 Axis
kuhlii
Bawean deer, Kuhl's deer |
Taxonomy | Description
| Reproduction | Ecology
| Behavior | Distribution
| Conservation | Remarks
| Literature |
| Taxonomy
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Axis kuhlii [Müller, 1840].
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Citation: In Temminck, Verh. Nat. Gesch. Nederland. Bezitt.,
Zool., Zoogd. Indisch. Archipel., p. 45[1840].
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Type locality: "Java en Borneo", but is found only on Bawean Island,
Indonesia.
The taxonomic record (above) is taken from Wilson and Reeder (1993).
Axis kuhlii is included in the subgenus Hyelaphus [Sundevall,
1846] along with the closely related hog deer and Calamian deer (Whitehead,
1993). Indeed, although recognized as a species here, the Bawean deer
is considered by some authors to be a subspecies of A. porcinus (see
Wilson and Reeder, 1993). A. kuhlii is a monotypic species
(Whitehead, 1993). There are no synonyms.
General Characteristics
The Bawean deer is poorly studied, and few scientific measurements of this
species are available. There is no consensus about either
adult weight or total length, although values of 50-60 kg and 140 cm respectively
seem more likely than those provided by Kurt (1990) as they refer directly
to A. kuhlii. Shoulder height is 65-70 cm.
Reported measurements for Bawean deer (Axis kuhlii) |
| Source |
Adult Weight |
Head & Body Length |
Shoulder Height |
Tail Length |
Blouch and
Atmosoedirdjo, 1987 |
- |
140 cm  |
65 cm |
- |
| Geist, 1998 |
- |
- |
65 cm |
- |
| Kurt, 1990 |
36-50 kg
("like hog deer") |
105-115 cm
("like hog deer") |
70 cm |
20 cm
("like hog deer") |
| Lydekker, 1915 |
- |
- |
~68.5 cm |
- |
| Whitehead, 1993 |
50-60 kg |
- |
68-70 cm |
- |
The coat is short, smooth, and soft, and generally a light teakish-brown
color (Sitwell, 1970). Although the pelage is a relatively uniform
brown from a distance, each hair is banded with yellow, giving a more grizzled
appearance up close (Sitwell, 1970). The few distinctive markings are
limited to the head and neck, where there is a light throat patch or 'bib',
and usually a whitish eye-ring (Blouch and Atmosoedirdjo, 1987; Geist, 1998).
The light lip area is separated from the face by a darker band. The
moderately long tail is bushy, being brown above and white on the underside
(Lydekker, 1915; Sitwell, 1970; Whitehead, 1993). White hairs are also
present around the groin (Sitwell, 1970).
The size and shape of the Bawean deer are virtually identical to the hog
deer (A. porcinus), although the legs of A. kuhlii are noticeably
shorter (Geist, 1998). Like many other inhabitants of dense forest,
the body is lower at shoulder than at the hip, giving the Bawean deer a crouched
appearance (Blouch and Atmosoedirdjo, 1987). The face is short compared
to that of the hog deer (Lydekker, 1915; Whitehead, 1993). The inflated
tympanic bullae of the skull are found only among Hyelaphid deer (A.
kuhlii, A. porcinus, and A. calamianensis) (Geist,
1998). The ears are small and pointed, and are densely haired on their
exterior surface (Lydekker, 1915).
Only the males bear the slender three-tined antlers (Sitwell, 1970).
Similar in form to those of the hog deer, the antlers have a brow tine
(found just above the base) and a forked main beam (Blouch and Atmosoedirdjo,
1987). However the antlers of A. kuhlii are much shorter than
those of A. porcinus, generally not growing longer than head (Lydekker,
1915; Whitehead, 1993). The longest main beam recorded by Blouch and
Atmosoedirdjo (1987) was 47 cm in length, while the leading trophy in Rowland
Ward's Record of Big Game has a length of only 24.8 cm, with a 7.3 cm basal
circumference, and a 27.6 cm inside span (Whitehead, 1993). The antlers
are supported by relatively long pedicels, permanent outgrowths of bone from
the forehead (Whitehead, 1993). Bucks may be found with hard antlers
at all times of the year (Blouch and Atmosoedirdjo, 1987)
Ontogeny and Reproduction
Whitehead (1993) reports a seasonal rut in September and October, although
males may be found in breeding condition (i.e., with hard antlers) throughout
the year (Blouch and Atmosoedirdjo, 1987). The gestation period is
225-230 days, after which a single fawn is born - cases of twins are known
but very rare (Blouch and Atmosoedirdjo, 1987; Whitehead, 1993). The
majority of births occur from February to June, although occasional births
may occur in other months (Blouch and Atmosoedirdjo, 1987). In captive
specimens, breeding occurs year-round with females maintaining an interbirth
interval of 9 months (Blouch and Atmosoedirdjo, 1978). However, in
the wild, it is unlikely that a female could raise more than one fawn per
year (Blouch and Atmosoedirdjo, 1987).
Geist (1998) reported that fawns are only faintly and sparsely spotted and
lose their spots very quickly, while Sitwell (1970) observed a fawn on Bawean
at least three months of age with a row of white spots along either side
of the spine. Males begin to grow antlers at about one year of age
(Sitwell, 1970).
Ecology
A. kuhlii is an inhabitant of upland forests, rather than low-lying
marshy grasslands like the closely related hog deer (Geist, 1998). Forests
with dense undergrowth are used for shelter, providing a refuge in which
the deer sleep and rest during the day (Blouch and Atmosoedirdjo, 1978).
The optimal cover appears to be brushy growth of woody plants no more
than 3 meters high (Blouch and Atmosoedirdjo, 1987). In areas disturbed
by humans, Bawean deer spend the day in forests on steep slopes that are
inaccessible to teak loggers; clearings predominanted by grasses such as
lalang (Imperata cylindrica), although rarely over 5 hectares in size,
are used extensively by Bawean deer at night. This is especially true
during the first months after a fire, when the vegetation is grazed heavily
(Blouch and Atmosoedirdjo, 1978). Individuals are occasionally sighted
on the beach in the southwestern part of the island; otherwise they are rarely
seen (Blouch and Atmosoedirdjo, 1987)
Secondary forest appears to be ideal Bawean deer habitat, supporting densities
of 19.2 deer per square kilometer. Such habitat is characterized by
tree species such as Ficus variegata, Macarange tanarius, and
Anthrocephalus indicus which form an overstory under which shrubs
such as Leea indica, Ficus spp., Antidesma montanus,
and Garcinia celebica grow. Teak (Tectona grandis) forests
with understory, primary forest, and areas with teak and lalang support densities
of 3.3 to 7.4 deer per square kilometer, while regions dominated by
Melastoma polyanthum and Eurya nitida brush, Rombok (Merremia
peltata), disturbed primary forest, and teak without understory support
only 0.9-2.2 deer per square kilometer (Blouch and Atmosoedirdjo, 1987).
A. kuhlii is usually solitary, although pairs made up of a doe and
fawn or a buck following a doe are also encountered (Blouch and Atmosoedirdjo,
1978). During nightly foraging in open clearings, Bawean deer may encounter
other conspecifics, although Blouch and Atmosoedirdjo (1978) state that this
cannot be considered a true "congregation". These open clearings, while
used extensively for feeding, are also the center of social activity, with
courting, challenging, fighting, and mating all occurring outside of the
dense forest (Blouch and Atmosoedirdjo, 1987). Trails made by deer
are commonly found leading from forest sites to feeding areas in brushy teak
and secondary growth at lower elevations (Blouch and Atmosoedirdjo, 1978).
The Bawean deer has no natural predators except large pythons (a python was
found by Blouch and Atmosoedirdjo (1978) with an adult deer in its stomach).
However, pythons are not common, and likely have little impact on the
deer population (Blouch and Atmosoedirdjo, 1987). It is possible that
wild pigs and macaques sometimes kill young fawns, although no evidence has
been found to support this (Blouch and Atmosoedirdjo, 1978; Blouch and
Atmosoedirdjo, 1987). Feral dogs are currently the greatest cause of
mortality to this species, being responsible for 9 out of 11 deaths examined
by Blouch and Atmosoedirdjo (1987) between October 1977 and May 1979.
Blouch and Atmosoedirdjo (1987) observed Bawean deer feeding on 39 plant
species, the bulk of which are forbs (15 species) and grasses (14 species).
Young lalang grass (Imperata cylindrica) is a major food source
for A. kuhlii, both because of its abundance and its apparent palatability
to this species - however, old lalang is never eaten. The grasses
Paspalum conjugatum and Axonopus compressus also appear to
be preferred, and although these species are rarer than lalang they are eaten
in all stages of growth. Of the forbs, Lygodium circinnatum,
Musa spp., Tridax procumbens, Pericampus glaucus, and
Euphorbia geniculata are commonly fed upon. Browsing was observed
on eight species of woody plants, but was primarily confined to young leaves
and twigs of Ficus and rombok (Merremia
peltata). These food sources are generally available throughout
the year on Bawean, and are so abundant that conspicuous signs of feeding
are rarely found. When in season, the fruits of Irvingia malayana
and Elaeocarpus glaber are eaten in large quantities. Bawean
deer frequently enter agricultural fields at the edge of forest at night,
feeding on the young leaves of corn and cassava, but also on grasses and
forbs growing among the crops (Blouch and Atmosoedirdjo, 1987). Each
deer deposits approximately 13 fecal pellet groups per day, a number which
has been used to estimate population numbers (Blouch and Atmosoedirdjo, 1978).
Behavior
Blouch and Atmosoedirdjo (1978, 1987) have conducted the most
behavioral research on this species. Except where noted,
the information presented here is from these two papers.
The Bawean deer is primarily nocturnal, emerging from dense cover just after
dark (around 1800 hours) and being active intermittently throughout the night.
Peaks of activity occur approximately every two hours, usually separated
by retreats into cover. As the night progresses, foraging periods become
shorter and rests become longer, until the animals retire back into dense
cover at sunrise. An individual deer may return to the same general
hiding place for several days.
A. kuhlii communicates extensively with vocalizations, primarily using
short, sharp barks which likely have several functions. Both sexes
create these sounds, although the calls of does are slightly higher pitched
than barks made by males. Commonly, one call consists of five to ten
barks strung together, audible to humans up to 100 meters away. If
a pair of deer are surprised and separated, one or both will bark one to
three time after a few minutes in an attempt to reestablish contact.
If a mother is separated from her fawn, the does will call, to which the
youngster responds with a high pitched squeak audible only at a short distance.
The most sustained barks are performed by males as a challenge to rivals
- one male was observed barking 95 times within a fifteen minute period.
This challenge-barking is often accompanied by foot stamping, which
is audible to humans 40 to 50 meters away, and by snorts. Other bucks
will approach a deer giving this call and answer it with a similar bark.
Such challenges may escalates into a fight among males in which the
antlers are used, especially if a receptive female is in the vicinity.
Because the deer respond to and approach vocalizing conspecifics, humans
can incite individuals to approach by imitating this call with a whistle.
Although highly vocal amongst themselves, Bawean deer do not appear to have
an alarm call. When mildly alarmed, Bawean deer do not vocalize, but
rather sneak quietly into cover in an attempt to escape undetected.
Likewise, if approached in hiding, individuals frequently remain still
in an attempt to remain unnoticed, or move quietly away from the potential
threat. If startled, Bawean deer will sprint for a short distance,
and then move on quietly. The body form, with low shoulders and a higher
rump, is conducive to moving through dense undergrowth, which these deer
do with a crouching gait (Sitwell, 1970). They are very wary, and appear
to avoid contact with humans, although with protection this appears to be
changing.
Other forms of communication are poorly studied. Bucks will rub their
antlers against small saplings, shredding off the bark, which Blouch and
Atmosoedirdjo (1987) suggest may serve as visual communication. Captive
deer regularly rub objects with their preorbital glands, which likely functions
in scent marking.
Distribution
A. kuhlii is endemic to Bawean, a 200-220 km2 island located
in the Javan Sea between Java and Kalimantan (Blouch and Atmosoedirdjo, 1978;
Geist, 1998). Two wild populations exist on the island, one found among
the island's central mountain range, and other based around Mount Bulu in
the southwestern quadrant of the island. Restricted to this tiny island,
A. kuhlii has the most restricted range of any extant deer species
(Blouch and Atmosoedirdjo, 1987).
Countries: Indonesia (IUCN, 2002).
Range Map (Redrawn from Blouch and Atmosoedirdjo, 1987)
Conservation Status
The Bawean deer is classified as endangered (D) by the IUCN (2006; last
reviewed in 1996), and is on CITES Appendix I (CITES, 2006). The
wild population is estimated at about 300 individuals (Blouch and
Atmosoedirdjo, 1987). Major threats to this species include habitat
loss, with secondary forest habitat being converted to teak plantations and
agricultural areas, and predation by feral dogs (Blouch and Atmosoedirdjo,
1978). Hunting by humans, while previously a pressure on the population
for hundreds of years, stopped in 1977 (Blouch and Atmosoedirdjo, 1987).
Remarks
Axis (Latin) is said to be Pliny's name for the chital (Axis
axis), although some records show it as "an unknown wild animal in India".
Dr. H. Kuhl (1796-1821) was a German naturalist who was in the
East Indies in 1820 and 1821.
Some confusion has resulted from this species' type locality, "Java en Borneo"
- no known populations of this deer live off of Bawean island (Wilson and
Reeder, 1993). This species was supposedly discovered by Salomon
Müller in 1836 in Tuban, a small town on the northern coast of Java,
where the local governor kept a small herd in his garden. Only after
the formal description was made was the true range of this diminutive deer
revealed (Sitwell, 1970).
The Bawean deer may have been derived from a Pleistocene Javan Axis
species (Axis oppenoorthi or A. lydekkeri) at a time when Bawean
was connected to Java via a land bridge (Blouch and Atmosoedirdjo, 1987;
Geist, 1998). It has also been suggested that this deer was introduced
to the island by early European settlers, although this seems doubtful (Sitwell,
1970).
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Local names (from Whitehead, 1993)
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Menjangan Bawean [Bahasa Java]
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Rusa bawean [Indonesia]
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Uncal Bawean [Bahasa Sunda]
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French
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Cerf-conchon de l'ile Bawean, Cerf de Bawean (Kurt, 1990; Whitehead,
1993)
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German
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Kuhhirsch, Bawean-Schweinshirsch, Bawean Hirsch (Kurt, 1990; Whitehead,
1993)
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Spanish
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Ciervo de Kuhl, Ciervo porquerizo de Kuhl (IUCN, 2002)
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Literature
Cited
Blouch, R. A., and A. Atmosoedirdjo. 1978. Preliminary report
on the status of the Bawean deer (Axis kuhli). In Threatened
Deer: Proceedings of a Working Meeting of the Deer Specialist Group of the
Survival Service Commission. Morges, Switzerland: IUCN. pp. 49-55.
Blouch, R. A., and S. Atmosoedirdjo. 1987. Biology of the Bawean
deer and prospects for its management. In Biology and Management
of the Cervidae. Edited by C. M. Wemmer. Washington, D.
C.: Smithsonian Institution Press. pp. 320-327.
CITES (Convention on the International Trade in Endangered Species of Wild
Flora and Fauna). 2006. Appendix I, II, and III as adopted by
the Conference of the Parties, valid from 14 June 2006. Available online
at http://www.cites.org/
Geist, V. 1998. Three-pronged Old World deer. In
Deer of the World: Their Evolution, Behaviour, and Ecology. By
Valerius Geist. Mechanicsburg, PA: Stackpole Books, 1998. pp.
55-80.
IUCN (International Union for Conservation of Nature and Natural Resources).
2006. 2006 IUCN Red List of Threatened Species. Available
online at http://www.redlist.org/
Kurt, F. 1990. Axis deer (Genus Axis). In
Grzimek's Encyclopedia of Mammals. Edited by S. P. Parker. New York:
McGraw-Hill. pp. 142-143, 148.
Lydekker, R. 1915. Catalogue of the Ungulate Mammals in the British
Museum of Natural History, Volume IV. London: the Order of the
Trustees of the British Museum.
Sitwell, N. 1970. Bawean Island Expedition. Animals. 12: 389-393.
Whitehead, K. G. 1993. The Whitehead Encyclopedia of Deer.
Stillwater, MN: Voyageur Press, Inc.
Wilson, D. E., and D. M. Reeder [editors]. 1993. Mammal Species of the World
(Second Edition). Washington: Smithsonian Institution Press.
Available online at
http://nmnhwww.si.edu/msw/
Additional Resources
Blouch, R. 1980. Kuhl’s Deer, Bawean Island, Indonesia. Tiger
Paper 7(1):22–23.
Blower, J. 1975. Report on a visit to Pulau Bawean. Nature
Conservation and Wildlife Management Project, ISN/73/013.
Dubey, G. K., A. C. Bandopadhyay, and K. N. P. Rao. 1994. A mixed
cell sarcoma of muzzle in a spotted deer (Axis kuhli). Indian
Journal of Animal Sciences; 64(10): 1044-1045.
Grimwood, I. 1976. Hunting a deer to extinction. Oryx 13:294.
Groves, C. P., and P. Grubb. 1987. Relationships of living deer.
In Biology and Management of the Cervidae. Edited by
C. M. Wemmer. Washington, D. C.: Smithsonian Institution Press. pp.
2l-59.
Gunawan, D. and I. Kustanto. 1994. Survey on larger mammals in
Bawean Island. Deer Specialist Group News; Newsletter 12 (June 1994):
10-11. Available online at
http://iibce.edu.uy/citogenetica/deer/DSGNews12.pdf
Haltenorth, T. 1963. Klassifikation der Saugetiere: Artiodactyla I. Handbuch
der Zoologie, 8(32):1-167.
Lachenmeier, K. and R. Melisch. 1996. Der Bawean Hirsch (Axis
kuhli): Status und Schutz einer wenig bekannten Hirschart Indonesiens
[The Bawean deer (Axis kuhli): status and protection of a little known
deer species of Indonesia]. Zoologische Gesellschaft fuer Arten und
Populationsschutz E. V. Mitteilungen; 12(1): 23-25.
Mayze, R. J., and G. I. Moore. 1990. The Hog Deer. Croydon,
Australia: Australian Deer Research Foundation.
Nowak, R. M. [editor]. 1991. Walker's Mammals of the World (Fifth Edition).
Baltimore: The Johns Hopkins University Press.
Oryx. 1978. Hunting ban helps rare deer. Oryx 14: 311.
Ratajszczak, R. 1976. Some remarks on Kuhl's deer Cervus
kuhlii. Przeglad Zoologiczny 20 (4): 454-455 1976
****van Bemmel, A. C. 1944. The taxonomic position of Cervus
kuhlii M¨ ull. et Schl. Treubia; 1: 149-155.
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