Bubalus quarlesi [Ouwens, 1910].
- Citation: Bull. Dépt. Agric. Indes Néerl., 38:7.
- Type locality: Indonesia, Sulawesi, mountains of C Toradja Dist.
- Citation: Bull. Dépt. Agric. Indes Néerl., 38:7.
The taxonomic record (above) is taken from Wilson and Reeder (1993). The taxonomy of the anoas (also known as dwarf buffalos) is still uncertain - including whether there are multiple species or just multiple morphotypes. Two putative species (the lowland anoa, Bubalus depressicornis, and the mountain anoa, B. quarlesi) are presently recognized "in the absence of significant opposing evidence" (Burton, Hedges, and Mustari, 2005). Genetic work has shown species-level variation among anoas, although the interpretation of the results is confounded by potential hybridization among captive animals and a significant overlap among supposedly diagnostic features. B. quarlesi belongs to the subgenus Anoa [Hamilton-Smith, 1827], which some authors argue merits full generic rank (see Nowak, 1991). B. quarlesi is monotypic (Wilson and Reeder, 1993); Burton, Hedges, and Mustari (2005) include Anoa anoa as a synonym to B. quarlesi.
|Reported measurements for mountain anoa (Bubalus quarlesi)|
|Source||Adult Weight||Head & Body Length||Shoulder Height||Tail Length|
|Buchholtz, 1990||150 kg||150 cm||70 cm||24 cm|
|Burton, Hedges, and Mustari, 2005||< 150 kg||122-153 cm
|< 75 cm||14.6-17.8%
of body length
Up to 27 cm
|Feer, 1994||-||-||70 cm||-|
|Groves, 1969||56 kg
of body length
There are usually no markings on the face, and there is never any crescent of white on the throat (Groves, 1969; Burton, Hedges, and Mustari, 2005). The ears are small and relatively narrow (Buchholtz, 1990). Skull length has been reported as 244-290 mm (see Burton, Hedges, and Mustari, 2005).
Both male and female mountain anoa possess a pair of straight, conical horns (Groves, 1969; Buchholtz, 1990). The horns of adults can be distinguished from those of B. depressicornis by having a round (rather than triangular) cross-section, although the horn characteristics of juveniles of both species are similar (Groves, 1969; Burton, Hedges, and Mustari, 2005). The horns are smooth, without any marked ridges or external keel (Groves, 1969; Buchholtz, 1990). Relatively short, horn lengths for both sexes vary between 14.6-19.9 cm (Groves, 1969).
This species has no natural predators (Schreiber et al., 1999). B. quarlesi feeds on plants in both forest and open areas, including those created by avalanches and characterized by secondary plant succession (Sugiharta, 1994). Sugiharta (1994) noted 27 species of plants which were fed upon by B. quarlesi in Lore Lindu National Park in central Sulawesi. These included seven grass and grass-like species (Axonopus compressus, Oplesmenus corpositus, Panicum trichoides, Paspalum conjugatum, Carex filicina, Carex speciosa, and Carex verticillata), three ferns (Diplazium repandum, Neprolepis radicans, and Sphaerostephanos unitus), seedlings of three tree species (Castanopsis acuminatissima, Elaeocarpus moribundus, and Calophyllum soulattri), and several forbs (Calamus sp., Homalomena sp., Elaphantopus scaber, Ageratum conyzoides, Blumea balsamifera, Desmodium scorpiurus, Salvia hispanica, Pandanus furcatus, Rubus moluccanus, Solanum involucratum, Elastostema lineolatum, Tetrastigma pedunculare, Deparia patersonii, and Globa strobilifera). E. lineolatum and the three species of fern are fed upon most frequently (Sugiharta, 1994). Dung samples analysed by Green (in Whitten et al., 1987) were composed of 38% woody plants, 9% ferns, 16% broad-leaved plants, 17% grasses and sedges, 5% other monocots, and 13% moss. Anoa are reported to visit mineral-rich hot springs and salt licks - the National Research Council (1983) reports they may drink seawater, which might fulfil their mineral needs in areas without licks or springs.
Feer (1994) conducted a behavioral study of the mountain anoa, observing a captive pair and their two offspring (one male and one female, both immature) at the Krefeld Zoo, Germany. The results of this study are generalized here, but with caveats regarding the low sample size (four individuals) and non-natural habitat.
Defecation and urination are performed separately in B. quarlesi. Both males and females adopt the same posture when defecating, with the back hunched, hind legs crouched, and tail raised. Urination postures differ between the sexes, with males merely moving their hind legs back slightly, while females lower their rump by bending and spreading their hind legs, elevating the tail so it is more or less horizontal.
The adult male performed several behaviors which Feer suggested might serve to reinforce territorial ownership or the dominance rank within the group. These behaviors included using the horns to thrash bushes and branches, digging up soil with the horns, and scraping the ground with the front feet (typically after smelling the area, and followed about half the time by urinating over the scraped up soil). The hierarchy between individuals was reinforced with dominant individuals pushing the forehead or tapping the horns on the flank or rump of a less dominant animal, resting the chin or cheek on the rump, or placing one (rarely both) forelegs on the body. On the approach of a dominant animal or in response to contact, a submissive animal would adopt a posture with the head stretched low and tilted forward, with the horns pressed close to the neck. A more extreme submissive behavior involved the less dominant animal lying down, pressing the neck and chin to the ground.
Indirect aggressive encounters typically involved posturing and ritualized displays, with one animal lowering the head so that the muzzle was near the ground and forehead nearly vertical. If this posture is returned by the other animal, the two anoas face each other, but rarely approach. This "standoff" behaviour was seen in 18% of aggressive encounters, and is likely a threat used to assert a point but avoid an actual fight. In more aggressive situations, the head is bent further, with the forehead close to the ground and the horns pointed towards the adversary. Aggressive charges are directed straight at the adversary, with the head in a normal position (at the height of the withers) or raised with the ears pointing behind. This is accompanied by short, soft grunts or moos. If the opponent does not flee, the charge ends with a sharp blow. This may be followed by using the forehead to hit the shoulder or flank, or by using the horns to strike at the opponent's belly or flank with upward strokes. Among the four anoa in Feer's study, the adults were more likely to initiate indirect encounters (64%) than full-contact ones, while the two younger animals used direct contact with much greater frequency (95%). Aggressive behaviours initiated by the adult male were directed primarily towards the young male, while the adult female displayed these same behaviours primarily to the adult male. Parallel displays - seen in the majority of the Bovinae - were not observed by Feer, although they may occur in the presence of two adult males.
Males test the receptiveness of females to breeding by using "flehmen" (lip-curl) to expose the scent of fresh urine to the Jacobson's organ. The flehmen position in this species was usually held for seven seconds. (Males occasionally flehmen after licking their penis, while females may perform the behavior to a diverse array of olfactory stimuli). If the female is receptive, the male follows with his head stretched forward at the height of the shoulder and the neck horizontal - a position which allows the male to lick the female's vulva to induce urination. Stamping of the forefeet and raising the neck so that the throat comes into contact with the rump of the female escalate the pursuit. After a continued sexual parade interspersed with testing the female's urine, the male places his chin on the rump of the female and then mounts. During copulation, the head and neck of the male are pressed against the back and shoulders of the female, with the front legs pressed against her flanks. Throughout courting, both males and females make shorts moos.
Solitary play was seen in the form of galloping around or chasing at objects, including birds. Play behaviour with conspecifics was elicited by galloping towards them with wild leaps, or with a shaking of the head with a presentation of the horns. This would lead to play fighting, consisting of head-to-head shoving matches with horns interlocked and heads bent to the ground (and thus facing backwards). Occasionally, the anoas dropped onto their "knees" (carpal-metacarpal joints) when sparring. While fighting, the animals occasionally tried to disengage and butt the flanks of their adversary with their forehead or horns. In doing so, the pair of individuals would end up parallel to each other, circling around. Play combat was sometimes interrupted by face-offs (with the animals standing face to face with their heads lowered to the ground) and by rapid chases. Most play behaviour was observed between the two young individuals. Playful advances by a young male to an adult male elicited a response only rarely, while the adult female (his mother) responded more playfully.
Countries: Indonesia (Sulawesi) (Semiadi et al., 2008).
Anoa de montagne (Buchholtz, 1990, Semiadi et al., 2008)
- Anoa de Quarle (Semiadi et al., 2008)
- Berganoa (Buchholtz, 1990)
- Anoa de montaña (Semiadi et al., 2008)