The giraffe's distinguishing characteristics are its extremely long neck and legs, horn-likeossicones, and spotted coat patterns. Its scattered range extends fromChad in the north toSouth Africa in the south and fromNiger in the west toSomalia in the east. Giraffes usually inhabitsavannahs andwoodlands. Their food source is leaves, fruits, and flowers of woody plants, primarilyacacia species, which they browse at heights most other ground-basedherbivores cannot reach.Lions,leopards,spotted hyenas, andAfrican wild dogs mayprey upon giraffes. Giraffes live in herds of related females and their offspring or bachelor herds of unrelated adult males but aregregarious and may gather in large groups. Males establish social hierarchies through "necking", combat bouts where the neck is used as a weapon.Dominant males gain mating access to females, which bear sole responsibility for rearing the young.
The giraffe has intrigued various ancient and modern cultures for its peculiar appearance and has often been featured in paintings, books, and cartoons. It is classified by theInternational Union for Conservation of Nature (IUCN) asvulnerable to extinction. It has beenextirpated from many parts of its former range. Giraffes are still found in manynational parks andgame reserves, but estimates as of 2016 indicate there are approximately 97,500 members ofGiraffa in the wild. More than 1,600 were kept in zoos in 2010.
"Camelopard" (/kəˈmɛləˌpɑːrd/) is anarchaic English name for the giraffe; it derives from theAncient Greekκαμηλοπάρδαλις (kamēlopárdalis), fromκάμηλος (kámēlos), "camel", andπάρδαλις (párdalis), "leopard", referring to its camel-like shape and leopard-like colouration.[4][5]
Taxonomy
Evolution
The giraffe is one of only two living genera of the family Giraffidae in the orderArtiodactyla, the other being theokapi.[6] They areruminants of the cladePecora, along withAntilocapridae (pronghorns),Cervidae (deer),Bovidae (cattle, antelope, goats and sheep) andMoschidae (musk deer). A 2019genome study (cladogram below) finds that Giraffidae are asister taxon to Antilocapridae, with an estimatedsplit of over 20 million years ago.[7]
The family Giraffidae was once much more extensive, with over 10 fossilgenera described.[6] The elongation of the neck appears to have started early in the giraffelineage. Comparisons between giraffes and their ancient relatives suggest vertebrae close to the skull lengthened earlier, followed by lengthening of vertebrae further down.[8] One early giraffid ancestor wasCanthumeryx, which has been dated variously to have lived25 to 20 million years ago, 17–15 mya or 18–14.3 mya and whose deposits have been found in Libya. This animal resembled anantelope and had a medium-sized, lightly built body.Giraffokeryx appeared 15–12 mya on theIndian subcontinent and resembled an okapi or a small giraffe, and had a longer neck and similarossicones.[6]Giraffokeryx may have shared a clade with more massively built giraffids likeSivatherium andBramatherium.[8]
The extinct giraffidSamotherium (middle) in comparison with theokapi (below) and giraffe. The anatomy ofSamotherium appears to have shown a transition to a giraffe-like neck.[9]
Giraffids likePalaeotragus,Shansitherium andSamotherium appeared 14 mya and lived throughout Africa and Eurasia. These animals had broader skulls with reduced frontal cavities.[6][8]Paleotragus resembled the okapi and may have been its ancestor.[6] Others find that the okapi lineage diverged earlier, beforeGiraffokeryx.[8]Samotherium was a particularly importanttransitional fossil in the giraffe lineage, as the length and structure of itscervical vertebrae were between those of a modern giraffe and an okapi, and its neck posture was likely similar to the former's.[9]Bohlinia, which first appeared in southeastern Europe and lived 9–7 mya, was likely a direct ancestor of the giraffe.Bohlinia closely resembled modern giraffes, having a long neck and legs and similar ossicones and dentition.[6]
Bohlinia colonised China and northern India and produced theGiraffa, which, around7 million years ago, reached Africa.Climate changes led to the extinction of the Asian giraffes, while the African giraffes survived and radiated into new species. Living giraffes appear to have arisen around1 million years ago in eastern Africa during thePleistocene.[6] Some biologists suggest the modern giraffes descended fromG. jumae;[10] others findG. gracilis a more likely candidate.G. jumae was larger and more robust, whileG. gracilis was smaller and more slender.[6]
The changes from extensive forests to more openhabitats, which began 8 mya, are believed to be the main driver for the evolution of giraffes.[6] During this time, tropical plants disappeared and were replaced by aridC4 plants, and a drysavannah emerged across eastern and northern Africa and western India.[11][12] Some researchers have hypothesised that this new habitat, coupled with a different diet, including acacia species, may have exposed giraffe ancestors to toxins that caused higher mutation rates and a higher rate of evolution.[13] The coat patterns of modern giraffes may also have coincided with these habitat changes. Asian giraffes are hypothesised to have had more okapi-like colourations.[6]
The giraffe genome is around 2.9 billionbase pairs in length, compared to the 3.3 billion base pairs of the okapi. Of the proteins in giraffe and okapi genes, 19.4% are identical. Thedivergence of giraffe and okapi lineages dates to around 11.5 mya. A small group ofregulatory genes in the giraffe appears responsible for the animal's height and associated circulatory adaptations.[14][15]
Species and subspecies
Map showing "Approximate geographic ranges, fur patterns, andphylogenetic relationships between some giraffe subspecies based onmitochondrial DNA sequences. Coloured dots on the map represent sampling localities. The phylogenetic tree is amaximum-likelihoodphylogram based on samples from 266 giraffes. Asterisks along branches correspond tonode values of more than 90%bootstrap support. Stars at branch tips identifyparaphyletichaplotypes found in Maasai and reticulated giraffes".[16]
Carl Linnaeus originally classified living giraffes as one species in 1758. He gave it thebinomial nameCervus camelopardalis.Mathurin Jacques Brisson coined the generic nameGiraffa in 1762.[17] During the 1900s, various taxonomies with two or three species were proposed.[18] A 2007 study on the genetics of giraffes using mitochondrial DNA suggested at least six lineages could be recognised as species.[16] A 2011 study using detailed analyses of the morphology of giraffes, and application of thephylogenetic species concept, described eight species of living giraffes.[19] A 2016 study also concluded that living giraffes consist of multiple species. The researchers suggested the existence of four species, which have not exchanged genetic information between each other for one to two million years.[20]
A 2020 study showed that, depending on the method chosen, different taxonomic hypotheses recognizing from two to six species can be considered for the genusGiraffa. That study also found that multi-species coalescent methods can lead to taxonomic over-splitting, as those methods delimit geographic structures rather than species. The three-species hypothesis, which recognisesG. camelopardalis,G. giraffa, andG. tippelskirchi, is highly supported byphylogenetic analyses and also corroborated by most population genetic and multi-species coalescent analyses.[21] A 2021whole genome sequencing study suggests the existence of four distinct species and seven subspecies,[22] which was supported by a 2024 study of cranial morphology.[23] A 2024 study found a higher amount of ancientgene flow than expected between populations.[24]
Thecladogram below shows the phylogenetic relationship between the four species and seven subspecies based on a 2021 genome analysis.[22] The eight lineages correspond to eight traditional subspecies in the one-species hypothesis. The Rothschild giraffe is subsumed intoG. camelopardalis camelopardalis.
Giraffa
Giraffa camelopardalis
G. camelopardalis antiquorum (Kordofan giraffe)
G. c. camelopardalis (Nubian giraffe)
G. c. peralta (West African giraffe)
(northern giraffe)
G. reticulata
(no subspecies)
(reticulated giraffe)
G. tippelskirchi
G. tippelskirchi tippelskirchi (Masai giraffesensu stricto)
G. t. thornicrofti (Luangwa or Thornicroft's giraffe)
(Masai giraffe sensu lato)
G. giraffa
G. giraffa angolensis (Angolan giraffe)
G. g. giraffa (South African giraffe)
(southern giraffe)
The following table compares the different hypotheses for giraffe species. The description column shows the traditional nine subspecies in the one-species hypothesis.[1][25]
TheKordofan giraffe (G. c. antiquorum) has a distribution which includes southernChad, theCentral African Republic, northernCameroon, and the northeasternDemocratic Republic of the Congo.[1] Populations in Cameroon were formerly included inG. c. peralta, but this was incorrect.[26] Compared to the Nubian giraffe, this subspecies has smaller and more irregular spotting patterns. Its spots are present on insides of the legs, sometimes below thehocks. A median lump is present in males.[27]: 51–52 Some 2,000 are believed to remain in the wild.[1] Considerable confusion has existed over the status of this subspecies andG. c. peralta in zoos. In 2007, all allegedG. c. peralta inEuropean zoos were shown to be, in fact,G. c. antiquorum.[26] With this correction, about 65 are living in zoos.[28]
G. c. reticulata (only in three-species hypothesis)
TheNubian giraffe (G. c. camelopardalis), is found in easternSouth Sudan and southwesternEthiopia, in addition toKenya andUganda.[1] It has sharp-edgedchestnut-coloured spots surrounded by mostly white lines, while undersides lack spotting. A lump is prominent in the middle of the male's head.[27]: 51 Around 2,150 are thought to remain in the wild, with another 1,500 individuals belonging to theRothschild'secotype.[1] With the addition of Rothschild's giraffe to the Nubian subspecies, the Nubian giraffe is very common in captivity, although the original phenotype is rare — a group is kept atAl Ain Zoo in theUnited Arab Emirates.[30] In 2003, this group numbered 14.[31]
Rothschild's giraffe (G. c. rothschildi) may be an ecotype ofG. camelopardalis. Its range includes parts of Uganda andKenya.[1] Its presence in South Sudan is uncertain.[32] This giraffe has large dark patches with normally well-defined edges but sometimes split. The dark spots may also have swirls of pale colour within them. Spotting rarely reaches below the hocks and rarely to the hooves. This ecotype may also develop five "horns".[27]: 53 Around 1,500 individuals are believed to remain in the wild,[1] and more than 450 are living in zoos.[28] According to genetic analysis circa September 2016, it isconspecific with theNubian giraffe (G. c. camelopardalis).[20]
TheWest African giraffe (G. c. peralta) isendemic to southwestern Niger.[1] This animal has a lighterpelage (fur) than other subspecies,[33]: 322 with red lobe-shaped blotches that reach under the hocks. The ossicones are more erect than in other subspecies, and males have well-developed median lumps.[27]: 52–53 It is the most endangered subspecies withinGiraffa, with 400 individuals remaining in the wild.[1] Giraffes in Cameroon were formerly believed to belong to this species, but are actuallyG. c. antiquorum. This error resulted in some confusion over its status in zoos, but in 2007 it was established that all "G. c. peralta" kept in European zoos are actuallyG. c. antiquorum. The same 2007 study found that the West African giraffe was more closely related to Rothschild's giraffe than the Kordofan, and its ancestor may have migrated from eastern to northern Africa and then west as theSahara Desert spread. At its largest,Lake Chad may have acted as a boundary between the West African and Kordofan giraffes during theHolocene (before 5000 BC).[26]
Thereticulated giraffe (G. c. reticulata) is native to northeastern Kenya, southern Ethiopia, andSomalia.[1] Its distinctive coat pattern consists of sharp-edged, reddish-brown polygonal patches surrounded by thin white lines. Spots may or may not extend under the hocks, and a median lump is present in males.[27]: 53 An estimated 8,660 individuals remain in the wild,[1] and based onInternational Species Information System records, more than 450 are living in zoos.[28] A 2024 study found that the reticulated giraffe is the result ofhybridisation between northern and southern giraffe lineages.[24]
TheAngolan giraffe (G. c. angolensis)[36] occurs in northernNamibia, southwesternZambia, centralBotswana, westernZimbabwe, southern Zimbabwe and, since mid-2023, again in Angola.[37][1][38][39] A 2009 genetic study on this subspecies suggested the northernNamib Desert andEtosha National Park populations form a separate subspecies.[40] This subspecies is white with large brown blotches with pointed or cut edges. The spotting pattern extends throughout the legs but not the upper part of the face. The neck and rump patches tend to be fairly small. The subspecies also has a white ear mark.[27]: 51 About 13,000 animals are estimated to remain in the wild,[1] and about 20 are living in zoos.[28]
TheSouth African giraffe (G. c. giraffa) is found in northernSouth Africa, southern Botswana, northern Botswana and southwesternMozambique.[1][38][39] It has a tawny background colour marked with dark, somewhat rounded patches "with some fine projections". The spots extend down the legs, growing smaller as they do. The median lump of males is relatively small.[27]: 52 A maximum of 31,500 are estimated to remain in the wild,[1] and around 45 are living in zoos.[28]
TheMasai giraffe (G. c. tippelskirchi) can be found in central and southern Kenya and inTanzania.[1] Its coat patterns are highly diverse, with spots ranging from mostly rounded and smooth-edged to oval-shaped and incised or loped-edged.[42] A median lump is usually present in males.[27]: 54 [43] A total of 32,550 are thought to remain in the wild,[1] and about 100 are living in zoos.[28]
Thornicroft's giraffe (G. c. thornicrofti) is restricted to theLuangwa Valley in eastern Zambia.[1] It has notched and somewhatstar-shaped patches which and may or may not extend across the legs. The median lump of males is modestly sized.[27]: 54 No more than 550 remain in the wild,[1] with none in zoos.[28] It was named afterHarry Scott Thornicroft.[36]
The first extinct species to be described wasGiraffa sivalensis from Pakistan, the holotype of which was reevaluated as a vertebra of separate species within the genus that was initially described as a fossil of the living giraffe.[44] Another extinct speciesGiraffa punjabiensis is known from Pakistan.[45] Four other valid extinct species ofGiraffa known from Africa areGiraffa gracilis,Giraffa jumae,Giraffa pygmaea andGiraffa stillei.[8] "G."pomeli from Algeria and Tunisia is not a species of Giraffinae, but a species of Palaeotraginae related toMitilanotherium.[46]
Fully grown giraffes stand 4.3–5.7 m (14–19 ft) tall, withmales taller than females.[47] The average weight is 1,192 kg (2,628 lb) for an adult male and 828 kg (1,825 lb) for an adult female.[48] Despite its long neck and legs, its body is relatively short.[49]: 66 The skin is mostly gray[48] or tan,[50] and can reach a thickness of 20 mm (0.79 in).[51]: 87 The 80–100 cm (31–39 in) long[36] tail ends in a long, dark tuft of hair and is used as a defense against insects.[51]: 94
Thecoat has dark blotches or patches, which can be orange,chestnut, brown, or nearly black, surrounded by light hair, usually white orcream coloured.[52] Male giraffes become darker as they grow old.[43] The coat patternhas been claimed to serve ascamouflage in the light and shade patterns of savannah woodlands.[36] When standing among trees and bushes, they are hard to see at even a few metres distance. However, adult giraffes move about to gain the best view of an approaching predator, relying on their size and ability to defend themselves rather than on camouflage, which may be more important for calves.[6] Each giraffe has a unique coat pattern.[53][54] Calves inherit some coat pattern traits from their mothers, and variation in some spot traits is correlated with calf survival.[42] The skin under the blotches may regulate the animal's body temperature, being sites for complex blood vessel systems and largesweat glands.[55] Spotless or solid-colour giraffes are very rare, but have been observed.[56][57]
The fur may give the animal chemical defense, as its parasite repellents give it a characteristic scent. At least 11 mainaromatic chemicals are in the fur, althoughindole and3-methylindole are responsible for most of the smell. Because males have a stronger odour than females, it may also have asexual function.[58]
Head
Closeup of the head of a Southern giraffe
Both sexes have prominent horn-like structures calledossicones, which can reach 13.5 cm (5.3 in). They are formed fromossifiedcartilage, covered in skin, and fused to theskull at theparietal bones.[43][51]: 95–97 Beingvascularised, the ossicones may have a role inthermoregulation,[55] and are used in combat between males.[59] Appearance is a reliable guide to the sex or age of a giraffe: the ossicones of females and young are thin and display tufts of hair on top, whereas those of adult males tend to be bald and knobbed on top.[43] A lump, which is more prominent in males, emerges in the middle of the skull.[17] Males developcalcium deposits that form bumps on their skulls as they age.[52] Multiplesinuses lighten a giraffe's skull.[51]: 103 However, as males age, their skulls become heavier and more club-like, helping them become more dominant in combat.[43] Theoccipital condyles at the bottom of the skull allow the animal to tip its head over 90 degrees and grab food on the branches directly above them with the tongue.[51]: 103, 110 [17]
With eyes located on the sides of the head, the giraffe has a broadvisual field from its great height.[51]: 85, 102 Compared to otherungulates, giraffe vision is morebinocular and the eyes are larger with a greaterretinal surface area.[60] Giraffes may see in colour,[51]: 85 and their senses of hearing andsmell are sharp.[52] The ears are movable.[51]: 95 The nostrils are slit-shaped, possibly to withstand blowing sand.[61] The giraffe'stongue is about 45 cm (18 in) long. It is black, perhaps to protect against sunburn, and cangrasp foliage and delicately pick off leaves.[51]: 109–110 The upper lip is flexible and hairy to protect against sharp prickles.[17] The upper jaw has a hardpalate instead of front teeth. Themolars andpremolars are wide with lowcrowns on the surface.[51]: 106
Neck
The giraffe has an extremely elongated neck, which can be up to 2.4 m (7.9 ft) in length.[62] Along the neck is a mane made of short, erect hairs.[17] The neck typically rests at an angle of 50–60 degrees, though juveniles are closer to 70 degrees.[51]: 72–73 The long neck results from a disproportionate lengthening of thecervical vertebrae, not from the addition of more vertebrae. Each cervical vertebra is over 28 cm (11 in) long.[49]: 71 They comprise 52–54 per cent of the length of the giraffe'svertebral column, compared with the 27–33 percent typical of similar large ungulates, including the giraffe's closest living relative, theokapi.[13] This elongation largely takes place after birth, perhaps because giraffe mothers would have a difficult timegiving birth to young with the same neck proportions as adults.[63] The giraffe's head and neck are held up by large muscles and anuchal ligament, which are anchored by longthoracic vertebrae spines, giving them a hump.[17][64][36]
The giraffe's neck vertebrae haveball and socket joints.[49]: 71 The point of articulation between the cervical and thoracic vertebrae of giraffes is shifted to lie between the first and second thoracic vertebrae (T1 and T2), unlike in most other ruminants, where the articulation is between the seventh cervical vertebra (C7) and T1.[13][63] This allows C7 to contribute directly to increased neck length and has given rise to the suggestion that T1 is actually C8, and that giraffes have added an extra cervical vertebra.[64] However, this proposition is not generally accepted, as T1 has othermorphological features, such as an articulatingrib, deemed diagnostic of thoracic vertebrae, and because exceptions to the mammalian limit of seven cervical vertebrae are generally characterised by increasedneurological anomalies and maladies.[13]
There are several hypotheses regarding the evolutionary origin and maintenance of elongation in giraffe necks.[59]Charles Darwin originally suggested the "competingbrowsers hypothesis", which has been challenged only recently. It suggests that competitive pressure from smaller browsers, likekudu,steenbok andimpala, encouraged the elongation of the neck, as it enabled giraffes to reach food that competitors could not. This advantage is real, as giraffes can and do feed up to 4.5 m (15 ft) high, while even quite large competitors, such as kudu, can feed up to only about 2 m (6 ft 7 in) high.[65] There is also research suggesting that browsing competition is intense at lower levels, and giraffes feed more efficiently (gaining more leaf biomass with each mouthful) high in the canopy.[66][67] However, scientists disagree about just how much time giraffes spend feeding at levels beyond the reach of other browsers,[10][59][65][68] and a 2010 study found that adult giraffes with longer necks actually suffered higher mortality rates under drought conditions than their shorter-necked counterparts. This study suggests that maintaining a longer neck requires more nutrients, which puts longer-necked giraffes at risk during a food shortage.[69]
Another theory, thesexual selection hypothesis, proposes that long necks evolved as a secondarysexual characteristic, giving males an advantage in "necking" contests to establish dominance and obtain access to sexually receptive females.[10] In support of this theory, some studies have stated that necks are longer and heavier for males than females of the same age,[10][59] and that males do not employ other forms of combat.[10] However, a 2024 study found that, while males have thicker necks, females actually have proportionally longer ones, which is likely because of their greater need to find more food to sustain themselves and their dependent young.[70] It has also been proposed that the neck serves to give the animal greater vigilance.[71][72]
Legs, locomotion and posture
Right hind leg of a Masai giraffe at theSan Diego Zoo
The front legs tend to be longer than the hind legs,[51]: 109 and males have proportionally longer front legs than females, which gives them better support when swinging their necks during fights.[70] The leg bones lack first, second and fifthmetapodials.[51]: 109 It appears that asuspensory ligament allows the lanky legs to support the animal's great weight.[73] Thehooves of large male giraffes reach 31 cm × 23 cm (12.2 in × 9.1 in) in diameter.[51]: 98 Thefetlock of the leg is low to the ground, allowing the hoof to better support the animal's weight. Giraffes lackdewclaws and interdigital glands. While the pelvis is relatively short, theilium has stretched-out crests.[17]
A giraffe has only twogaits: walking and galloping. Walking is done by moving the legs on one side of the body, then doing the same on the other side.[43] When galloping, the hind legs move around the front legs before the latter move forward,[52] and the tail will curl up.[43] The movements of the head and neck provide balance and control momentum while galloping.[33]: 327–29 The giraffe can reach a sprint speed of up to 60 km/h (37 mph),[74] and can sustain 50 km/h (31 mph) for several kilometres.[75] Giraffes would probably not be competent swimmers as their long legs would be highly cumbersome in the water,[76] although they might be able to float.[77] When swimming, the thorax would be weighed down by the front legs, making it difficult for the animal to move its neck and legs in harmony[76][77] or keep its head above the water's surface.[76]
A juvenile giraffe walking in Malawi
A giraffe rests by lying with its body on top of its folded legs.[33]: 329 To lie down, the animal kneels on its front legs and then lowers the rest of its body. To get back up, it first gets on its front knees and positions its backside on top of its hindlegs. It then pulls the backside upwards, and the front legs stand straight up again. At each stage, the animal swings its head for balance.[51]: 67 If the giraffe wants to reach down to drink, it either spreads its front legs or bends its knees.[43] Studies in captivity found the giraffe sleeps intermittently around 4.6 hours per day, mostly at night. It usually sleeps lying down; however, standing sleeps have been recorded, particularly in older individuals. Intermittent short "deep sleep" phases while lying are characterised by the giraffe bending its neck backwards and resting its head on the hip or thigh, a position believed to indicateparadoxical sleep.[78]
Internal systems
Scheme of path of the recurrent laryngeal nerve in giraffe
In mammals, the leftrecurrent laryngeal nerve is longer than the right; in the giraffe, it is over 30 cm (12 in) longer. These nerves are longer in the giraffe than in any other living animal;[79] the left nerve is over 2 m (6 ft 7 in) long.[80] Each nerve cell in this path begins in thebrainstem and passes down the neck along thevagus nerve, then branches off into the recurrent laryngeal nerve which passes back up the neck to the larynx. Thus, these nerve cells have a length of nearly 5 m (16 ft) in the largest giraffes.[79] Despite its long neck and large skull, thebrain of the giraffe is typical for an ungulate.[81] Evaporative heat loss in the nasal passages keep the giraffe's brain cool.[55] The shape of the skeleton gives the giraffe a smalllung volume relative to its mass. Its long neck gives it a large amount ofdead space, though this is limited by its narrow windpipe. The giraffe also has a hightidal volume, so the balance of dead space and tidal volume is much the same as other mammals. The animal can still provide enough oxygen for its tissues, and it can increase itsrespiratory rate and oxygen diffusion when running.[82]
Reticulated giraffe bending down to drink in Kenya. The circulatory system is adapted to deal with blood flow rushing down its neck.
The giraffe'scirculatory system has several adaptations to compensate for its great height.[14] Its 11 kg (25 lb) and 60 cm (2 ft)heart must generate approximately double the blood pressure required for a human to maintain blood flow to the brain. As such, the wall of the heart can be as thick as 7.5 cm (3.0 in).[52] Giraffes have relatively high heart rates for their size, at 150 beats per minute.[49]: 76 When the animal lowers its head, the blood rushes down fairly unopposed and arete mirabile in the upper neck, with its largecross-sectional area, prevents excess blood flow to the brain. When it raises again, the blood vessels constrict and push blood into the brain so the animal does not faint.[83] Thejugular veins contain several (most commonly seven) valves to prevent blood flowing back into the head from theinferior vena cava andright atrium while the head is lowered.[84] Conversely, the blood vessels in the lower legs are under great pressure because of the weight of fluid pressing down on them. To solve this problem, the skin of the lower legs is thick and tight, preventing too much blood from pouring into them.[36]
Giraffes haveoesophageal muscles that are strong enough to allow regurgitation of food from the stomach up the neck and into the mouth forrumination.[49]: 78 They havefour-chambered stomachs, which are adapted to their specialized diet.[17] Theintestines of an adult giraffe measure more than 70 m (230 ft) in length and have a relatively small ratio ofsmall tolarge intestine.[85] The giraffe has a small, compact liver.[49]: 76 In fetuses there may be a smallgallbladder that vanishes before birth.[17][86][87]
Behaviour and ecology
Habitat and feeding
AMasai giraffe extending its tongue to feed, in Tanzania.
A giraffe in Malawi eating leaves from a tree
Giraffes usually inhabit savannahs and openwoodlands. They prefer areas dominated byAcacieae,Commiphora,Combretum andTerminalia trees overBrachystegia which are more densely spaced.[33]: 322 The Angolan giraffe can be found in desert environments.[88] Giraffes browse on the twigs of trees, preferring those of the subfamily Acacieae and the generaCommiphora andTerminalia,[89] which are important sources of calcium and protein to sustain the giraffe's growth rate.[6] They also feed on shrubs, grass and fruit.[33]: 324 A giraffe eats around 34 kg (75 lb) of plant matter daily.[43] When stressed, giraffes may chew on large branches, stripping them of bark.[33]: 325 Giraffes are also recorded tochew old bones.[51]: 102
During the wet season, food is abundant and giraffes are more spread out, while during the dry season, they gather around the remaining evergreen trees and bushes.[89] Mothers tend to feed in open areas, presumably to make it easier to detect predators, although this may reduce their feeding efficiency.[68] As aruminant, the giraffe first chews its food, then swallows it for processing and then visibly passes the half-digested cud up the neck and back into the mouth to chew again.[49]: 78–79 The giraffe requires less food than many other herbivores because the foliage it eats has more concentrated nutrients and it has a more efficient digestive system.[89] The animal's faeces come in the form of small pellets.[17] When it has access to water, a giraffe will go no more than three days without drinking.[43]
Giraffes have a great effect on the trees that they feed on, delaying the growth of young trees for some years and giving "waistlines" to particularly tall trees. Feeding is at its highest during the first and last hours of daytime. Between these hours, giraffes mostly stand and ruminate. Rumination is the dominant activity during the night, when it is mostly done lying down.[43]
Social life
Gathering of female South African giraffes inTswalu Kalahari Reserve, South Africa. These animals commonly gather in herds.
Giraffes usually form groups that vary in size and composition according to ecological, anthropogenic, temporal, and social factors.[90] Traditionally, the composition of these groups had been described as open and ever-changing.[91] For research purposes, a "group" has been defined as "a collection of individuals that are less than a kilometre apart and moving in the same general direction".[92] More recent studies have found that giraffes have long-lasting social groups or cliques based on kinship, sex or other factors, and these groups regularly associate with other groups in larger communities or sub-communities within afission–fusion society.[93][94][95][96] Proximity to humans can disrupt social arrangements.[93] Masai giraffes in Tanzania sort themselves into different subpopulations of 60–90 adult females with overlapping ranges, each of which differ in reproductive rates and calf mortality.[97]Dispersal is male biased, and can include spatial and/or social dispersal.[98] Adult female subpopulations are connected by males into super communities of around 300 animals.[99]
The number of giraffes in a group can range from one up to 66 individuals.[90][96] Giraffe groups tend to be sex-segregated[96] although mixed-sex groups made of adult females and young males also occur.[92] Female groups may bematrilineally related.[96] Generally, females are more selective than males when deciding which individuals of the same sex they associate with.[95] Particularly stable giraffe groups are those made of mothers and their young,[92] which can last weeks or months.[100] Young males also form groups and will engage in playfights. However, as they get older, males become more solitary but may also associate in pairs or with female groups.[96][100] Giraffes are notterritorial,[17] but they havehome ranges that vary according to rainfall and proximity to human settlements.[101] Male giraffes occasionally roam far from areas that they normally frequent.[33]: 329
Early biologists suggested giraffes were mute and unable to create enough air flow to vibrate theirvocal folds.[102] This has been proved to the contrary; they have been recorded tocommunicate using snorts, sneezes, coughs, snores, hisses, bursts, moans, grunts, growls and flute-like sounds.[43][102] During courtship, males emit loud coughs. Females call their young by bellowing. Calves will emit bleats, mooing and mewing sounds.[43] Snorting and hissing is associated with vigilance.[103] During nighttime, giraffes appear tohum to each other.[104] There is some evidence that giraffes useHelmholtz resonance to createinfrasound.[105] They also communicate with body language.Dominant males display to other males with an erect posture; holding the chin and head up while walking stiffly and displaying their side. The less dominant show submissiveness by dropping the head and ears, lowering the chin and fleeing.[43]
Reproduction and parental care
Angolan giraffes mating in Namibia
Reproduction in giraffes is broadlypolygamous: a few older males mate with the fertile females.[92] Females can reproduce throughout the year and experienceoestrus cycling approximately every 15 days.[106][107] Female giraffes in oestrus are dispersed over space and time, so reproductive adult males adopt a strategy of roaming among female groups to seek mating opportunities, with periodic hormone-induced rutting behaviour approximately every two weeks.[108] Males prefer young adult females over juveniles and older adults.[92]
Male giraffes assess female fertility by tasting the female's urine to detect oestrus, in a multi-step process known as theflehmen response.[92][100] Once an oestrous female is detected, the male will attempt to court her. When courting, dominant males will keep subordinate ones at bay.[100] A courting male may lick a female's tail, lay his head and neck on her body or nudge her with his ossicones. During copulation, the male stands on his hind legs with his head held up and his front legs resting on the female's sides.[43]
Giraffegestation lasts 400–460 days, after which a single calf is normally born, although twins occur on rare occasions.[106] The mother gives birth standing up. The calf emerges head and front legs first, having broken through thefetal membranes, and falls to the ground, severing theumbilical cord.[17] A newborn giraffe is 1.7–2 m (5 ft 7 in – 6 ft 7 in) tall.[47] Within a few hours of birth, the calf can run around and is almost indistinguishable from a one-week-old. However, for the first one to three weeks, it spends most of its time hiding,[109] its coat pattern providing camouflage. The ossicones, which have lain flat in the womb, raise up in a few days.[43]
Female Angolan giraffe with calf
Mothers with calves will gather in nursery herds, moving or browsing together. Mothers in such a group may sometimes leave their calves with one female while they forage and drink elsewhere. This is known as a "calving pool".[109] Calves are at risk of predation, and a mother giraffe will stand over them and kick at an approaching predator.[43] Females watching calving pools will only alert their own young if they detect a disturbance, although the others will take notice and follow.[109] Allo-sucking, where a calf will suckle a female other than its mother, has been recorded in both wild and captive giraffes.[110][111] Calves first ruminate at four to six months and stopnursing at six to eight months. Young may not reach independence until they are 14 months old.[51]: 49 Females are able to reproduce at four years of age,[43] whilespermatogenesis in males begins at three to four years of age.[112] Males must wait until they are at least seven years old to gain the opportunity to mate.[43]
Necking
Here, male South African giraffes engage in low intensity necking to establish dominance, inIthala Game Reserve,Kwa-Zulu-Natal, South Africa.
Male giraffes use their necks asweapons in combat, a behaviour known as "necking". Necking is used to establish dominance, and males that win necking bouts have greaterreproductive success.[10] This behaviour occurs at low or high intensity. In low-intensity necking, the combatants rub and lean on each other. The male that can keep itself more upright wins the bout. In high-intensity necking, the combatants will spread their front legs and swing their necks at each other, attempting to land blows with their ossicones. The contestants will try to dodge each other's blows and then prepare to counter. The power of a blow depends on the weight of the skull and the arc of the swing.[43] A necking duel can last more than half an hour, depending on how well matched the combatants are.[33]: 331 Although most fights do not lead to serious injury, there have been records of broken jaws, broken necks, and even deaths.[10]
After a duel, it is common for two male giraffes to caress and court each other. Such interactions between males have been found to be more frequent than heterosexual coupling.[113] In one study, up to 94 percent of observed mounting incidents took place between males. The proportion ofsame-sex activities varied from 30 to 75 percent. Only one percent of same-sex mounting incidents occurred between females.[114]
Giraffes have high adult survival probability,[115] and an unusually long lifespan compared to other ruminants, up to 38 years.[116] Adult female survival is significantly correlated with the number of social associations.[117] Because of their size, eyesight and powerful kicks, adult giraffes are mostly safe from predation,[43] withlions being their only major threats.[51]: 55 Calves are much more vulnerable than adults and are also preyed on byleopards,spotted hyenas andwild dogs.[52] A quarter to a half of giraffe calves reach adulthood.[115][118] Calf survival varies according to the season of birth, with calves born during thedry season having higher survival rates.[119]
The local, seasonal presence of large herds of migratorywildebeests andzebras reduces predation pressure on giraffe calves and increases their survival probability.[120] In turn, it has been suggested that other ungulates may benefit from associating with giraffes, as their height allows them to spot predators from further away. Zebras were found to assess predation risk by watching giraffes and spend less time looking around when giraffes are present.[121]
Red-billed oxpeckers on a giraffe, Zambia
Some parasites feed on giraffes. They are oftenhosts forticks, especially in the area around the genitals, which have thinner skin than other areas. Tick species that commonly feed on giraffes are those of generaHyalomma,Amblyomma andRhipicephalus.Red-billed andyellow-billed oxpeckers clean giraffes of ticks and alert them to danger. Giraffes host numerous species of internal parasites and are susceptible to various diseases. They were victims of the (now eradicated) viral illnessrinderpest.[17] Giraffes can also suffer from a skin disorder, which comes in the form of wrinkles,lesions or rawfissures. As much as 79% of giraffes have symptoms of the disease inRuaha National Park, but it did not cause mortality inTarangire and is less prevalent in areas with fertile soils.[122][123][124]
Human relations
Cultural significance
With its lanky build and spotted coat, the giraffe has been a source of fascination throughout human history, and its image is widespread in culture. It has represented flexibility, far-sightedness, femininity, fragility, passivity, grace, beauty and the continent of Africa itself.[125]: 7, 116
Giraffes were depicted in art throughout the African continent,.[125]: 45–47 TheKiffians were responsible for a life-size rock engraving of two giraffes, dated 8,000 years ago, that has been called the "world's largest rock art petroglyph".[125]: 45 [126] Judging from evidence including incised clay pots, archaeologists now believe that, in the ancientKushite societies located inNubia, or what is now northern Sudan, giraffes may have featured in popular religion and women's religion, though not in elite or royal religion, and may have also had a connection to beliefs about the sun.[127] TheTugen people of modern Kenya used the giraffe to depict their god Mda.[128] TheEgyptians gave the giraffe its ownhieroglyph; 'sr' inOld Egyptian and 'mmy' in later periods.[125]: 49 How the giraffe got its height has been the subject of variousAfrican folktales.[10]
Giraffes have a presence in modernWestern culture.Salvador Dalí depicted them with burning manes in some surrealist paintings. Dali considered the giraffe to be a masculine symbol. A flaming giraffe was meant to be a "masculine cosmic apocalyptic monster".[125]: 123 Several children's books feature the giraffe, including David A. Ufer'sThe Giraffe Who Was Afraid of Heights,Giles Andreae'sGiraffes Can't Dance andRoald Dahl'sThe Giraffe and the Pelly and Me. Giraffes have appeared in animated films as minor characters inDisney'sDumbo andThe Lion King, and in more prominent roles inThe Wild and theMadagascar films.Sophie the Giraffe has been a popularteether since 1961. Another famous fictional giraffe is the Toys "R" Us mascotGeoffrey the Giraffe.[125]: 127
The giraffe has also been used for some scientific experiments and discoveries. Scientists have used the properties of giraffe skin as a model forastronaut andfighter pilot suits because the people in these professions are in danger of passing out if blood rushes to their legs.[49]: 76 Computer scientists have modeled the coat patterns of several subspecies usingreaction–diffusion mechanisms.[129] Theconstellation ofCamelopardalis, introduced in the 17th century, depicts a giraffe.[125]: 119–20 TheTswana people of Botswana traditionally see the constellationCrux as two giraffes—Acrux andMimosa forming a male, andGacrux andDelta Crucis forming the female.[130]
Painting of a giraffe imported to China during theMing dynasty, in which it was identified with the mythologicalQilin
Captivity
The Egyptians were among the earliest people to keep giraffes in captivity and shipped them around the Mediterranean.[125]: 48–49 The giraffe was among the many animals collected anddisplayed by the Romans. The first one in Rome was brought in byJulius Caesar in 46 BC.[125]: 52 With thefall of the Western Roman Empire, the housing of giraffes in Europe declined.[125]: 54 During theMiddle Ages, giraffes were known to Europeans through contact with the Arabs, who revered the giraffe for its peculiar appearance.[52]
Individual captive giraffes were given celebrity status throughout history. In 1414, a giraffe fromMalindi wastaken to China by explorerZheng He and placed in aMing dynasty zoo. The animal was a source of fascination for the Chinese people, who associated it with the mythicalQilin.[125]: 56 TheMedici giraffe was a giraffe presented toLorenzo de' Medici in 1486. It caused a great stir on its arrival inFlorence.[131]Zarafa, another famous giraffe, was brought fromEgypt to Paris in the early 19th century as a gift forCharles X of France. A sensation, the giraffe was the subject of numerous memorabilia or "giraffanalia".[125]: 81
Giraffes have become popular attractions in modernzoos, though keeping them is difficult as they prefer large areas and need to eat large amounts of browse. Captive giraffes in North America and Europe appear to have a higher mortality rate than in the wild, the most common causes being poor husbandry, nutrition, and management.[51]: 153 Giraffes in zoos displaystereotypical behaviours, particularly the licking of inanimate objects and pacing.[51]: 164 Zookeepers may offer various activities to stimulate giraffes, including training them to take food from visitors.[51]: 167, 176 Stables for giraffes are built particularly high to accommodate their height.[51]: 183
Exploitation
Giraffes were probably common targets for hunters throughout Africa.[33]: 337 Different parts of their bodies were used for different purposes.[17] Their meat was used for food. The tail hairs wereflyswatters, bracelets, necklaces, and threads. Shields, sandals, and drums were made using the skin, and the strings of musical instruments were from the tendons.[17][33]: 337 InBuganda, the smoke of burning giraffe skin was traditionally used to treat nosebleeds.[33]: 337 TheHumr people ofKordofan consume the drink Umm Nyolokh, which is prepared from theliver andbone marrow of giraffes.Richard Rudgley hypothesised that Umm Nyolokh might containDMT.[132] The drink is said to cause hallucinations of giraffes, believed to be the giraffes' ghosts, by theHumr.[133]
Conservation status
In 2016, giraffes were assessed asVulnerable from a conservation perspective by theInternational Union for Conservation of Nature (IUCN).[1] In 1985, it was estimated there were 155,000 giraffes in the wild. This declined to over 140,000 in 1999.[134] Estimates as of 2016 indicate there are approximately 97,500 members ofGiraffa in the wild.[135][136] The Masai and reticulated subspecies areendangered,[137][138] and the Rothschild subspecies isnear threatened.[32] The Nubian subspecies iscritically endangered.[139] In 2025, the IUCN currently accepted four species of giraffe with seven subspecies.[1][140]
The primary causes for giraffe population declines arehabitat loss and direct killing forbushmeat markets. Giraffes have beenextirpated from much of their historic range, includingEritrea,Guinea,Mauritania andSenegal.[1] They may also have disappeared fromAngola,Mali, andNigeria, but have been introduced toRwanda andEswatini.[1][139] As of 2010[update], there were more than 1,600 in captivity atSpecies360-registered zoos.[28] Habitat destruction has hurt the giraffe. In theSahel, the need for firewood and grazing room for livestock has led todeforestation. Normally, giraffes can coexist with livestock, since they avoid direct competition by feeding above them.[36] In 2017, severe droughts in northern Kenya led to increased tensions over land and the killing of wildlife by herders, with giraffe populations being particularly hit.[141]
Protected areas like national parks provide important habitat and anti-poaching protection to giraffe populations.[1] Community-based conservation efforts outside national parks are also effective at protecting giraffes and their habitats.[142][143] Private game reserves have contributed to the preservation of giraffe populations in eastern and southern Africa.[36] The giraffe is a protected species in most of its range. It is thenational animal of Tanzania,[144] and is protected by law,[145] and unauthorised killing can result in imprisonment.[146] The UN-backed Convention of Migratory Species selected giraffes for protection in 2017.[147] In 2019, giraffes were listed under Appendix II of the Convention on International Trade in Endangered Species (CITES), which means international trade including in parts/derivatives is regulated.[148]
Translocations are sometimes used to augment or re-establish diminished or extirpated populations, but these activities are risky and difficult to undertake using the best practices of extensive pre- and post-translocation studies and ensuring a viable founding population.[149][150] Aerial survey is the most common method of monitoring giraffe population trends in the vast roadless tracts of African landscapes, but aerial methods are known to undercount giraffes. Ground-based survey methods are more accurate and can be used in conjunction with aerial surveys to make accurate estimates of population sizes and trends.[151]
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