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Equus kiang (Perissodactyla: Equidae)
Author(s): Antoine St-Louis and Steeve D. Côté
Source: Mammalian Species, Number 835:1-11. 2009.
Published By: American Society of Mammalogists
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Equus kiang (Perissodactyla: Equidae)
ANTOINE ST-LOUIS AND STEEVE D. CO
De´partement de Biologie, Universite´ Laval, Que´bec, Que´bec G1V 0A6, Canada; email@example.com
´tudes Nordiques, Universite´ Laval, Que´ bec, Que´bec G1V 0A6, Canada
Abstract: Equus kiang Moorcroft, 1841, is an equid commonly called the kiang or Tibetan wild ass and is the only equid living
on the Tibetan Plateau. It is the largest of the wild asses, with a distinct dark-brown coat on the back, and 1 of the 7 species of
Equus. It is endemic to the high-elevation rangelands of China (Tibet, Xinjiang, Qinghai, and Gansu), India (Ladakh and
Sikkim), Pakistan (Khunjerab National Park), and Nepal (Mustang). It inhabits open steppes and rolling hills sparsely
vegetated with grasses and sedges. E. kiang is vulnerable to illegal poaching and competition with domestic livestock, and data
are insufficient to accurately estimate its total abundance. DOI: 10.1644/835.1.
Key words: China, equid, kiang, Ladakh, Qinghai, Tibet, Tibetan wild ass,
Published 27 May 2009 by the American Society of Mammalogists
Synonymy completed 12 June 2008 www.mammalogy.org
Equus kiang Moorcroft, 1841
Kiang or Tibetan Wild Ass
Equus kiang Moorcroft in Moorcroft and Trebeck, 1841:312.
Type locality ‘‘eastern parts of the country [5Ladakh],’’
state of Jammu and Kashmir, India.
Asinus equioides Hodgson, 1842:287. Nomen nudum (Grubb
Asinus polyodon Hodgson, 1847a:469. Type locality ‘‘Tibet;’’
restricted to the region north of Sikkim border (Groves
and Maza´k 1967:352).
Asinus hemionus Gray, 1852:272. Type locality ‘‘Thibet
Asinus kyang: Kinloch, 1869:13. Name combination and
incorrect subsequent spelling of Equus kiang Moorcroft,
Equus hemionus kiang Lydekker, 1904a:432. Type locality
‘‘Tibet and Ladak [5Ladakh].’’
Equus kiang holdereri Matschie, 1911:29. Type locality
‘‘[southwestern shore of] Kukunor’’ vide Groves and
Microhippus tafeli Matschie, 1924:68. Type locality ‘‘Tosson
Nor, Tibet’’ vide Ellerman and Morrison-Scott
Hemionus kiang: Trumler, 1959:17. Name combination.
Hemionus nepalensis Trumler, 1959:24. Type locality ‘‘Nepal.’’
CONTEXT AND CONTENT. Order Perissodactyla, suborder
Hippomorpha, family Equidae, subfamily Equinae. The
genus Equus has 7 living species: E. asinus,E. burchellii,E.
caballus,E. grevyi,E. hemionus,E. kiang,andE. zebra
(Bennett 1980; key in Grinder et al. 2006). E. quagga is
now extinct (Groves 1974). Three subspecies of E. kiang
have been identified, mainly on the basis of geographical
location, body size, and color differences (Groves and
Maza´k 1967; Neumann-Denzau and Denzau 2003; Shah
E. k. holdereri Matschie, 1911:29. See above (tafeli Matschie,
1924, is a synonym—Groves and Maza´k 1967); vernac-
ular name is eastern kiang.
E. k. kiang Moorcroft, 1841:312. See above (equioides
Hodgson, 1842, kyang Kinloch, 1869, and hemionus
Fig. 1.—An adult male Equus kiang (subspecies kiang) in summer
coat from eastern Ladakh, India (33u189N, 78u009E). Used with
permission of the photographer EJ. Van Gruisen.
MAMMALIAN SPECIES 835:1–11
kiang Lydekker, 1904, are synonyms); vernacular name
is western kiang.
E. k. polyodon Hodgson, 1847a:469. See above (kiang
nepalensis Trumler, 1959, is a synonym); vernacular
name is southern kiang.
Current population assessments commonly recognize
these 3 subspecies (Shah 2002). However, different opinions
on their validity exist in the literature. Wang (2002) only
recognized E. k. kiang and E. k. holdereri; Schaller (1998)
questioned subspecific status because the distribution of E.
kiang is continuous across its range and morphological
differences are small, which suggests a cline rather than 3
subspecies. Little genetic information exists for extant
populations of E. kiang (Schaller 1998).
NOMENCLATURAL NOTES. Taxonomy of wild asses and
hemiones, or ‘‘half-asses,’’ continues to generate debate.
Although it is now accepted that Equus kiang is a distinct
species, older literature often placed it as a subspecies of E.
hemionus (Asiatic wild ass—cf. E. h. kiang). In this account,
only information that could be specifically attributed to E.
kiang is included. For example, information on E. hemionus
from Tibet could be assumed to be about E. kiang, but
information on E. hemionus without any geographic
reference was not used.
There was historical confusion with identification of E.
kiang. In an extensive review of early discoveries in Central
Asia and Tibet, Hedin (1922) referred to the French
Franciscan friar William de Rubruck as the 1st European
to mention E. kiang from his travels in Central Asia during
the 13th century. However, because William de Rubruck
only traveled in Mongolia and never went to Tibet, he only
could have seen E. h. hemionus (Mongolian kulan) or E. h.
luteus (Gobi kulan), unless the range of E. kiang extended
into Mongolia in the 13th century, an unlikely proposition.
William Moorcroft identified E. kiang as a distinct species
in a letter to John Fleming in 1822 but spelled it Equus
Kia`ng (Moorcroft 1827). It appears that this source
constitutes the 1st mention of E. kiang as a species. More
recently, Trumler (1959) proposed the name Hemionus kiang
sinensis for a 4th subspecies based on a fossil equid
discovered by P. Teilhard de Chardin in the ‘‘Bassin du
Sjara-Osso-Gol’’ in northern China (Trumler 1959:23).
However, this name appears unavailable because it is not
clear whether it is associated to E. kiang or another related
In the recent literature, morphometric comparisons
between E. kiang and E. hemionus, based on tooth patterns
and skull measurements, revealed a close relationship
between the 2 forms, which led Eisenmann (1980, 1986) to
propose classifying E. kiang as a subspecies of E. hemionus.
A different opinion was expressed by Groves and Maza´k
(1967) and Groves (1974), who argued that the allopatric
separation between E. kiang and E. hemionus and their
biological and morphological differences were important
enough to consider E. kiang as a separate species. In their
view, morphological differences between the 6 recognized
subspecies of E. hemionus are small compared with
differences between E. kiang and the E. hemionus group.
The name kiang is Tibetan (spelled rkyang in Tibet—Huber
2005), but its meaning remains unclear (Gotch 1979; Groves
1974; Prejevalsky 1876). E. kiang is named skiang in central
Ladakh (Humbert-Droz and Dawa 2004; Pfister 2004) and
zang yelu in China (Smith and Xie 2008).
Equus kiang is the largest of the wild asses (Schaller
1998; Fig. 1). The closest relative of E. kiang is E. hemionus,
the Asiatic wild ass (Eisenmann 1980, 1986; Groves and
Maza´k 1967). E. kiang has a darker coat than E. hemionus
and is stockier, with shoulder height about 5% higher and
body mass about 30% greater than E. hemionus (Groves and
Maza´k 1967). The rump in E. kiang is narrow and descends
steeply, whereas the rump in E. hemionus is more rounded
(Groves and Maza´k 1967). Hooves of E. kiang are broader
and rounder than in other wild asses and similar to those of
E. caballus (Lydekker 1904b). There is a distinctive white
wedge behind the shoulder in E. kiang, which reaches high
on the back. The white color on the throat is more
prominent in E. kiang than in E. hemionus. Legs of E. kiang
are never striped (Groves and Maza´k 1967).
There are pronounced differences in the spring molting
pattern between E. kiang and E. hemionus.InE. kiang, hair
from underneath the belly is the last to be shed, whereas in
E. hemionus hair from the belly is shed very early during
molting (Maza´k 1962). The complete molt is achieved in
about 80 days in E. kiang, whereas it is completed in 40–45
days in E. hemionus (Maza´k 1962). The mane is longer in E.
kiang than in E. hemionus, and the dorsal stripe is about 13%
narrower (Groves and Maza´k 1967). The tail of adult E.
kiang has hairs starting at its base, a character only found in
young individuals of other wild ass species (Groves and
The lower jaw is more massive in E. kiang than in E.
hemionus with a convex lower border, and upper incisors are
more vertically implanted (Groves and Maza´k 1967).
Although generally similar, tooth patterns differ slightly
between the 2 species. The metaconid–metastylid valley of
the lower teeth is more penetrating, and the hypoconulid is
larger and more prominent in E. kiang than in E. hemionus,
similar to horses (Groves and Maza´k 1967). In early
descriptions of E. kiang, Hodgson (1847a, 1847b) reported
that dP1, often referred to as the ‘‘wolf tooth,’’ was an
exclusive characteristic of the species. However, it appears,
that this extra premolar occurs in only some E. kiang and
also is found in adults of other equid species (Eisenmann
1980; Groves and Maza´k 1967).
2 MAMMALIAN SPECIES 835—Equus kiang
Equus kiang has a chestnut-brown coat covering the
upper part of the thighs, back, upper flanks, upper part of
the hind legs, dorsal part of the neck, cheeks, and forehead;
undersides, throat, and insides of the ears are all white
(Fig. 1). The rostrum also is white, turning gray around the
mouth and nostrils. The mane and edges and tips of the ears
are black. A dark brown dorsal stripe extends from the mane
to the tip of the tail (Schaller 1998). Legs are generally white
but can be light brown on the front. A thin, dark stripe
separates hooves from legs. The coat is darker in winter, and
paler and more reddish in summer. The summer coat is
short, with hairs 14–16 mm long. Winter hairs are thicker
and longer, reaching 35–46 mm (Groves 1974; Maza´k 1962).
Equus kiang has a large head, a convex nose, and a thick
rostrum (Fig. 2). The snout is short (Bennett 1980). The
body is relatively short, with long limbs and broad horselike
hooves (Groves 1974). Chestnuts—bare skin growths
located above the knees—are present on the forelegs only.
Among the 3 recognized subspecies, E. k. holdereri is the
largest, with a long nasalia, toothrow, and diastema, and its
coat has a lighter brown color than E. k. kiang.E. k. kiang
has a comparatively short nasalia, toothrow, and diastema,
and its coat is dark brown, especially in winter. E. k.
polyodon is the smallest of the 3 subspecies, with a color
pattern similar to that of E. k. holdereri (Groves and Maza´k
1967). Adult E. kiang have the following body measurements
(cm): shoulder height, 132–142; body length, 182–214; length
of tail, 32–45; length of hind foot, 41–54; and length of ear,
22 (Smith and Xie 2008).
Equus kiang is endemic to the Tibetan Plateau of Central
Asia at elevations of 2,700–5,300 m (Schaller 1998; Shah
2002; Fig. 3). The western limit of its distribution lies on the
border of Khunjerab National Park in Pakistan (75u199E),
and its eastern limit occurs in Qinghai, China (102u549E). To
Fig. 2.—Dorsal, ventral, and lateral views of skull and lateral view
of mandible of a male Equus kiang (Field Museum of Natural
History specimen 182) from Tibet. Greatest length of skull is
568 mm. Used with permission of the photographer B.
Fig. 3.—Geographic distribution of Equus kiang. The bold line
delineates the species distribution. The distribution areas of the
proposed subspecies are indicated as follows: 1, E. k. holdereri;2,E.
k. kiang;3,E. k. polyodon;?5uncertain subspecies identification
(modified from Schaller 1998 and used with permission of
835—Equus kiang MAMMALIAN SPECIES 3
the south, E. kiang is limited mainly by the Himalayas
(27u539N) and to the north by the Kunlun-Arjin Shan
Mountains in Xinjiang, Qinghai, and Gansu, along the
northern edge of the Tibetan Plateau (40u209N). Approxi-
mately 95% of the population of E. kiang occurs in China,
mainly in Tibet but also in Xinjiang, Qinghai, and Gansu
(Schaller 1998). Outside of China, E. kiang occurs mainly in
the Ladakh region of India, and small numbers are found in
Sikkim (Fox et al. 1991; Shah 2002). E. kiang is reported in
the Dolpo and Mustang regions of Nepal in small numbers
(Paklina and van Orden 2003; Schaller 1998; Shah 2002). In
Pakistan, the species probably occurs only sporadically,
coming from Xinjiang (G. B. Schaller, pers. comm.).
Because of the geographical extent and remoteness of
the Tibetan Plateau, wildlife surveys are difficult to conduct,
and no intensive survey of E. kiang has been undertaken
over its entire range. Numbers of E. kiang were estimated in
several regions of its range, but most of those censuses were
carried out .15 years ago and used different methods, thus
making any estimation of actual population sizes difficult.
Available rough estimates are: Tibet: 37,000–48,000 individ-
uals; Xinjiang: 4,500–5,500 individuals; Qinghai and Gansu:
15,000 individuals (Schaller 1998); Ladakh: 1,500–2,000
individuals (Fox et al. 1991); Sikkim: 74–120 individuals
(Shah 2002); and Pakistan: 15–25 individuals (Rasool 1992:
Shafiq and Ali 1998). Available numbers for Nepal range
from 37 to 500 individuals (Shah 2002; Sharma et al. 2004).
Total abundance of E. kiang was estimated at 60,000–70,000
individuals by Schaller (1998) and 200,000 individuals by
Gao and Gu (1989). It is difficult to know which estimate is
more reliable because of the lack of statistical testing in both
attempts. Gao and Gu (1989) may have overestimated
numbers of E. kiang in the Arjin Mountain Reserve of
Xinjiang (Schaller 1998), so their rangewide evaluation also
appears doubtful (Schaller 1998; Shah 2002).
Wildlife surveys conducted on the Tibetan Plateau after
1999 have not resulted in a new population estimate for E.
kiang, but they illustrate local trends in population changes
(e.g., Bhatnagar et al. 2006; Fox and Ba˚rdsen 2005; Schaller
et al. 2005, 2007). For example, Schaller et al. (2005)
documented an increase of E. kiang in Tibet since their
surveys in the early 1990s (see ‘‘Conservation’’ section).
Intensive survey efforts are still required to achieve a more
accurate evaluation of numbers of E. kiang in the wild.
Despite extensive literature on fossil horses, little
paleontological information is available for Equus kiang.
E. kiang and the extinct Pleistoene species E. conversidens
share common features that distinguish them from E.
hemionus and the extinct Pleistocene stilt-legged horse, E.
calobatus (Bennett 1980). Similarities in bone size and
proportions of limbs and dental morphology exist between
E. kiang and the extinct Plio-Pleistocene E. sivalensis from
India and Pakistan (Forste´n 1986). A fossilized right
metatarsal attributed to Equus cf. E. kiang was found in
Pleistocene deposits at Gold Run Creek, Yukon Territory,
Canada (Harington and Clulow 1973). A left metatarsal
assigned to Equus cf. E. kiang also was found at Lost
Chicken Creek, Alaska (Harington 1980). Environmental
conditions similar to those prevailing on the Central Asian
steppes and the Bering land bridge may have allowed E.
kiang to colonize North America (Harington and Clulow
1973). This hypothesis has been questioned by other authors
because no such bones have been found in northeastern
Siberia; the North American fossils may have belonged to
the strictly New World ‘‘stilt-legged’’ Equus (V. Eisenmann,
FORM AND FUNCTION
Equus kiang is slightly sexually dimorphic. Female body
mass is 250–300 kg, whereas males may weigh 350–400 kg
(Scha¨fer 1937). The skull of E. kiang is characterized by a
large and short muzzle, high facial height, and narrow
occipital breadth (Eisenmann 1980; Fig. 2). Skull measure-
ments (mm 6SD) of adult specimens are: greatest length,
506.7 619.0 (n543); basilar length, 449.2 621.6 (43);
diastema length, 82.6 68.3 (41); maximal palatal length,
229.2 610.9 (46); muzzle length, 111.0 67.5 (50); skull
height, 94.2 65.4 (46); facial width, 144.3 68.4 (47);
minimal muzzle width, 42.8 63.7 (49); frontal width, 203.9
611.0 (49); length of upper toothrow, 161.0 66.1 (36);
orbital height, 57.1 62.6 (47); and orbital length, 61.6 63.0
(47—Eisenmann 1980). Additional skull measurements can
be found in Groves and Maza´k (1967), Eisenmann (1980),
Groves and Willoughby (1981), and Feng et al. (1986).
Postcranial measurements are available in Groves and
Maza´k (1967) and Groves and Willoughby (1981).
Dental formula of E. kiang is i 3/3, c 0–1/0–1, p 3–4/3, m
3/3, total 36–42 (Hodgson 1847a). Canine and 4th premolars
may be present or absent. Like other equids, E. kiang has
long molar and premolar teeth showing complex enamel
patterns on the grinding surface. Three subgroups of Equus
can be discriminated based on lower cheek-tooth patterns:
zebrines, caballines, and asinines and hemionines (Eisen-
mann 1986). E. kiang belongs to the latter group,
characterized by shallow ectoflexids and shallow V- or U-
shaped linguaflexids (Eisenmann 1986; McFadden 1992),
but these characters can vary geographically (McFadden
1992). In northern populations of E. kiang, the linguaflexids
tend to be U-shaped, but they tend to be more V-shaped in
southern populations (McFadden 1992). Eisenmann (1980,
1986) noted various characteristics of upper cheek teeth:
caballine fold in 36% of premolars and 4% on M1 and M2
and absent on M3; 3 or 4 folds on P3 and P4; 47% of M3
with isolated hypoglyphs; 27% of M3 with open postfos-
4 MAMMALIAN SPECIES 835—Equus kiang
settes; and the wolf tooth (dP1) in 30% of the specimens. On
P3 and P4, the protocone has a slight fold at its roots
(Groves and Maza´k 1967).
ONTOGENY AND REPRODUCTION
Mating season in Equus kiang occurs from late July to
the end of August (Schaller 1998). Various lengths of
gestation have been reported: 11 months (Schaller 1998), 7–
10 months (Hayssen et al. 1993), 355 days (Shah 2002), and
365 days (Groves and Willoughby 1981). Females give birth
to 1 offspring in summer between mid-July and mid-August
every 2 years, although no data on marked animals exist
(Schaller 1998; Fig. 4). Newborns are #90 cm at the
shoulder and weigh #36 kg (Pohle 1991); they can walk a
few hours after birth (Denzau and Denzau 1999). Little is
known about age of sexual maturity or 1st reproduction. In
the Gobi Desert of Mongolia, female E. hemionus breed for
the 1st time at 3 or 4 years (Feh et al. 2001; Schaller 1998). E.
kiang may be comparable (Schaller 1998). A captive male
did not show any sign of sexual maturity at 3.5 years of age
(Groves 1974). It is likely that females can come into estrus
shortly after giving birth because mating of females
accompanied by newborns has been observed (Pfister
2004). This is the case in other arid-adapted species, such
as Gre´vy’s zebra (E. grevyi—Rubenstein 1994).
Infanticide occurs in captive E. kiang (Berger 1986). In
the wild, a male attempting to mate with a female also
inflicted severe injuries to her newborn foal (B. Humbert-
Droz, pers. comm.). Infanticide has often been observed in
other equid species (Cameron et al. 2003; Duncan 1982;
Linklater et al. 1999). Induced abortion has been observed in
wild horses (Berger 1983), but whether it occurs in E. kiang is
Population characteristics.—Although population esti-
mates of Equus kiang are difficult to obtain, density
estimates conducted in specific locations and repeated in
time are more reliable, and thus can help understand
population trends at a regional scale. Densities of E. kiang
in the southeastern part of the Chang Tang Nature Reserve
were 0.07 individuals/km
in 1991 and 0.19 individuals/km
in 2003, indicating a recent population increase (Schaller et
al. 2005). Surveys conducted between 1999 and 2002 along a
different route across the Chang Tang Nature Reserve gave
densities of 0.48–0.82 individuals/km
(Fox and Ba˚rdsen
2005). In Ladakh, densities were estimated at 0.25 individ-
(Fox et al. 1991). A survey in Ladakh in 2000 gave
comparable figures, with densities of 0.03–0.86 individuals/
—Bhatnagar et al. 2006).
Surveys in the Hanle Valley of eastern Ladakh between 2001
and 2004 yielded densities of 0.56 individuals/km
gar et al. 2006). Other available estimates are: Lhasa–
Golmud road (Qinghai), 0.1 individuals/km
(Schaller et al.
1991); Yeniugou (Qinghai), 0.8 individuals/km
Miller 1995); and Xinjiang, 0.3 individuals/km
1998). Human presence appears to negatively impact
densities of E. kiang. In the Chang Tang Nature Reserve,
densities were 1.06–1.53 individuals/km
under low human
influence and 0.88 individuals/km
under medium human
influence (Fox and Ba˚rdsen 2005).
Adult E. kiang have been reported to live up to 20 years
in the wild, based on comparative tooth wear with zebras
(Schaller 1998). Young of the year comprised 11% of the
individuals censused in Chang Tang Nature Reserve in 1985–
1993, and survival of offspring seemed low in some years
(Schaller 1998). Mortality rates are poorly known. Illegal
hunting may be an important cause of mortality because
nomadic communities often hunt for subsistence (Huber 2005;
Schaller 1998). Blizzards also may cause mortality in some
occasions due to starvation (Schaller 1998).
Space use.—Three major vegetation formations cover
the distributional range of Equus kiang: alpine meadow,
alpine steppe, and desert steppe (Miller and Schaller 1996;
Schaller 1998). Alpine meadows are generally found at
elevations ,4,500 m in regions where precipitation exceeds
400 mm/year (Schaller 1998); alpine meadows occur mostly
in the eastern part of the Chang Tang Nature Reserve and
Qinghai. To the west, alpine meadows are associated with
riparian habitat along streams and lakes. Vegetation in
meadows is dominated by short sedges (e.g., Kobresia)and
various forbs and covers about 25–30% of the ground
(Miller and Schaller 1996; Schaller 1998). The growing
season starts earlier in riparian areas than in other habitats
because water is usually present for a longer period. Alpine
steppe is the most widely found vegetation formation in
western Tibet and occupies most of the southern one-half of
the Chang Tang Nature Reserve (Schaller 1998). Alpine
Fig. 4.—A group of female Equus kiang with 1 foal in summer from
eastern Ladakh, India. Used with permission of the photographer E
J. Van Gruisen.
835—Equus kiang MAMMALIAN SPECIES 5
steppe occurs at elevations of 4,000–5,000 m in regions with
low precipitation and is generally devoid of any sod layer
(Schaller 1998). Vegetation is scarce with a cover of about
15%, and is mostly represented by Stipa, other grasses such
as Festuca and Poa, and sedges such as Carex moorcroftii
(Miller and Schaller 1996; Schaller 1998). Desert steppe
occurs at elevations .5,000 m and is mostly found in the
northern one-half of the Chang Tang Nature Reserve and in
Xinjiang. Plant composition is similar to the that of alpine
steppe along with cushion plants and dwarf shrubs such as
Ceratoides compacta, but vegetation cover is much lower,
averaging only 5% (Miller and Schaller 1996; Schaller 1998).
Equus kiang mostly inhabits alpine steppes and alpine
meadows and rarely occurs in desert steppes (Schaller 1998).
The species typically occurs in plains and hills in generally
broad and open valleys and basins (Harris and Miller 1995;
Schaller 1998). Harris and Miller (1995) found E. kiang
mostly in xeric south-facing basins. In eastern Ladakh, most
groups were found in wide valleys (79%), and a lesser
proportion was observed in hills (19%—Bhatnagar et al.
2006). E. kiang may use flat and open areas as escape terrain
where it can flee easily, a common antipredator strategy
among wild equids (Harris and Miller 1995; Schaller 1998).
In the Hanle Valley of eastern Ladakh, most groups (74%)
were observed on alpine meadows (Bhatnagar et al. 2006).
Schaller (1998) and Bhatnagar et al. (2006) suggested that
these meadows represented critical habitats for E. kiang in
summer and winter.
No regular migration patterns among populations of E.
kiang have been observed (Schaller 1998). However, E. kiang
makes seasonal movements between different habitat types,
often dispersing in small groups into hilly terrain in summer
and concentrating in basins and flat terrain during winter
(Schaller 1998). It has been suggested that these movements
are linked to availability of relatively high-quality forage
such as Stipa (Schaller 1998). In summer, groups have often
been observed making daily movements, shifting from
meadows and flat basins in early morning to higher-
elevation terrain later in the day and returning to lower
terrain after sunset (Denzau and Denzau 1999). This
movement pattern also occurs in other wild equids such as
feral horses in the Great Plains, United States, probably as a
strategy to avoid heat (Berger 1986).
Diet.—Equids are hind-gut fermenters, and the majority
of microbial digestion occurs in the cecum (Janis 1976). This
allows equids to feed on coarse and fibrous forage with a
high stem-to-leaf ratio (Duncan 1992; Janis 1976; McNaugh-
ton 1985). Equus kiang mostly feeds on graminoids (Harris
and Miller 1995; Schaller 1998; Fig. 5). On the Chang Tang
Plateau, the summer diet of E. kiang was approximately 65%
Stipa, followed by Kobresia,Carex,Poa,Elymus, and small
amounts of a few forbs and shrubs (Schaller 1998). In the
same area, the proportion of Stipa in winter diets increased
to .90% (Schaller 1998). In Qinghai, the diet of E. kiang was
95% Stipa (Harris and Miller 1995). Forbs were rarely eaten,
so legumes were usually consumed in lower proportion than
their relative availability (Harris and Miller 1995). Occa-
sionally, E. kiang has been observed digging Oxytropis roots,
mostly in fall and winter when little dry grass is available
(Schaller 1998). In desert steppe, shrubs such as Ceratoides
may be important in the diet (Schaller 1998). Hay (1859:356)
reported seeing E. kiang ‘‘feeding almost entirely on the
roots of a species of Artemisia, or Worm-wood.’’
Water requirements of E. kiang are poorly documented.
Ungulates living in arid environments are often dependent
on water sources, and this may restrict their movements and
habitat use (Berger 1986; Duncan 1992). On the Tibetan
Plateau, however, water sources are generally scarce and
mostly frozen in winter (Schaller 1998). Occasionally, E.
kiang drinks from waterholes, lakes, and streams, but this
behavior is not common (Denzau and Denzau 1999; Schaller
1998). It seems that most of the ungulate species, including
E. kiang, of the Tibetan Plateau fulfill their water
requirements from vegetation and snow (Schaller 1998).
Diseases and parasites.—Little information is available
on diseases of Equus kiang. A neonate in captivity died from
omphalitis (Benirschke 2008), and hemorrhages have been
found in the digestive tract of a dead foal in the wild
(Schaller 1998). The following parasites are known to occur
in E. kiang:Strongylus,Trichonema,Ascaris,Skrjabinema,
Anoplochephala perfoliata, and Paranoplocephala mamillana
(Feng et al. 1986). Oestrid fly (Gasterophilus) infestations
also are reported in E. kiang (Feng et al. 1986; Schaller
Interspecific interactions.—Distribution of Equus kiang
overlaps with the range of 6 other wild ungulate species:
Tibetan gazelle (Procapra picticaudata), chiru or Tibetan
antelope (Pantholops hodgsonii—Leslie and Schaller 2008),
blue sheep (Pseudois nayaur—Wang and Hoffmann 1987),
Fig. 5.—A group of Equus kiang feeding in a mixed vegetation
patch of grasses and sedges in the alpine steppe of eastern Ladakh,
India. Photograph by S. D. Coˆte´.
6 MAMMALIAN SPECIES 835—Equus kiang
wild yak (Bos mutus), Tibetan argali (Ovis ammon hodg-
soni—Fedosenko and Blank 2005), and white-lipped deer
(Prezwalskium albirostris—Harris and Miller 1995; Schaller
1998). In Chang Tang, open basins are used by P. hodgsonii
and P. picticaudata alongside E. kiang (Schaller 1998). E.
kiang has a high degree of spatial co-occurrence with P.
picticaudata, and to a lesser extent with domestic yaks (Bos
grunniens) in eastern Ladakh close to the Tibet border
(Namgail et al. 2008). In the Rupshu area of Ladakh, E.
kiang and O. ammon are often seen in the same habitats but
rarely interact (Fox et al. 1991; Hay 1859).
The predominantly grass-based diet of E. kiang overlaps
little with diets of sympatric wild ungulates on the Tibetan
Plateau (Harris and Miller 1995; Schaller 1998). Domestic
sheep, goats, yaks, and horses are kept by pastoral herders in
most of the range of E. kiang (Goldstein and Beall 1989;
Miller and Schaller 1996). In Yeniugou, Qinghai, E. kiang
has a high degree of dietary overlap with domestic sheep
(Harris and Miller 1995). Wild ungulates of the Tibetan
Plateau likely coevolved over a long period and may thus
partition their ecological niche, whereas domestic livestock,
because they were introduced only recently, may have a
greater potential of interspecific competition with wild
ungulates (Namgail et al. 2008; Schaller 1998).
Among potential predators, Tibetan wolf (Canis lupus
chanco) and snow leopard (Uncia uncia) may occasionally
prey on young and old individuals, but overall predation is
unlikely to be an important limiting factor in populations of
E. kiang (J. Van Gruisen, pers. comm.; Schaller 1998).
Remains of E. kiang were absent in scats of C. lupus (n5
384), U. uncia (n5193), and Ursus arctos (Tibetan brown
bear—n548) from Qinghai and Tibet (Schaller 1998).
Observations from Mongolia report E. hemionus in scats of
C. lupus (Feh et al. 2001).
Miscellaneous.—According to the International Species
Information System, there were 114 individuals of Equus
kiang in captivity in May 2008, all E. k. holdereri
(International Species Information System 2008). Although
attempts to domesticate E. kiang have occurred (Hay 1859),
they have never been successful (Groves 1974; Tegetmeier
and Sutherland 1895).
Grouping behavior.—Equus kiang is often found alone or
in small groups but also may form herds of several hundred
individuals (Foggin 2000; Paklina and van Orden 2003;
Scha¨fer 1937; Schaller 1998). Herds of 500–1,000 animals
were reported in the 1800s (Rockhill 1895), and large
aggregations still occur today. Schaller (1998) observed
aggregations of up to 261 individuals in the Chang Tang
Nature Reserve. More than 500 E. kiang were observed in 4
or 5 herds on the edge of the Kekexili range in Qinghai
(Foggin 2000). In Ladakh, groups ranged from 1 to 74
¯52.8, n5365—Bhatnagar et al. 2006).
Group size appears to vary seasonally. Individuals of E.
kiang are dispersed in summer and tend to congregate in fall
and winter after rut. Average group size was 6.8 individuals
in summer and 10.9 in winter in the Chang Tang Nature
Reserve (Schaller 1998). In Qinghai and Tibet, 13.6% of the
animals observed were alone in summer, but it dropped to
2.5% in September–December (Schaller 1998). In Novem-
ber–December 2006, 2.2% of the animals observed in the
Chang Tang Nature Reserve were solitary, and 73% were in
groups of 2–10 individuals (n548—Schaller et al. 2007).
During surveys conducted in September and October 1998 in
southwestern Tibet, 1.1% of E. kiang were solitary, 1.6%
were in pairs, and 97.3% were in groups of 3–160 individuals
(Paklina and van Orden 2003).
Two distinct types of social organizations have been
described in wild equids (Klingel 1975, 1977). In the 1st type,
permanent groups contain a few males and several females and
young, and males are not territorial. This type of social
organization occurs in equids from temperate ecosystems such
as E. burchellii (Burchell’s zebra), E. zebra (mountain zebra),
and E. caballus przewalskii (Przewalski’s horse—Moehlman
1998). In the 2nd type, the only permanent associations occur
between mother and foal. Only temporary groups are formed,
comprising several females and their young or only males
(bachelor groups); adult males are often territorial. This type
of social organization has been documented in equids adapted
to arid environments, such as E. grevyi,E. asinus africanus
(African wild ass—Grinder et al. 2006), and E. hemionus
(Klingel 1977; Moehlman 1988; Rubenstein 1989). In those
environments where food is often scarce, resource needs
between males and females may be too different to maintain a
tight social structure (Rubenstein 1989, 1994).
Equus kiang seems to display the 2nd type of social
organization (Groves and Willoughby 1981; Schaller 1998).
Groups of E. kiang are not permanent but seem to be formed
of temporary aggregations; only female and foal remain
together at all times (Fig. 4). Young adult males typically
form bachelor groups, and older males are often solitary and
show territorial behavior (Denzau and Denzau 1999;
Schaller 1998). Solitary males often herd female or family
groups that pass within their territory (Schaller 1998).
Defecating at dung piles is common in territorial equids
(Klingel 1977; Moehlman 1998) and occurs in E. kiang
(Denzau and Denzau 1999). In Ladakh, territory size varies
from 0.5 to 5.0 km
(Denzau and Denzau 1999). Territorial
defense by males involves aggressive chases toward intruding
males (Denzau and Denzau 1999; Schaller 1998). These
chases are often preceded by threatening behavior when a
defending male approaches an intruder with its head up, ears
laid back, and tail horizontal, often grunting or making a
‘‘wheezing bray’’ at the same time (Schaller 1998:272). One
of us (AS-L) observed chases that lasted .20 min. Aggres-
sive fighting between adult males also occurs, involving
835—Equus kiang MAMMALIAN SPECIES 7
kicking and biting the neck, mane, and tail; such males often
have scars and wounds resulting from aggressive interactions
(Denzau and Denzau 1999).
Reproductive behavior.—Male Equus kiang tend females
by trotting around them and chasing those that wander from
the group, often with their head down and their ears laid
back (Schaller 1998). Such chases typically precede mating,
and copulation is often followed by the male trotting away
with its head held high and obliquely (Pfister 2004).
Communication.—The vocalization of Equus kiang has
been described as a ‘‘shrieking bray’’ (Lydekker 1904b:584).
Hay (1859) noted that it was distinct from neighing of the
horse and braying of the ass. E. kiang makes a snort puff
when alarmed (Pfister 2004).
Two diploid numbers (2n) are exhibited by members of
Equus kiang. Karyotypes exhibit either 52 or 51 chromosomes
and each karyotype possesses 92 chromosomal arms (Ryder
and Chemnick 1990). This polymorphism of diploid number
occurs in both male and female E. kiang and is attributed to a
Roberstonian translocation (Ryder and Chemnick 1990). The
X chromosome is submetacentric; the Y chromosome is
acrocentric (Ryder and Chemnick 1990).
Equus kiang is often considered a subspecies of E.
hemionus (Groves 1974; Groves and Maza´k 1967; Pohle
1991). Nevertheless, genetic analyses demonstrate that E.
kiang is a distinct species and that divergence from E.
hemionus occurred approximately 500,000 years ago (Ryder
and Chemnick 1990). Genetic variation among the 3
subspecies of E. kiang has not been investigated.
Hybrids between E. kiang and E. caballus,E. asinus,E.
hemionus,andE. burchellii have been reported in captivity
(Gray 1972; Hay 1859; Kinloch 1869). A female hybrid
between E. kiang and E. caballus, born in the Jardin des
Plantes, Paris, France, lived for .36 years (Gray 1972).
Another female hybrid between E. kiang and E. burchellii
lived there for .32 years (Gray 1972). Hybrids between
captive E. kiang and captive E. hemionus are reported for
both E. h. hemionus and E. h. khur (Gray 1972). In the wild,
geographical barriers likely prevented hybridization of E.
kiang with adjacent populations of E. h. hemionus and E. h.
khur (Groves and Maza´k 1967). No evidence exists that
hybrids of E. kiang can reproduce. Genetic analyses have
established that E. a. asinus (domestic donkey) did not
originate from E. kiang or E. hemionus, but from E. a.
africanus (Beja-Pereira et al. 2004).
Equus kiang is fully protected in China and India (China
Class I; Indian Wildlife Protection Act 1972 Schedule I—
Shah 2002). In Pakistan, E. kiang is listed on the national red
list as Critically Endangered (Shafiq and Ali 1998). E. kiang
is listed on the World Conservation Union (IUCN) Red List
as Lower Risk/Least Concern (LR/LC—Baillie and Groom-
bridge 1996). Each subspecies is designated separately by
World Conservation Union: Lower Risk/Least Concern for
E. k. holdereri and Data Deficient for E. k. kiang and E. k.
polyodon (Baillie and Groombridge 1996). E. kiang is in
Appendix II of the Convention on International Trade in
Endangered Species of Wild Fauna and Flora (Shah 2002).
Historically, E. kiang and other wildlife species were
hunted by nomads on the Tibetan Plateau for their
subsistence (Huber 2005). However, religious beliefs prohib-
ited killing equids in ancient civilizations of Tibet, and
although it could have occasionally been a necessity, E.
kiang was not traditionally hunted on a wide scale (Bellezza
2008; Harris 2008; Huber 2005). Hunting of E. kiang
drastically increased around the mid-20th century due to
easier access to remote locations with roads and availability
of the modern rifle (Leslie and Schaller 2008; Schaller et al.
2005). E. kiang was heavily hunted in China during the
socioeconomic changes and the great famine of 1958–1961
(Harris 2008; Schaller 1998; Schaller et al. 2005). In
southeastern Qinghai, for example, populations of E. kiang
have been diminished, and they are now absent or scattered
(Schaller 1998). In Ladakh, populations of E. kiang were
greatly reduced during the 1962 war with China (Bhatnagar
et al. 2006).
During the last decade, the Chinese government took
successful wildlife protection measures. Illegal hunting has
been strictly controlled, and a 600,000-km
was established in the Chang Tang region (Harris 2008;
Leslie and Schaller 2008; Schaller et al. 2005). E. kiang is
now abundant enough that there is no immediate threat to
its persistence, and populations are reported to be increasing
(Schaller et al. 2005). In the southeastern part of the Chang
Tang Nature Reserve, populations of E. kiang have
increased since the early 1990s to the extent that they are
now perceived by pastoralists as serious competitors with
livestock (Fox et al. 2004; Schaller et al. 2005; Tsering et al.
2006). In Nyima County, Tibet, for example, E. kiang was
reported to damage pastures in early spring when they dig
for grass roots (Tsering et al. 2006). Although harvesting of
E. kiang does not occur at a wide scale, they are often chased
from pastures on motorbikes (Tsering et al. 2006). Herders
and local authorities have requested permission to cull E.
kiang to reduce their potential negative impact on pastures,
but culling was denied by the Chinese government because
of the national protection status of E. kiang (Tsering et al.
In remote regions of Chang Tang, changes in rangeland
use policy are increasing human densities and movements in
key wildlife areas, along with increasing livestock numbers
(Fox and Tsering 2005; Schaller et al. 2005). Rangelands in
some locations are turned into private ranches owned by
8 MAMMALIAN SPECIES 835—Equus kiang
several families (G. B. Schaller, pers. comm.), likely causing
an intensification of pasture use and deteriorating wildlife
habitats (Fox and Tsering 2005). Rangelands are often
fenced from wildlife, which not only prevents E. kiang access
to key resources but also may cause injuries and mortalities
(Bhatnagar et al. 2006; Schaller et al. 2005). Conflicts with
domestic livestock may be the biggest challenge for the long-
term conservation of E. kiang (Fox et al. 2004; Fox and
Tsering 2005; Schaller et al. 2005; Tsering et al. 2006).
Whether these conflicts are real or perceived depends on the
location (Bhatnagar et al. 2006; Tsering et al. 2006). Large
numbers of E. kiang may compete with livestock, but this
situation probably occurs mainly at a local scale. For
example, Bhatnagar et al. (2006) estimated that throughout
Ladakh, populations of E. kiang use only 10–11% of the
total forage consumed, whereas the other 89–90% is used by
Increased human densities in remote areas of the
Tibetan Plateau augment the threats to E. kiang and other
wildlife species. Mining activities occur in parts of Qinghai
and Xinjiang, and the easier access to remote regions by
road facilitates illegal hunting (Fox and Tsering 2005;
Schaller et al. 2005; Tsering et al. 2006). In some places of
Qinghai, poaching continues and is accepted by the nomads,
who see E. kiang as a nuisance to their livestock (G. B.
Schaller, pers. comm.). Since early 2006, male E. kiang have
been poached for their penis, which is reputed to enhance
men’s virility when eaten (Tsering et al. 2006).
Long-term conservation of E. kiang lies mainly in the
resolution of potential conflicts with domestic livestock,
minimizing habitat degradation from pastoralists and
human development (e.g., fences, road, and mining), and
control of poaching. Because conflicts mostly occur at local
scales, solutions also must be implemented at that scale. For
example, different management and conservation strategies
could be considered inside and outside wildlife reserves. Pilot
projects could be initiated in specific places to reduce the
number and impact of E. kiang, but these changes would 1st
need to be accompanied with policy changes and ecological
studies (Harris 2008; G. B. Schaller, pers. comm.). Data are
needed on the dynamics of populations of E. kiang and
resource use to identify at which scale and to what extent
competition with livestock occurs, during which season, and
for what resource components. Moreover, genetic analyses
are needed to ascertain the status of the 3 named subspecies
of E. kiang. If recognized as such, E. k. polyodon may require
specific conservation measures because of its low abundance
and restricted distribution (Neumann-Denzau and Denzau
We thank C. Barrette, J. Mainguy, G. B. Schaller, and
an anonymous reviewer for their constructive comments on
earlier versions of the manuscript; V. Eisenmann for helping
with the osteology section and fossil records; A. D. Jungblut
for translating German texts; and Xiao-Mian Xie for
translating Chinese texts. We are indebted to J. Van Gruisen
for providing photographs of E. kiang in the wild (copyright
J. Van Gruisen), B. Patterson of the Field Museum of
Chicago for providing skull photographs of E. kiang,andG.
B. Schaller for permission to reproduce the distribution map.
This manuscript benefited from fruitful communications
with J. V. Bellezza, G. Denzau, B. Humbert-Droz, O.
Pfister, A. Shrieber, and J. Van Gruisen. Finally, the staff of
the scientific library of Universite´ Laval provided invaluable
help to find rare literature. Funding to A. St-Louis during
writing was granted through a doctoral scholarship from the
Natural Sciences and Engineering Research Council of
Canada and the Natural Sciences and Engineering Research
Council of Canada Discovery Grant to SDC.
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Associate editors of this account were DAVID M. LESLIE,JR. and
PAMELA R. OWEN.COLIN GROVES and IGOR PAVLINOV reviewed the
synonymy. Editor was MEREDITH J. HAMILTON.
835—Equus kiang MAMMALIAN SPECIES 11