Bos gaurus Hamilton-Smith, 1827 Wild Gaur B [os]. Bubalus guavera Kerr, 1792:339. Type locality ��� Ceylon.��� Bos frontalis Lambert, 1804:57. Type locality ��� India;��� first use of the current name combination of the binomial for the domestic form (Gentry et al. 2004; International Commission on Zoological Nomenclature 2003). Bos gavaeus Colebrooke, 1808:512, 516. Type locality ���mountains that form the eastern boundary of the provinces of Aracan [Burma], Chittagong (Chitgozou) [Bangladesh], Tipara [India], and Syilhet [Bangladesh].��� Bos sylhetanus Cuvier, 1824:2. Type locality ���vivans �� la m��nagerie de Barracpour; au pied des montagnes du Sylhet.��� Bos gaurus Hamilton-Smith, 1827a:399. No type locality given; first use of the current name combination of binomial for the wild form (Gentry et al. 2004; International Commission on Zoological Nomenclature 2003). B [os. (Bison)] gaurus: Hamilton-Smith, 1827b:373. Name combination. B [os. (Bison)] gavaeus: Hamilton-Smith, 1827b:375. Name combination. Bos gour Hardwicke, 1828:231. Type locality ���Mountainous District of Ramgurh [= Ramgarh, Jharkhand, India], and Table-land of Sirgoojas [= Surguja, Chhattisgarh, India].��� Bos gayaeus Hardwicke, 1828:232. Incorrect subsequent spelling of Bos gavaeus Colebrooke, 1808. Bison gaurus: Jardine, 1836a:251. Name combination. Bison sylhetanus: Jardine, 1836b:257. Name combination. [Bos (Bibos)] subhemachalus Hodgson, 1837:499. Type locality ���saul [= sal] forest of Nipal [= Nepal].��� [Bos (Bibos)] cavifrons Hodgson, 1837:747 Replacement name for Bibos subhemachalus Hodgson, 1837. Bos gareus Gray, 1843:151. Incorrect subsequent spelling of Bos gaurus Hamilton-Smith, 1827. Bos gaur Sundevall, 1844:201. Incorrect subsequent spelling of Bos gaurus Hamilton-Smith, 1827. Bibos frontatus Gray 1846:230. Incorrect subsequent spelling of Bos frontalis Lambert, 1804. [Gaveus] frontalis: Hodgson, 1847:706. Name combination and incorrect subsequent spelling of Bos frontalis Lambert, 1804. [Gaveus] gayoeus: Hodgson, 1847:706. Name combination. [Gaveus] sylhetanus: Hodgson, 1847:706. Name combination. Bibos concavifrons Roulin 1849:619. Incorrect subsequent spelling of Bibos cavifrons Hodsgon, 1837. Gavaeus frontalis: Horsfield, 1851:179. Name combination. Bibos asseel Horsfield, 1851:181. Type locality ���South-eastern Frontier of Bengal and Silhet.��� Bos frontalis domesticus Fitzinger, 1860:387. Name combination. Gavaeus gaurus: Blyth, 1860:284. Name combination. Bos (Bibos) frontalis: Lydekker, 1898:32. Name combination. Gauribos brachyrhinus Heude, 1901:3, 4. Type locality ��� Pursat, station situ��e sur un affluent des grand lacs du Cambodge [= on one of the tributaries of the grand lakes of Cambodia],��� based on lectotype selection by Braun et al. (2001:652). Gauribos laosiensis Heude, 1901:3. Type locality ���la chaine qui s��pare le Laos du Tonkin, vers la province de Camoun [= Annamite Mountain, separating Laos and Cambodia],��� based on lectotype selection by Braun et al. (2001:652). Gauribos sylvanus Heude, 1901:4. Type locality ���for��ts des Mois [= Moi forests],��� Vietnam, based on holotype selection by Braun et al. (2001:652). Gauribos mekongensis Heude, 1901:5. Type locality ��� Krati��,��� Cambodia, based on lectotype selection by Braun et al. (2001:652). Uribos platyceros Heude, 1901:5. Type locality ���Tourane de bassins des rivi��res de Hu��,��� Vietnam, based on lectotype selection by Braun et al. (2001:652). Bubalibos annamiticus Heude, 1901:3, 6. Type locality ���Hu�� [Province],��� Vietnam, based on lectotype selection by Braun et al. (2001:652). Bos? leptoceros Heude, 1901:7. Type locality ���Kampot, au bord du golfe de Siam [= Cambodia on shores of the Gulf of Siam].��� Bibos discolor Heude, 1901:3, 8. No type locality given. Bibos sondaicus Heude, 1901:3, 8. No type locality given. Bibos longicornis Heude, 1901:9. No type locality given. Bibos? fuscicornis Heude, 1901:9. Type locality ���D��a sur la rivi��re de Vinh.��� Bos gaurus readi Lydekker, 1903:266. Type locality ��� Burma.��� Bos gaurus frontalis: Lydekker, 1912:177 Name combination. CONTEXT AND CONTENT. Order Artiodactyla, suborder Ruminantia, infraorder Pecora, family Bovidae, subfamily Bovinae, tribe Bovini. Lydekker (1907) classified 3 subspecies of Bos gaurus: B. gaurus gaurus (Bangladesh, India, and Nepal), B. gaurus readei (Burma and China), and B. gaurus hubbacki (Malaysia). Lydekker classified the 3 subspecies using only 5��� 6 specimens, and all of the morphological differences that he relied on have since been proved to be incorrect. More recently, based on skull and horn measurements, 2 subspecies were proposed (Groves 2003; Groves and Grubb 2011): B. gaurus gaurus in India and Nepal (and possibly Bangladesh) and B. gaurus laosiensis in Myanmar (Burma), Lao PDR, Vietnam, Cambodia, Thailand, and West Malaysia (and presumably southern China). This new classification was also based on a small sample size of skulls. Thus far, no genetic analyses have yet conclusively corroborated the existence of subspecies. Therefore, the evidence to split B. gaurus into subspecies remains inconclusive. NOMENCLATURAL NOTES. Although Wilson and Reeder (2005) list Bos diardii Temminck, 1838, and Bos frontatus Temminck, 1938, as synonyms of Bos gaurus Hamilton-Smith, 1827a, and Bos frontalis Temminck, 1838 as synonyms of Bos frantalis Lambert, 1804 ���while using Bos frontalis to refer to both the wild and domestic forms���they clarify that references for both these synonyms were ���nomen nudum.��� Kerr (1792) included Bos bubalus guavera in his list of Mammalia of the Animal Kingdom based on the description by Knox (1681:21) of a wild buffalo with white legs called gauvera in Ceylon. Pennant (1792:31), also referring to Knox (1681), described a subspecies of buffalo from Ceylon with ���legs that are white one-half way from the hoofs��� as gauvera in his ���History of quadrupeds.��� Colebrooke (1808) provided the first detailed description and measurements of wild and domestic form and called the animal gayal, which is one of the common names for the domestic form. Hardwicke (1827) coined his Bos gour synonym based on the detailed description by Traill (1824), who described an animal he thought was known as gour in India. F. Cuvier (1824) provided the first colored illustrations of male and female B. gaurus that were based on the description of wild B. gaurus by M. Alfred Duvaucel. Common names of the wild form in other languages include tadok (Adi); peeoug (Burmese); kaati, kaadu kona, kaadu yemme (Kannada); kulga, gameya (Kannada in Uttara Kannada district); duddu (Kannada in Northern Udupi district); meuay (Lao); seladang (Malay); Da-E-Ni, Xffi+ (Mandarin); raangawa (Marathi); kattu pothu (Malayalam); gauri gai (Nepali); kaatu maadu (Tamil); B�� T��t (Vietnamese); adavi dunna (Telugu); krating ��������������� (Thai); and moo (Gonds). DIAGNOSIS Species in the subfamily Bovinae are large and have stout bodies, hollow horns, relatively short legs, long tails with a terminal tuft of hair, broad muzzles, and no facial, pedal, or inguinal glands (Blanford 1888; Lydekker 1913). Five genera in Bovinae (Grubb 2005) are currently considered in the tribe Bovini: Bison, Bos, Bubalus, Pseudoryx, and Syncerus. Species in the Bovini tribe are distinguishable by their smooth horns that are strongly keeled and spirally twisted, although not being regularly ridged (Figs. 1���3; Grubb 2005; Groves and Grubb 2011). Extant species of Bovini are further distinguished from other Bovinae species by their low, wide skulls, internal sinuses in the frontals extending into the horn cores, a short braincase and widened occiput, molars with larger basal pillars and complicated central cavities, and upper molars that are strongly hypsodont (Lydekker 1913; Gentry 1992; Grubb 2005; Groves and Grubb 2011). Except for the yak (Bos mutus), which is hairy and has 14 dorsal and 5 lumbar vertebrae (Leslie and Schaller 2009), Bos gauru s shares its traits of not being hairy and having 13 dorsal and 6 lumbar vertebrae with the other Bos and Bubalus wild cattle species of Asia. Dewlaps that hang under the neck and chin and a dorsal ridge are prominent features that distinguish Bos males from Bubalus males. Dewlaps and the dorsal ridge are prominent distinguishing features of adult male B. gaurus (Fig. 1), and these traits are shared with the males of other Southeast Asian Bos species, such as the banteng (B. javanicus) and the now presumed extinct kouprey (B. sauveli). Both sexes in all 3 species have white lower legs, and females of both species are primarily brown and males are primarily black. Bos javanicus, however, has a white patch on its rump that B. gaurus does not, and reported weights suggest that B. javanicus is smaller in size than B. gaurus. Adult male B. gaurus are further distinguishable from adult male B. javanicus by their very muscular appearance, and the brown pelage of female B. javanicus is a shade lighter than the brown pelage of female B. gaurus. GENERAL CHARACTERS Bos gaurus is a sexually dimorphic species (Figs. 1 and 2), and differences between the sexes begin to be noticeable after the age of 2 years (Ahrestani and Prins 2011). Young calves (0���2 months) have light orange-brown body coats and do not have white stockings. The white stockings (all 4 legs, starting just above the knee in both sexes, are white) develop from the age of 3 months (Fig. 1). Males grow rapidly to attain large sizes (> 900 kg) and develop a black pelage with age; adult males, generally> 5 years are referred to as black bulls. Females are smaller in size (B. gaurus, the older is the individual (Fig. 4). Horns of males are thicker and extend outwards first before curving inward, which results in the horns on males being further apart from each other (Figs. 3 and 4). Female horns, in contrast, extend outward a lot less, and are thus closer to one another, and have a spiral curvature that makes the horns point at each other; the inward curvature of female horns begins by the age of 2 years (Figs. 3 and 4). Horns on both sexes appear to grow throughout the lifetime of an individual, and it is not uncommon to see old females with horns that are nearly touching. A muscular elevated dorsal ridge and dewlaps that hang under the neck and chin easily distinguish adult males from females. Frontals and parietals of the skull are in a single plane and are similar to other Bos species (Fig. 5). A more detailed description of the differences in morphological characteristics between the sexes across age classes is presented in Ahrestani and Prins (2011). Reported weights of adult males shot in the wild were as follows: 590, 782, 864, 864, 931, and 941 kg (Dunbar-Brander 1923; Meinertzhagen 1939; Morris 1947). Reported weights of adult females shot in hunts were as follows: 440 kg (excluding blood���Schaller 1967) and 703 kg (Meinertzhagen 1939). The skull of a male B. gaurus has been recorded to weigh about 21 kg (Robison 1941). Based on measurements of 9 captive males and 14 captive females, Ahrestani and Prins (2011) reported a maximum shoulder height of 175 cm for males and 148 cm for females. Other records of shoulder height of males include those killed in hunts: 145 cm (Cameron 1929), 176 cm (Inverarity 1889), 178 cm (Forsyth 1889), and 197 cm (Pillay 1952). Length from nose to root of the tail of a male���s body was reported to be 249 cm (Cameron 1929) and 284 cm (Dunbar- Brander 1923). Length of tail was 86 cm (Dunbar-Brander 1923) and 89 cm (Cameron 1929). The range of length of a male���s horn is 61���96 cm (Forsyth 1889; Inverarity 1889; Baker 1903; Cameron 1929; Robison 1941; Hundley 1952; Pillay 1952; Schaller 1967), and the spread (between the widest outside points) is 89���134 cm (Inverarity 1889; Baker 1903; Cameron 1929; Robison 1941; Hundley 1952; Pillay 1952). For males, the circumference of horns at the base has been measured to be 43���58 cm (Baker 1903; Pillay 1952), and ear length is about 23 cm. Hind foot length is about 55 cm (Cameron 1929); the proportion of the length of legs to its body mass is probably one of the smallest in the animal kingdom. The eyes of a B. gaurus are normally colored brown. The eyes, however, sometimes appear blue under certain light conditions because of the presence of tapetum lucidum, a membrane behind the retina that makes the eyes of several animals shine in the dark. DISTRIBUTION The global distribution of Bos gaurus, both historically and in present time, has been restricted to southern and southeastern Asia (Fig. 6), which includes Bangladesh, Bhutan, Cambodia, China (Yunnan and southern Tibet), India, Lao PDR, Malaysia (Peninsular Malaysia), Myanmar (Burma), Nepal, Sri Lanka (extinct), Thailand, and Vietnam. In the last century, the overall distribution of B. gaurus has shrunk by> 80%, and today, B. gaurus is mainly found in protected areas (Schaller 1967; Choudhury 2002; Ahrestani and Karanth 2014). Although the elevational range of habitats that B. gaurus occupies is wide, sea level to 2,700 m, it is found more on hills than on plains. Early natural history accounts of B. gaurus report that B. gaurus prefers hilly areas, particularly during the dry season. However, it is unclear if this behavior is driven by some ecological need or if it is a function of the fact that the majority of all remaining habitat available to B. gaurus is hilly terrain as nearly all suitable habitat in the plains have been lost to agriculture (Schaller 1967). More than 80% of the global population of B. gaurus is found in India, distributed over 3 widely separated geographical regions: the Western Ghats, central India, and northeastern India (Choudhury 2002). The largest population of B. gaurus (3,000 ���5,000) in the world is found in the 5,520 km 2 Nilgiri Biosphere Reserve, southern India. Currently, in India, populations of B. gaurus in the Western Ghats are secure; populations in central India are less secure, while populations in northeastern India are vulnerable (Ahrestani and Karanth 2014). Bos gaurus is most probably extinct in Bangladesh; no records have been reported since the 1970s (Khan 1985). It is possible that individuals from Mizoram and Tripura in India occasionally cross over into Bangladesh (Choudhury 2002). The overall population of B. gaurus in Nepal, confined mainly to Chitwan National Park and Parsa Wildlife Reserve, is understood to be less than 500 individuals, but is considered to be stable. In Bhutan, B. gaurus is apparently found all along the southern foothill zone, mostly in protected areas with a few recent sightings outside protected areas. The status of B. gaurus in Myanmar is poorly understood, though during a national tiger survey from over a decade ago, B. gaurus was camera trapped in 11 of 15 sites, with a high rate of capture in 5 of these sites (Lynam 2003). The overall population of B. gaurus in Thailand could be in excess of 1,000 individuals. The outlook for B. gaurus in northern Thailand is currently favorable because an effective antipoaching campaign and reforestation program over the last couple of decades has led to several increasing populations, most notably in Khao Yai National Park, Huai Kha Khaeng, and Thung Yai Naresuan wildlife sanctuaries. Forests are highly fragmented in southern Thailand, and it is assumed that B. gaurus has been largely extirpated in this region. Nevertheless, it may survive along the Malaysian border, where the human population is low and forest fragments are larger because of an ongoing insurgency in that region. Across the border, the B. gaurus population within mainland Malaysia was estimated to be around 500 in 1994; it is suspected that this population has now reduced by 50%. The outlook for B. gaurus in Malaysia is grim, and it possible that it survives at a viable population only in Taman Negara (peninsular Malaysia���s largest national park���Lynam et al. 2007). In China, B. gaurus occurs in Yunnan and southeastern Tibet (Ahrestani and Karanth 2014). Although the exact status of the populations in these regions is unknown, it is understood that the overall population of B. gaurus in China does not exceed 200 (H. Jianlin, pers. comm.). A report from nearly 20 years stated that B. gaurus had been extirpated from much of Yunnan province (Xiang and Santiapillai 1993). In Cambodia, B. gaurus was widespread until the 1960s, after which the overall country population has decreased by nearly 90%. The largest population can be found in eastern Cambodia (Mondulkiri Province), and recent protection measures may have stabilized this population (Timmins and Rattanak 2001; Tordoff et al. 2005). B. gaurus in Lao DPR was estimated to be about 1,000 individuals in the 1990s (Byers et al. 1995); however, since then, multiple populations in Lao have been extirpated, and it is estimated that the current overall population in Lao is no more than 500 individuals. In Vietnam, the current status of B. gaurus in unknown, and it is thought that populations that remain are in serious decline. The majority of B. gaurus in Vietnam are confined to Cat Tien National Park (Polet and Ling 2004; Nguyen 2009). FOSSIL RECORD The first fossil record attributed to Bos gaurus included a partial skull and horns found in the older alluvium of the Narmada (= Narbada) River in central India (Spilsbury 1840). This fossil record, however, was not dated. In general, the origins of Bos and the relationships of fossils that have been attributed to this genus remain problematic, which when combined with the poor fossil record from Africa for the time period 7���10 million years ago has made it difficult to confirm whether or not the Bovini tribe originated in southern Asia (Bibi et al. 2009). 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