American Society of Mammalogists

Giraffa camelopardalis
Author(s): Anne Innis Dagg
Source: Mammalian Species, No. 5, Giraffa camelopardalis (Jan. 19, 1971), pp. 1-8
Published by: American Society of Mammalogists
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MAMMALIAN SPECIES No. 5, pp. 1-8, 3 figs.

Giraffa camelopardalis.By Anne Innis Dagg

Published 19 January 1971 by The American Society of Mammalogists

Giraffa Brunnich, 1772 pointed out that reticulata de Winton, 1899, was a junior
homonym of reticulata Weinland, 1863, but requested vali-
Giraffa Briinnich, 1772:36. Type species Cervus camelopardalis dation of de Winton's name and suppression of Weinland's
Linnaeus, 1758, by monotypy. Giraffa Brisson, 1762, al- by the International Commission on Zoological Nomen-
though frequently cited, is in a work that is not consistently clature under their plenary powers, because Weinland's
binomial and is therefore unavailable for purposes of no- type locality was within the range of antiquorum rather
menclature unless specifically validated by the International than that of reticulata as these names have been used re-
Commission on Zoological Nomenclature, a step that has cently. The Commission has not yet (May 1970) acted on
not been taken, although some have advocated it. the request. The name australis Rhoads may also be a
Camelopardalis Schreber, 1784:pl. 255. Type species Camelo- synonym.
pardalis giraffa Schreber, 1784, by monotypy. G. c. rothschildi Lydekker, 1903:122. Type locality, as given
Orasius Oken, 1816:744. Type species Cervus camelopardalis originally, Uasin Gishu Plateau east of Lake Baringo,
Linnaeus, 1758, by monotypy. Ruled unavailable, because Kenya, corrected to west of Lake Baringo (Lydekker, 1908)
in a work not consistently binomial, by the International (cottoni Lydekker a synonym).
Commission (Opinion 417, 1956, Bull. Zool. Nomenclature G. c. tippelskirchi Matschie, 1898:78, see above (schillingsi
14:1). Matschie a synonym).
Trachelotherium Gistl, 1848:81. Proposed as a replacement G. c. thornicrofti Lydekker, 1911:484. Type locality Petauke,
name for Camelopardalis Schreber, 1784. Eastern Province, Northern Rhodesia (now Zambia).
G. c. angolensis Lydekker, 1903:121. Type locality by the
CONTEXT AND CONTENT. Order Artiodactyla, Sub- Cunene River 240 kms. southwest of Humbe, Angola (in-
order Ruminantia, Infraorder Pecora, Family Giraffidae, Sub- fumata Noack a synonym according to Ansell).
family Giraffinae. There are only two living giraffids, the G. c. giraffa Schreber, 1784:pl. 255, see above (capensis Les-
okapi, Okapia johnstoni, and the giraffe, Giraffa camelopardalis.
son, australis Swainson, maculata Weinland, and wardi
Lydekker, are synonyms).
Giraffa camelopardalis Linnaeus, 1758
Cervus camelopardalis Linnaeus, 1758:66. Type locality "Sennar
DIAGNOSIS AND GENERAL CHARACTERS. The
and Aethiopia." genus is monospecific. Height is 5 to 6 m., females smaller
than males; legs and neck each over 1.5 m. long. Slope of
Camelopardalis Giraffa Schreber, 1784:pl. 255 only, rather than back makes forelegs appear longer than the hind legs, which
Boddaert, 1785:133 as usually cited. Type locality of they marginally are. Two unbranched permanent horns, in
Schreber not given, Boddaert gave "Cape of Good Hope," both sexes, are fused to the skull above the fronto-parietal
but no giraffe have been found that far south; here re- suture and are covered with skin; a median "horn" anterior
stricted to Warmbad (as mapped in fig. 3), where Brink to this is more developed in males than in females; the facial
(1761) encountered and described giraffe just north of region of the skull of old males is also covered with other
the Orange River. irregular bony growths. Skull is up to 73 cm. long; molars
Cameleopardalis antiquorum Jardine, 1835. Type locality "Sen- brachydont, upper molars without inner accessory columns;
nar and Darfour," restricted to Baggar el Homer, Kordofan, canine teeth bilobed or trilobed. Hide has a buff color base
about 10?N and 28?E by Harper (1940). on which are spread darker brown spots varying in shape from
Camelopardalis aethiopica Ogilby, 1837:134. Type locality un- irregular jagged blotches to smooth polygons; spots large on
designated, by inference Ethiopia. body but small on head and upper legs. Tail is long and has
long, black, coarse terminal hairs; neck maned with short,
Camelopardalis capensis Lesson, 1842:168. Type locality Cape brown, stiff hairs.
of Good Hope.
Giraffa senaariensis Trouessart, 1898:902. Type locality inter- DISTRIBUTION. Giraffe are widespread in Africa
preted to be south of Sennaar, Anglo-Egyptian Sudan by where there are scattered trees or bushes. Their range includes
Allen (1939:468). semi-arid regions but not deserts, rain forests, or mountain
Giraffa tippelskirchi Matschie, 1898:78. Type locality Lake ranges. Within historic times their distribution has been much
Eyassi, southeast of Victoria Nyanza, Tanganyika Territory reduced, because of increasing aridity and increased human
(now Tanzania). population pressures, and their numbers everywhere have de-
clined because of hunting, farming and periodic epidemics of
Giraffa schillingsi Matschie, 1898:79. Type locality Taveta,
Kenya. rinderpest. None now survive in North Africa but some may
have remained in Morocco as late as A.D. 600 (Schomber and
Giraffa infumata Noack, 1908:356. Type locality Barotse, mid- Kock, 1961:137). Recently, especially with the advent of
dle Zambesi region, Northern Rhodesia (now Zambia).
drugging techniques, giraffe have been transplanted to areas
Giraffa hagenbecki Knottnerus-Meyer, 1910:800. Type locality of Africa (not shown in fig. 3) where they were not previously
Gallaland, southern Abyssinia. found, e.g. Orange Free State (Griesel, 1961), Natal, Swaziland
CONTEXT AND CONTENT. Contextnoted in generic
summary above. Nine subspecies are recognized (Ansell, 1968)
as follows:
G. c. camelopardalis (Linnaeus, 1758:66), see above (biturigum
Duvernoy, aethiopica Ogilby, typica Bryden, and perhaps
congoensis Lydekker, are synonyms).
G. c. antiquorum (Jardine), 1835:187, see above (senaariensis
Trouessart a synonym).
G. c. peralta Thomas, 1898:40. Type locality near Lokoja, at A B C D
the junction of the Niger and Benue rivers in Nigeria,
probably north of the confluence (Happold, 1969). FIGURE1. Patterns of trunk spots (all at about the same scale)
G. c. reticulata de Winton, 1899:212. Type locality the Loroghi of giraffe belonging to three adjacent subspecies (from Dagg,
Mts. in Kenya (hagenbecki Knottnerus-Meyer and nigres- 1968): A, G. c. rothschildi; B, G. c. reticulata; C and D,
cens Lydekker are synonyms). Mertens (1968 a and b) G. c. tippelskirchi.

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2

FIGURE 3. Recent distribution of the giraffe in subsaharan

Africa, within historic times (outlined with dotted lines, after
Krumbiegel, 1939), at present (outlined with solid lines), and
at three fossil sites (marked X). Type localities of the nine
recognized subspecies are shown by triangles: A, G. c. camelo-
pardalis; B, G. c. antiquorum; C, G. c. peralta; D, G. c.
reticulata; E, G. c. rothschildi; F, G. c. tippelskirchi; G, G. c.
thornicrofti; H, G. c. angolensis; and I, G. c. giraffa.

et al. (1962). The teeth and bones of the giraffe are especially
large but not otherwise greatly different from those of other
pecoran species. The dental formula is 0/3, 0/1, 3/3, 3/3 = 32.
The molars are brachydont and rugose. The row of front teeth
is wide, in part because of the broad lobed canines, which
makes the combing action used while browsing on leaves par-
ticularly effective. The skull has extensive air spaces, but,
even with these spaces, its weight may reach 13 kg. in old
males (Dagg, 1965). Dorsal neck muscles and ligaments attach
to two enlarged bony areas at the back of the adult skull which
FIGURE 2. Top: Skull of adult male giraffe. Middle: Upper are sometimes referred to as occipital, posterior, or mizzen
view of mandible (note the lobed canine teeth). Bottom: horns (Thomas, 1901). The occipital condyles allow the head
Section of skull of adult male showing extensive pneumatic to be raised through an angle of more than 90?; this allows
sinuses. All photos by J. B. Foster. free movement for browsing or fighting. The various sutures in
the skull close at different times so that a skull of an animal
up to at least six years of age can be aged fairly accurately
(Kirk, 1966) and Saanane Island in Lake Victoria (Achard (Singer and Bone, 1960). They also found that giraffe could
and McCulloch, 1967). be aged by sequences of eruption and wear of the teeth. The
giraffe has only 7 cervical vertebrae, each very elongate, but
FOSSIL RECORD. The giraffids are believed to have the first of the 14 (rarely 13) thoracic vertebrae is modified
evolved in the early or middle Miocene in central Asia to resemble a cervical vertebra too (Lankester, 1908). The
(Churcher, 1970). Specimens thought to belong to Giraffa dorsal spines of the anterior thoracics are long, forming the
camelopardalis are known only from Pleistocene deposits of prominent hump on the upper back. They serve for the attach-
Israel and Africa itself. The Israeli records are for the lower ments of the large muscles and the nuchal ligament which sup-
Pleistocene of the central Jordan Valley (Haas, 1966) and for port the neck and head. The thoracics are followed by 5 lum-
the Villafranchian at Bethlehem (Hooijer, 1958). The African bar, 4 to 5 fused sacral and 16 to 20 caudal vertebrae. The
records are for the Villafranchian from Tchad (Abadie et al., pelvis is shorter than in most ruminants, and the upper ends
1959), for the middle Pleistocene of Olduvai, Tanzania (Hop- of the ilia are more expanded (Flower, 1870). The legs are
wood, 1936) and for the lower Pleistocene from Omo, near highly specialized; the first, second and fifth metapodials are
Lake Rudolph (Arambourg, 1947). However, Leakey (1965) usually absent. The joints and ligaments of the hind leg at
noted that the lower and middle Pleistocene specimens of least are very similar to those of antelope (Heinze, 1965). The
Giraffa from Africa may not be of the species G. camelopardalis. hooves are low posteriorly so that the fetlock nearly touches
the ground; this may increase the foot's ability to support a
FORM. Lochte (1952) described five different types of large weight. The footprint is about 31 by 23 cm. in an adult
hair found on giraffe. Broman (1938a) explored one of these male and slightly less in a female. There are no interdigital
types, from the mane, in detail. The mane, which is about 12 glands or lateral hooves. The musculature of the giraffe has
cm. long in the adult, can be identified microscopically in a been broadly described by Owen (1841), Joly and Lavocat
fetus only 28 cm. long. The arrangement of hairs produces (1843) and Murie (1872). More recently, smaller muscle
areas of hair whorls, feathering, and crests which seem to be groups have been studied in depth: the neck and shoulder
correlated with positions or areas in which movement occurs muscles by Chaine (1902), Rothschild and Neuville (1911),
in any particular region of the body (Kidd, 1900, 1903; Lan- Zuckerman and Kiss (1932), Willemse (1958) and Angermeyer
kester, 1907; Rothschild and Neuville, 1911), although there is (1966); the foreleg muscles by Bego (1960); and the hind
also much individual variation. The female has 4 mammae, leg muscles by Heinze (1964, 1965). The circulatory system is
Shortridge's (1934) widely quoted count of 2 mammae being specialized to cope with the high blood pressure necessary in
an error. The milk of giraffe has been analyzed by Stephan the giraffe to pump blood to the brain. The arteries of the
(1925), Greed (1960), Ben Shaul (1962) and Aschaffenburg neck have thick walls composed largely of elastic tissue al-

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 3

though in the carotids the muscle fibres increase in prominence was 200 mm. Hg. when she was standing but only 175 mm. Hg.
towards the head (Franklin and Haynes, 1927). The blood when she lowered her head (Goetz and Budtz-Olsen, 1955).
pressure at the brain is regulated by a rete mirabile into which The mechanism that controlled the blood pressure so effec-
the external carotids divide (Lawrence and Rewell, 1948). The tively was not related to a high viscosity or high protein con-
blood pressure in the carotid may reach 353 (systolic) to 303 tent of the blood, as these values in the female giraffe were
(diastolic) mm. Hg. in an experimental animal (Goetz and not unlike those in man. Recently information has been ob-
Keen, 1957; Goetz et al., 1960). Some large veins are exten- tained via radiotelemetry in free-ranging giraffe (Van Citters
sively valved, to help counteract the effects of gravity (Amoroso et al., 1966). An animal while resting had a low blood pres-
et al., 1947). The veins of the legs are very thick and mus- sure reading from the carotid artery of 150 (systolic) to 105
cular and have tiny lumens. In general, the vessels from vari- (diastolic) mm. Hg. When galloping, these values increased
ous parts of the body have wide variations in structure cor- to 230/125 mm. Hg. The pulse rates ranged from 60 to 170
responding to the prevailing hydrostatic pressure (Goetz and beats per minute. The blood flow in the carotid artery ranged
Budtz-Olsen, 1955; Goetz and Keen, 1957; Goetz et al., 1960). between 50 cm.`/sec. in a prone giraffe and 35 cm.3/sec. in a
The heart of a wild female weighed 12 kg. (Goetz and Budtz- standing one. The peak systolic blood velocity was 60 cm./sec.;
Olsen, 1955), far more than those of captive giraffe. The wall during diastole the velocity fell to nearly 40 cm./sec. (Van
of her left ventricle was 7.5 cm. thick. The atrioventricular Citters et al., 1968). Radiotelemetric thermometry has been
node is large in the giraffe and the Purkinje cells and fibres used to determine the deep body temperature of a giraffe
are particularly well developed (Castigli and Sacchi, 1941). A (Bligh and Harthoorn, 1965). At low ambient temperatures
heart bone up to 2.5 cm. long is present in adults (Crisp, the deep body temperature in the lower neck of a 400 kg.
1864b). There are twice as many blood cells per cubic mm. female was remarkably thermostable at between 37.75?C and
in the giraffe as in man, but otherwise the blood is not par- 39.1?C. This thermostability may be relaxed at high ambient
ticularly distinctive (Goetz and Budtz-Olsen, 1955). Nor did temperatures, as in eland Taurotragus oryx (Taylor and Lyman,
these men find the fetal circulation especially noteworthy. The 1967), and this would help the giraffe keep cool while conserv-
spleen is not unlike that of other ruminants (Crisp, 1853; L6nn- ing water. Backhaus (1959) studied color vision in a captive
berg, 1900; Retterer and Neuville, 1916; and Derscheid and giraffe. He offered food in variously colored containers and
Neuville, 1925). The brain of the giraffe is relatively small judged that the animal could distinguish between red, orange,
compared to the size of the animal, the weight ratio is about yellow, yellow-green and violet.
1:800, which is similar to that of the horse. The anatomy of
the brain is detailed by Black (1915), Friant (1952, 1954) ONTOGENY AND REPRODUCTION. Following cop-
and Kato (1963). The total length of the nerves is immense ulation the fertilized egg implants without delay. The gestation
in the giraffe, but their arrangement is much as in other un- period averages 15 months (the mean of 35 pregnancies of
gulates. Because the trachea of the giraffe is so long, the captive females, Backhaus, 1961), but varies considerably,
giraffe has a large respiratory dead space where no exchange from 394 days for a small animal that died shortly after birth
of gases can take place. This is partly balanced by a large (Lang, 1955) to 488 days (Robinson et al., 1965). In two male
total lung capacity. Patterson et al. (1957) calculated that the fetuses, 28 cm. in body length and about 3 months of age, no
resting tidal volume of the giraffe is about 4 liters per breath hair was visible macroscopically (Broman, 1938b), the main
and that these animals, like men, use about 'io to 1/12 of their horns were indicated by two mounds above the skull 8 mm. in
total lung capacity under basal conditions. A giraffe breathes diameter and 5 mm. high, and the testes had already descended
only about 8 to 10 times a minute (Robin et al., 1960). The into the scrotal sacs. In a female fetus, 60 cm. long and about
vocal cords are of the simple form present in most ruminants 4 months old, also described by Broman, hairs were evident
(Burne, 1917). They enable giraffe to make a variety of on various parts of the head, especially on both lips and on
sounds although they seldom do so. The digestive system is the upper eyelid, and on the tip of the tail. The main horn
complex in the giraffe, which eats only leaves and twigs. The anlagen were 2 cm. in diameter and 5 mm. high while the
prehensile lips are extensively haired as a protection against anterior horn was represented as a minute bump. The incisors
thorns. The dark-coloured tongue is up to 54 cm. long and canines were visible through the gums. The 4 teats were
(Schneider, 1951) which increases the reach of the animal papillae 3 to 4 mm. high and 1.5 mm. thick. Beddard (1906)
appreciably. Both it and the inside of the mouth are exten- noted in an 8 month fully haired fetus that the hairs were all
sively covered with papillae (Sonntag, 1922). The salivary whitish so that there were no spots. The horn cartilages could
glands are large as in all ruminants but the palatine tonsils be felt through the skin above the skull. The hooves were soft
are small (Burne, 1917; Hett, 1928). The long muscular and quite pointed. Many parturitions have been observed in
oesophagus leads to the rumen, the first of the four stomach captivity. The forelegs and head of the young appear first,
compartments which are all modified for the diet of a "selec- after the rupture of the fetal membranes. The young is slowly
tive feeder" (Hofmann, 1968). In the rumen the food is pushed outward as the mother strains while standing with her
thoroughly moistened by gastric juices and partly fermented hind legs spread apart. The umbilical cord breaks about 30
before being shaped into boluses that are rechewed as the cm. from the abdomen as the young falls to the ground (Gijzen,
animal ruminates. There are no special stomach cells for water 1958) and at this time the animal first begins to breathe (Har-
retention (Fox, 1938). The intestines of an adult are about rison Matthews in Slijper, 1958). The process lasts from 20
80 m. long. Beyond the ileo-caecal junction there are various minutes to several hours (Gijzen, 1958). Only one young is
crypts that increase the surface where mucous is secreted (Neu- born at a time. The only sure record for twinning is from the
ville, 1922). The caecum itself is large. The feces are elimi- San Francisco Zoo where two stillborn twins, together weighing
nated from the anus as acorn-like pellets about 4 cm. long. only 41 kg., were born in 1943 (Reuther, pers. comm.). Back-
The liver shows few signs of lobulation (Neuville, 1914). The haus (1961) recorded the dates of calving of 53 females in
hepatic ducts unite to form a long common bile duct which captivity and found that these occurred throughout the year.
opens in the duodenum about 30 cm. beyond the pylorus (Cave, Giraffe also breed throughout the year in the wild, although
1950). A gall bladder may or may not be present; Cave (1950) different seasonal peaks in different areas and in different
found only 2 in 19 dissections of giraffe but Kobara and years for the same area have been noted-in the Kruger Na-
Kamiya (1965) found one in each of 2 female adults they tional Park from October to January (Stevenson-Hamilton,
examined. The kidneys were described by Wrobel (1965). 1947), from February to April (Brynard and Pienaar, 1960),
The reproductive systems resemble those of other ungulates. for September and October (Pienaar, 1963) and for February
The placenta is typically polycotyledonary, with large, many- to March and August to October (Fairall, 1968). Ansell (1960)
branched villi (Mossman, 1937; Ludwig, 1962). Wilkinson and found no definite calving seasons for giraffe in Zambia, al-
de Fremery (1940) detected gonadotropic hormones in the though he states that October and March have been mentioned
urine of a pregnant female. The giraffe lacks preorbital, in- by other authors. In Rhodesia, Dasmann and Mossman (1962)
guinal and interdigital glands (Pocock, 1910). The pituitary reported that most births occurred in May during the dry
is quite large, especially the adenohypophysis (Hanstrim, 1953; season. In the Nairobi National Park and Serengeti Park most
Kladetzky, 1954). Frank (1960) discussed the parathyroid and young were also born during the dry months (J. B. Foster and
thyroid glands of the giraffe. G. Schaller, pers. comm.), although parturitions occurred near
Nairobi in every month. In captivity 78% of 27 calvings took
FUNCTION. Few physiological experiments have been place in the daytime, but these times may have been influenced
carried out on giraffe except on the circulatory system. To by the presence of human observers (Backhaus, 1961). The
study this, wild giraffe have been drugged so that apparatus neonate is about 2 m. tall at birth and weighs about 54.5 kg.
could be fastened to individuals to determine their blood pres- (mean of 16 captive neonates). It has a relatively shorter neck
sures. The maximum blood pressure in the carotid artery at than the adult which it holds in a more erect position. Its
the base of the brain in a female surrounded by scaffolding spots tend to be paler than those of older animals. The hair

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4

is short and slightly woolly. The horn cartilages lie flat at main near their mothers. In 1967 the resident giraffe popula-
first under the skin but become upright between two hours tion of the Nairobi National Park was estimated at 31% adult
(Benchley, 1946) and four weeks (Sigel, 1886) after birth and female, 25% adult male, 12% in their fourth and fifth years,
fuse to the skull during ossification. Their position is indicated 8% in their third year, 10% in their second year and 14% in
by two prominent tufts of black hair. In the Nairobi National their first year. These percentages have been quite stable for
Park the male to female ratio of newborn calves was 31:30 seven years. The average life span of a giraffe was about six
(Foster, pers. comm.) but Bourliire (1961) found that of 117 years, but some lived more than 25 years. In Wankie and
captive neonates 61.5% were males. Calves can stand and suck Garamba National Parks, Dasmann (pers. comm.) and Back-
at 10 minutes after birth, and walk and run soon afterwards haus (1961) respectively estimated about 30% of the giraffe
(Gijzen, 1958). They can look after themselves within a few were immature and 20% of these were calves. In South Africa,
months. They supplement their milk diet with leaves after a Innis found only 12% of the giraffe were immature and half
few weeks although they may suck for over a year. In captivity of these were calves. The proportion of females was low in
even adults will suck, although the mother will not allow them this population. Giraffe are usually found on lightly forested
to do so unless her own calf is already sucking (Dagg, 1970). areas, always near trees or bushes. They are often sedentary
The young grow at least 1 m. during their first year and ap- animals, seldom moving great distances except when forced to
proach their full size at 4, although they may continue to grow by fire or drought (Innis, 1958). In Nairobi National Park,
slightly until they are 7 or 8 years old (Owen, 1841; Paulus, Foster (pers. comm.) found that the bulls were more restless
1943; Reventlow, 1949). At this time a wild female stands than the cows, often walking from herd to herd presumably
about 4.3 m. and a wild male about 5.3 m. Captive giraffe to find a female in heat. One male was spotted 50 km. from
seldom exceed 5.0 m. with a shoulder height in males of about the Park. The giraffe in the Park are not all permanently
3.3 m. The average adult weight is considered to be 800 kg. established, because although there are about 80 giraffe there
(Stewart and Zaphiro, 1963) although a large male may weigh at one time, these may be any of 241 individuals. Giraffe do
1220 kg. or more (Crile, 1941). The females become sexually not show territorial behavior. In the Nairobi National Park,
mature at 31/, years and the males at about 41/2 years of age Foster (pers. comm.) calculated the average home ranges for
(Gijzen, 1958). The female thereafter comes into heat about the 10 most commonly seen males and females at 62 and 85 sq.
every 14 days (Lang, 1955) and can conceive while she is km. respectively.
lactating. She can breed until she is at least 20 years old Various diseases have been reported in wild giraffe, the
(Ried, 1958). A captive male that had sired many giraffe most virulent of which is rinderpest. This disease first swept
was no longer breeding at 24 years of age (Dagg, 1970). In through Africa in the 1890's, killing hundreds of giraffe and
captivity the oldest giraffe attained 28 years of age (King, other ruminants (Grzimek, 1956a). There have been scattered
1947) but giraffe in the wild not uncommonly live at least up outbreaks in Africa since then, including a severe epidemic
to 26 years (Foster, pers. comm.). The mortality of adult in northern Kenya in 1960 where about 40% of the giraffe died
giraffe is not great in the wild, but in their first year of life (Simon, 1962). Earlier in this same area an epidemic of gas-
as many as 68% of those born may die (Foster, pers. comm.). troenteritis killed some giraffe (Percival, 1924) and others died
during a brief outbreak of anthrax in southern Africa (Pienaar,
ECOLOGY. Lions are by far the major predators of 1961). This disease was suppressed by disinfecting the water
giraffe (Wright, 1960; Brynard and Pienaar, 1960; Pienaar, sources responsible for the spread of the infection (Payne,
1963; Kruuk and Turner, 1967) although on rare occasions a 1961). Giraffe eat from a variety of species of plants, many of
lion itself may be killed by a giraffe (R. Douglas of Tan- them acacias. Lists of species eaten are given by Innis (1958),
ganyika Safaris, Arusha, pers. comm.; Campbell, 1951). Cheetah Van der Schijff (1959), Brynard and Pienaar (1960), and
and crocodiles kill giraffe very rarely-0.28% each of 675 Foster (1966). Giraffe eat the leaves, but sometimes also the
giraffe kills in the Kruger National Park over a 25 year period pods, fruit and twigs of these trees and bushes. They are not
(Pienaar, 1969). The other kills were all by lion. Various deterred by the presence of long thorns and spines on their food
tickbirds live commensally with giraffe-the buffalo weaver plants but they may be by ants (Brown, 1960). Throughout
Texor niger (FitzSimons, 1920) and the red-billed and yellow- most of Africa at present, the giraffe is a protected species al-
billed ox-peckers Buphagus erythrorhynchus and B. africanus though a few may be shot on special licenses for museums or
(Attwell, 1966). These birds call out at the approach of a sport. Undoubtedly a number are also poached each year.
man and probably of other predators too (Attwell, 1966). They Giraffe are found in zoos throughout the world, where they
also serve a giraffe by searching through its hair for ticks, breed readily. Some captive mothers refuse to suckle their
which they eat. They sometimes peck at wounds, so aggravat- young, but these calves can be raised using bottled milk (Zell-
ing them, but perhaps they compensate for this by devouring mer, 1960; Savoy, 1966). The Africans have always used parts
maggots in the area. of giraffe-the flesh for meat; the skin for shields, sandals and
Ticks usually infest giraffe, particularly on the relatively drums; the tendons for stringed musical instruments and
thin-skinned area of the genitalia. The common species include thread; and the tail hairs for bracelets, flyswitches and thread.
the bont tick Anmblyommahebraeum, blue tick Boophilus de- With the coming of the white man many more giraffe were
coloratus, brown tick Rhipicephalus appendiculatus (Innis, killed and the species eliminated in areas where it conflicted
1958), Rhipicephalus camelopardalis (Walker and Wiley, 1959) with man. Now it is fully protected in many national parks.
and Hyalomma aegyptium (Lonnberg, 1912). On giraffe im- Giraffe were formerly captured in stockades (Ogrizek, 1954;
ported into New York the following ticks were found: Am- Stanton, 1955) or by lassoes (Webb, 1954) but now they are
blyomma gemma, Boophilus decoloratus, Hyalomma albiparma- usually drugged by darts (Pienaar et al., 1966). Drugs and
turn and Rhipicephalus pulchellus (Becklund, 1968). Internal tranquillizers were used in the circulation experiments discussed
parasites are more numerous. Species that have been identified earlier. They have also been used to allow Foster (pers. comm.)
are Cooperia pectinata, C. punctata, Haemonchus contortus, H. to put a radio transmitter collar onto a giraffe to study home
mitchelli, Trichuris globulosa (Ortlepp, S. Afr. govn't., pers. range. In most of his work, Foster has identified individuals
comm.), Trichuris giraffae, Parabronema skrjabini, Monodon- by photographing the left side of each animal's neck. The ar-
tella giraffae (Leiper, 1935), Trichocephalus gracilis (Cobbold, rangements of the spots are unique in each individual, so that
1860), Uncinaria smithi (Weidman, 1918), Haemonchus sp., it can always be identified once photographed.
Trichostrongylus sp. (Sloan, 1965), Fasciola gigantica (Cob-
bold, 1855), Taenia saginata = Cysticercus bovis of Schwartz, BEHAVIOR. Dagg (1970) has studied tactile encounters
1928, Moniezia nullicollis (Yamaguti, 1959), Echinococcus sp. between individuals in a herd of 18 captive giraffe. The two
(Crisp, 1864a) and Rhinoestrus sp. (Laurence, 1961). Dinnik sexually-active males were the most active in initiating such
and Sachs (1968) found protostrongylid larvae in the droppings encounters, and the four calves the least active. Nosing and
of wild giraffe in Tanzania. licking were the most common types (53% together of the 1590
Giraffe have little competition for food, since the tallest encounters), followed by rubbing and hitting (18% and 14%).
bovid cannot browse above a height of 2.5 m. Even when the Sniffing genitalia and "urine testing" were infrequent and usu-
giraffe browses on low bushes it often chooses one with many ally done by a bull to a cow. Necking matches were rare and
thorns that other ruminants avoid. By far the best data for always involved a male and a female although in the wild they
population structure and dynamics of giraffe have been col- are restricted to males (Coe, 1967). These types of behavior
lected by Foster (1966 and pers. comm.) from the approxi- are apparently not seasonal, either in captivity or in the wild.
mately 80 giraffe of the Nairobi National Park. Apparently Mating was observed twice and was similar to mating in the
the population increases by nearly 25% each year but of these wild (Innis, 1958), with the male following a receptive female
young about 68% die in their first year. The young often and mounting her when she stood still a number of times over
wander away from their mothers at an early age but these a several hour period. The giraffe were all tolerant of each
young do not have a greater mortality than the young that re- other generally, the dominant male sometimes allowing the

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 5

second male to make sexual advances to a female in heat. cluded that the number, area and shape of the spots on an
During the day most giraffe fed in early morning and in the animal are inherited, but not in the simple manner suggested
late afternoon, although some at least also browsed after dark by Spinage (1968). She also concluded that horn development
(Innis, 1958). In the wild they spent more time hunting for is not under close genetic control, despite the emphasis on the
food during the dry season than during the growing season, horns in many early taxonomic studies.
and less time resting and cudding (Innis, 1958). Giraffe com-
municate indirectly with other giraffe and other species by REMARKS. Not all of the works cited were seen by the
either standing motionless and staring at possible danger or author.
stampeding from it. Either reaction sooner or later attracts the
attention of other animals. Giraffe can communicate by vocal LITERATURE CITED
noises, but they do so rarely. Occasionally a giraffe will grunt Abadie, J., J. Barbeau, and Y. Coppens. 1959. Une faune
when alarmed, annoyed, or hungry and a female and her calf de vertebres villafranchiens au Tchad. C. R. Acad. Sci.
may call to each other with a moo or low. Giraffe do not move (Paris) 248:3328-3330.
far, generally. In South Africa, browsing giraffe averaged 0.21
km./hr. (Innis, 1958). Giraffe have only two gaits: walking Achard, P. L., and B. McCulloch. 1967. Creation of a zoo
and galloping at speeds up to 56 km./hr. (Arbuthnot, 1954). in Tanzania Saanane Island Game Reserve. Int. Zoo
These have been analysed by Dagg and De Vos (1968a and b) Yearbook 7:235-240.
and Bourdelle (1934) and compared with the gaits of other Allen, G. M. 1939. A checklist of African mammals. Bull.
Mus. Comp. Zool. 83:1-763.
pecoran species. The walk of the giraffe is notable because
an individual swings the two legs on one side of its body for- Amoroso, E. C., O. G. Edholm, and R. E. Rewell. 1947. Ve-
ward almost in unison. The long legs, relatively short trunk nous valves in the giraffe, okapi, camel and ostrich. Proc.
and large inertia of the body influenced the evolution of this Zool. Soc. London 117:435-440.
gait. The movement of the long, heavy neck of the giraffe is Angermeyer, M. 1966. Zur kenntnis der Halsmuskeln und
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gallop and in the jumping of fences (Dagg, 1962 b and d). 177:188-200.
The neck is also used to express various emotions in the giraffe Anon. 1910. Hagenbeck's giraffe. Field 116:353.
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with its nose pointing upward. If it is alert or nervous, it holds liminary identification for African mammals, Dr. J. A. J.
its head more nearly perpendicular than usual, with its mouth Meester [ed.]. Smithsonian Institution, Washington, D.C.
pointing to the ground and its nostrils open wide. If it is Arambourg, C. 1947. Contributions a l'etude geologique et
preparing to flee it turns its body at an angle to the disturb- paleontologique du bassin du Lac Rodolphe et de la basse
ance. When it shakes its head, it holds its neck about 45? vallee de l'Omo. 2nd. part, Paleontologie, Mission Sci.
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