takes about 3 to 4 weeks.
Pubic lice are transmitted from person to person, most
often by sexual contact.
Diseases Transmitted by Lice
Three diseases of public health importance that are

associated with lice are louse-borne (epidemic) ty-
phus, trench fever, and epidemic relapsing fever.
Pediculus humanus humanus is probably the sole vec-
tor of the organisms that cause these diseases. Louse- Fig. 21-21. An assassin or kissing bug (Family Reduvi-
borne (epidemic) typhus has been recognized since idae), which is the vector of American trypanosomiasis.
ancient times and occurs mainly in Europe; the Afri- Photograph: Courtesy of Ed Rowton, Walter Reed Army
can highlands, especially Burundi, Rwanda, and Institute of Research, Silver Spring, Md.
489
usually painless but may cause urticaria if the host Stinging Insects
becomes sensitive to the injected saliva. Chagas’
disease, or American trypanosomiasis, is one of the Allergic reactions to insect stings are a common and
most important arthropod-borne diseases in tropical often a serious medical problem: estimates of
America; it is caused by Trypanosoma cruzi, which is anaphylaxis in the general population range from
transmitted by various species of Triatominae. The 0.3% to 3%.69 Insects that sting are members of the
four principal vectors of Chagas’ disease in Central order Hymenoptera. There are two major subgroups:
and South America are Panstrongylus megitus, Rhodnius vespids (eg, yellow jacket, hornet, wasp) and apids
prolixus, Triatoma infestans, and Triatoma dimidiata. (eg, honeybee, bumblebee); ants belong to a third
The most important vector in Mexico is T barberi, subgroup. In most parts of the United States, yellow
while T gerstaeckeri and T proctata are important jackets are the principle cause of allergic reactions to
species in the southwestern United States, and T insect stings.
sanguisuga is found throughout the United States.68 The stinging apparatus originates in the abdomen
The victims of this disease are mainly humans, dogs, of the female insect. It consists of a sac containing
and to a varying extent other domestic and wild venom attached to a barbed stinger. A sting occurs
animals. The infection mainly occurs in rural areas when the stinger is inserted into tissue, the sac
of the tropics and subtropics, but may occur in contracts, and venom is deposited into tissue.
suburban and urban areas around poorly constructed
houses. Bees and Wasps
Both sexes of triatomid bugs bite, and they take
their blood meal at night and hide in any cracks or Although Africanized honeybees (Apis mellifera
crevices when not feeding.3 The pearl-like eggs are scutellata), the so-called “killer bees,” have received
laid singly around the adult habitat. The incubation much publicity, their venom is no more allergenic or
period varies, depending on the species and toxic than that of European honeybees (Apis mellifera
temperature. The nymphs typically go through five mellifera).69 However, Africanized bees are much more
developmental instars. Nymphs camouflage them- easily provoked and more aggressive than European
selves with dust particles or other debris. In temperate honeybees. This behavior can lead to massive stinging
regions, some species overwinter in the egg stage, incidents. These bees are expected to keep moving
others as adults, and still others as nymphs. northward in the United States, although they do not
survive well in colder areas.
Bedbugs Reaction to honeybee stings range from slight pain
and swelling to much more serious symptoms,
Bedbugs have been associated with humans for including anaphylaxis. In the United States, deaths
centuries. Bedbugs have been occasionally found from all hymenopterous insects (bees, wasps, yellow
infected with anthrax, plague, and typhus disease jackets, and ants) average between 40 and 50 per
organisms, but they are not considered an important year.70 A single bee sting is seldom fatal unless its
vector of these diseases. 5 They are extremely victim is hypersensitive and has a severe allergic
annoying; their bites produce small hard swellings reaction. All persons should know whether or not they
or wheals that are often confused with flea bites. are hypersensitive to bee and wasp stings (Figure 21-
The bedbug’s principal medical importance is the
itching and inflammation associated with its bite.
Two bedbug species attack humans: Cimex lectu-
larius (in temperate regions) and Cimex hemipterus
(in tropical areas).2
Bedbugs are dorso-ventrally flattened, reddish
brown, wingless insects approximately 5 mm long.
They lay eggs in wall cracks, furniture, bedding, and
other sheltered places. The eggs hatch in about 6
days, and if a suitable host is available, the young
bugs begin feeding on blood. Immature bedbugs
look like adults except they are yellowish white. The
nymphs mature into adults in 30 to 45 days. Adults
live for 6 to 8 months and may survive for several
days without food. The adults lack hind wings, and Fig. 21-22. A wasp. Photograph: Courtesy of Ed Rowton,
the forewings are reduced to two small pads. Walter Reed Army Institute of Research, Silver Spring, Md.
490
22). Hypersensitive persons should carry insect sting Fire Ants

kits when frequenting an area where interaction with
bees and wasps may occur. The fire ant, Solenopsis invicta, is a nonwinged
Honeybees are often encountered in swarms, which hymenopteran. It is responsible for an ever-increasing
can remain in an area a few hours or days. Bee swarms number of allergic reactions in the United States and
are only temporary and will move away as soon as throughout its range in North and South America.
the bees find a new location to colonize. However, With its jaws, the fire ant attaches itself to a person. It
situations may occur that require the immediate then pivots around its head and stings with the
elimination of bee swarms. Swarms may be eliminated abdominal stinger at multiple sites in a circular
with approved insecticides or by using high-volume pattern. Within 24 hours, a sterile pustule develops
sprays of a 5% solution of liquid dishwashing soap.70 that is diagnostic of a fire ant sting.
CLASS ARACHNIDA
The class Arachnida includes such diverse forms in ticks and some mites, a hypostome. In general,
as ticks, mites, spiders, and scorpions and is found arachnids are predatory or parasitic, although many
almost exclusively in terrestrial habitats throughout mites are plant feeders or scavengers.
the temperate and tropical regions of the world. The Like many of the medically important insects, ticks
most important of the arachnids—ticks, mites, and and some mites are important as mechanical and
spiders—lack distinct body segmentation; scorpions, biological vectors of bacteria, rickettsia, viruses, and
pseudoscorpions, and a few others are obviously seg- protozoa, which they transmit to humans and other
mented. The body is divided into two parts, the animals by their bites.
cephalothorax (comprosed of the combined head and
thorax) and the abdomen. In ticks and mites, the Ticks and Mites
cephalothorax and abdomen are strongly fused, giv-
ing the body a sac-like form. These arthropods have Ticks
four pairs of legs, at least in the adult stage; larval
ticks and mites have only three pairs of legs. None There are three families of ticks, of which two
of the arachnids possess antennae or wings. The contain species capable of transmitting pathogens to
mouthparts lack mandibles and usually consist of a humans: the hard ticks (family Ixodidae) and the soft
pair of piercing chelicereae and the pedipalpi, and ticks (family Argasidae) (Figure 21-23). Hard ticks are
a. Hard Tick b. Soft Tick

Capitulum
Capitulum
Palpus Not
Visible
Visible
Basis Capituli
Eye
Dorsal Shield
Leg
Leg
Festoons
Tapered Blunt
Anteriorly Anteriorly
Fig. 21-23. The characteristics of female hard and soft ticks.

Source: Centers for Disease Control and Prevention. Vector-borne Disease Control Self-study Course 3013-G. Atlanta:
CDC; 1994: 14.
491
oval with a sac-like body and vary in length from 2 to

10 mm when unfed, although the engorged female
may be at least twice this size. They possess a hard
dorsal plate called a scutum (hence the name “hard
ticks”), and their mouthparts point anteriorly and are
visible from above. Soft ticks are leathery and
nonscutate. They are about the same size as the hard
ticks. Their mouthparts point downward and are not
visible from above.
Ticks feed by using their mouthparts to cut a small
hole in the host epidermis; they insert the hypostome
into the cut, thereby attaching themselves to the host.
Blood flow is presumably maintained with the aid of
an anticoagulant from the tick’s salivary glands. Some
hard ticks also secrete a “cement” to secure their Fig. 21-24. Various life stages of the Ixodes scapularis tick.
attachment to the host. Blood-fed female ticks are The larva has six legs and the nymph and adult have
capable of enormous expansion. eight. The scale is in millimeters.
In the United States, more vector-borne diseases Photograph: Courtesy of Colonel Philip Lawyer, MS, US
are transmitted by ticks than by any other arthropod.71 Army.
Documented cases of ticks and tick-borne diseases
affecting military personnel are increasing.72 Ticks only sporadic cases of tick-borne relapsing fever are
generally affect military operations in two ways: (1) diagnosed, but isolated epidemics have occurred in
directly, by tick bite and the accompanying psychological such places as the Grand Canyon.78
stress, and (2) indirectly, by disease transmission. Tick-borne relapsing fever has an incubation period
Various bacteria, rickettsiae, viruses, and protozoa are of 4 to 18 days, after which the illness characteristically
transmitted to people via tick bites. begins abruptly with high fever, chills, tachycardia,
Lyme Disease. Lyme disease or Lyme borreliosis headache, myalgias, arthralgias, abdominal pain, and
has emerged as the most common vector-borne malaise. If the disease is untreated, the primary fever
disease in the United States. It also occurs in Eurasia breaks in 3 to 6 days and is followed by an afebrile
and Australia. In the United States, 16,273 cases were interval of approximately 8 days. Without treatment,
reported in 1999 73 and nearly 50,000 cases were 3 to 5 relapses typically occur. The severity of illness
reported from 1982 through 1992.74 Most human generally decreases with each relapse. Death is rare
infections of Borrelia burgdorferi in the United States and limited mainly to infants and the elderly.
occur during the months of May through August, Antibiotics are an effective treatment.
when tick activity is at its peak. Tularemia. Tularemia is caused by the bacillus
The causative agent of Lyme disease is the Francisella tularensis. The most important reservoirs
spirochete B burgdorferi, and it is transmitted by Ixodes for F tularensis are rabbits, hares, and ticks. The many
ticks. 75 Ixodes scapularis (previously known as I modes of transmission include tick bite, direct contact
dammini) (Figure 21-24) in the East and upper Midwest with infected animal tissues, inhalation of aerosolized
and I pacificus in the West are the primary vectors of organisms, ingestion of contaminated meat or water,
this disease in the United States. Ixodes ricinus is the and bites of infected mammals, deer flies, or
vector in eastern, central, and western Europe and I mosquitoes.77 The three major tick vectors of tularemia
persulcatus in Japan, eastern Russia, and China.76 in the United States are Amblyomma americanum
Tick-borne Relapsing Fever. Tick-borne re- (Figure 21-25) in the southeastern and south central
lapsing fever is caused by Borrelia species other than United States, Dermacentor andersoni in the West, and
B burgdorferi and is the one significant disease D variabilis in many parts of the country. Tick bites
transmitted by soft ticks. In the United States, the account for about 50% of the tularemia cases in the
geographic distribution of this disease is limited to United States. The incubation period of this disease
remote, undisturbed natural areas in the West. The is usually 3 to 5 days. The severity of tick-borne
tick Ornithodoros hermsi transmits B hermsi in tularemia is highly variable and ranges from mild,
forested mountain areas, generally above an afebrile, self-limited disease to cases of fulminating
elevation of 900 m. In semi-arid plains areas, Borrelia septic shock.79
turicatae is transmitted by O turicata ticks and B Rocky Mountain Spotted Fever. Rocky Moun-
parkeri is transmitted by O parkeri ticks.77 In general, tain spotted fever is caused by Rickettsia rickettsia
492
antimicrobial agents. Death is usually associated with

missed or delayed diagnosis and mistreatment.
Human Monocytic Ehrlichiosis. Ehrlichia chaffeensis
is the sole causative agent of human monocytic
ehrlichiosis in the United States.82 This disease was
first described in 1987 in a middle-aged man in
Arkansas. 83 An outbreak of this disease also oc-
curred in an Army Reserve unit.84 Most of the 250
reported cases have occurred in the central and
south Atlantic states. Recently, the ticks D variabilis
and Am americanum have been identified as potential
vectors.77
Ehrlichiosis generally presents as a nonspecific
febrile illness that resembles Rocky Mountain
Fig. 21-25. Amblyomma americanum, the tick that trans- spotted fever without the rash. Approximately 90%
mits tularemia and human monocytic ehrlichiosis. The of patients have a history of a tick bite in the 3-week
large tick at the top of the slide with the white spot is the period preceding the onset of illness. Characteristic
female, the nymph is in the middle, and male is at the bottom. clinical features include high fever and headache, and
Photograph: Courtesy of James Occi, Biological Sciences, other common symptoms include malaise, nausea,
195 University Avenue, Newark, NJ 07102. vomiting, myalgia, and anorexia. Acute complications,
including acute renal failure, encephalopathy,
respiratory failure, and death, are rare. Tetracycline
and is transmitted by Dermacentor variabilis (Figure (or its analogues) is an effective treatment.
21-26) in the eastern United States and D andersoni in Colorado Tick Fever. Colorado tick fever (“moun-
the western United States. Although Rocky Mountain tain fever”) is caused by an RNA virus and is
spotted fever is widely distributed across the United presumably transmitted by the tick D andersoni.
States, most infections are now acquired in the south Ground squirrels are the main vertebrate reservoir
Atlantic coastal region and the western and south- for this disease. The disease occurs in 11 Rocky
central states.80 Three hundred sixty-five cases were Mountain states. 85 Most human infections are
reported in the United States in 1998.81 This disease, if caused by tick bites, but the virus has also been
properly diagnosed, is easily treated with appropriate transmitted by blood transfusion.77
Babesiosis. Babesiosis is a malaria-like illness
caused by a protozoan parasite that invades
erythrocytes. Babesia microti has been implicated in
babesiosis acquired in the northeastern United
States, and B equi is thought to be the causative
agent of the disease in California.
Babesiosis is usually transmitted by the bite of I
scapularis and I pacificus ticks. The white-footed
mouse, Peromyscus leucopus, is the principal reservoir
for B microti in the northeastern United States.
Tick Paralysis. Five tick species (D andersoni, D
variabilis, Am americanum, Am maculatum, and I
scapularis) can cause tick paralysis in humans. A
neurotoxin that is produced in the tick’s salivary
glands is believed to cause the paralysis.86 This
toxin, which is usually transmitted by an engorged,
gravid female tick, either blocks the release of
acetylcholine at the synapse or inhibits motor-
stimulus conduction.77
Diagnosis is made by finding an embedded tick,
Fig. 21-26. Dermacentor variabilis, the dog tick. usually on the scalp. After removal of the tick,
Photograph: Courtesy of Ed Rowton, Walter Reed Army symptoms generally resolve within several hours
Institute of Research, Silver Spring, Md. or days. If untreated, tick paralysis can be fatal.
493
Boutonneuse Fever. An outbreak of boutonneuse dermatitis or transmit diseases. The larval body is
fever in US Army personnel deployed to Botswana less than 1 mm long and more or less round. As
was reported in 1996.87 More than 30% of the 169 adults, they are about 1 mm long, oval or, more
deployed soldiers sought medical attention for often, hour-glass shaped. They are most often bright
boutonneuse fever symptoms. Subsequent serologi- red. Larvae are six-legged and the only parasitic
cal tests indicated that 39 soldiers had boutonneuse form. The adults and nymphs are eight-legged and
fever. This disease is caused by Rickettsia conori and free-living. Chigger larvae do not burrow into the
other rickettsia species and is believed to be trans- skin or feed primarily on blood. The formation of a
mitted by ticks. This disease has previously been feeding tube, or stylostome, at the site of chigger
reported from travelers returning from Morocco, attachment is characteristic of chigger attack. It is
Kenya, Botswana, and South Africa. presumably the action of the mite’s digestive
fluid that causes the attachment site to itch after
Mites a few hours.
The life cycle of chigger mites is very complex
There are many free-living and predaceous mites, and includes six stages. 88 The female lays eggs
but some groups and families are exclusively singly on soil or litter, and the eggs hatch in about
parasitic. More than 200 families of mites are a week, exposing the six-legged larvae. The larval
recognized in the entomological literature, but only stage is the only parasitic stage. Larvae may attach
a few contain species that affect humans. Some themselves to many species of vertebrates. Engorged
mites are vectors of organisms that cause human larvae leave the host and pass through a quiescent
disease, and others cause dermatitis and allergic stage before they become nymphs and finally
reactions in humans. Most species of mites are so adults. The nymphs and adults are eight-legged and
small that they are barely visible to the naked eye. free living; they feed on insect eggs and other small
Their life cycles are often short (2 to 3 weeks), so soil invertebrates. The total life cycle takes about
mites can increase their numbers very rapidly under 60 days.
favorable conditions. For simplicity, the mites From a public health perspective, chiggers
discussed here are grouped into five categories: deserve attention because they are known to cause
chigger mites, house-dust mites, human biting dermatitis and, more important, serve as vectors of
mites, scabies mites, and follicle mites.5 scrub typhus. Chigger mites are found worldwide.
Chigger Mites. Larvae of mites belonging to the Among the chigger mites that cause dermatitis is the
family Trombiculidae are called “chigger mites” European species Trombicula autumnalis, also known
(Figure 21-27). There are approximately 20 different as the harvest mite, and T alfreddugesi, which occurs
species in the family that are known to cause in the United States and parts of Central and South
America. The chigger mite species that transmit
scrub typhus, Leptotrombidium deliense and L
fletcheri, are prevalent from New Guinea and the
coastal fringe of Queensland, Australia, through the
Philippines and China and westward through
Southeast Asia to Pakistan.2 L pallidum is reported
as the vector of scrub typhus in parts of Japan,
Korea, and the Primorye region of the former USSR.
Scrub typhus has caused significant casualties in
Asian-Pacific military operations.89 The American
6th Army in World War II lost more than 150,000
man-days during their operations in and around
Schouten Islands and Sansapor beach head in
Netherlands New Guinea.90 The causal organism is
Rickettsia tsutsugamushi. The disease has an incubation
period of 6 to 21 days and is discussed in detail in
Fig. 21-27. Chiggers, which are trombiculid mites, are Chapter 36.
vectors of scrub typhus. House Dust Mites. House dust mites, in the
Source: Walter Reed Army Institute of Research, Silver family Pyroglyphidae, do not feed directly on living
Spring, Md. tissue but can be found in skin dander, stored food
494
products, furniture, debris in household carpets, The etiologic agent is Rickettsia akari, which is
and areas that provide a variety of organic materials. primarily found in the house mouse, Mus muscularis,
Mites in the genus Dermatophagoides have been but may be transmitted to humans by accident.
associated with house dust allergy and climate The straw itch mite, Pyemotes tritici, belongs to
allergy.91 The European house dust mite, D pteronyssinus, the family Pyemotidae. It commonly attacks a
was implicated as an allergenic component of house variety of stored-grain insects and is highly toxic
dust to which most asthmatics are sensitive and that to humans. Dermatitis associated with P tritici is
elicits the strongest responses.2,92 The American house known as straw, hay, or grain itch. The infestation
dust mites D farinae, Euroglyphus maynei, and generally occurs after sleeping on straw mattresses,
Glycyphagus destructor have also been reported to working in grain fields during harvesting, or
cause allergies.93 coming in contact with various grains or materials
Adult mites are plump and have well developed infested with the mites.
chelicerae and suckers. Their color varies from white Scabies Mites. The scabies mites belong to the
to light tan. The life cycle takes about a month to family Sarcoptidae. The family includes the genera
complete, and adults may survive up to 2 months Sarcoptes, Notoedres, and Trixacarus, each producing
under optimum conditions. They are most abundant a particular type of dermatosis (Figure 21-28).
in home environments that are warm with high Sarcoptes scabiei causes scabies, also known as 7-year
humidity.94 itch or Norwegian itch. The mite most often is found
Biting Mites. The majority of the human-biting on skin between the fingers, at the bend of knees
mites belong to the families Dennanyssidae, and elbows, on the penis, on the breasts, and on
Macronyssidae, and Sarcoptidae. They are generally the shoulder blades. Rash and itching are not
ectoparasites of poultry, wild birds, and rodents but experienced in newly infested persons for up to a
may also attack humans, causing skin disorders and month after infestation. The rash and itching are
discomfort. The more medically important mites are directly associated with the burrowing of mites into
the scabies mite and the hair follicle mite, which the skin. Human scabies occurs worldwide.
are discussed separately. Follicle Mites. Follicle mites, Demodex folliculorum
The tropical rat mite, Ornithonyssus bacoti, has and Demodex brevis, live in hair follicles and seba-
been associated with debilitation, retarded growth, ceous glands, respectively.95 They are mainly found
and high mortality in colonies of research mice. around the eyelids, nose, and other facial areas.
When the rats die or abandon their nests, the mites Most commonly, the infestation is benign but may
can travel considerable distances and bite humans, result in the loss of eye lashes or in granulomatous
causing a sharp itching pain that may lead to
development of dermatitis in sensitive individuals.
The tropical fowl mite, Ornithonyssus bursa, is
primarily an ectoparasite of poultry (it cannot exist
for more than 10 days apart from its avian host),
but it may bite humans. The bite causes only slight,
temporary irritation.2 Ornithonyssus sylviarum, the
northern fowl mite, is a widespread parasite of
poultry in New Zealand and Australia. The
crawling of these mites is known to cause itching
in personnel working on heavily infested farms. The
chicken mite, Dermanyssus gallinae, commonly
found on domestic fowl, pigeons, English sparrows,
starlings and other birds, is considered one of the
most common species causing dermatitis in hu-
mans, especially those working in poultry houses
and on farms.
The house mouse mite, Liponyssoides sanguineus,
is primarily an ectoparasite of mice but has been Fig. 21-28. A scabies mite.
reported to feed on rats and other rodents and will Photograph: Courtesy of Richard G. Robbins, PhD, De-
readily attack humans. This mite is known to fense Pest Management Information Analysis Center,
transmit the rickettsial pox pathogen to humans. Armed Forces Pest Management Board.
495
Fig. 21-29. The ventral view of the follicle mite Demodex Fig. 21-30. The brown recluse spider (Loxosceles
folliculorum. reclusa)
Source: Uniformed Services University of the Health Sci- Photograph: Courtesy of Ed Rowton, Walter Reed Army
ences, Bethesda, Md. Institute of Research, Silver Spring, Md.
acne. 96 The follicle mite is an elongated species spider) found in most temperate, tropical, and
(Figure 21-29). subtropical regions, can be particularly toxic and
sometimes fatal. Other species (eg, the brown
Spiders and Scorpions recluse spider, Loxosceles reclusa, Figure 21-30) can
cause considerable necrosis at the site of the bite.
There are several thousand species of spiders The bite of almost any large spider, although not
worldwide and the size range is considerable, the necessarily poisonous, may be contaminated with
smallest being a few millimeters in length and the pathogenic bacteria and may result in extensive and
largest having a leg span of up to 20 cm. Spiders dramatic secondary infection.
use their mouthparts to capture their prey and Scorpions are found in many parts of the tropics
sometimes cause paralysis by injecting venom. The and warm temperate regions. They are found in
venom of a few species is particularly toxic to both humid and arid areas. Adults have eight legs,
humans. large pedipalps, and a flexible tail with a needle-
Large, hairy tarantulas found in many tropical like stinger. They typically exhibit cryptic behavior,
and Mediterranean countries are rarely dangerous, sheltering under rocks, logs, and other debris. All
although their large size is commonly frightening. scorpions inject a paralyzing poison into their prey
In contrast, the small widow spiders (ie, Lactrodectus and feed on the fluid content. All these animals should
species, which includes L mactans, the black widow be presumed to be dangerous and are to be avoided.
SUMMARY
Arthropods and the diseases they transmit have are immunologically naïve to the diseases they
had devastating effects on US and foreign military carry. One of the maxims of war is to know your
forces in the past and there is great potential for enemy. This applies to arthropod enemies as well
these effects to be felt in the future. Arthropod- at to human enemies. The more preventive medicine
borne disease and injury can severely affect training professionals know regarding arthropods of
and can render entire units temporarily or per- military medical importance, the more likely they
manently ineffective. Deployed military personnel are to be able to implement countermeasures to
are particularly vulnerable to arthropods and often protect service members.
496
Acknowledgment
The authors wish to thank Major (retired) Louis Rutledge, Dr. (Captain) Pollie Rueda, Dr. Michael
Turell, Dr. Curtis Hayes, Colonel (retired) William Bancroft, Lieutenant Colonel Jeffrey Gambel, and
Colonel (retired) Don Johnson for their insightful review of the chapter. Dr. Edgar Rowton, Dr. Richard
Wilkerson, and Medical Audio-Visual Services personnel from the Letterman Army Institute of
Research and the Walter Reed Army Institute of Research were very helpful with photographs and
illustrations.
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501

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MILITARY PREVENTIVE MEDICINE:

MOBILIZATION AND DEPLOYMENT

RAJ K. GUPTA, PH D; LEON L. ROBERT JR., PHD; AND PHILLIP G. LAWYER, PH D

Scientific name Common name Scientific name Common name

Family Macronyssidae Fowl mites Family Cimicidae Bed bugs

Scientific name Common name Scientific name Common name

Dengue Hemorrhagic Fever

Species Locality Species Locality

Ae lineatopennis collected from larvae in the field Eastern

inside human or animal dwellings.

Diseases Transmitted by Sand Flies

The Leishmaniases. The public health impact of

has been reported in at least two patients infected

exclusively by day. susceptible host during a subsequent blood meal.

Fig. 21-16. (a) a horse fly (Tabanus punctifer) and (b) a

diseases. They have observed that a rise in enteric

Crab lice are only found on humans and die within

Diseases Transmitted by Lice

Three diseases of public health importance that are

22). Hypersensitive persons should carry insect sting Fire Ants

a. Hard Tick b. Soft Tick

Fig. 21-23. The characteristics of female hard and soft ticks.

oval with a sac-like body and vary in length from 2 to

antimicrobial agents. Death is usually associated with

1. Peterson RKD. Insects, disease, and military history. Am Entomologist. 1995;41:147–161.

9. Clements AN. The Physiology of Mosquitoes. Oxford: Pergamon Press; 1963.

10. Gillett JD. Mosquitoes. London: Weidendield and Nicolson; 1971.

39. Outbreak of Rift Valley fever—Yemen, August–Ocotber, 2000. MMWR. 2000;49:1065–1066.

58. Hertig M. Phlebotomus and Carrion’s disease. Am J Trop Med. 1942;22(Suppl):1–80.

69. Reisman RE. Insect stings. N Engl J Med. 1994;331:523–527.

88. Sasa M. Biology of chiggers. Ann Rev Entomol. 1961;6:221–244.

93. Wharton GW. House dust mites. J Med Entomol. 1976;12:577–621.

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