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Bos frontalis and Bos gaurus (Artiodactyla: Bovidae)

  • Frontier Wildlife Conservation

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Bos frontalis Lambert, 1804 and Bos gaurus Hamilton-Smith, 1827 are the domestic and wild forms, respectively, of the bovid commonly called the gaur. It is the world's largest cattle species. Bos gaurus is endemic to south and southeastern Asia, and today, the majority of its population occurs in India. It is sexually dimorphic, with adult males having a distinctive dorsal ridge and often a dewlap. Although B. gaurus consumes numerous browse species, it is primarily a grazer. Except for older males, all other B. gaurus are nearly always found in herds. It is classified as a Vulnerable species, and in 2011-2012, the first reintroductions of B. gaurus occurred in central India.
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MaMMalian SpecieS 50(959):34–50
Bos frontalis and Bos gaurus (Artiodactyla: Bovidae)
Farshid s. ahrestani
Foundation for Ecological Research, Advocacy and Learning, Auroville Post, Tamil Nadu 605 101, India;
Abstract: Bos frontalis Lambert, 1804 and Bos gaurus Hamilton-Smith, 1827 are the domestic and wild forms, respectively, of the
bovid commonly called the gaur. It is the world’s largest cattle species. Bos gaurus is endemic to south and southeastern Asia, and
today, the majority of its population occurs in India. It is sexually dimorphic, with adult males having a distinctive dorsal ridge and
often a dewlap. Although B. gaurus consumes numerous browse species, it is primarily a grazer. Except for older males, all other
B. gaurus are nearly always found in herds. It is classified as a Vulnerable species, and in 2011–2012, the first reintroductions of
B. gaurus occurred in central India.
Key words: Asia, Bovini, cattle, gaur, gayal, mithun, ruminant, ungulate
Synonymy completed 7 July 2016
Version of Record, first published online August 17, 2018, with fixed content and layout in compliance with Art. ICZN.
Nomenclatural statement—A life science identifier (LSID) number was obtained for this publication: urn:lsid:zoobank.
Bos frontalis Lambert, 1804
Domestic Gaur
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 domes-
tic form (Gentry et al. 2004; International Commission on
Zoological Nomenclature 2003).
Bos gavæus Colebrooke, 1808:512, 516. Type locality “moun-
tains 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 à la
ménagerie 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
B[os. (Bison)] gavæus: Hamilton-Smith, 1827b:375. Name
Bos gour Hardwicke, 1828:231. Type locality “Mountainous
District of Ramgurh [= Ramgarh, Jharkhand, India], and
Table-land of Sirgoojas [= Surguja, Chhattisgarh, India].”
Bos gayæus Hardwicke, 1828:232. Incorrect subsequent spelling
of Bos gavæus Colebrooke, 1808.
Bison gaurus: Jardine, 1836a:251. Name combination.
Bison sylhetanus: Jardine, 1836b:257. Name combination.
© The Author(s) 2018. Published by Oxford University Press on behalf of American Society of Mammalogists.
All rights reserved. For permissions, please e-mail:
Fig. 1.—Adult male Bos gaurus with conspicuous dorsal ridge and dew-
laps from Nagarahole Tiger Reserve, southern India. Used with permis-
sion of the photographer K. Varma (http://www.
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2 MAMMALIAN SPECIES 50(959)—Bos frontalis and Bos gaurus
[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,
[Gaveus] gayœus: Hodgson, 1847:706. Name combination.
[Gaveus] sylhetanus: Hodgson, 1847:706. Name combination.
Bibos concavifrons Roulin 1849:619. Incorrect subsequent spell-
ing 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.
Gavæus gaurus: Blyth, 1860:284. Name combination.
Bos (Bibos) frontalis: Lydekker, 1898:32. Name combination.
Gauribos brachyrhinus Heude, 1901:3, 4. Type locality
“Pursat, station située 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.
Gauribos laosiensis Heude, 1901:3. Type locality “la chaine
qui sépare 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 “foréts des
Mois [= Moi forests],” Vietnam, based on holotype selec-
tion by Braun et al. (2001:652).
Gauribos mekongensis Heude, 1901:5. Type locality “Kratié,”
Cambodia, based on lectotype selection by Braun et al.
Uribos platyceros Heude, 1901:5. Type locality “Tourane de
bassins des rivières de Hué,” Vietnam, based on lectotype
selection by Braun et al. (2001:652).
Bubalibos annamiticus Heude, 1901:3, 6. Type locality “Hué
[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
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 “Dûa sur la
rivière 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 pro-
posed (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 cor-
roborated 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)
Fig. 2.—Adult female Bos gaurus with typical brown pelage and
smooth, spiral-shaped horns from BR Hills, southern India. Used with
permission of the photographer K. Varma (http://www.http://kalyan-
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50(959)—Bos frontalis and Bos gaurus MAMMALIAN SPECIES 3
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 illus-
trations 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, 大额牛 (Mandarin); raangawa
(Marathi); kattu pothu (Malayalam); gauri gai (Nepali); kaatu
maadu (Tamil); Bò Tót (Vietnamese); adavi dunna (Telugu);
krating กระทง (Thai); and moo (Gonds).
Species in the subfamily Bovinae are large and have stout
bodies, hollow horns, relatively short legs, long tails with a ter-
minal 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 extend-
ing 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
gaurus 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 promi-
nent 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 pre-
sumed extinct kouprey (B. sauveli). Both sexes in all 3 species
have white lower legs, and females of both species are prima-
rily brown and males are primarily black. Bos javanicus, how-
ever, 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.
Bos gaurus is a sexually dimorphic species (Figs. 1 and 2), and
differences between the sexes begin to be noticeable after the age
Fig. 3.—Differences in thickness, shape, and curvature of horns, and thickness of neck between female and male Bos gaurus from Mysore Zoo,
southern India. Photographer F. S. Ahrestani.
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4 MAMMALIAN SPECIES 50(959)—Bos frontalis and Bos gaurus
of 2 years (Ahrestani and Prins 2011). Young calves (0–2 months)
have light orange-brown body coats and do not have white stock-
ings. 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 (< 600 kg) and have a
brown pelage (Figs. 1 and 2). Both sexes have a light brown “boss”
between their horns. Horns in both sexes are black in their early
stages of development and, with advancing age, begin to whiten
from the base up. The greater the proportion of white on the horns
of a 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 indi-
vidual, 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 (exclud-
ing 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 cap-
tive 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.
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 over-
all 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
Fig. 4.—Age-specific differences in the size and shape of the head and horns, and the black:white ratio on horns of (a) female and (b) male Bos
gaurus. (Reprinted from F. S. Ahrestani [2009] Ph.D. dissertation titled “Asian Eden: large herbivore ecology in India”).
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50(959)—Bos frontalis and Bos gaurus MAMMALIAN SPECIES 5
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 agricul-
ture (Schaller 1967).
More than 80% of the global population of B. gaurus is
found in India, distributed over 3 widely separated geographi-
cal 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 km2 Nilgiri
Biosphere Reserve, southern India. Currently, in India, popula-
tions 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 sight-
ings 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 antipoach-
ing 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 popula-
tions in Lao have been extirpated, and it is estimated that the
current overall population in Lao is no more than 500 individu-
als. 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).
The first fossil record attributed to Bos gaurus included a par-
tial skull and horns found in the older alluvium of the Narmada
Fig. 5.—Dorsal, ventral, and lateral views of skull and lateral view
of mandible of adult male Bos gaurus (American Museum of Natural
History, New York, [AMNH] specimen 54468). Greatest length of skull
is 683 mm. Photographer F. S. Ahrestani.
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6 MAMMALIAN SPECIES 50(959)—Bos frontalis and Bos gaurus
(= 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).
Pilgrim (1939), in great detail, attributed the different
Bovinae (wild cattle) fossils that had been found in India from
the early 19th century to various genera, namely Proamphibos,
Hemibos, Bubalus, Bucapra, Proleptobos, Leptobos, Platybos,
Bison, and Bos. This splitting of Bos-like fossils into multiple
genera was based mostly on partial fossils of not more than a
couple of specimens found each in the Siwalik Hills (Hemibos);
Padri, Kangra, and Hoshiarpur in the Siwaliks, Jhansi Ghat in the
Narmada (= Narbada) valley, and Pemganga River in Hyderabad
(Bos); and Pinjor in the Upper Siwaliks (Bison and Platybos).
In general, fossils of Bos-like species found in India have
been dated as belonging to the Upper Pliocene, i.e., 3.5–5 mil-
lion years ago (Pilgrim 1939; Chauhan 2008). The majority of
fossils of oxen species found in India from the late Quaternary
(Pleistocene) period have been attributed mainly to Bos nama-
dicus, the species considered to be the wild stock of the extant
domestic Bos indicus (Grigson 1985; Chauhan 2008). Fossil
records of Bos namadicus that were attributed to middle–late
Pleistocene (32,000 BP) were found mainly in the Godavari
Valley (Badam et al. 1984; Badam and Jain 1988). Fossils found
in the Narmada valley in central India, which were also dated
Fig. 6.—Distribution of Bos gaurus endemic to southern and southeastern Asia. Map redrawn from International Union for Conservation of Nature
and Natural Resources Redlist with modifications and courtesy of Srinivas Vaidyanathan, Foundation for Ecological Research, Advocacy and
Learning, Puducherry, India.
Fig. 7.Bos gaurus herd with adult females and calves from Bandipur Tiger Reserve, southern India. Used with permission of the photographer
M. N. Naveen.
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50(959)—Bos frontalis and Bos gaurus MAMMALIAN SPECIES 7
to the Pleistocene, were classified as belonging to Bibos gaurus
(Badam and Grigson 1990).
Besides India, abundant skulls, horns, mandibles, and teeth
of Bibos (Bos) gaurus (and supposedly also of a similar spe-
cies Bibos geron) belonging to the Pleistocene were found in
different areas in China beginning in the end of the 19th century
(Colbert et al. 1953). These areas included the regions of Sichuan
(= Szechuan—Matsumoto 1915), Yanjinggou (= Yenchingkou—
Matthew et al. 1923; Young 1932, 1939), and the Bailong cave,
Hubei (Wang et al. 2015).
A distinguishing feature of a male Bos gaurus is its dorsal
ridge, which is “formed by a row of single bones springing from
the back bone immediately behind the junction of each pair of
ribs, of which B. gaurus has 13 pairs” (Inverarity 1889:295). The
highest point of the dorsal ridge is above the 5th–6th rib, after
which the ridge reduces in height and ends abruptly at the last
rib. The dental formula is i 0/4, c 0/0, p 3/3, m 3/3, total 32
(A. Bourgeois, pers. comm.), which is the same as that of cattle.
B. gaurus secretes an oily chemical through its skin, which had
been noticed by early naturalists who had proposed that it acted
as an insect repellent (Hubback 1937). The oil, identified as
5-(1-hydroxynonyl)-2-tetrahydrofuranpentanoic acid, has been
named bovidic acid (Ishii et al. 2004), and 2 studies have found it
to have mosquito repellent properties (Tran and Chauhan 2007;
Phillips et al. 2015).
Mean heart rates for 5 B. gaurus ranged from 49.3 to
57.7 beats/min, and mean body temperatures for 2 B. gaurus
were 38.2°C and 38.8°C (Thomas et al. 1996). Short-duration
adverse stimuli caused brief 3-fold increase in heart rate,
but baseline rates returned after the stressors were removed.
Moving B. gaurus to novel environments or pairing them with
nonaffiliates also increased heart rates (Thomas et al. 1996).
Body temperature was not affected by short-term stressors, but
it was positively correlated with ambient temperature (Thomas
et al. 1996).
Bos gaurus is a ruminant and, among all extant true rumi-
nants, is only second to the giraffe in body mass. Observations
by multiple naturalists and field biologists suggest that
B. gaurus depends less on sight (despite being capable of
seeing at night) and more on smell and hearing to detect dan-
ger (Inverarity 1889; Dunbar-Brander 1923; Hubback 1937;
Schaller 1967; Krishnan 1972).
Johnston et al. (1994) found that ovarian oocytes of imma-
ture Bos gaurus were capable of in vitro maturation and fer-
tilization with thawed homologous spermatozoa and that the
resulting embryos were capable of advancing to blastocysts in
culture and of producing live-born offspring after embryo trans-
fer. Specifically, the embryos developed to the blastocyst stage in
7 days, and a live-born calf was delivered 308 days after transfer-
ring the embryos to Holstein recipients.
Primiparity (i.e., age at first birth) for captive B. gaurus is
3 years, which is similar to what has been observed for other
Bovini species (Ahrestani et al. 2011). Ovulation has been found
to occur at 19- to 22-day intervals, and onset of ovulation can
be predicted based on measured temperature spikes (Thomas
et al. 1996). Although no data exist for B. gaurus, the average
length of the estrous cycle in the domestic form Bos frontalis
has been found to be 22 days, and the average duration of how
long female B. frontalis remain in heat was found to be 45 days
(Giasuddin et al. 2003). Volume of semen in a single ejaculation
ranges from 0.2 to 11.0 ml, and pH level in the semen ranges
from 6.58 to 8.42 (Iswadi et al. 2016).
Data from Omaha Zoo, United States, and Mysore Zoo,
India, suggest that the gestation period for B. gaurus is between
9 and 10 months, about 280 days (Ahrestani and Prins 2011).
This is similar in duration to the 9-month gestation period that
we know for domestic cattle and that was reported for B. gaurus
by Hubback (1937). In captivity, B. gaurus has been known to
give birth every year, and an 18-year-old B. gaurus was known
to have given birth with no complications (Ahrestani et al. 2011).
Given their 9- to 10-month gestation period, there is no reason
not to assume that free-ranging B. gaurus give birth every year
too (Ahrestani et al. 2011).
Schaller (1967:181) reported that “single births are the rule,
there being no records of twins” for B. gaurus. Thus far, no
evidence has been found to contradict this statement; no twins
were recorded in 319 births of B. gaurus in European zoos until
2011 (A. Bourgeois, pers. comm.), in more than 150 births
(1968–2006) in Omaha Zoo, United States, and in 41 births
(1994–2006) in Mysore Zoo, India (F. S. Ahrestani, in litt.).
A study of 13 B. frontalis females in Bangladesh found the mean
daily milk yield and lactation length were 305 ml and 117 days,
respectively (Giasuddin et al. 2003). Although calves as old as
180–240 days have been observed suckling (F. S. Ahrestani, in
litt.), it is understood that calves rarely suckle beyond the age of
6 months (Schaller 1967). The longest continuous suckling by a
calf has been recorded to be about 9 min (Schaller 1967).
Bos gaurus appears to breed throughout the year. In a study
from southern India, Ahrestani et al. (2012b) observed B. gaurus
calves throughout the year, with no discernable peak, similar
to what Inverarity (1889) reported for India, Peacock (1933)
reported for Burma, and Weigum (1972) reported for Malaysia.
There have been, however, others that have suggested mating
and calving seasons in different regions. For example, Dunbar-
Brander (1923) reported that the majority of mating in central
India occurred in December–January, with the majority of calves
born in August–September; Stebbing (1911) and Sanderson
(1912) not only reported similar months but also noted that
calves were born in April–June; Hubback (1937) reported see-
ing young in Malaysia at all times except from October to
December; Morris (1937) reported that the peak rutting period in
southern India was from November to March; K. U. Karanth (in
litt.) found peak rutting activity in June–September (wet season)
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8 MAMMALIAN SPECIES 50(959)—Bos frontalis and Bos gaurus
in Nagarahole, southern India; and Schaller (1967:180), while
having observed calves throughout the year, reported that “the
frequency of rutting call and other aspects of sexual behavior
reached a peak in March and April” in Kanha, India.
Population characteristics.—The majority of density
(individuals/km2) estimates of free-ranging Bos gaurus are
from India, and these estimates vary by location and time.
The highest densities of B. gaurus have been found in the
southern region of the Western Ghats, which include: densi-
ties as high as 10–15 in the backwaters of the Kabini dam in
2001–2002 (Madhusudan 2004); 14.4 (± 3.8 SE) in Mudumalui
Tiger Reserve in 1988–1992 (Varman and Sukumar 1995);
12.3 (5.6–16.4, 95% CI) in the Anamalai Tiger Reserve in
2001–2004 (Kumaraguru et al. 2011); densities in Nagarahole
Tiger Reserve have been consistently high over 20 years: 5.6
(± 1.8 SE) between 1987 and 1990 (Karanth and Sunquist
1995), 4.5 (± 0.8 SE) in the mid-1990s (Karanth and Nichols
1998), and 5.1 (± 1.3 SE) individuals/km2 in 2011 (Ahrestani
and Karanth 2014); 5.3 (± 1.5 SE) in Bandipur Tiger Reserve in
2011; and estimates from Bhadra Tiger Reserve in 2011 were
1.0 (± 0.4 SE) (Ahrestani and Karanth 2014), down from 1.48
(± 0.6 SE) recorded in 2000 (Jathanna et al. 2003). Densities
of 0.7 (± 0.2 SE) in the mid-1990s (Karanth and Nichols 1998)
and densities of 0.3 (0.1–1.1, 95% CI) in 1998–1999 (Biswas
and Sankar 2002) were recorded in Pench National Park, cen-
tral India. The population density in Huai Kha Khaeng Wildlife
Sanctuary, western Thailand, was reported to be 1.8 (1.3–2.3,
95% CI) in 1998 (Srikosamatara 1993).
The ratio of males:females at birth is only available for
captive populations and has been recorded as 0.86 (n = 41) in
Mysore Zoo, 1.36 (n = 180) in Omaha Zoo, and 1.29 (n = 16)
from the National Zoological Park, Calcutta, India (Reed 1959;
Ahrestani et al. 2011). Although these reports do not provide a
definite ratio, based on data of other Bovini species, Ahrestani
et al. (2011) concluded that there was no reason not to assume
that the sex ratio at birth for B. gaurus is parity. Given that the
difference between males and females till the age of 2 years
is minimal, it is not surprising that the survival rates for both
sexes were found to be similar till the age of 2 years: among 109
medium-sized and large-sized calves sexed in Kanha National
Park, India, 55 were male and 54 were female (Schaller 1967);
and in the Bandipur–Mudumalai landscape, India, among 282
B. gaurus sexed below the age of 1 year, 140 were males and
142 female, and among 153 B. gaurus sexed between the age of
1 and 2 years, 76 were males and 77 females (Ahrestani et al.
The ratio of adult females:males in B. gaurus popula-
tions is often 2:1, and sometimes 4:1. In southern India, the
adult male:female ratio has been reported as being 25:100
(Ashokkumar et al. 2010), 39:100 (Ramesh et al. 2012), and
33:100 (Ahrestani et al. 2011) in Mudumalai Tiger Reserve;
18:100 (Karanth and Sunquist 1992) in Nagarahole Tiger
Reserve; 45:100 in Parambikulam Tiger Reserve (Vairavel
1998); in central India ratios of 60:100 in Pench National Park
(Sankar et al. 2002) and 50:100 (Belsare et al. 1984) and 80:200
(Schaller 1967) from Kanha National Park; and in northeastern
India a ratio of 58:100 has been reported from Trishna Wildlife
Sanctuary (Dasgupta et al. 2008). Also, survival rates of females
appear to be higher than those of males. Analyzing the survival
of 72 males and 58 females from birth to death in Omaha Zoo,
United States, Ahrestani et al. (2011) found that after the age of 2,
survival of females was higher than males in captive conditions.
Despite the difference in survival between the sexes, the
maximum-recorded lifespan for both sexes in captivity has
been found to be the same: about 24 years for a captive female
(Crandall 1964) and 23.6 years for a captive male (Ahrestani
et al. 2011). Considering data from populations of other free-
ranging Bovini species, a female B. gaurus that reaches the age
of 20 years may be expected to produce 8–10 calves in her life-
time (Ahrestani et al. 2011).
Space use.—Today, Bos gaurus is primarily confined to
forested areas (dry and moist deciduous, semi-evergreen, and
evergreen) and hilly terrain (< 2,500 m) in India and across
much of its distribution in Southeast Asia (Choudhury 2002;
Karanth et al. 2009; Ahrestani et al. 2012a). This appears to
be a function of the forested habitat available in the protected
areas to which the species is more-or-less confined; there is
an acute lack of protected plains and grassland habitats in
India and southeastern Asia. For example, Schaller (1967:178)
suggested nearly 50 years ago that “the apparent preference
of B. gaurus for hilly terrain may in part be due to the con-
version of much of an earlier habitat in the plains into fields,
whereas the hills have until recent years been left relatively
undisturbed.” Naturalists have observed the preference for hilly
terrain for more than a century (Forsyth 1889; Inverarity 1889;
Dunbar-Brander 1923), but there is no evidence to support the
oft-mentioned claim that B. gaurus moves to hilly terrain to
escape insects.
In Thailand, a population of B. gaurus recovering after
poaching selected deciduous over evergreen forest (Steinmetz
et al. 2010). In Lao PDR, Vietnam, and Cambodia, B. gaurus is
found in evergreen forests, montane forests, and in both open and
closed lowland forests, with the species frequenting grassy open-
ings in closed canopy forests (Duckworth et al. 1999; Timmins
and Rattanak 2001). In Malaysia, B. gaurus has been associated
with habitats dominated by secondary vegetation, such as jungle
clearings, abandoned fields of shifting cultivators, forest fringes,
and openings along rivers (Hubback 1937; Foenander 1952;
Stevens 1968; Weigum 1972; Conry 1989). Multiple naturalists
have reported observing B. gaurus foraging along stream banks
in India (Forsyth 1889; Inverarity 1889; Dunbar-Brander 1923;
Krishnan 1972). It is possible that low-lying regions near rivers
and drainage lines—areas that typically retain the last remaining
herbaceous layer in the dry season—act as key foraging habi-
tats for B. gaurus in the dry season. B. gaurus have historically
and even today use mineral licks throughout their distributional
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50(959)—Bos frontalis and Bos gaurus MAMMALIAN SPECIES 9
range (Inverarity 1889; Hubback 1937; Ogilvie 1954; Schaller
1967; Krishnan 1972; Weigum 1972; Steinmetz 2004).
Bos gaurus forages day and night and seems to prefer to
forage at night, dawn, and dusk in areas with warm climate.
During the hottest periods of the day, B. gaurus often retreats
to forested areas to chew cud, often done while lying down
(Forsyth 1889; Krishnan 1972; F. S. Ahrestani, in litt.). During
his seminal study, Schaller (1967) observed that the major-
ity of B. gaurus were under shelter of trees by 0700 h, were
rarely seen in the open after 0800 h, and emerged from forested
patches to open grasslands in the evenings. A radio-telemetry
study in Pench Tiger Reserve, India, recorded the mean daily
(daytime) movement of an adult male to be 1.8 km in summer
and 1.3 km in the rainy (monsoon) season and that of an adult
female to be 1.2 km in the summer and 1.4 km in the rainy sea-
son (Sankar et al. 2002).
Bos gaurus has been found to range over areas that vary
from 8 to 169 km2. A free-ranging B. gaurus herd monitored for
2 years (1967–1969) in Taman Negara National Park, Malaysia,
ranged over 8 km2, mainly along the Tembeling River (Weigum
1972). Another telemetry study from Malaysia of 3 B. gaurus
found a yearling male that ranged over 30 km2 (November 1978–
May 1979), a yearling female that ranged over 52 km2 (October
1977–December 1978), and an adult male (about 5 years old) that
ranged over 137 km2 (January–November 1978; Conry 1989).
Despite the availability of primary forest within 1.5 km of their
home ranges, these 3 B. gaurus that were monitored in Malaysia
primarily used disturbed and early seral habitats created by log-
ging and agricultural development, secondary forest, and agri-
cultural estates. Furthermore, areas within 500 m of agricultural
fields, areas within 500 m of major rivers, and areas below 61
m elevation were used disproportionately with respect to their
availability in the ranges of these individuals. These 3 B. gaurus
individuals also frequently used areas adjacent to human settle-
ments, and all their ranges included at least 1 salt lick.
In Pench Tiger Reserve, India, the ranges of an adult male
and female—radio-collared and monitored for nearly a year
(1997–1998)—were 12.6 and 7.3 km2 in summer and 7.6 and
13.8 km2 in winter, respectively (Sankar et al. 2002). Among
the 19 B. gaurus reintroduced in 2011 to Bandhavgarh Tiger
Reserve, India, 3 adult males and 9 adult females were radio-
collared and monitored from January 2011 to January 2012. This
reintroduced herd utilized an area of 290 km2 in summer, 137
km2 in the rainy (monsoon) season, and 155 km2 in winter. The
ranges of individual males in this reintroduced herd varied from
135 to 142 km2, and the ranges of individual females varied from
32 to 169 km2 (Sankar et al. 2013).
Bos gaurus has shown site fidelity spanning a few days to
sometimes years. In Nagarahole, India, an adult female, recog-
nized because of a horn deformity, remained within a 30 km2
area for over a decade (K. U. Karanth, pers. comm.). It is also
common to see the same solitary males in an area for months
(Inverarity 1889; F. S. Ahrestani, in litt.). B. gaurus, however,
also makes local seasonal migrations: a year-long study across
200 km2 in Bandipur, India, found B. gaurus to be confined to
moist deciduous forests in the dry season and to dry decidu-
ous forests in the wet season (Ahrestani et al. 2012a); in both
Garumara Wildlife Sanctuary, West Bengal, and the Rairakhol
region, Orissa, India, B. gaurus was found to be absent during
the dry season (July–October, Garumara—Guin and Pal 1982;
March–July, Rairakhol—Imam 1985), but was present during
the rest of the year. It is likely that such seasonal migrations are
driven by B. gaurus trying to satisfy the nutritional requirements
of its large body mass.
Diet.—Although Bos gaurus eats a diverse array of plant
species and plant parts, a growing body of evidence suggests
it is primarily a grazer. This is consistent with Hofmann and
Stewart (1972) classifying Bovini species as bulk feeders of
grasses, a classification that was made considering the large
rumen size, the stomach structure, and the feeding habits of
Bovini species such as the African buffalo Syncerus caffer
and American bison Bison bison (currently recognized as Bos
bison). In general, other Bovini species in Asia that are closely
related to B. gaurus, such as the water buffalo Bubalus arnee,
are also considered grazers. Over a century ago, naturalists
such as Inverarity (1889) claimed that B. gaurus was mainly
a grazer. Since then, evidence to support this claim has grown.
For example, the mean proportion of grasses found in the rumen
contents of 4 B. gaurus autopsied in Kanha was 85% (range
66–100%, Schaller 1967); grasses made up 66%, browse 26%,
and herbs and other plants 8% of the diet of B. gaurus studied
in Nepal (Chetri 2003, 2006); graminoids (grasses, sedges, and
bamboo) accounted for > 60% of epidermal fragments found
in their feces in a multisite study from central India (Gad and
Shyama 2011); and an isotopic analysis of their feces from
southern India showed that B. gaurus was primarily a grazer
throughout the year (Ahrestani et al. 2012a).
Leaves and shoots of bamboo, the world’s largest grass spe-
cies, are sought after by B. gaurus throughout its range: the 3
dominant bamboo species in southern India (Bambusa arundina-
cea, Dendrocalamus strictus, and Oxytenanthera monostigma)
are all eaten (Krishnan 1972; K. U. Karanth, pers. comm.), and
in Thailand, B. gaurus has been found to preferentially eat bam-
boo shoots in the wet season and bamboo leaves in the dry sea-
son (Prayurasiddhi 1997; Steinmetz 2004).
Bos gaurus, however, eats far more species than just grasses.
For example, Gad and Shyama (2011) found that B. gaurus fed
on 7 grass species, 5 herb species, 8 shrub species, and 12 tree
species; in Nepal, B. gaurus has been found to feed on 49 plant
species (23 grasses, 17 browse species, and 9 herbs and others—
Chetri 2003, 2006) and in Malaysia 89 different plant species, of
which only 38 were grass species were reported (Weigum 1972);
in Kanha, B. gaurus have been found to feed on 7 shrub spe-
cies, 6 grass species, 3 vine species, 4 forb species, fruits from
2 tree species, and leaves from at least 17 tree species (Schaller
1967); and Dunbar-Brander (1923) recorded B. gaurus feeding
on more than 40 plant species. In some areas, the proportion
of browse in their diet might vary by season; for example, in a
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10 MAMMALIAN SPECIES 50(959)—Bos frontalis and Bos gaurus
4-year study, Gad and Shyama (2011) found that the proportion
of browse consumed by B. gaurus increased from 15% in winter
to 40% in summer.
Although B. gaurus feeds on different plant parts, it pri-
marily eats leaves. For example, in their histologic study, Gad
and Shyama (2011) found that 87% of B. gaurus diet composed
of leaves. B. gaurus feeds primarily by extending its tongue to
curl around leaves and twigs, which it pulls into its mouth and
bites off with a tug (Schaller 1967; Krishnan 1972). Multiple
studies have found B. gaurus feeding on fruits (Dunbar-Brander
1923; Schaller 1967; Krishnan 1972; Chetri 2003), and its eat-
ing of shoots is generally associated with its foraging on bam-
boo (Forsyth 1889; Krishnan 1972; Gad and Shyama 2011).
B. gaurus has also been observed eating the bark of trees,
including Adina cordifolia (Dunbar-Brander 1923), teak Tectona
grandis (Pasha et al. 2002), and cashew Anacardium occidentale
(Gad and Shyama 2011) in India and Holarrhena antidysentrica
and Wendlandia natoniana in Malaysia (Ogilvie 1954). The eat-
ing of bark, which has often been recorded in summer, is under-
stood to be in response to either a shortage of forage or mineral
resources, or both.
Diseases and parasites.—The cattle diseases rinderpest,
foot-and-mouth, and anthrax have been reported as affecting
Bos gaurus populations from the very earliest reports of the
species. Rinderpest has been the most common and widespread
of the 3 diseases: rinderpest was reported to have affected
B. gaurus populations in India as early as the 19th century
(Inverarity 1889; Baker 1890); the disease affected populations
in central India in the early part of the 20th century (Dunbar-
Brander 1923; Stewart 1927); and Schaller (1967:181)
reported that “a virulent epidemic of rinderpest killed many
B. gaurus in the Kanha Park area in the years 1925–26.” In
southern India, rinderpest has been reported to have killed
hundreds of B. gaurus in the 1st one-half of the 20th century
(Anderson 1954), to have all but wiped out B. gaurus from
Mudumalai in 1968 (Krishnan 1972), and to have nearly wiped
out the B. gaurus population in Bandipur in 1989 (D. V. Girish,
pers. comm.). In northeastern India, rinderpest epidemics
killed several hundred B. gaurus in Raimora, Assam, in 1967,
in the Rairakhol region, Orissa, in 1972, and in Berbera and
Dhuanali, Orissa, in 1973 (Imam 1985). Although epidemics of
foot-and-mouth disease have never been reported for B. gaurus
populations, Morris (1949) reported shooting an adult male
that had foot-and-mouth disease; a case of foot-and-mouth dis-
ease was reported from Hyderabad, southern India (Ali 1953);
4 Bos frontalis contracted foot-and-mouth disease and died
within 10 days in October 1990 in Calcutta Zoo (Choudhury
2002); in March 2007, a male afflicted with foot-and-mouth
disease was observed in Bandipur (F. S. Ahrestani, in litt.); and
2 B. gaurus died of foot-and-mouth disease in Bannerghatta
Park, India, after a major outbreak of the disease in domestic
cattle in neighboring villages (Chandranaik et al. 2015). Thus
far, there has been only 1 report of a B. gaurus dying from
anthrax (Peacock 1933).
Mycobacterium paratuberculosis was found in “histopath-
ological sections of the intestine mucous” during the autopsy
of an adult male B. gaurus that fell ill and subsequently died
in Palamau Tiger Reserve (Lal and Ashraf 1993). In Kanha,
a B. gaurus female autopsied was lightly infested by nema-
todes (Oesophagostomum radiatum) in the large intestine and
trematodes (Gastrothylax crumenifer) in the rumen, and ticks
(Boophilus microplus) were found on 2 adults that were checked
for ectoparasites (Schaller 1967). Histologic examination of 2
B. gaurus that died in the Oklahoma City Zoo, United States,
showed that they had Sarcosporidiosis, an infection caused
by the intracellular protozoan parasite Sarcocystis (Welch and
Zimmer 1981).
Interspecific interactions.—There have been no reports
of Bos gaurus fighting with other herbivores, and in general,
B. gaurus is considered to be tolerant of other herbivores. While
old bachelor males are tolerant of the presence of humans,
females and herds are generally shy of humans, avoiding con-
tact as much as possible. There have been, however, reports of
humans being charged by B. gaurus, and even fatalities from
such attacks. Reports of such encounters have been related
to B. gaurus charging after being shot or wounded (Forsyth
1889), after being inadvertently surprised by people in forests
(Inverarity 1889; Morris 1952, 1953), after being injured by
tigers (K. U. Karanth, pers. comm.), and when disturbed by
close-up flash photography (F. S. Ahrestani, in litt.).
Bos gaurus calves are preyed on by leopards Panthera par-
dus, dhole Cuon alpinus, and tigers, although adults are preyed
on primarily by tigers (Ahrestani et al. 2011). Leopards and tigers
typically kill calves and females by asphyxiation using bites to
their throats; 88% of 33 B. gaurus that were recorded killed by
tigers were found with bites to their throats in Nagarahole, India
(Karanth and Sunquist 1995). Males, however, have a thick neck
and often a dewlap, which makes killing them by asphyxiation
difficult. Tigers have been known to overcome this by biting the
hock, thereby hamstringing and bringing down male B. gaurus
(Dunbar-Brander 1923).
Records of tigers preying on calves and adults, including sol-
itary males, date back to nearly a century ago (Dunbar-Brander
1923). Reports from different areas suggest that this behavior
has not changed: the cause of death of B. gaurus in Kanha in
1964 was attributed mainly to tiger predation (Schaller 1967);
a carcass of an adult female B. gaurus killed by a tiger, and the
remains of B. gaurus were regularly found in tiger feces within
Eravikulam National Park, India (Rice 1986); analysis of scats
of the primary predators in Mudumalai Tiger Reserve, India,
showed that B. gaurus made up 0.19% of dhole diet and about
5% of tiger diet (Johnsingh 1983); and adult female and adult
male B. gaurus comprised 23% and 15% of tiger kills, respec-
tively, in Nagarahole, India (Karanth and Sunquist 1995, 2000).
Bos gaurus has been observed facing the direction of a
perceived threat with a raised head, raised muzzle, and flared
nostrils (Schaller 1967; Krishnan 1972; Belsare et al. 1984;
F. S. Ahrestani, in litt.). In such situations, males and females
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50(959)—Bos frontalis and Bos gaurus MAMMALIAN SPECIES 11
often emit short, deep, and loud grunts and snorts that have
been described as follows: “a violent and rapid expulsions
of air through the nostrils” (Hubback 1937:275); “a trumpet-
like blast of air through the nose, accompanied occasionally
by a growling sound—a harsh, rolling bru-u-u-u” (Schaller
1967:186); and “a call spelled pff-hong; the pff is the noise
made by the rush of air past the lips before the note is struck”
(Dunbar-Brander 1923:151). These snorts or grunts often
“raise every head in the herd” (Dunbar-Brander 1923:151; F. S.
Ahrestani, in litt.). If a herd has calves, the females in the herd
are known to bunch together in a muskox fashion, i.e., backs
to each other and facing outwards, and have been observed
leaping and kicking their hind legs, snorting and tossing their
horns, and holding their heads low and advancing, sometimes
rapidly toward detected leopards and tigers and other perceived
threats (Krishnan 1972; Johnsingh 1983; Karanth 1984; Tyabji
1989; F. S. Ahrestani, in litt.).
When fleeing from danger, which it often does, B. gaurus
has been observed giving “a series of rather stiff-legged bounds,
some 2 to 8 in number, with their forelegs brought down hard in
unison to produce a series of distinct thumps. After thumping the
ground, they usually trot off without making further sounds other
than the usual commotion of a large animal moving through the
forest” (Schaller 1967:186). Finally, males are capable of kill-
ing tigers: corpses of a dead tigress and a male were both found
after what was evidently a fight to death (Blackburn 1934), and
a radio-collared tiger that was found dead was apparently killed
by a B. gaurus (K. U. Karanth, pers. comm.).
The mithun/gayal/mithan Bos frontalis, the domestic form
of Bos gaurus, is found in northeastern India, Bhutan, Myanmar,
and China. B. frontalis looks very similar to B. gaurus: the males
have small dewlaps and a pronounced dorsal ridge, and both
sexes have white lower legs. However, B. frontalis differs from
B. gaurus by being slightly smaller in size and by the shape of
its horns, i.e., horns of both male and female B. frontalis extend
outwards and are straight, with only a gentle curve inwards. In
contrast, the horns of female B. gaurus barely extent outwards
and curve significantly inwards, and though the horns of male
B. gaurus extend outwards, they curve noticeably inwards at
their ends. The domestication of B. frontalis might be as old as
the Indus Valley civilization (Clutton-Brock 1987), but B. fron-
talis are rarely milked, are rarely used as draught or plough ani-
mals, and are often allowed to range free, though some choose
to return to village pens for the night. B. frontalis is primarily
used as a status symbol, to barter for goods, pay for brides, and
is often eaten after being sacrificed for various occasions, includ-
ing weddings, burials, prayers to ward off misfortune etc.
Bos gaurus males have been found mating with B. frontalis
females, and there have been reports of B. frontalis × B. gaurus
hybrids (Baker 1890; Gee 1964; Simmons 1984). Beginning
in 1983, a B. gaurus male was interbred with Sahiwal Friesian
dairy heifers on a Malaysian Government, Department of
Veterinary Services farm, and these hybrids outgrew other dairy
calves (Kamalludin 2009). B. frontalis and cattle have been
deliberately interbred in Bhutan, and all cases of interbreeding
between B. gaurus and B. frontalis and between B. gaurus and
cattle have resulted in only a few fertile females and always in
infertile males (Simmons 1984).
Initial attempts to breed B. gaurus in captivity failed
(Forsyth 1889). The first successful and consistent breeding of
B. gaurus occurred at the National Zoological Park, Washington,
D.C., United States, when a male and female—obtained from
Mysore, India, in 1937—produced 13 offspring (1940–1957) till
the male died (Crandall 1964). Since then, there have been over
319 B. gaurus births in European zoos, an equal number have
been born in US zoos, and there have been over 50 births in
Indian zoos. Today, the largest captive B. gaurus population is
in Omaha Zoo (there over 70 individuals housed in 2 subpopula-
tions), and across India, there are over 70 in captivity. Many cap-
tive populations have a high degree of inbreeding. For example,
all 23 B. gaurus that were present in Mysore Zoo in 2005 were
bred from 3 founder individuals that were captured 15 years ago
in the wild. The male:female ratio in inbred captive populations
has been skewed toward males, 62:38, in contrast to the 46:54
male:female ratio found in captive populations that were not
inbred (Hintz and Foose 1982).
The 1st group of Bos gaurus individuals relocated from
Kanha Tiger Reserve to Bandhavgarh Tiger Reserve, India, in
2011 were immobilized before relocation using a combination
of etorphine hydrochloride and azaperone, and the 2nd group of
individuals relocated in 2013 were immobilized before reloca-
tion using a combination of thiafentanil and ketamine (Nigam
et al. 2014; P. Nigam, in litt.).
Grouping behavior.Bos gaurus is predominantly a herd-
ing animal (Fig. 7). Females of all ages and males below the age
of 3 years are nearly always found in herds, and adult males are
found both within herds and alone (Inverarity 1889; Hubback
1937; Morris 1937; Schaller 1967). Herds of only adult males
are uncommon, though in 2007, 16 males (> 4 years) were
once observed together in Mudumalai, India (F. S. Ahrestani,
in litt.). Given the absence of detailed studies of B. gaurus, it is
unclear how herds are formed, maintained, change over time,
are organized socially and hierarchically, and how individuals
in a herd are related to each other.
The majority of solitary males are old, black bulls, which
suggests that the solitary nature of males increases with advanc-
ing age. As solitary adult males appear to be fearless of man,
and mostly anything else too, they are a common sight in areas
with high densities of B. gaurus. For example, 137 of 385 (35%)
encounters of B. gaurus over a 12-month period (2006–2007) in
the Bandipur–Mudumalai landscape were of solitary adult males
(F. S. Ahrestani, in litt.), and in other studies from Mudumalai,
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12 MAMMALIAN SPECIES 50(959)—Bos frontalis and Bos gaurus
48% of males observed in 2008–2009 (Ramesh et al. 2012) and
52% of all B. gaurus observed in 1976–1978 were solitary adult
males (Johnsingh 1983).
Black bulls, however, are not always solitary. Of 200 herds
tallied in Kanha, about 50% had a black bull (Schaller 1967),
and 56 of 147 herds encountered in the Bandipur–Mudumalai
landscape had a black bull (F. S. Ahrestani, in litt.). Herds often
include 2–3 black bulls and sometimes even include 6–8 adult
males (Schaller 1967; F. S. Ahrestani, in litt.). The structure of
herds with adult males, especially black bulls, is most proba-
bly fluid; there is, however, not enough information to under-
stand when adult males join herds, how long they remain in a
herd, when they leave a herd, and how such behavior changes
with age.
Mean herd size is about 6–7 and varies by study and loca-
tion: 7.8 (± 0.51 SE) in Mudumalai, India (Ashokkumar et al.
2010; Ramesh et al. 2012); 4 (± 0.21 SE) in Trishna Wildlife
Sanctuary, India (Dasgupta et al. 2008); 4.6 (± 0.29 SE) in Pench
Tiger Reserve, India (Sankar et al. 2002); 6 in Parambikulam
Tiger Reserve, India (Vairavel 1998); 6.9 in Nargarahole Tiger
Reserve, India (Karanth and Sunquist 1992); 6 in Palamau
Wildlife Sanctuary, India (Sahai 1977); 6.5 (Belsare et al. 1984)
and about 9 (Schaller 1967) in Kanha Tiger Reserve, India; 5.5
(± 5.2 SD) in Vietnam (Nguyen 2009); about 11 in Malaysia
(Hubback 1937); and 10–20 in Burma (Peacock 1933). Large
(> 50 individuals) aggregations of B. gaurus have also been
observed (Sanderson 1912; Mustill 1938; Johnsingh 1983; F. S.
Ahrestani, in litt.). These large aggregations appear to be tem-
porary composite herds, i.e., multiple smaller herds coming
together to form 1 large herd, most likely to exploit a favorable
forage resource in an area.
Although understood to be an uncommon occurrence, males
do fight each other (Inverarity 1889; Krishnan 1972; Schaller
1967). One such confrontation involved a pair of males fight-
ing next to a B. gaurus herd, with the pair displaying at each
other, thumping the ground and snorting repeatedly, the younger
of the 2 repeatedly threatening the older combatant with horns
held low, and the 2 males making contact with their heads mul-
tiple times, pushing against each other, and once twisting their
entwined horns (Johnson 1986). The confrontation finally ended
by the older male managing to push the younger male down a
slope, and the younger male was found limping a couple of days
later. Another report describes “2 bison fighting, shoving each
other about, and when disengaged, swinging their heads from
side to side with a twisting motion, and in so doing giving each
other terrific blows on the horns” (Dunbar-Brander 1923:150).
Schaller (1967) observed that the bony boss (the forehead
between the horns) of the males absorbed much of the head-
butting in male–male confrontations.
Adult males have been observed making a lateral (oblique)
end-on display, which appears to be a nonphysical method
to establish dominance (Krishnan 1972). This display, first
described by Schaller (1967:192), is “when a bull B. gaurus is
seen standing still with its feet together and back held somewhat
hunched, in the presence of another bull.” Such lateral displays
can last several minutes. Adult males also threaten each other
by lowering their heads and walking up to opponents, laterally
sweeping their horns, emitting low moans and grumbles, snort-
ing, thumping the ground with hooves, and sometimes running
or walking around in circles while swiping at undergrowth with
their lowered horns (Dunbar-Brander 1923; Krishnan 1972;
Belsare et al. 1984; F. S. Ahrestani, in litt.). Although older
black bulls do not commonly spar with each other, younger
(3–5 years) adult males spar by facing each other with lowered
heads (Fig. 8), locking horns, and then twisting their locked
heads from side to side (Schaller 1967; F. S. Ahrestani, in litt.).
Schaller observed 81 sparring incidents during his study, the
majority of which (48%) were between young adult males, 10%
were between young adult males and adult females, and 14%
were between adult females. There are also reports from over a
century ago of adult males sparring with their horns and butting
heads (Inverarity 1889).
Reproductive behavior.—Adult males and females have
been observed licking each other, and it is assumed such
Fig. 8.—Two (3- and 6-year-old) captive male Bos gaurus sparring in Mysore Zoo, southern India. Photographer F. S. Ahrestani.
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50(959)—Bos frontalis and Bos gaurus MAMMALIAN SPECIES 13
behavior is related to courtship. Schaller (1967:198) reported
that “cows and bulls lick each other’s necks, shoulders, and
rump, occasionally for as long as 10 min without interruption,
and bulls also lick each other at times, in 5 out of 7 instances
the subordinate individual licked the individual of higher
rank.” B. gaurus males exhibit flehmen, i.e., raising the muzzle
and curling the lips after licking a female’s vulva, or sniffing
a female’s urine or feces, a habit that is common among ungu-
lates (Schaller 1967). Males have also been observed tend-
ing to females—a behavior originally described for American
bison (McHugh 1958)—which is when a male stands beside a
female while she grazes and then follows her step for step as
she moves and grazes (Schaller 1967).
Females separate from their herd to give birth alone and have
been known to stay away for as long as 4 days (Sanderson 1912).
Mothers tend to thoroughly lick calves directly after their birth
(Krishnan 1972). Calves below the age of 2 months have been
found alone lying motionless in undergrowth with their necks
stretched out close to the ground that appear to be attempts to
remain concealed (Inverarity 1889; Hubback 1937; Ahrestani
et al. 2011). Calves and yearlings (< 18 months) are always seen
in herds with adult females, and a young calf (< 3 months) is
nearly always found at the heels of an adult female, presumably
its mother.
Communication.—Older (black) bulls make a call, pre-
sumably a rutting call, that is so unique that it distinguishes an
adult male Bos gaurus from other animals as much as its other
prominent features do, such as the dorsal ridge and dewlap.
When this call is heard, it is common to hear other black bulls
responding with the same call. The black bull begins the call
by keeping its head low, and as the call progresses, it raises
its head slowly and stretches out its neck, pointing its muz-
zle upwards with lips partly open and the whites of its eyes
showing with the effort. The call has been described by many
naturalists in their own words: “as the most absurd piping or
whistling sound, more like the call of a bird than anything else,
and absurd because such a strange sound emanates from an
animal so large and powerful” (Dunbar-Brander 1923:151); “a
most peculiar sound, a cross between the bugling of a wapiti
(elk) and the trumpeting of an elephant, but at the same time a
melodious sound that carries a long way” (Hubback 1937:275);
“a song, because of its musical qualities that is not low but
can be heard a mile away” (Schaller 1967:195); and “a long
drawn, resonant, low and high pitched at the same time, not
necessarily loud if heard at close quarters, but has the abil-
ity to be heard one-half a mile away” (Krishnan 1972:335).
The call has been described in detail as follows: “a clear, reso-
nant u-u-u-u-u about one to 3 seconds long, either constant in
pitch or slightly rising and falling; this note may be followed
by a second one somewhat lower in tone, by a third one still
lower, and so forth, giving the impression of someone prac-
ticing musical scales; and as many as ten seconds sometimes
elapse between the 1st and last note (Schaller 1967:195).” In
contrast to adult males, females, besides snorting during fear
and threatening behavior, are only known to moo, sounding
just like cattle (Dunbar-Brander 1923; Krishnan 1972; F. S.
Ahrestani, in litt.). It is still unclear, however, if the mooing
is a mother communicating to its calf or is communication for
some other reason.
Miscellaneous behavior.—Similar to cattle, Bos gaurus-
grooms itself by licking its hide; rubbing its neck, rump, and sides
against trees; or scratching itself with the tips of its horns. When
B. gaurus lies down, it generally does so with its legs folded
under its body, but it will occasionally also lie flat on its side with
legs stretched out. B. gaurus generally lies down under the shade
of trees and within swards of tall grass, where it is difficult for it
to be detected in the flickering shadow of trees (Forsyth 1889).
The amplification of the cytochrome b(Cyt b) gene, used
as a mitochondrial DNA genetic maker, produced a 154- and
603-bp fragment in the DNA sequence of a Bos gaurus from
Malaysia (Romaino et al. 2014). Testing 130 cattle micro-
satellite markers on a panel of 11 individual B. gaurus from
Vietnam found amplification of 117 markers (90%) with a total
of 264 alleles (Nguyen et al. 2007). Of the 117 makers, 68
were polymorphic that had 2–6 alleles per locus, and 3 cattle
Y chromosome microsatellite markers (INRA124, INRA126,
and BM861) were specific to B. gaurus (Nguyen et al. 2007).
Sequencing the entire genome of an individual, the domes-
tic form Bos frontalis detected 23,828,562 single-nucleotide
polymorphisms (SNPs) and identified 16,901 breed-specific
nonsynonymous SNPs among 6,167 genes (Mei et al. 2016).
Annotation of these SNPs showed that 78.2% of the SNPs
were located in intergenic regions; 21.1% were located in
genic regions, including intronic regions, splicing sites, exonic
regions, and untranslated regions; and the remaining 0.7% were
located in upstream or downstream regions. This sequenc-
ing found the homozygous/heterozygous ratio to be 1:0.8, and
Nei’s unbiased mean heterozygosity and the mean allele num-
ber across loci were 0.23 and 2.2, respectively (Mei et al. 2016).
Another cytogenetic study of the domestic form B. frontalis
found the karyotype of the female B. frontalis (n = 4) comprised
58 chromosomes, including 54 acrocentric and 4 large submeta-
centric chromosomes (Qu et al. 2012).
A genetic analysis of a B. gaurus female at Toronto Zoo
detected a chromosome anomaly in an individual that had
2n = 57 chromosomes (an extra submetacentric, but 2 less acro-
centric chromosomes) instead of the normal 2n = 58 found in
B. gaurus, which consists of 27 pairs of acrocentric chromo-
somes, 1 pair of submetacentric chromosomes, and the submeta-
centric sex chromosomes (Mastromonaco et al. 2004).
The genotype data collected from 117 successfully ampli-
fied microsatellites used to assess the genetic diversity within
the reaming Bos gaurus population in Vietnam—estimated to
be no more than 500 individuals—found the mean polymor-
phic information content (PIC) to be 0.252 (range: 0.083–
0.767) and mean heterozygosity (Ho) to be 0.269 (range:
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14 MAMMALIAN SPECIES 50(959)—Bos frontalis and Bos gaurus
0.091–0.909—Nguyen et al. 2007). Multiple studies in the
last decade have genetically investigated whether B. frontalis
is a domestic version of B. gaurus, and the majority of these
studies have found evidence that this is indeed the case. These
studies include sequencing the Cyt b genes of 33 B. frontalis
from Myanmar and Bhutan and finding B. gaurus haplotypes
in 28 of these individuals (Tanaka et al. 2011); sequencing the
16S rRNA gene in the mitochondrial DNA of mithun from
Bhutan that demonstrated phylogenetic proximity to B. gaurus
(Dorji et al. 2010); and sequencing the Cyt b genes of 11 gayal
from Yunan, China, found 6 haplotypes that clustered around
B. gaurus and other domestic cattle species such as Bos tau-
rus and Bos indicus (Li et al. 2008). In contradiction to the
above studies, a study that sequenced the Cyt b genes from 28
B. frontalis from Yunnan, China, 13 B. frontalis from Arunachal
Pradesh, India, and 1 B. gaurus from Yunnan, China, found the
genome sequence of B. frontalis to differ from the B. gaurus
(Baig et al. 2013). This individual study, however, does not
provide enough conclusive evidence to override all the other
genetic evidence and the detailed morphological comparisons
that collectively strongly suggest that the B. frontalis is indeed
the domestic derivative of B. gaurus.
Bos gaurus is one of the first mammals to have been
cloned. Somatic cells from the skin of a male were successfully
electrofused with enucleated oocytes from domestic females
(Lanza et al. 2000). One of these embryos successfully devel-
oped in a surrogate domestic female. The calf that was success-
fully delivered, however, developed a fatal bacterial infection
2 days after birth and died on 8 January 2001.
The global Bos gaurus population is estimated to be 13,000–
30,000 and is projected to decline by 30% over the next 3 decades
(Duckworth et al. 2016). B. gaurus is listed as “Vulnerable” by the
International Union for Conservation of Nature, and its greatest
threats are loss of habitat, being poached for its meat and horns,
and contracting fatal diseases from overlapping cattle popula-
tions. Country-wide populations in Bangladesh, Cambodia,
China, Laos, Malaysia, Vietnam, and Thailand have declined
by over 70% in the last 2–3 decades, although the decline in
India and Nepal have been considerably lower. Recent poaching,
mainly for meat and horns, has decimated the B. gaurus popula-
tion in Malaysia, and there is a fear that the hunters responsible
for this catastrophe could be soon targeting populations in other
countries. Conservation action in Thailand has recently shown
that it is possible to stall the decline of B. gaurus populations
prevalent across southeastern Asia. On a positive note, multiple
protected areas in southern India have recorded increasing trends
in their B. gaurus populations over the last couple of decades.
In 1995, the last remaining herd of B. gaurus left Bandhavgarh
Tiger Reserve. To restore B. gaurus in Bandhavgarh, 19 indi-
viduals in January 2011—5 males (3 subadults, 2 adults)
and 14 females (1 yearling, 5 subadults, and 7 adults)—were
translocated from Kanha Tiger Reserve to Bandhavgarh. Since
the reintroduction, the newly established B. gaurus population
in Bandhavgarh has been thriving and multiplying.
I am grateful to S. Vaidyanathan, Senior Scientist at the
Foundation for Ecological Research, Advocacy and Learning, India,
for developing the distributional map of Bos gaurus. I am grateful
to E. Westwig for helping with my photography of B. gaurus skulls
at the American Museum of Natural History. I thank K. Varma and
M. N. Naveen for letting me use their photographs.
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... The mighty gaur (Bos gaurus), also known as the Indian bison, is the largest species of wild cattle and is at risk of becoming endangered in the near future. The gaur can attain up to 198 cm shoulder height [1], weigh up to 900 kg, has white stockings on the legs, and possesses spiral-shaped horns [2] (Fig. 1a) to protect itself against predators such as tigers. It is native to South and Southeast Asia and listed as vulnerable on the International Union for Conservation of Nature (IUCN) Red List [3]. ...
... Based on morphological characteristics the species is classified into three subspecies, B. gaurus gaurus, B. gaurus readei and B. gaurus hubbacki [2]. B. gaurus gaurus is mainly found in India, Nepal and Bangladesh; B. gaurus readei inhabits China and Myanmar; B. gaurus hubbacki is mainly found in Malaysia and occurs in two distinct forms, one with well-developed dewlap and one without [6]. ...
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Background The gaur ( Bos gaurus ) is the largest extant wild bovine species, native to South and Southeast Asia, with unique traits, and is listed as vulnerable by the International Union for Conservation of Nature (IUCN). Results We report the first gaur reference genome and identify three biological pathways including lysozyme activity, proton transmembrane transporter activity, and oxygen transport with significant changes in gene copy number in gaur compared to other mammals. These may reflect adaptation to challenges related to climate and nutrition. Comparative analyses with domesticated indicine ( Bos indicus ) and taurine ( Bos taurus ) cattle revealed genomic signatures of artificial selection, including the expansion of sperm odorant receptor genes in domesticated cattle, which may have important implications for understanding selection for male fertility. Conclusions Apart from aiding dissection of economically important traits, the gaur genome will also provide the foundation to conserve the species.
... Further challenges to the assessment of the association between domestication and neuropeptide gene changes stem from the domestication classification of some species and the genome assembly quality of other species. With respect to domestication assignments, for example, the gayl (Bos frontalis) is often considered the domestic form of the wild gaur (Bos gaurus) [67], however the degree of domestication is highly variable among the former group. With regards to sequence quality, the virtually identical nucleotide or protein sequences predicted from all camelid genomes available in this study invalidates the conclusion of adaptive introgression of endothelin 3 (EDN3) in South American camelids [6]. ...
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The impact of evolution and domestication processes on the sequences of neuropeptide prohormone genes that participate in cell–cell signaling influences multiple biological process that involve neuropeptide signaling. This information is important to understand the physiological differences between Cetartiodactyla domesticated species such as cow, pig, and llama and wild species such as hippopotamus, giraffes, and whales. Systematic analysis of changes associated with evolutionary and domestication forces in neuropeptide prohormone protein sequences that are processed into neuropeptides was undertaken. The genomes from 118 Cetartiodactyla genomes representing 22 families were mined for 98 neuropeptide prohormone genes. Compared to other Cetartiodactyla suborders, Ruminantia preserved PYY2 and lost RLN1. Changes in GNRH2, IAPP, INSL6, POMC, PRLH, and TAC4 protein sequences could result in the loss of some bioactive neuropeptides in some families. An evolutionary model suggested that most neuropeptide prohormone genes disfavor sequence changes that incorporate large and hydrophobic amino acids. A compelling finding was that differences between domestic and wild species are associated with the molecular system underlying ‘fight or flight’ responses. Overall, the results demonstrate the importance of simultaneously comparing the neuropeptide prohormone gene complement from close and distant-related species. These findings broaden the foundation for empirical studies about the function of the neuropeptidome associated with health, behavior, and food production.
... Gaur utilize agricultural fringes mostly at night to feed and leave to adjacent secondary forest habitat early morning to avoid conflict (Chaiyarat et al., 2021). Gaur are generally shy and avoid contact as much as possible (Ahrestani, 2018). ...
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Aim: To predict the distribution of suitable habitats for Malayan gaur (Bos gaurus) at a highly fragmented forest area in Peninsular Malaysia and to identify the potential connectivity between suitable habitat patches. Methodology: Maximum entropy (MaxEnt) approach was used to predict the distribution of suitable habitats of the Malayan gaur. Gaur presence-only data and six environmental variables were collated for the habitat suitability modeling, and area under curve (AUC) value was used to estimate the performance of the model. The resulting model was then used to derive a potential connectivity map through least-cost analysis using Corridor Designer toolbox in ArcGIS 10.4. Results: The AUC value of the habitat suitability model was 0.84. Distance from urban areas indicated the highest relative contribution to the model (26.9%), followed by distance from water body (24.2%) land use (18.0%) elevation (14.3%), slope (14.0%) and lithology (2.6%). Predicted suitable habitats for gaur were found mostly in lowland forest areas, especially in the vicinity of rivers within forest reserves. A total of five wildland blocks were derived from the habitat suitability model, and several potential corridor swaths were identified connecting the wildland blocks. Interpretation: The absence of gaur occurrence in suitable habitats suggest that fragmented habitats greatly affected gaur distribution and population. Road network and agricultural lands are the major barriers of gaur movement as they are very sensitive towards disturbances and conflict. Thus, this research proposes potential connectivity at a regional scale for Malayan gaur for use in future planning in conservation, management and development.
... Ungulate species have been found to have different food and feeding habits 43 . Gaur, have been described as grazers 42,44,45 , browsers 46 and generalists 47 depending on habitat types. ...
The presence of gaur (Bos gaurus) at the border of Khao Yai National Park (KYNP) in Thailand has resulted in a dramatic increase in the number of individuals' crop feeding. This study examines the feeding adaptations of gaur at the edge of the protected area and assesses whether gaur response to increased nutrient availability in crop plants compared to natural forage. During the day, gaur mostly utilized forest areas in KYNP and entered the agricultural areas at night. Gaur ate 43 natural forage species. Natural forage species contain high levels of crude protein and lipid, but they are found in small quantities and scattered areas when compared to crop plants, especially Zea mays L., that are available in large quantity and are heavily foraged on by gaur. However, greater understanding of the electivity index and nutrition of forage species along the edge of the protected area can be used to reduce the gaur-human conflict by keeping gaur in KYNP. Reducing the large monoculture areas that is the food sources of gaur along the edge may reduce or prevent gaur leaving the park and can be applied to advance conservation actions. Human-wildlife coexistence at the edge of protected areas can create problems that are referred to as human-wildlife conflicts 1. In general, specialist species are more affected by habitat modification than are generalist species. Moreover, some species are able to change to forage on food species that are more readily available when their preferred forage species are scarce 2 , thereby using crops as an alternative food source. Some crops are attractive to wild animals and provide both energy and nutrition 3. However, this subject is poorly studied, especially in the large bovids of tropical environments. Gaur (Bos gaurus), family Bovidae (Fig. 1), is globally vulnerable 4 , and protected under the Thai Reserved and Protected Animals Act, B.C.2562 5. Gaur are distributed in scattered areas of Bhutan,
... Ungulate species have been found to have different food and feeding habits 43 . Gaur, have been described as grazers 42,44,45 , browsers 46 and generalists 47 depending on habitat types. ...
Full-text available
The presence of gaur ( Bos gaurus ) at the border of Khao Yai National Park (KYNP) in Thailand has resulted in a dramatic increase in the number of individuals’ crop feeding. This study examines the feeding adaptations of gaur at the edge of the protected area and assesses whether gaur response to increased nutrient availability in crop plants compared to natural forage. During the day, gaur mostly utilized forest areas in KYNP and entered the agricultural areas at night. Gaur ate 43 natural forage species. Natural forage species contain high levels of crude protein and lipid, but they are found in small quantities and scattered areas when compared to crop plants, especially Zea mays L., that are available in large quantity and are heavily foraged on by gaur. However, greater understanding of the electivity index and nutrition of forage species along the edge of the protected area can be used to reduce the gaur-human conflict by keeping gaur in KYNP. Reducing the large monoculture areas that is the food sources of gaur along the edge may reduce or prevent gaur leaving the park and can be applied to advance conservation actions.
Full-text available
The gaur ( Bos gaurus ) is found throughout mainland South and Southeast Asia but is listed as an endangered species in Thailand with a decreasing population size and a reduction in suitable habitat. While gaur have shown a population recovery from 35 to 300 individuals within 30 years in the Khao Phaeng Ma (KPM) Non-Hunting Area, this has caused conflict with villagers along the border of the protected area. At the same time, the ecotourism potential of watching gaurs has boosted the local economy. In this study, 13 mitochondrial displacement-loop sequence samples taken from gaur with GPS collars were analyzed. Three haplotypes identified in the population were defined by only two parsimony informative sites (from 9 mutational steps of nucleotide difference). One haplotype was shared among eleven individuals located in different subpopulations/herds, suggesting very low genetic diversity with few maternal lineages in the founder population. Based on the current small number of sequences, neutrality and demographic expansion test results also showed that the population was likely to contract in the near future. These findings provide insight into the genetic diversity and demography of the wild gaur population in the KPM protected area that can inform long-term sustainable management action plans.
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Significance We undertook an ancient genomic DNA investigation of large animal remains dated ∼5,200 y B.P. from the Tibetan Plateau. We provide compelling evidence that the present-day low-latitude tropical inhabitants Bos gaurus and Dicerorhinus sumatrensis once roamed as far north as the margin of the northeastern Tibetan Plateau (NETP) during the late Neolithic, pushing the historical gaur distribution from ∼29°N to ∼34°N. Further multidisciplinary exploration indicates that a high summer temperature in the late Neolithic might have facilitated the northward expansion of these tropical animals to the NETP, which enriched the biodiversity of wildlife and contributed to the exploration of the Tibetan Plateau as one of the last habitats for hunting game in East Asia.
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The gaur is the largest extant cattle species and distributed across South and Southeast Asia. Around 85% of its current global population resides in India, however there has been a gradual decrease in the gaur population over the last two decades due to various anthropogenic activities. Mitochondrial genome is considered as an important tool for species identification and monitoring the populations of conservation concern and therefore it becomes an obligation to sequence the mitochondrial genome of Indian gaur. We report here for the first time 16,345 bp mitochondrial genome of four Indian gaur sequenced using two different approaches. Mitochondrial genome consisted of 13 protein-coding genes, 2 rRNA genes, 22 tRNA genes, and a control region. Among the 37 genes, 28 were positioned on the H-strand and 9 were positioned on the L-strand. The overall base composition appeared to be 33.5% A, 27.2% T, 25.9% C and 13.4% G, which yielded a higher AT content. The phylogenetic analysis using complete mitochondrial genome sequences unambiguously suggested that gaur is the maternal ancestor of domestic mithun. Moreover, it also clearly distinguished the three sub species of B. gaurus i.e. B. gaurus gaurus, B. gaurus readei and B. gaurus hubbacki. Among the three sub species, B. gaurus gaurus was genetically closer to B. gaurus readei as compared to B. gaurus hubbacki. The findings of our study provide an insight into the genetic structure and evolutionary history of Indian gaur.
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The primary prey of tigers across much of South‐East Asia has been depleted, reducing the ability of already limited habitat to support tigers. To better understand the extent to which two of the largest prey species, gaur (Bos gaurus) and banteng (Bos javanicus), contribute to the tiger's diet, we estimated the average size of these species killed by tigers. This information is needed to more accurately calculate biomass of these species in the tiger's diet and to devise strategies to increase tiger carrying capacity where habitat is fragmented and limited in west‐central Thailand. We used temporally clumped locations of 24 satellite radio‐collared tigers to identify their kill sites and obtained mandibles from 82 gaur and 79 banteng. Kills were aged by teeth eruption sequence, sectioning the M1 molar and counting cementum annuli. Of all gaur killed, 45.2% were adults; of all banteng killed, 55.7% were adults. The average weight of banteng killed was 423.9 kg, which was similar to the 397.9 kg average weight for gaur. The mean weight of both prey species is 3.5–4.5 times greater than the predicted 1:1 preferred prey to predator ratio. In the absence of medium‐sized prey, killing these larger animals may be especially critical for female tigers provisioning nearly independent young when male offspring are already larger than the mother. This is the first study to present data on the average weights of gaur and banteng killed in South‐East Asia, and these results suggest that these are key prey species to target in tiger prey recovery efforts.
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Gayal (Bos frontalis) is a semi-wild and endangered bovine species that differs from domestic cattle (Bos taurus and Bos indicus), and its genetic background remains unclear. Here, we performed whole-genome sequencing of one Gayal for the first time, with one Red Angus cattle and one Japanese Black cattle as controls. In total, 97.8 Gb of sequencing reads were generated with an average 11.78-fold depth and >98.44% coverage of the reference sequence (UMD3.1). Numerous different variations were identified, 62.24% of the total single nucleotide polymorphisms (SNPs) detected in Gayal were novel, and 16,901 breed-specific nonsynonymous SNPs (BS-nsSNPs) that might be associated with traits of interest in Gayal were further investigated. Moreover, the demographic history of bovine species was first analyzed, and two population expansions and two population bottlenecks were identified. The obvious differences among their population sizes supported that Gayal was not B. taurus. The phylogenic analysis suggested that Gayal was a hybrid descendant from crossing of male wild gaur and female domestic cattle. These discoveries will provide valuable genomic information regarding potential genomic markers that could predict traits of interest for breeding programs of these cattle breeds and may assist relevant departments with future conservation and utilization of Gayal.
Names Genus: Bos Linnaeus, 1758 Species: Gaur Bos gaurus C. H. Smith, 1827 Names in other languages: French: Gaur; German: Gaur; Spanish: Gaur; Italian: Gaur; Adi: Tadok; Burmese: Peeoug; Kannada: Kaati, Kaadu kona, Kaadu yemme; Kannada-in Uttara Kannada District kulga: Gameya; Northern Udupi district, Kannada: Duddu; Lao: Meuay; Malay: Seladang; Mandarin: Da-E-Ni, ???; Marathi: Raangawa; Malayalam: Kattu pothu; Nepali: Gauri gai; Tamil: Kaatu maadu; Vietnamese: Bò Tót; Telugu: Adavi dunna; Thai: Krating ?????; Gonds: Moo. Other common names: Indian bison Note: Although Bos frontalis (Lambert 1804), is often used as the scientific name for the gaur (see Grubb 2005), the International Commission of Zoological Nomenclature (2003) ruled that Bos gaurus be adopted as the name for the gaur despite Bos gaurus being antedated by Bos frontalis. Bos frontalis is used as the scientific name for the mithun, the domestic form of the gaur. Taxonomy The gaur is a member of the Bovini clade (tribe), one of three clades (the other two being Boselaphini and Tragelaphini) recognized to be part of the larger clade (subfamily) Bovinae; the Bovinae along with the Antilopinae comprise the family Bovidae (Bibi 2007). Members of the bovine lineage were present by 8.9 Ma, appear to have originated in Asia, south of the Himalayas, and are understood to have expanded their ancestral dietary niche to include roughage in more open environments by diversifying and extending their range in response to the vegetation changes in the late Miocene (Bibi 2007). Using 15 complete or partially sequenced autosomal genes, MacEachern et al. (2009) identified three lineages in the Bovini clade after it split from the Boselaphini and Tragelaphini clades: The buffalo clade (Bubalus and Syncerus species), the banteng Bos javanicus and gaur clade, and the cattle clade.
Biotelemetry transmitters were implanted in adult female gaur (Bos gaurus) at the Bronx Zoo/Wildlife Conservation Park to establish baseline heart rate and body temperature value ranges in unrestrained animals and to document changes in these physiological functions when animals were exposed to different environmental conditions. Heart rate and body temperature were monitored using similar, but distinct, telemetry systems. Mean heart rates for five gaur ranged from 49.3 to 57.7 beats/min, and the mean body temperatures for two animals were 38.2°C and 38.8°C. Short-duration adversive stimuli caused brief three-fold increases in heart rate, but baseline rates returned once the stressors were removed. Moving gaur to novel environments or pairing them with nonaffiliates also resulted in heart rate increases. Body temperature was not affected by short-term stressors but was positively correlated with ambient temperature. The onset of ovulation may be predictable based on temperature spikes exhibited by the gaur at 19-22 day intervals.
The Malayan gaur (Bos gaurus hubbacki) or Seladang is classified as vulnerable by the International Union for Conservation of Nature and Natural Resources (IUCN). The Malayan gaur is mainly distributed in the tropical woodlands of Peninsular Malaysia and Southern Thailand. The aim of this study was to collect, analyze and cryopreserve the semen of wild Malayan gaur. Transrectal massage (TM) and electroejaculation (EEJ) technique was applied in semen collection of the Malayan gaur. The semen was then cryopreserved in liquid nitrogen using slow freezing technique. Makler counting chamber was used to evaluate sperm concentration and motility, while the sperm viability and morphology of fresh and post-thaw sperm was determined using eosin-nigrosin staining protocol. As a result, we have successfully collected the Malayan gaur semen using EEJ technique. Sperm motility, viability and morphological changes of the post-thaw semen of Malayan gaur were found undesirable due to the complication of the cryopreservation process. On the basis of current study it can be concluded that Malayan gaur bulls semen can be obtain by EEJ with no evidence of rectal trauma. Optimization of the process of cryopreservation for Malayan gaur sperm is needed to maintain the cryoviability of the good sperm quality. The data generated in this study would be useful in conservation of genetic diversity program for Malayan gaur.