The giraffe (Giraffa) is an African artiodactyl mammal, the tallest living terrestrial animal and the largest ruminant. It is traditionally considered to be one species, Giraffa camelopardalis, with nine subspecies. However, the existence of up to nine extant giraffe species has been described, based upon research into the mitochondrial and nuclear DNA, as well as morphological measurements of Giraffa. Seven other species are extinct, prehistoric species known from fossils.
|Masai giraffe (G. c. tippelskirchi) at the Mikumi National Park, Tanzania|
|Range map of extant Giraffa subspecies|
The giraffe's chief distinguishing characteristics are its extremely long neck and legs, its horn-like ossicones, and its distinctive coat patterns. It is classified under the family Giraffidae, along with its closest extant relative, the okapi. Its scattered range extends from Chad in the north to South Africa in the south, and from Niger in the west to Somalia in the east. Giraffes usually inhabit savannahs and woodlands. Their food source is leaves, fruits and flowers of woody plants, primarily acacia species, which they browse at heights most other herbivores cannot reach.
Giraffes may be preyed on by lions, leopards, spotted hyenas and African wild dogs. Giraffes live in herds of related females and their offspring, or bachelor herds of unrelated adult males, but are gregarious and may gather in large aggregations. Males establish social hierarchies through "necking", which are combat bouts where the neck is used as a weapon. Dominant males gain mating access to females, which bear the sole responsibility for raising the young.
The giraffe has intrigued various cultures, both ancient and modern, for its peculiar appearance, and has often been featured in paintings, books, and cartoons. It is classified by the International Union for Conservation of Nature as vulnerable to extinction, and has been extirpated from many parts of its former range. Giraffes are still found in numerous national parks and game reserves but estimates as of 2016 indicate that there are approximately 97,500 members of Giraffa in the wild. More than 1,600 were kept in zoos in 2010.
The name "giraffe" has its earliest known origins in the Arabic word zarāfah (زرافة), perhaps borrowed from the animal's Somali name geri. The Arab name is translated as "fast-walker". In early Modern English the spellings jarraf and ziraph were used, probably directly from the Arabic, and in Middle English orafle and gyrfaunt, gerfaunt. The Italian form giraffa arose in the 1590s. The modern English form developed around 1600 from the French girafe.
"Camelopard" // is an archaic English name for the giraffe; it derives from the Ancient Greek καμηλοπάρδαλις (kamēlopárdalis), from κάμηλος (kámēlos), "camel", and πάρδαλις (párdalis), "leopard", referring to its camel-like shape and leopard-like colouration.
Living giraffes were originally classified as one species by Carl Linnaeus in 1758. He gave it the binomial name Cervus camelopardalis. Morten Thrane Brünnich classified the genus Giraffa in 1762. The species name camelopardalis is from Latin.
|Cladogram based on a 2003 study by Hassanin and Douzery.|
The giraffe is one of only two living genera of the family Giraffidae in the order Artiodactyla, the other being the okapi. The family was once much more extensive, with over 10 fossil genera described. Their closest known relatives may have been the extinct deer-like climacocerids. They, together with the family Antilocapridae (whose only extant species is the pronghorn), have been placed in the superfamily Giraffoidea. These animals may have evolved from the extinct family Palaeomerycidae which might also have been the ancestor of deer.
The elongation of the neck appears to have started early in the giraffe lineage. Comparisons between giraffes and their ancient relatives suggest that vertebrae close to the skull lengthened earlier, followed by lengthening of vertebrae further down. One early giraffid ancestor was Canthumeryx which has been dated variously to have lived 25–20 million years ago (mya), 17–15 mya or 18–14.3 mya and whose deposits have been found in Libya. This animal was medium-sized, slender and antelope-like. Giraffokeryx appeared 15 mya in the Indian subcontinent and resembled an okapi or a small giraffe, and had a longer neck and similar ossicones. Giraffokeryx may have shared a clade with more massively built giraffids like Sivatherium and Bramatherium.
Giraffids like Palaeotragus, Shansitherium and Samotherium appeared 14 mya and lived throughout Africa and Eurasia. These animals had bare ossicones and small cranial sinuses and were longer with broader skulls. Paleotragus resembled the okapi and may have been its ancestor. Others find that the okapi lineage diverged earlier, before Giraffokeryx. Samotherium was a particularly important transitional fossil in the giraffe lineage as its cervical vertebrae was intermediate in length and structure between a modern giraffe and an okapi, and was more vertical than the okapi's. 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.
Bohlinia entered China and northern India in response to climate change. From there, the genus Giraffa evolved and, around 7 mya, entered Africa. Further climate changes caused the extinction of the Asian giraffes, while the African giraffes survived and radiated into several new species. Living giraffes appear to have arisen around 1 mya in eastern Africa during the Pleistocene. Some biologists suggest the modern giraffes descended from G. jumae; others find G. gracilis a more likely candidate. G. jumae was larger and more heavily built while G. gracilis was smaller and more lightly built. The main driver for the evolution of the giraffes is believed to have been the changes from extensive forests to more open habitats, which began 8 mya. During this time, tropical plants disappeared and were replaced by arid C4 plants, and a dry savannah emerged across eastern and northern Africa and western India. 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. 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.
The giraffe genome is around 2.9 billion base 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. The two species are equally distantly related to cattle, suggesting the giraffe's unique characteristics are not because of faster evolution. The divergence of giraffe and okapi lineages dates to around 11.5 mya. A small group of regulatory genes in the giraffe appear to be responsible for the animal's stature and associated circulatory adaptations.
Species and subspecies
The IUCN and most giraffe scientists currently recognise only one species of giraffe with nine subspecies. During the 1900s, various taxonomies with 2 or 3 species were proposed. In 2001, a two-species taxonomy was proposed. A 2007 study on the genetics of giraffes, suggested six species. A 2011 study using detailed analyses of the morphology of giraffes, and application of the phylogenetic species concept, described eight species of living giraffes. 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 1 to 2 million years. Since then, a response to this publication has been published, highlighting seven problems in data interpretation, and concludes "the conclusions should not be accepted unconditionally".
A 2020 study showed that depending on the method chosen, different taxonomic hypotheses recognizing from two to six species can be considered for the genus Giraffa. 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 recognises G. camelopardalis, G. giraffa, and G. tippelskirchi, is highly supported by phylogenetic analyses and also corroborated by most population genetic and multi-species coalescent analyses. A 2021 whole genome sequencing study suggests the existence of four distinct species and seven subspecies.
G. attica, also extinct, was formerly considered part of Giraffa but was reclassified as Bohlinia attica in 1929. There are also seven extinct species of giraffe, listed as the following:
- †Giraffa gracilis
- †Giraffa jumae
- †Giraffa priscilla
- †Giraffa punjabiensis
- †Giraffa pygmaea
- †Giraffa sivalensis
- †Giraffa stillei
|One species taxonomy||Three species taxonomy||Four species taxonomy||Eight species taxonomy||Description||Image|
|Giraffe (G. camelopardalis)||Northern giraffe (G. camelopardalis)||Northern giraffe (G. camelopardalis)||Kordofan giraffe (G. antiquorum)||The Kordofan giraffe (G. c. antiquorum) has a distribution which includes southern Chad, the Central African Republic, northern Cameroon, and the northeastern DR Congo. Populations in Cameroon were formerly included in G. c. peralta, but this was incorrect. Compared to the Nubian giraffe, this subspecies has smaller and more irregular spotting patterns. Its spots may be found below the hocks and the insides of the legs. A median lump is present in males.:51–52 Some 2,000 are believed to remain in the wild. Considerable confusion has existed over the status of this subspecies and G. c. peralta in zoos. In 2007, all alleged G. c. peralta in European zoos were shown to be, in fact, G. c. antiquorum. With this correction, about 65 are kept in zoos. The formerly recognised subspecies G. c. congoesis is now considered part of the Kordofan giraffe.|
|Nubian giraffe including Rothschild's giraffe (G. camelopardalis) also known as Baringo giraffe or Ugandan giraffe||The Nubian giraffe (G. c. camelopardalis), is found in eastern South Sudan and southwestern Ethiopia, in addition to Kenya and Uganda. It has sharply defined chestnut-coloured spots surrounded by mostly white lines, while undersides lack spotting. The median lump is particularly developed in the male.:51 Around 2,150 are thought to remain in the wild, with another 1,500 individuals belonging to the Rothschild's ecotype. 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 at Al Ain Zoo in the United Arab Emirates. In 2003, this group numbered 14.
Rothschild's giraffe (G. c. rothschildi) may be an ecotype of G. camelopardalis. Its range includes parts of Uganda and Kenya. Its presence in South Sudan is uncertain. This giraffe has large dark patches that usually have complete margins, but may also have sharp edges. The dark spots may also have paler radiating lines or streaks within them. Spotting does not often reach below the hocks and almost never to the hooves. This ecotype may also develop five "horns".:53 Around 1,500 individuals believed to remain in the wild, and more than 450 are kept in zoos. According to genetic analysis circa September 2016, it is conspecific with the Nubian giraffe (G. c. camelopardalis).
|West African giraffe (G. peralta), also known as Niger giraffe or Nigerian giraffe||The West African giraffe (G. c. peralta) is endemic to southwestern Niger. This animal has a lighter pelage than other subspecies,:322 with red lobe-shaped blotches that reach below the hocks. The ossicones are more erect than in other subspecies and males have well-developed median lumps.:52–53 It is the most endangered subspecies within Giraffa, with 400 individuals remaining in the wild. Giraffes in Cameroon were formerly believed to belong to this species, but are actually G. 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 actually are G. 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 to its current range with the development of the Sahara Desert. At its largest, Lake Chad may have acted as a barrier between the West African and Kordofan giraffes during the Holocene (before 5000 BC).|
|Reticulated giraffe (G. reticulata), also known as Somali giraffe||The reticulated giraffe (G. c. reticulata) is native to northeastern Kenya, southern Ethiopia, and Somalia. Its distinctive coat pattern consists of sharp-edged, reddish-brown polygonal patches divided by a network of thin white lines. Spots may or may not extend below the hocks, and a median lump is present in males.:53 An estimated 8,660 individuals remain in the wild, and based on International Species Information System records, more than 450 are kept in zoos.|
|Southern giraffe (G. giraffa)||Southern giraffe (G. giraffa)||Angolan giraffe (G. angolensis), also known as Namibian giraffe||The Angolan giraffe (G. c. angolensis) is found in northern Namibia, southwestern Zambia, Botswana, and western Zimbabwe. A 2009 genetic study on this subspecies suggested the northern Namib Desert and Etosha National Park populations form a separate subspecies. This subspecies has large brown blotches with edges that are either somewhat notched or have angular extensions. 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 patch.:51 About 13,000 animals are estimated to remain in the wild; and about 20 are kept in zoos.|
|South African giraffe (G. giraffa) also known as Cape giraffe||The South African giraffe (G. c. giraffa) is found in northern South Africa, southern Botswana, southern Zimbabwe, and southwestern Mozambique. It has dark, somewhat rounded patches "with some fine projections" on a tawny background colour. The spots extend down the legs and get smaller as they do. The median lump of males is less developed.:52 A maximum of 31,500 are estimated to remain in the wild, and around 45 are kept in zoos.|
|Masai giraffe (G. tippelskirchi)||Masai giraffe (G. tippelskirchi)||Masai giraffe (G. tippelskirchi), also known as Kilimanjaro giraffe||The Masai giraffe (G. c. tippelskirchi) can be found in central and southern Kenya and in Tanzania. It has distinctive, irregular, jagged, star-like blotches which extend to the hooves. A median lump is usually present in males.:54 A total of 32,550 are thought to remain in the wild, and about 100 are kept in zoos.|
|Thornicroft's giraffe ("G. thornicrofti", after Harry Scott Thornicroft), also known as Luangwa giraffe, or Rhodesian giraffe||Thornicroft's giraffe (G. c. thornicrofti) is restricted to the Luangwa Valley in eastern Zambia. The patches are notched and somewhat star-shaped, and may or may not extend across the legs. The median lump of males is underdeveloped.:54 No more than 550 remain in the wild, with none kept in zoos.|
Appearance and anatomy
Fully grown giraffes stand 4.3–5.7 m (14.1–18.7 ft) tall, with males taller than females. The tallest recorded male was 5.88 m (19.3 ft) and the tallest recorded female was 5.17 m (17.0 ft) tall. The average weight is 1,192 kg (2,628 lb) for an adult male and 828 kg (1,825 lb) for an adult female with maximum weights of 1,930 kg (4,250 lb) and 1,180 kg (2,600 lb) having been recorded for males and females, respectively. Despite its long neck and legs, the giraffe's body is relatively short.:66 Located at both sides of the head, the giraffe's large, bulging eyes give it good all-round vision from its great height.:25 Giraffes see in colour:26 and their senses of hearing and smell are also sharp. The animal can close its muscular nostrils to protect against sandstorms and ants.:27
The giraffe's prehensile tongue is about 45 cm (18 in) long. It is purplish-black in colour, perhaps to protect against sunburn, and is useful for grasping foliage, as well as for grooming and cleaning the animal's nose.:27 The upper lip of the giraffe is also prehensile and useful when foraging, and is covered in hair to protect against thorns. The tongue and inside of the mouth are covered in papillae.
The coat has dark blotches or patches (which can be orange, chestnut, brown, or nearly black in colour) separated by light hair (usually white or cream in colour.) Male giraffes become darker as they age. The coat pattern has been claimed to serve as camouflage in the light and shade patterns of savannah woodlands. 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. Each individual giraffe has a unique coat pattern. Giraffe calves inherit some coat pattern traits from their mothers, and variation in some spot traits are correlated with neonatal survival. The skin underneath the blotches may serve as windows for thermoregulation, being sites for complex blood vessel systems and large sweat glands.
The skin of a giraffe is mostly gray, or tan. Its thickness allows the animal to run through thorn bushes without being punctured.:34 The fur may serve as a chemical defence, as its parasite repellents give the animal a characteristic scent. At least 11 main aromatic chemicals are in the fur, although indole and 3-methylindole are responsible for most of the smell. Because the males have a stronger odour than the females, the odour may also have sexual function. Along the animal's neck is a mane made of short, erect hairs. The one-metre (3.3-ft) tail ends in a long, dark tuft of hair and is used as a defense against insects.:36
Skull and ossicones
Both sexes have prominent horn-like structures called ossicones, which are formed from ossified cartilage, covered in skin and fused to the skull at the parietal bones. Being vascularized, the ossicones may have a role in thermoregulation, and are also used in combat between males. 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 end in knobs and tend to be bald on top. Also, a median lump, which is more prominent in males, emerges at the front of the skull. Males develop calcium deposits that form bumps on their skulls as they age. A giraffe's skull is lightened by multiple sinuses.:70 However, as males age, their skulls become heavier and more club-like, helping them become more dominant in combat. The upper jaw has a grooved palate and lacks front teeth.:26 The giraffe's molars have a rough surface.:27
Legs, locomotion and posture
The front and back legs of a giraffe are about the same length. The radius and ulna of the front legs are articulated by the carpus, which, while structurally equivalent to the human wrist, functions as a knee. It appears that a suspensory ligament allows the lanky legs to support the animal's great weight. The foot of the giraffe reaches a diameter of 30 cm (12 in), and the hoof is 15 cm (5.9 in) high in males and 10 cm (3.9 in) in females.:36 The rear of each hoof is low and the fetlock is close to the ground, allowing the foot to provide additional support to the animal's weight. Giraffes lack dewclaws and interdigital glands. The giraffe's pelvis, though relatively short, has an ilium that is outspread at the upper ends.
A giraffe has only two gaits: walking and galloping. Walking is done by moving the legs on one side of the body at the same time, then doing the same on the other side. When galloping, the hind legs move around the front legs before the latter move forward, and the tail will curl up. The animal relies on the forward and backward motions of its head and neck to maintain balance and the counter momentum while galloping.:327–29 The giraffe can reach a sprint speed of up to 60 km/h (37 mph), and can sustain 50 km/h (31 mph) for several kilometres.
A giraffe rests by lying with its body on top of its folded legs.: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 knees and spreads its hind legs to raise its hindquarters. It then straightens its front legs. With each step, the animal swings its head.:31 Studies in captivity find that 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 indicate paradoxical sleep. If the giraffe wants to bend down to drink, it either spreads its front legs or bends its knees. Giraffes would probably not be competent swimmers as their long legs would be highly cumbersome in the water, although they could possibly float. 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 or keep its head above the surface.
The giraffe has an extremely elongated neck, which can be up to 2–2.4 m (6.6–7.9 ft) in length, accounting for much of the animal's vertical height.:29 The long neck results from a disproportionate lengthening of the cervical vertebrae, not from the addition of more vertebrae. Each cervical vertebra is over 28 cm (11 in) long.:71 They comprise 52–54 per cent of the length of the giraffe's vertebral column, compared with the 27–33 percent typical of similar large ungulates, including the giraffe's closest living relative, the okapi. This elongation largely takes place after birth, perhaps because giraffe mothers would have a difficult time giving birth to young with the same neck proportions as adults. The giraffe's head and neck are held up by large muscles and a strengthened nuchal ligament, which are anchored by long dorsal spines on the anterior thoracic vertebrae, giving the animal a hump.
The giraffe's neck vertebrae have ball and socket joints.:71 In particular, the atlas–axis joint (C1 and C2) allows the animal to tilt its head vertically and reach more branches with the tongue.:29 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 most other ruminants where the articulation is between the seventh cervical vertebra (C7) and T1. 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. However, this proposition is not generally accepted, as T1 has other morphological features, such as an articulating rib, deemed diagnostic of thoracic vertebrae, and because exceptions to the mammalian limit of seven cervical vertebrae are generally characterised by increased neurological anomalies and maladies.
There are several hypotheses regarding the evolutionary origin and maintenance of elongation in giraffe necks. The "competing browsers hypothesis" was originally suggested by Charles Darwin and challenged only recently. It suggests that competitive pressure from smaller browsers, such as kudu, steenbok and impala, 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. 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. However, scientists disagree about just how much time giraffes spend feeding at levels beyond the reach of other browsers, 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.
Another theory, the sexual selection hypothesis, proposes that the long necks evolved as a secondary sexual characteristic, giving males an advantage in "necking" contests (see below) to establish dominance and obtain access to sexually receptive females. In support of this theory, necks are longer and heavier for males than females of the same age, and the former do not employ other forms of combat. However, one objection is that it fails to explain why female giraffes also have long necks. It has also been proposed that the neck serves to give the animal greater vigilance.
In mammals, the left recurrent 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; the left nerve is over 2 m (6 ft 7 in) long. Each nerve cell in this path begins in the brainstem and passes down the neck along the vagus 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. The structure of a giraffe's brain resembles that of domestic cattle.:31 It is kept cool by evaporative heat loss in the nasal passages. The shape of the skeleton gives the giraffe a small lung volume relative to its mass. Its long neck gives it a large amount of dead space, in spite of its narrow windpipe. These factors increase the resistance to airflow. Nevertheless, the animal can still supply enough oxygen to its tissues and it can increase its respiratory rate and oxygen diffusion when running.
The circulatory system of the giraffe has several adaptations for its great height. Its heart, which can weigh more than 11 kg (25 lb) and measures about 60 cm (2 ft) long, 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). Giraffes have unusually high heart rates for their size, at 150 beats per minute.:76 When the animal lowers its head the blood rushes down fairly unopposed and a rete mirabile in the upper neck, with its large cross sectional area, prevents excess blood flow to the brain. When it raises again, the blood vessels constrict and direct blood into the brain so the animal does not faint. The jugular veins contain several (most commonly seven) valves to prevent blood flowing back into the head from the inferior vena cava and right atrium while the head is lowered. 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.
Giraffes have oesophageal muscles that are unusually strong to allow regurgitation of food from the stomach up the neck and into the mouth for rumination.:78 They have four chambered stomachs, as in all ruminants, and the first chamber has adapted to their specialized diet. The intestines of an adult giraffe measure more than 70 m (230 ft) in length and have a relatively small ratio of small to large intestine. The liver of the giraffe is small and compact.:76 A gallbladder is generally present during fetal life, but it may disappear before birth.
Behaviour and ecology
Habitat and feeding
Giraffes usually inhabit savannahs and open woodlands. They prefer Acacieae, Commiphora, Combretum and open Terminalia woodlands over denser environments like Brachystegia woodlands.:322 The Angolan giraffe can be found in desert environments. Giraffes browse on the twigs of trees, preferring trees of the subfamily Acacieae and the genera Commiphora and Terminalia, which are important sources of calcium and protein to sustain the giraffe's growth rate. They also feed on shrubs, grass and fruit.:324 A giraffe eats around 34 kg (75 lb) of foliage daily. When stressed, giraffes may chew the bark off branches. Although herbivorous, the giraffe has been known to visit carcasses and lick dried meat off bones.:325
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. Mothers tend to feed in open areas, presumably to make it easier to detect predators, although this may reduce their feeding efficiency. As a ruminant, 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.:78–79 It is common for a giraffe to salivate while feeding.:27 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. The animal's faeces come in the form of small pellets. When it has access to water, a giraffe drinks at intervals no longer than three days.
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 trees that are too tall. 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.
Giraffes are usually found in groups that vary in size and composition according to ecological, anthropogenic, temporal, and social factors. Traditionally, the composition of these groups had been described as open and ever-changing. 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." More recent studies have found that giraffes have long-term social associations and may form groups or pairs based on kinship, sex or other factors, and these groups regularly associate with one another in larger communities or sub-communities within a fission–fusion society. Social relationships can be disrupted by proximity to humans. Masai giraffes of Tanzania live in distinct social subpopulations that overlap spatially, but have different reproductive rates and calf survival rates.
The number of giraffes in a group can range from 1 up to 66 individuals. Giraffe groups tend to be sex-segregated although mixed-sex groups made of adult females and young males also occur. Female groups may be matrilineally related. In general, females are more selective than males in who they associate with in regards to individuals of the same sex. Particularly stable giraffe groups are those made of mothers and their young, which can last weeks or months. 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. Giraffes are not territorial, but they have home ranges that vary according to rainfall and proximity to human settlements. Male giraffes occasionally wander far from areas that they normally frequent.:329
Although generally quiet and non-vocal, giraffes have been heard to communicate using snorts, sneezes, coughs, snores, hisses, bursts, moans, grunts and flute-like sounds. During courtship, males emit loud coughs. Females call their young by bellowing. Calves will emit snorts, bleats, mooing and mewing sounds. During nighttime, giraffes appear to hum to each other above the infrasound range for purposes which are unclear. Giraffes may also communicate visually with actions such as stamping, head tossing, and mane-waving.
Reproduction and parental care
Reproduction in giraffes is broadly polygamous: a few older males mate with the fertile females. Females can reproduce throughout the year and experience oestrus cycling approximately every 15 days. Female giraffes in oestrous 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. Males prefer young adult females over juveniles and older adults.
Male giraffes assess female fertility by tasting the female's urine to detect oestrus, in a multi-step process known as the flehmen response. Once an oestrous female is detected, the male will attempt to court her. When courting, dominant males will keep subordinate ones at bay. A courting male may lick a female's tail, rest 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.
Giraffe gestation lasts 400–460 days, after which a single calf is normally born, although twins occur on rare occasions. The mother gives birth standing up. The calf emerges head and front legs first, having broken through the fetal membranes, and falls to the ground, severing the umbilical cord. The mother then grooms the newborn and helps it stand up.:40 A newborn giraffe is 1.7–2 m (5.6–6.6 ft) tall. Within a few hours of birth, the calf can run around and is almost indistinguishable from a one-week-old. However, for the first 1–3 weeks, it spends most of its time hiding; its coat pattern providing camouflage. The ossicones, which have lain flat while it was in the womb, become erect within a few days.
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". Adult males play almost no role in raising the young,:337 although they appear to have friendly interactions. Calves are at risk of predation, and a mother giraffe will stand over her calf and kick at an approaching predator. Females watching calving pools will only alert their own young if they detect a disturbance, although the others will take notice and follow.
The length time in which offspring stay with their mother varies, though it can last until the female's next calving. Likewise, calves may suckle for only a month:335 or as long as a year. Females become sexually mature when they are four years old, while males become mature at four or five years. Spermatogenesis in male giraffes begins at three to four years of age. Males must wait until they are at least seven years old to gain the opportunity to mate.:40
Male giraffes use their necks as weapons in combat, a behaviour known as "necking". Necking is used to establish dominance and males that win necking bouts have greater reproductive success. This behaviour occurs at low or high intensity. In low intensity necking, the combatants rub and lean against each other. The male that can hold itself more erect 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 get ready to counter. The power of a blow depends on the weight of the skull and the arc of the swing. A necking duel can last more than half an hour, depending on how well matched the combatants are.:331 Although most fights do not lead to serious injury, there have been records of broken jaws, broken necks, and even deaths.
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. In one study, up to 94 percent of observed mounting incidents took place between males. The proportion of same-sex activities varied from 30 to 75 percent. Only one percent of same-sex mounting incidents occurred between females.
Mortality and health
Giraffes have high adult survival probability, and an unusually long lifespan compared to other ruminants, up to 38 years. Because of their size, eyesight and powerful kicks, adult giraffes are usually not subject to predation, although lions may regularly prey on individuals up to 550 kg (1,210 lb). Giraffes are the most common food source for the big cats in Kruger National Park, comprising nearly a third of the meat consumed, although only a small portion of the giraffes were probably killed by predators, as a majority of the consumed giraffes appeared to be scavenged. Nile crocodiles can also be a threat to giraffes when they bend down to drink. Adult female survival is significantly correlated with gregariousness, the average number of other females she is seen associating with. Calves are much more vulnerable than adults and are additionally preyed on by leopards, spotted hyenas and wild dogs. A quarter to a half of giraffe calves reach adulthood. Calf survival varies according to the season of birth, with calves born during the dry season having higher survival rates.
The local, seasonal presence of large herds of migratory wildebeests and zebras reduces predation pressure on giraffe calves and increases their survival probability. 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 glean information on predation risk from giraffe body language and spend less time scanning the environment when giraffes are present.
Some parasites feed on giraffes. They are often hosts for ticks, especially in the area around the genitals, which has thinner skin than other areas. Tick species that commonly feed on giraffes are those of genera Hyalomma, Amblyomma and Rhipicephalus. Giraffes may rely on red-billed and yellow-billed oxpeckers to clean them of ticks and alert them to danger. Giraffes host numerous species of internal parasite and are susceptible to various diseases. They were victims of the (now eradicated) viral illness rinderpest. Giraffes can also suffer from a skin disorder, which comes in the form of wrinkles, lesions or raw fissures. In Tanzania, it appears to be caused by a nematode, and may be further affected by secondary infections. As much as 79% of giraffes show signs of the disease in Ruaha National Park, but it did not cause mortality in Tarangire and is less prevalent in areas with fertile soils.
Relationship with humans
Humans have interacted with giraffes for millennia. They were depicted in art throughout the African continent, including that of the Kiffians, Egyptians, and Kushites.:45–47 The Kiffians 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".:45 How the giraffe got its height has been the subject of various African folktales. The Tugen people of modern Kenya used the giraffe to depict their god Mda. The Egyptians gave the giraffe its own hieroglyph, named 'sr' in Old Egyptian and 'mmy' in later periods.:49 They also kept giraffes as pets and shipped them around the Mediterranean.:48–49 The giraffe was also known to the Greeks and Romans, who believed that it was an unnatural hybrid of a camel and a leopard or a panther and called it camelopardalis.:50 The giraffe was among the many animals collected and displayed by the Romans. The first one in Rome was brought in by Julius Caesar in 46 BC and exhibited to the public.:52 With the fall of the Western Roman Empire, the housing of giraffes in Europe declined.:54 During the Middle Ages, giraffes were known to Europeans through contact with the Arabs, who revered the giraffe for its peculiar appearance.
Individual captive giraffes were given celebrity status throughout history. In 1414, a giraffe was shipped from Malindi to Bengal. It was then taken to China by explorer Zheng He and placed in a Ming dynasty zoo. The animal was a source of fascination for the Chinese people, who associated it with the mythical Qilin.:56 The Medici giraffe was a giraffe presented to Lorenzo de' Medici in 1486. It caused a great stir on its arrival in Florence. Zarafa, another famous giraffe, was brought from Egypt to Paris in the early 19th century as a gift from Muhammad Ali of Egypt to Charles X of France. A sensation, the giraffe was the subject of numerous memorabilia or "giraffanalia".:81
Giraffes continue to have a presence in modern culture. Salvador Dalí depicted them with burning manes in some of his surrealist paintings. Dali considered the giraffe to be a symbol of masculinity, and a flaming giraffe was meant to be a "masculine cosmic apocalyptic monster".:123 Several children's books feature the giraffe, including David A. Ufer's The Giraffe Who Was Afraid of Heights, Giles Andreae's Giraffes Can't Dance and Roald Dahl's The Giraffe and the Pelly and Me. Giraffes have appeared in animated films, as minor characters in Disney's The Lion King and Dumbo, and in more prominent roles in The Wild and in the Madagascar films. Sophie the Giraffe has been a popular teether since 1961. Another famous fictional giraffe is the Toys "R" Us mascot Geoffrey the Giraffe.:127 The giraffe has also been used for some scientific experiments and discoveries. Scientists have looked at the properties of giraffe skin when developing suits for astronauts and fighter pilots:76 because the people in these professions are in danger of passing out if blood rushes to their legs. Computer scientists have modeled the coat patterns of several subspecies using reaction–diffusion mechanisms.
The constellation of Camelopardalis, introduced in the seventeenth century, depicts a giraffe.:119–20 The Tswana people of Botswana traditionally see the constellation Crux as two giraffes – Acrux and Mimosa forming a male, and Gacrux and Delta Crucis forming the female.
Exploitation and conservation status
In 2010, giraffes were assessed as Least Concern from a conservation perspective by the International Union for Conservation of Nature (IUCN), but the 2016 assessment categorized giraffes as Vulnerable. In 1999, it was estimated that over 140,000 giraffes existed in the wild, but estimations as of 2016 indicate that there are approximately 97,500 members of Giraffa in the wild, down from 155,000 in 1985. The Masai and reticulated subspecies are endangered, and the Rothschild subspecies is near threatened. The Nubian subspecies is critically endangered. The primary causes for giraffe population declines are habitat loss and direct killing for bushmeat markets. Giraffes have been extirpated from much of their historic range including Eritrea, Guinea, Mauritania and Senegal. They may also have disappeared from Angola, Mali, and Nigeria, but have been introduced to Rwanda and Swaziland. As of 2010[update], there were more than 1,600 in captivity at Species360-registered zoos.
Protected areas such as national parks provide important habitat and anti-poaching protection to giraffe populations. Community-based conservation efforts outside national parks are also effective at protecting giraffes and their habitats. Private game reserves have contributed to the preservation of giraffe populations in southern Africa. The giraffe is a protected species in most of its range. It is the national animal of Tanzania, and is protected by law, and unauthorised killing can result in imprisonment. The UN backed Convention of Migratory Species selected giraffes for protection in 2017. In 2019, giraffes were listed under Appendix 2 of the Conference of the Parties to the UN Convention on International Trade in Endangered Species (CITES), which will now document international trade in giraffe parts, such as hides, bones and meat. 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. 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.
Giraffes were probably common targets for hunters throughout Africa. Different parts of their bodies were used for different purposes. Their meat was used for food. The tail hairs served as flyswatters, bracelets, necklaces, and thread. Shields, sandals, and drums were made using the skin, and the strings of musical instruments were from the tendons. The smoke from burning giraffe skins was used by the medicine men of Buganda to treat nose bleeds. The Humr people of Kordofan consume the drink Umm Nyolokh, which is prepared from the liver and bone marrow of giraffes. Richard Rudgley hypothesised that Umm Nyolokh might contain DMT. The drink is said to cause hallucinations of giraffes, believed to be the giraffes' ghosts, by the Humr. In the 19th century, European explorers began to hunt them for sport. Habitat destruction has hurt the giraffe. In the Sahel, the need for firewood and grazing room for livestock has led to deforestation. Normally, giraffes can coexist with livestock, since they do not directly compete with them. In 2017, severe droughts in northern Kenya have led to increased tensions over land and the killing of wildlife by herders, with giraffe populations being particularly hit.
- Muller, Z.; Bercovitch, F.; Brand, R.; Brown, D.; Brown, M.; Bolger, D.; Carter, K.; Deacon, F.; Doherty, J. B.; Fennessy, J.; Fennessy, S.; Hussein, A.A.; Lee, D.; Marais, A.; Strauss, M.; Tutchings, A. & Wube, T. (2016). "Giraffa camelopardalis". IUCN Red List of Threatened Species. 2016: e.T9194A136266699. doi:10.2305/IUCN.UK.2016-3.RLTS.T9194A136266699.en.
- "Giraffe". Online Etymology Dictionary. Retrieved 1 November 2011.
- Peust, C. (2009). "Some Cushitic Etymologies". In Dolgopolʹskiĭ, A.; Takács, G.; Jungraithmayr, H. (eds.). Semito-Hamitic Festschrift for A.B. Dolgopolsky and H. Jungraithmayr. Reimer. pp. 257–60. ISBN 978-3-496-02810-9.
- Kingdon, J. (1997). The Kingdon Field Guide to African Mammals. Academic Press. pp. 339–44. ISBN 978-0-12-408355-4.
- Őrsi, Tibor (2006). French Linguistic Influence in the Cotton Version of Mandeville's Travels. Tinta Könyvkiadó. p. 113. ISBN 9789637094545.
- "Definition of CAMELOPARD". m-w.com. Encyclopædia Britannica: Merriam-Webster. Retrieved 3 September 2014.
- "Definition of camelopard". Dictionary of Medieval Terms and Phrases. Retrieved 3 September 2014.
- Dagg, A. I. (1971). "Giraffa camelopardalis" (PDF). Mammalian Species. 5 (5): 1–8. doi:10.2307/3503830. JSTOR 3503830.
- "camelopardalis". A Latin Dictionary, Perseus Digital Library. Retrieved 23 November 2011.
- Hassanin, A.; Douzery, E. J. P. (2003). "Molecular and morphological phylogenies of Ruminantia and the alternative position of the Moschidae". Systematic Biology. 52 (2): 206–28. doi:10.1080/10635150390192726. PMID 12746147.
- Mitchell, G.; Skinner, J. D. (2003). "On the origin, evolution and phylogeny of giraffes Giraffa camelopardalis". Transactions of the Royal Society of South Africa. 58 (1): 51–73. doi:10.1080/00359190309519935. S2CID 6522531.
- Danowitz, M.; Vasilyev, A.; Kortlandt, V.; Solounias, V. (2015). "Fossil evidence and stages of elongation of the Giraffa camelopardalis neck". Royal Society Open Science. 2 (10): 150393. Bibcode:2015RSOS....250393D. doi:10.1098/rsos.150393. PMC 4632521. PMID 26587249.
- Danowitz, M.; Domalski, R.; Solounias, N. (2015). "The cervical anatomy of Samotherium, an intermediate-necked giraffid". Royal Society Open Science. 2 (11): 150521. Bibcode:2015RSOS....250521D. doi:10.1098/rsos.150521. PMC 4680625. PMID 26716010.
- "Giraffa (giraffe)". The Paleobiology Database. Retrieved 13 September 2016.
- Simmons, R. E.; Scheepers, L. (1996). "Winning by a Neck: Sexual Selection in the Evolution of Giraffe" (PDF). The American Naturalist. 148 (5): 771–86. doi:10.1086/285955. S2CID 84406669. Archived from the original (PDF) on 23 August 2004.
- Janis, Christine M. (1 January 1993). "Tertiary Mammal Evolution in the Context of Changing Climates, Vegetation, and Tectonic Events". Annual Review of Ecology and Systematics. 24: 467–500. doi:10.1146/annurev.ecolsys.24.1.467. JSTOR 2097187.
- Ramstein, Gilles; Fluteau, Frédéric; Besse, Jean; Joussaume, Sylvie (24 April 1997). "Effect of orogeny, plate motion and land–sea distribution on Eurasian climate change over the past 30 million years". Nature. 386 (6627): 788–795. Bibcode:1997Natur.386..788R. doi:10.1038/386788a0. S2CID 4335003.
- Badlangana, L. N.; Adams, J. W.; Manger, P. R. (2009). "The giraffe (Giraffa camelopardalis) cervical vertebral column: A heuristic example in understanding evolutionary processes?". Zoological Journal of the Linnean Society. 155 (3): 736–57. doi:10.1111/j.1096-3642.2008.00458.x.
- Agaba, M.; Ishengoma, E.; Miller, W. C.; McGrath, B. C.; Hudson, C. N.; Bedoya, R. O. C.; Ratan, A.; Burhans, R.; Chikhi, R.; Medvedev, P.; Praul C. A.; Wu-Cavener, L.; Wood, B.; Robertson, H.; Penfold, L.; Cavener, D. R. (2016). "Giraffe genome sequence reveals clues to its unique morphology and physiology". Nature Communications. 7: 11519. Bibcode:2016NatCo...711519A. doi:10.1038/ncomms11519. PMC 4873664. PMID 27187213.
- Brown, D. M.; Brenneman R. A.; Koepfli, K.-P.; Pollinger, J. P.; Milá, B.; Georgiadis, N. J.; Louis Jr., E. E.; Grether, G. F.; Jacobs, D. K.; Wayne R. K. (2007). "Extensive population genetic structure in the giraffe". BMC Biology. 5 (1): 57. doi:10.1186/1741-7007-5-57. PMC 2254591. PMID 18154651.
- Linnaeus, Carl (1758). Systema Naturæ.
- Muller, Zoe; et al. (2016). "Giraffa camelopardalis (Giraffe)". IUCN Red List of Threatened Species. 2016. Retrieved 2 May 2017.old-form url
- Gippoliti, S. (2018). "Impacts of taxonomic inertia for the conservation of African ungulate diversity: an overview". Biological Reviews. 93 (1): 115–130. doi:10.1111/brv.12335. PMID 28429851. S2CID 5189968.
- Russell, Seymour (2001). Patterns of subspecies diversity in the giraffe, Giraffa camelopardalis (L. 1758) : comparison of systematic methods and their implications for conservation policy. PHD Thesis, University of Kent at Canterbury (PhD). University of Kent at Canterbury.
- Groves, Colin; Grubb, Peter (1 November 2011). Ungulate Taxonomy. JHU Press. ISBN 9781421400938.
- Fennessy, Julian; Bidon, Tobias; Reuss, Friederike; Kumar, Vikas; Elkan, Paul; Nilsson, Maria A.; Vamberger, Melita; Fritz, Uwe; Janke, Axel (2016). "Multi-locus Analyses reveal four giraffe species instead of one". Current Biology. 26 (18): 2543–2549. doi:10.1016/j.cub.2016.07.036. PMID 27618261. S2CID 3991170.
- Bercovitch, Fred B.; Berry, Philip S.M.; Dagg, Anne; Deacon, Francois; Doherty, John B.; Lee, Derek E.; Mineur, Frédéric; Muller, Zoe; Ogden, Rob; Seymour, Russell; Shorrocks, Bryan; Tutchings, Andy (2017). "How many species of giraffe are there?". Current Biology. 27 (4): R136–R137. doi:10.1016/j.cub.2016.12.039. ISSN 0960-9822. PMID 28222287. S2CID 3356310.
- Petzold, Alice; Hassanin, Alexandre (13 February 2020). "A comparative approach for species delimitation based on multiple methods of multi-locus DNA sequence analysis: A case study of the genus Giraffa (Mammalia, Cetartiodactyla)". PLOS ONE. 15 (2): e0217956. Bibcode:2020PLoSO..1517956P. doi:10.1371/journal.pone.0217956. ISSN 1932-6203. PMC 7018015. PMID 32053589.
- Coimbra, Raphael T.F.; Winter, Sven; Kumar, Vikas; Koepfli, Klaus-Peter; Gooley, Rebecca M.; Dobrynin, Pavel; Fennessy, Julian; Janke, Axel (2021). "Whole-genome analysis of giraffe supports four distinct species". Current Biology. doi:10.1016/j.cub.2021.04.033. ISSN 0960-9822. PMID 33957077.
- Swainson 1835. Camelopardalis antiquorum. Bagger el Homer, Kordofan, about 10° N, 28° E (as fixed by Harper, 1940)
- Hassanin, A.; Ropiquet, A.; Gourmand, B.-L.; Chardonnet, B.; Rigoulet, J. (2007). "Mitochondrial DNA variability in Giraffa camelopardalis: consequences for taxonomy, phylogeography and conservation of giraffes in West and central Africa". Comptes Rendus Biologies. 330 (3): 173–83. doi:10.1016/j.crvi.2007.02.008. PMID 17434121.
- Seymour, R. (2002) The taxonomic status of the giraffe, Giraffa camelopardalis (L. 1758), PH.D Thesis
- "Giraffa". ISIS. 2010. Archived from the original on 6 July 2010. Retrieved 4 November 2010.
- "Giraffe – The Facts: Current giraffe status?". Giraffe Conservation Foundation. Archived from the original on 19 March 2016. Retrieved 21 December 2010.
- "Exhibits". Al Ain Zoo. 25 February 2003. Archived from the original on 29 November 2011. Retrieved 21 November 2011.
- "Nubian giraffe born in Al Ain zoo". UAE Interact. Archived from the original on 20 March 2012. Retrieved 21 December 2010.
- Fennessy, S.; Fennessy, J.; Muller, Z.; Brown, M. & Marais, A. (2018). "Giraffa camelopardalis ssp. rothschildi". IUCN Red List of Threatened Species. 2018. Retrieved 25 August 2019.old-form url
- Fennessy, J.; Marais, A.; Tutchings, A. (2018). "Giraffa camelopardalis ssp. peralta". IUCN Red List of Threatened Species. 2018: e.T136913A51140803.
- Thomas 1908. Giraffa camelopardalis peralta Lokojya, Niger– Benue junction, Nigeria.
- Kingdon, J. (1988). East African Mammals: An Atlas of Evolution in Africa, Volume 3, Part B: Large Mammals. University of Chicago Press. pp. 313–37. ISBN 978-0-226-43722-4.
- Winton, W. E. de (1 November 1899). "XXXVIII.—On mammals collected by Lieut.-Colonel W. Giffard in the northern territory of the Gold Coast". Annals and Magazine of Natural History. 4 (23): 353–359. doi:10.1080/00222939908678212. ISSN 0374-5481.
- Brenneman, R. A.; Louis, E. E. Jr; Fennessy, J. (2009). "Genetic structure of two populations of the Namibian giraffe, Giraffa camelopardalis angolensis". African Journal of Ecology. 47 (4): 720–28. doi:10.1111/j.1365-2028.2009.01078.x.
- Rookmaaker, L. C. (1 June 1989). The Zoological Exploration of Southern Africa 1650–1790. CRC Press. ISBN 9789061918677.
- Matschie 1898 Giraffa tippelskirchi. Lake Eyasi, NW Tanzania.
- Estes, R. (1992). The Behavior Guide to African Mammals: including Hoofed Mammals, Carnivores, Primates. University of California Press. pp. 202–07. ISBN 978-0-520-08085-0.
- Pellow, R. A. (2001). "Giraffe and Okapi". In MacDonald, D. (ed.). The Encyclopedia of Mammals (2nd ed.). Oxford University Press. pp. 520–27. ISBN 978-0-7607-1969-5.
- Nowak, R.M. (1999). Giraffe Pages 1086–1089 in Walker's Mammals of the World. Volume 1. The Johns Hopkins University Press, Baltimore, USA and London, UK.
- Maisano, Sarah. "Giraffa camelopardalis giraffe". Animaldiversity.ummz.umich.edu.
- Owen-Smith, R.N. 1988. Megaherbivores: The Influence of Very Large Body Size on Ecology. Cambridge: Cambridge University Press.
- Dagg, A.I. and J. B. Foster (1976/1982): The Giraffe. Its Biology, Behavior, and Ecology. Krieger Publishing Company, Malabar, Florida (Reprint 1982 with updated supplementary material.)
- Skinner, J. D.; Smithers, R. H. M. (1990). The mammals of the southern African subregion. University of Pretoria. pp. 616–20. ISBN 978-0-521-84418-5.
- Swaby, S. (2010). "Giraffe". In Harris, T. (ed.). Mammal Anatomy: An Illustrated Guide. Marshall Cavendish. pp. 64–84. ISBN 978-0-7614-7882-9.
- Williams, E. (2011). Giraffe. Reaktion Books. ISBN 978-1-86189-764-0.
- Prothero, D. R.; Schoch, R. M. (2003). Horns, Tusks, and Flippers: The Evolution of Hoofed Mammals. Johns Hopkins University Press. pp. 67–72. ISBN 978-0-8018-7135-1.
- Lee, Derek E.; Cavener, Douglas R.; Bond, Monica L. (2 October 2018). "Seeing spots: quantifying mother-offspring similarity and assessing fitness consequences of coat pattern traits in a wild population of giraffes (Giraffa camelopardalis)". PeerJ. 6: e5690. doi:10.7717/peerj.5690. ISSN 2167-8359. PMC 6173159. PMID 30310743.
- Mitchell, G.; Skinner, J.D. (2004). "Giraffe thermoregulation: a review". Transactions of the Royal Society of South Africa. 59 (2): 49–57. doi:10.1080/00359190409519170. ISSN 0035-919X. S2CID 87321176.
- Langley, Liz (4 March 2017). "Do Zebras Have Stripes On Their Skin?". National Geographic. Retrieved 2 June 2020.
- Wood, W. F.; Weldon, P. J. (2002). "The scent of the reticulated giraffe (Giraffa camelopardalis reticulata)". Biochemical Systematics and Ecology. 30 (10): 913–17. doi:10.1016/S0305-1978(02)00037-6.
- Aridi, Rasha. "Scientists Report First Instances of Dwarf Giraffes". Smithsonian Magazine. Retrieved 10 January 2021.
- Reuters (9 January 2021). "Scientists surprised to discover two dwarf giraffes in Namibia, Uganda". The Hindu. ISSN 0971-751X. Retrieved 10 January 2021.
- Simmons, R. E.; Altwegg, R. (2010). "Necks-for-sex or competing browsers? A critique of ideas on the evolution of giraffe". Journal of Zoology. 282 (1): 6–12. doi:10.1111/j.1469-7998.2010.00711.x.
- MacClintock, D.; Mochi, U. (1973). A natural history of giraffes. Scribner. p. 30. ISBN 978-0-684-13239-6.
- Wood, C. (7 March 2014). "Groovy giraffes…distinct bone structures keep these animals upright". Society for Experimental Biology. Retrieved 7 May 2014.
- Garland, T; Janis, C. M. (1993). "Does metatarsal/femur ratio predict maximal running speed in cursorial mammals?" (PDF). Journal of Zoology. 229 (1): 133–51. doi:10.1111/j.1469-7998.1993.tb02626.x.
- Rafferty, John. P (2011). Grazers (Britannica Guide to Predators and Prey). Britannica Educational Publishing. p. 194. ISBN 978-1-61530-336-6.
- Tobler, I.; Schwierin, B. (1996). "Behavioural sleep in the giraffe (Giraffa camelopardalis) in a zoological garden". Journal of Sleep Research. 5 (1): 21–32. doi:10.1046/j.1365-2869.1996.00010.x. PMID 8795798. S2CID 34605791.
- Henderson, D. M.; Naish, D. (2010). "Predicting the buoyancy, equilibrium and potential swimming ability of giraffes by computational analysis". Journal of Theoretical Biology. 265 (2): 151–59. doi:10.1016/j.jtbi.2010.04.007. PMID 20385144.
- Naish, D. (January 2011). "Will it Float?". Scientific American. 304 (1): 22. Bibcode:2011SciAm.304a..22N. doi:10.1038/scientificamerican0111-22. ISSN 0036-8733.
- Taylor, M. P.; Wedel, M. J. (2013). "Why sauropods had long necks; and why giraffes have short necks". PeerJ. 1: e36. doi:10.7717/peerj.36. PMC 3628838. PMID 23638372.
- Van Sittert, S. J.; Skinner, J. D.; Mitchell, G. (2010). "From fetus to adult – An allometric analysis of the giraffe vertebral column". Journal of Experimental Zoology Part B: Molecular and Developmental Evolution. 314B (6): 469–79. doi:10.1002/jez.b.21353. PMID 20700891.
- Solounias, N. (1999). "The remarkable anatomy of the giraffe's neck" (PDF). Journal of Zoology. 247 (2): 257–68. doi:10.1111/j.1469-7998.1999.tb00989.x.
- du Toit, J. T. (1990). "Feeding-height stratification among African browsing ruminants" (PDF). African Journal of Ecology. 28 (1): 55–62. doi:10.1111/j.1365-2028.1990.tb01136.x. Archived from the original (PDF) on 10 November 2011. Retrieved 21 November 2011.
- Cameron, E. Z.; du Toit, J. T. (2007). "Winning by a Neck: Tall Giraffes Avoid Competing with Shorter Browsers". American Naturalist. 169 (1): 130–35. doi:10.1086/509940. PMID 17206591. S2CID 52838493.
- Woolnough, A. P.; du Toit, J. T. (2001). "Vertical zonation of browse quality in tree canopies exposed to a size-structured guild of African browsing ungulates" (PDF). Oecologia. 129 (1): 585–90. Bibcode:2001Oecol.129..585W. doi:10.1007/s004420100771. PMID 24577699. S2CID 18821024. Archived from the original (PDF) on 10 November 2011. Retrieved 7 March 2012.
- Young, T. P.; Isbell, L. A. (1991). "Sex differences in giraffe feeding ecology: energetic and social constraints" (PDF). Ethology. 87 (1–2): 79–89. doi:10.1111/j.1439-0310.1991.tb01190.x. Archived from the original (PDF) on 16 May 2013. Retrieved 2 February 2012.
- Mitchell, G.; van Sittert, S.; Skinner, J. D. (2010). "The demography of giraffe deaths in a drought". Transactions of the Royal Society of South Africa. 65 (3): 165–68. doi:10.1080/0035919X.2010.509153. hdl:2263/18957. S2CID 83652889.
- Mitchell, G.; van Sittert, S. J.; Skinner, J. D. (2009). "Sexual selection is not the origin of long necks in giraffes". Journal of Zoology. 278 (4): 281–86. doi:10.1111/j.1469-7998.2009.00573.x.
- Brownlee, A. (1963). "Evolution of the Giraffe". Nature. 200 (4910): 1022. Bibcode:1963Natur.200.1022B. doi:10.1038/2001022a0. S2CID 4145785.
- Williams, Edgar M. (2016). "Giraffe Stature and Neck Elongation: Vigilance as an Evolutionary Mechanism". Biology. 5 (3): 35. doi:10.3390/biology5030035. PMC 5037354. PMID 27626454.
- Wedel, M. J. (2012). "A monument of inefficiency: the presumed course of the recurrent laryngeal nerve in sauropod dinosaurs" (PDF). Acta Palaeontologica Polonica. 57 (2): 251–56. doi:10.4202/app.2011.0019. S2CID 43447891.
- Harrison, D. F. N. (1995). The Anatomy and Physiology of the Mammalian Larynx. Cambridge University Press. p. 165. ISBN 978-0-521-45321-9.
- Skinner, J. D.; Mitchell, G. (2011). "Lung volumes in giraffes, Giraffa camelopardalis" (PDF). Comparative Biochemistry and Physiology A. 158 (1): 72–78. doi:10.1016/j.cbpa.2010.09.003. hdl:2263/16472. PMID 20837156.
- Mitchell, G.; Skinner, J. D. (1993). "How giraffe adapt to their extraordinary shape". Transactions of the Royal Society of South Africa. 48 (2): 207–18. doi:10.1080/00359199309520271.
- Mitchell, G.; van Sittert, S. J.; Skinner, J. D. (2009). "The structure and function of giraffe jugular vein valves" (PDF). South African Journal of Wildlife Research. 39 (2): 175–80. doi:10.3957/056.039.0210. hdl:2263/13994. S2CID 55201969.
- Pérez, W.; Lima, M.; Clauss, M. (2009). "Gross anatomy of the intestine in the giraffe (Giraffa camelopardalis)" (PDF). Anatomia, Histologia, Embryologia. 38 (6): 432–35. doi:10.1111/j.1439-0264.2009.00965.x. PMID 19681830. S2CID 28390695.
- Cave, A. J. E. (1950). "On the liver and gall-bladder of the Giraffe". Proceedings of the Zoological Society of London. 120 (2): 381–93. doi:10.1111/j.1096-3642.1950.tb00956.x.
- Oldham-Ott, Carla K.; Gilloteaux, Jacques (1997). "Comparative morphology of the gallbladder and biliary tract in vertebrates: variation in structure, homology in function and gallstones". Microscopy Research and Technique. 38 (6): 571–79. doi:10.1002/(SICI)1097-0029(19970915)38:6<571::AID-JEMT3>3.0.CO;2-I. PMID 9330347.
- Fennessy, J. (2004). Ecology of desert-dwelling giraffe Giraffa camelopardalis angolensis in northwestern Namibia (PhD thesis). University of Sydney.
- Bond, Monica L.; Lee, Derek E.; Ozgul, Arpat; König, Barbara (27 August 2019). "Fission–fusion dynamics of a megaherbivore are driven by ecological, anthropogenic, temporal, and social factors". Oecologia. 191 (2): 335–347. Bibcode:2019Oecol.191..335B. doi:10.1007/s00442-019-04485-y. ISSN 1432-1939. PMID 31451928. S2CID 201732871.
- van der Jeugd, H. P.; Prins, H. H. T. (2000). "Movements and group structure of giraffe (Giraffa camelopardalis) in Lake Manyara National Park, Tanzania" (PDF). Journal of Zoology. 251 (1): 15–21. doi:10.1111/j.1469-7998.2000.tb00588.x. Archived from the original (PDF) on 6 November 2013.
- Pratt D. M.; Anderson V. H. (1985). "Giraffe social behavior". Journal of Natural History. 19 (4): 771–81. doi:10.1080/00222938500770471.
- Bond, Monica L.; König, Barbara; Lee, Derek E.; Ozgul, Arpat; Farine, Damien R. (2020). "Proximity to humans affects local social structure in a giraffe metapopulation". Journal of Animal Ecology. 90 (1): 212–221. doi:10.1111/1365-2656.13247. ISSN 1365-2656. PMID 32515083.
- Bercovitch, F. B.; Berry, P. S. M. (2013). "Herd composition, kinship and fission–fusion social dynamics among wild giraffe". African Journal of Ecology. 51 (2): 206–216. doi:10.1111/aje.12024.
- Carter, K. D.; Seddon, J. M.; Frèreb, C. H.; Carter, J. K. (2013). "Fission–fusion dynamics in wild giraffes may be driven by kinship, spatial overlap and individual social preferences". Animal Behaviour. 85 (2): 385–394. doi:10.1016/j.anbehav.2012.11.011. S2CID 53176817.
- VanderWaal, K. L.; Wang, H.; McCowan, B.; Fushing, H.; Isbell, L. A. (2014). "Multilevel social organization and space use in reticulated giraffe (Giraffa camelopardalis)". Behavioral Ecology. 25 (1): 17–26. doi:10.1093/beheco/art061.
- Bond, Monica L.; König, Barbara; Ozgul, Arpat; Farine, Damien R.; Lee, Derek E. (2021). "Socially Defined Subpopulations Reveal Demographic Variation in a Giraffe Metapopulation". The Journal of Wildlife Management. n/a (n/a). doi:10.1002/jwmg.22044. ISSN 1937-2817.
- Leuthold, B. M. (1979). "Social organization and behaviour of giraffe in Tsavo East National Park". African Journal of Ecology. 17 (1): 19–34. doi:10.1111/j.1365-2028.1979.tb00453.x.
- Knüsel, Mara; Lee, Derek; König, Barbara; Bond, Monica (March 2019). "Correlates of home range sizes of giraffes, Giraffa camelopardalis" (PDF). Animal Behaviour. 149: 143–151. doi:10.1016/j.anbehav.2019.01.017. S2CID 72332291.
- Baotic, A.; Sicks, F.; Stoeger, A. S. (2015). "Nocturnal "humming" vocalizations: adding a piece to the puzzle of giraffe vocal communication". BMC Research Notes. 8: 425. doi:10.1186/s13104-015-1394-3. PMC 4565008. PMID 26353836.
- del Castillo, Susan M.; Bashaw, Meredith J.; Patton, Marilyn L.; Rieches, Randy R.; Bercovitch, Fred B. (1 May 2005). "Fecal steroid analysis of female giraffe (Giraffa camelopardalis) reproductive condition and the impact of endocrine status on daily time budgets". General and Comparative Endocrinology. 141 (3): 271–281. doi:10.1016/j.ygcen.2005.01.011. ISSN 0016-6480. PMID 15804514.
- Bercovitch, Fred B.; Bashaw, Meredith J.; del Castillo, Susan M. (1 August 2006). "Sociosexual behavior, male mating tactics, and the reproductive cycle of giraffe Giraffa camelopardalis". Hormones and Behavior. 50 (2): 314–321. doi:10.1016/j.yhbeh.2006.04.004. ISSN 0018-506X. PMID 16765955. S2CID 45843281.
- Seeber, Peter A.; Duncan, Patrick; Fritz, Hervé; Ganswindt, André (23 October 2013). "Androgen changes and flexible rutting behaviour in male giraffes". Biology Letters. 9 (5): 20130396. doi:10.1098/rsbl.2013.0396. PMC 3971675. PMID 23925833.
- Langman, V. A. (1977). "Cow-calf relationships in giraffe (Giraffa camelopardalis giraffa)". Zeitschrift für Tierpsychologie. 43 (3): 264–86. doi:10.1111/j.1439-0310.1977.tb00074.x.
- Hall-Martin A. J.; Skinner J. D.; Hopkins B. J. (1978). "The development of the reproductive organs of the male giraffe, Giraffa camelopardalis" (PDF). Journal of Reproduction and Fertility. 52 (1): 1–7. doi:10.1530/jrf.0.0520001. PMID 621681.
- Coe, M. J. (1967). "'Necking' behavior in the giraffe". Journal of Zoology. 151 (2): 313–21. doi:10.1111/j.1469-7998.1967.tb02117.x.
- Bagemihl, B. (1999). Biological Exuberance: Animal Homosexuality and Natural Diversity. St. Martin's Press. pp. 391–93. ISBN 978-0-312-19239-6.
- Lee, D. E.; Strauss, M. K. L (1 January 2016). Reference Module in Earth Systems and Environmental Sciences. Elsevier. doi:10.1016/B978-0-12-409548-9.09721-9. ISBN 9780124095489.
- Marealle, Wilfred N; Eggen, Grethe Stavik; Røskaft, Eivin (26 April 2020). "Faecal Glucocorticoids Metabolite Response in Giraffes (Giraffa camelopardalis tippelskirchi) in Relation to Protected Area Management Objectives in Tanzania". East African Journal of Forestry and Agroforestry. 2 (1): 47–58. doi:10.37284/eajfa.2.1.142. ISSN 2707-4323.
- Müller, DW; Zerbe, P; Codron, D; Clauss, M; Hatt, JM (2011). "A long life among ruminants: giraffids and other special cases". Schweizer Archiv für Tierheilkunde. 153 (11): 515–519. doi:10.1024/0036-7281/a000263. PMID 22045457. S2CID 10687135.
- Hayward, Matt W.; Kerley, Graham (2005). "Prey preferences of the lion (Panthera leo)" (PDF). Journal of Zoology. 267 (3): 309–22. CiteSeerX 10.1.1.611.8271. doi:10.1017/S0952836905007508.
- Pienaar, U. de V. (1969). "Predator–prey relationships among the larger mammals of the Kruger National Park". Koedoe. 12 (1). doi:10.4102/koedoe.v12i1.753.
- Owen-Smith, N.; Mills, M. G. (2008). "Predator-prey size relationships in an African large-mammal food web" (PDF). Journal of Animal Ecology. 77 (1): 173–83. doi:10.1111/j.1365-2656.2007.01314.x. hdl:2263/9023. PMID 18177336.
- Bond, M. L.; Lee, D. E.; Farine, D. R.; Ozgul, A.; König, B. (10 February 2021). "Sociability increases survival of adult female giraffes". Proceedings of the Royal Society B: Biological Sciences. 288 (1944): 20202770. doi:10.1098/rspb.2020.2770. PMC 7893237. PMID 33563118.
- Lee, Derek E.; Bond, Monica L.; Kissui, Bernard M.; Kiwango, Yustina A.; Bolger, Douglas T. (11 May 2016). "Spatial variation in giraffe demography: a test of 2 paradigms". Journal of Mammalogy. 97 (4): 1015–1025. doi:10.1093/jmammal/gyw086. ISSN 0022-2372. S2CID 87117946.
- Lee, Derek Edward; Bond, Monica Louise; Bolger, Douglas Thomas (1 January 2017). "Season of birth affects juvenile survival of giraffe". Population Ecology. 59 (1): 45–54. doi:10.1007/s10144-017-0571-8. ISSN 1438-3896. S2CID 7611046.
- Lee, Derek E.; Kissui, Bernard M.; Kiwango, Yustina A.; Bond, Monica L. (2016). "Migratory herds of wildebeests and zebras indirectly affect calf survival of giraffes". Ecology and Evolution. 6 (23): 8402–8411. doi:10.1002/ece3.2561. ISSN 2045-7758. PMC 5167056. PMID 28031792.
- Schmitt, M. H.; Stears, K.; Shrader, A. M. (2016). "Zebra reduce predation risk in mixed-species herds by eavesdropping on cues from giraffe". Behavioral Ecology. 27 (4): 1073–1077. doi:10.1093/beheco/arw015.
- Lee, Derek E.; Bond, Monica L. (26 July 2016). "The Occurrence and Prevalence of Giraffe Skin Disease in Protected Areas of Northern Tanzania". Journal of Wildlife Diseases. 52 (3): 753–755. doi:10.7589/2015-09-247. PMID 27310168. S2CID 10736316.
- Bond, Monica L.; Strauss, Megan K. L.; Lee, Derek E. (16 August 2016). "Soil Correlates and Mortality from Giraffe Skin Disease in Tanzania". Journal of Wildlife Diseases. 52 (4): 953–958. doi:10.7589/2016-02-047. ISSN 0090-3558. PMID 27529292. S2CID 46776142.
- Muneza, Arthur B.; Montgomery, Robert A.; Fennessy, Julian T.; Dickman, Amy J.; Roloff, Gary J.; Macdonald, David W. (1 June 2016). "Regional variation of the manifestation, prevalence, and severity of giraffe skin disease: A review of an emerging disease in wild and captive giraffe populations". Biological Conservation. 198: 145–156. doi:10.1016/j.biocon.2016.04.014.
- "The Dabous Giraffe rock art petrograph". The Bradshaw Foundation. Retrieved 6 November 2011.
- Shorrocks, B. (2016). The Giraffe: Biology, Ecology, Evolution and Behaviour. Wiley. p. 3. ISBN 9781118587478.
- Ringmar, E. (2006). "Audience for a Giraffe: European Expansionism and the Quest for the Exotic" (PDF). Journal of World History. 17 (4): 353–97. doi:10.1353/jwh.2006.0060. JSTOR 20079397. S2CID 143808549.
- Walter, M.; Fournier, A.; Menevaux, D. (2001). Integrating shape and pattern in mammalian models in SIGGRAPH '01 (PDF). Proceedings of the 28th Annual Conference on Computer Graphics and Interactive Techniques. pp. 317–26. CiteSeerX 10.1.1.10.7622. doi:10.1145/383259.383294. ISBN 978-1-58113-374-5. S2CID 13488215. Archived from the original on 23 September 2015. Retrieved 16 November 2011.
- Clegg, A. (1986). "Some Aspects of Tswana Cosmology". Botswana Notes and Records. 18: 33–37. JSTOR 40979758.
- Matt McGrath (8 December 2016). "Giraffes facing 'silent extinction' as population plunges". BBC News. Retrieved 8 December 2016.
- "New bird species and giraffe under threat – IUCN Red List".
- Bolger, D.; Ogutu, J.; Strauss, M.; Lee, D.; Muneza, A.; Fennessy, J.; Brown, D. (2019). "Masai Giraffe". IUCN Red List of Threatened Species. 2019. doi:10.2305/IUCN.UK.2019-1.RLTS.T88421036A88421121.en.
- Muneza, A.; Doherty, J. B.; Hussein Ali, A.; Fennessy, J.; Marais, A.; O'Connor, D.; Wube, T. (2018). "Reticulated Giraffe". IUCN Red List of Threatened Species. 2018. doi:10.2305/IUCN.UK.2018-2.RLTS.T88420717A88420720.en.
- Wube, T.; Doherty, J. B.; Fennessy, J.; Marais, A. (2018). "Giraffa camelopardalis ssp. camelopardalis". IUCN Red List of Threatened Species. 2018. doi:10.2305/IUCN.UK.2018-2.RLTS.T88420707A88420710.en.
- Lee, Derek E. (2018). "Evaluating conservation effectiveness in a Tanzanian community wildlife management area". The Journal of Wildlife Management. 82 (8): 1767–1774. doi:10.1002/jwmg.21549. ISSN 1937-2817.
- Lee, Derek E; Bond, Monica L (26 February 2018). "Quantifying the ecological success of a community-based wildlife conservation area in Tanzania". Journal of Mammalogy. 99 (2): 459–464. doi:10.1093/jmammal/gyy014. ISSN 0022-2372. PMC 5965405. PMID 29867255.
- Knappert, J (1987). East Africa: Kenya, Tanzania & Uganda. Vikas Publishing House. p. 57. ISBN 978-0-7069-2822-8.
- Charles Foley; Lara Foley; Alex Lobora; Daniela De Luca; Maurus Msuha; Tim R. B. Davenport; Sarah M. Durant (8 June 2014). A Field Guide to the Larger Mammals of Tanzania. Princeton University Press. pp. 179–. ISBN 978-1-4008-5280-2.
- "National Symbols: National Animal". tanzania.go.tz. Tanzania Government Portal. Archived from the original on 18 January 2015. Retrieved 14 January 2015.
- "Chimpanzees among 33 breeds selected for special protection". BBC. 28 October 2017. Retrieved 30 October 2017.
- "Good News for Giraffes at CITES CoP18 > Newsroom". newsroom.wcs.org. Retrieved 16 November 2020.
- Muller, Zoe; Lee, Derek E.; Scheijen, Ciska P. J.; Strauss, Megan K. L.; Carter, Kerryn D.; Deacon, Francois (2020). "Giraffe translocations: A review and discussion of considerations". African Journal of Ecology. 58 (2): 159–171. doi:10.1111/aje.12727. ISSN 1365-2028.
- Lee, De; Fienieg, E; Van Oosterhout, C; Muller, Z; Strauss, M; Carter, Kd; Scheijen, Cpj; Deacon, F (27 February 2020). "Giraffe translocation population viability analysis". Endangered Species Research. 41: 245–252. doi:10.3354/esr01022. ISSN 1863-5407.
- Lee, Derek E.; Bond, Monica L. (9 June 2016). "Precision, accuracy, and costs of survey methods for giraffe Giraffa camelopardalis". Journal of Mammalogy. 97 (3): 940–948. doi:10.1093/jmammal/gyw025. ISSN 0022-2372. S2CID 87384776.
- Rudgley, Richard The Encyclopedia of Psychoactive Substances, pub. Abacus 1998 ISBN 0 349 11127 8 pps. 20–21.
- Ian Cunnison (1958). "Giraffe hunting among the Humr tribe". Sudan Notes and Records. 39.
- Qiu, Jane (22 June 2017). "Surge in wildlife killings is wiping out giraffes". Science. doi:10.1126/science.aan7000.
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