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The taxonomic status, distribution and conservation of the lowland anoa Bubalus depressicornis and mountain anoa Bubalus quarlesi

January 2005
Mammal Review 35(1):25 - 50

January 2005
35(1):25 - 50

DOI:10.1111/j.1365-2907.2005.00048.x

Authors:

James Burton

International Union for Conservation of Nature

Simon Hedges

Zoological Society of London

Abdul Haris Mustari

Bogor Agricultural University

1. The anoas are two species of dwarf buffalo, the lowland anoa Bubalus depressicornis and mountain anoa Bubalus quarlesi that are endemic to the island of Sulawesi, Indonesia. The classification of the subgenus Anoa within Bubalus is upheld by assessment of recent genetic and morphological research. The classification of anoas into two species is still debated, but with the absence of significant opposing evidence, this position is adopted here. 2. Information about the distribution of the two species is presented that adds to but largely supports existing reports. However, it is still uncertain whether the two putative species are sympatric or parapatric in their distribution. A review of anoa distribution from historical reports and recent field data (1990s to 2002) highlights their decline throughout Sulawesi, especially in the southern and north-eastern peninsulas. The decline has been attributed to local hunting for meat and habitat loss. Most populations are rapidly becoming fragmented, suggesting that the conservation of viable populations may eventually require management of metapopulations. 3. There is an urgent requirement for conservation efforts to: (i) protect anoas from hunting; (ii) prevent habitat loss in key sites; (iii) complete genetic studies to better determine the number of anoa taxa and Management Units and assess their distribution; and (iv) determine the status of the remaining anoa populations.

The location of survey areas and forest cover in Sulawesi. Protected areas are shaded in black. Forested areas are shaded in light grey according to surveys between 1989 and 1995 (MacKinnon, 1997).

…

. Summary of anoa taxonomy

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Distribution of lowland Bubalus depressicornis and mountain anoa Bubalus quarlesi after 1990 and altitude (m asl). The map symbols represent locations where data have been collected. Each symbol may represent between 1 and 19 data points from a single location (e.g. numerous skeletal remains, direct observations or indirect signs on different occasions or locations within the site). See Appendix 1 for key to survey areas.Key to criteria for anoa distribution: B. depressicornis and B. quarlesi – see Table 1; Unknown Bubalus species – if an individual showed a morphological characteristic from both species or there was no data on characteristics, then the individual was included in the data set as ‘Unknown Bubalus species’; Unconfirmed presence – secondary data indicating presence since 1990; Locally extinct? – anoa presumed extinct during the period since 1990, after extensive field surveys or area no longer suitable for anoa; Absent – anoa never reported from this area; status uncertain – near local extinction.Altitude key: white 0–200 m asl; light grey 200–1000 m asl; dark grey +1000 m asl. Altitude measured in metres above sea level. Data courtesy of E. Colijn.

…

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Mammal Rev

. 2005, Volume 35, No. 1, 25–50.

Printed in Great Britain

Mammal Review

35,

25–50

Blackwell Science, LtdOxford, UKMAMMammal Review0305-1838Blackwell Publishing Ltd, 2004

2005

12550

Review Article

Anoa taxonomy, distribution and conservationJ. A.

Burton, S. Hedges and A. H. Mustari

Correspondence: James Burton. E-mail: james.burton@ed.ac.uk

The taxonomic status, distribution and conservation of the

lowland anoa

Bubalus depressicornis

and mountain anoa

Bubalus quarlesi

J. A. BURTON*, S. HEDGES† and A. H. MUSTARI‡

Preclinical Veterinary Sciences, Royal (Dick) School of Veterinary Studies, University of

Edinburgh, Edinburgh EH9 1QH, UK,

†

Wildlife Conservation Society (Asia Program), 2300

Southern Boulevard, Bronx, NY 10460, USA,

‡

Faculty of Forestry, Bogor Agricultural University,

PO Box 168, Bogor 16001, Indonesia

ABSTRACT

The anoas are two species of dwarf buffalo, the lowland anoa

Bubalus depressicornis

and

mountain anoa

Bubalus quarlesi

that are endemic to the island of Sulawesi, Indonesia. The

classiﬁcation of the subgenus

Anoa

within

Bubalus

is upheld by assessment of recent genetic

and morphological research. The classiﬁcation of anoas into two species is still debated, but

with the absence of signiﬁcant opposing evidence, this position is adopted here.

Information about the distribution of the two species is presented that adds to but largely

supports existing reports. However, it is still uncertain whether the two putative species are

sympatric or parapatric in their distribution. A review of anoa distribution from historical

reports and recent ﬁeld data (1990s to 2002) highlights their decline throughout Sulawesi,

especially in the southern and north-eastern peninsulas. The decline has been attributed to

local hunting for meat and habitat loss. Most populations are rapidly becoming fragmented,

suggesting that the conservation of viable populations may eventually require management

of metapopulations.

There is an urgent requirement for conservation efforts to: (i) protect anoas from hunting;

(ii) prevent habitat loss in key sites; (iii) complete genetic studies to better determine the

number of anoa taxa and Management Units and assess their distribution; and (iv) determine

the status of the remaining anoa populations.

Keywords

: buffalo, conservation, dwarf buffalo, Indonesia, Sulawesi, Wallacea

INTRODUCTION

Anoas are dwarf buffaloes, endemic to the Indonesian island of Sulawesi (and offshore

islands). They are the smallest of the Bovini, standing about 1 m tall at the shoulder. Two

species are presently recognized, the lowland anoa

Bubalus depressicornis

and the mountain

anoa

Bubalus quarlesi

(Groves, 1969; Corbet & Hill, 1992; Wilson & Reeder, 1993). Both

anoa species are classiﬁed as Endangered by International Union for the Conservation of

Nature (IUCN) (IUCN, 2002), are legally protected under Indonesian law (Jahja, 1987) and

are included in Appendix I of the Convention on International Trade in Endangered Species

of Wild Fauna and Flora (CITES) (UNEP-WCMC, 2000). There is much debate over the

taxonomic status and distribution of these two species (Mohr, 1921; Bohlken, 1958; Dolan,

1965; Groves, 1969; Weise, 1979; Kakoi

et al

., 1994; Pitra, Furbass & Seyfert, 1997; Schreiber

J. A. Burton, S. Hedges and A. H. Mustari

Mammal Review

35,

25–50

et al

., 1999). Nevertheless, the anoas are important ‘ﬂagship species’ for conservation, being

the largest endemic mammals on the highly bio-diverse island of Sulawesi. Conservation of

the anoas to date has focused on the protection of forested areas and on captive breeding

(Nötzold, 1999). The captive population doubled in size in the 10 years prior to the stud-

book’s publication; there were 125 animals in European, North American and Asian zoos in

December 1998. However, management of the captive breeding programme is hampered by

the uncertain taxonomic status of the anoas, and the uncertain classiﬁcation of those animals

already in zoos (Nötzold, 1999). Protected areas on Sulawesi cover 2223 km

or 11.9% of the

island (total area 181 000 km

), and 82% of the area under protection is reportedly still

forested (Aden

et al

., 2001). Unfortunately, until recently little information has been available

about the status of the anoas in these areas. This paper summarizes current knowledge about

the taxonomy and distribution of the anoas, and presents new data collected by the authors

and their colleagues. The conservation requirements of the anoas are also assessed.

BIOGEOGRAPHY OF SULAWESI

Sulawesi is situated within the Wallacea bioregion, renowned for its high level of endemism.

Of the mammal species found on Sulawesi, 61% (98% excluding bats) are endemic, compared

to just 18% on the neighbouring island of Borneo and 58% in Papua New Guinea (Whitten,

Mustafa & Henderson, 1987). Thirty-four per cent of Sulawesi’s bird species are endemic,

compared to just 6% on the neighbouring island of Borneo and 52% in Papua New Guinea

(Whitten

et al

., 1987). Being at the boundary of the Asian and Australasian biogeographical

regions, Wallacea comprises a mix of the Asian and Australasian fauna and ﬂora, including

two cuscus species (

Ailurops ursinus

and

Strigocuscus celebensis

) from Australasia, and two

suids (babirusa

Babyrousa babyrussa

and Sulawesi warty pig

Sus celebensis

) and the tarsiers

(

Tarsius

spp.) from Asia. The anoas are thought to have originated from Asia, as they are

close relatives of the Asian water buffalo

Bubalus arnee

(Groves, 1969).

Fossil and subfossil remains of anoas have been found in the Sulawesi Recent Faunas of

Late Pleistocene to Holocene age (van den Bergh, de Vos & Sondaar, 2001), but not from

earlier deposits as suggested previously (Hooijer, 1950). Both species of anoas were believed

to be present throughout the island’s forests in historic times (Weber, 1890; Sarasin & Sarasin,

1901; Mohr, 1921; Harper, 1945; Groves, 1969). However, their date of arrival on Sulawesi

and geological history of the island is less well understood. A number of theories have been

proposed to explain the presence of two morphologically similar species on Sulawesi, and to

clarify the taxonomic uncertainty. These include environmental variation resulting from

altitude (Heller, 1889; Whitten

et al

., 1987); and two separate immigrations (Kakoi

et al

1994; Kikkawa

et al

., 1997). Schreiber

et al

. (1999) suggest a number of additional possibil-

ities. In essence, their theories suggests that two populations of anoas were geographically

separated for a period on two landmasses before Sulawesi was ﬁnally formed (Hall, 1996,

1998), or after being spilt into smaller islands by rises in sea level. The latter is known to have

occurred during the Pleistocene from pollen records found in the south-western peninsula

(Fooden, 1969). Alternatively, variation in climate may have reduced forest cover as well as

led to forest fragmentation during the Pleistocene period (Whitten

et al

., 1987). This could

have led to the isolation of anoa populations and subsequent divergence, thus giving rise to

the two species thought to occur today. Unfortunately, there are currently insufﬁcient data

to choose between these alternative explanations (van de Berg

et al

., 2001).

It is still unclear if the anoas currently live sympatrically, or whether they are separated,

for example, by altitude. However, the distribution of Sulawesi’s other mammal species may

provide insights into the distribution of the anoas, and help answer the question. Between

Anoa taxonomy, distribution and conservation

Mammal Review

35,

25–50

ﬁve and seven

Macaca

species are present on Sulawesi (Fooden, 1969; Albrecht, 1978; Bynum,

1999). Their distribution is thought to be parapatric (Bynum, 1999), although interbreeding

between some of these populations is known to occur in hybrid zones (Ciani

et al

., 1989;

Watanabe & Matsumura, 1991; Evans, Supriatna & Melnick, 2001). However, the regional

distribution of these species does not appear to follow the island-wide distribution pattern

of the anoa species and the ﬁne scale distribution of anoa species remains unclear; they may

live in localized forest patches at different altitudes or sympatrically. Distinct species or forms

are found in another group of primates, the tarsiers of Sulawesi (Niemitz

et al

., 1991). At

least two species occupy distinct regions within Sulawesi’s lowland forests,

Tarsius dianae

and

T. spectrum

, while a further species,

T. pumilus

, is found at high altitudes (Shekelle

et al

1997; Groves, 1998). This altitudinal separation of tarsier species is likely to lead to a

parapatric distribution resulting from the uninhabited altitudinal band between high- and

low-altitude species. Further species such as squirrels and other small mammals, divided by

boundaries of unknown origin, have been recorded on Sulawesi (Musser, 1987; Groves, 2001).

In comparison, the distribution of anoa species appears to be more localized than the above-

mentioned mammal species.

TAXONOMY AND SYSTEMATICS

Systematics

The systematic status of the anoas has been a source of much confusion ever since the ﬁrst

specimens were described. Despite the close similarity between the anoas and the Asian

water buffalo, several authors have placed the anoas in a specially erected genus,

Anoa

Smith (1827) (Mohr, 1921; Pilgrim, 1939; Hooijer, 1948, 1950). However, Dolan (1965)

considered

Anoa

to be a subgenus within

Bubalus

and Groves (1969) also concluded that the

anoas show all the specialized characters of

Bubalus

and few signiﬁcant or absolute differ-

ences. Groves advocated employing

Anoa

H. Smith (1827) as a subgenus to better indicate

the relationship between the anoas

Bubalus (Anoa) depressicornis

and

B. (A.) quarlesi

, and

the water buffalo

Bubalus (Bubalus) arnee

and the tamaraw

B. (B.) mindorensis.

Recent

analyses of the anoas’ haemoglobin

chain amino acid sequences (Kakoi

et al

., 1994), and

the cytochrome

region of their mtDNA (Tanaka

et al

., 1996), support the placing of

Anoa

as a subgenus within

Bubalus

. Tanaka

et al

. (1996) estimate a coalescence time (mean esti-

mated divergence time) for the present

Bubalus

species at around 2 million years, using the

divergence time of

Bos

and

Syncerus

for comparison (corroborated from fossil records)

(Savage & Russell, 1983).

Comparative studies of present day anoa skulls and fossil buffalo remains from between

3.20 and 1.78 million years ago (Mya) suggest that the anoas might be members of the most

plesiomorphic clade of wild cattle (Bovini: oxen, bison, yak and buffalo) (Geraads, 1992;

McDougall

et al

., 1992). Other studies of allozyme data and antigenic epitopes of the trans-

ferin molecule also support this theory (Hartl

et al

., 1988; Schreiber, Erker & Bauer, 1990).

However, recent analysis of nuclear DNA (partial nucleotide sequences of the aromatase

cytochrome P450 and lactoferrin genes) suggests that lowland anoa are most closely related

Boselaphus

not

Bubalus

(Pitra

et al

., 1997). Pitra and co-workers also argue that their

nuclear DNA data indicate that the

Anoa

and boselaphine lineage diverged from the ancestral

bovine lineage before the buffalo and cattle lineages, some 12.4 Mya. This contradicts the

hypothesis that

Anoa

diverged much more recently (

2 Mya) from the ancestral

Proam-

phibos–Hemibos

lineage leading to the living Asiatic buffaloes (Groves, 1981). If Pitra and

co-workers are correct,

Anoa

should probably be treated as a genus of Asiatic antelope (i.e.

included in an expanded Boselaphini tribe) rather than a subgenus within

Bubalus

. Never-

J. A. Burton, S. Hedges and A. H. Mustari

Mammal Review

35,

25–50

theless, as Pitra

et al

. (1997) noted, further studies are needed to test their conclusions.

Therefore,

Anoa

will here be treated as a subgenus of

Bubalus

following Groves (1969).

How many species?

Most classiﬁcations accept the existence of two forms, lowland anoa and mountain anoa,

and many authorities have treated these forms as two species, with or without additional

subspecies (Harper, 1945; Groves, 1969; Honacki, Kinman & Koeppl, 1982; Corbet & Hill,

1992; Wilson & Reeder, 1993). Nevertheless, uncertainty remains over the validity of these

two species. The English common names relate to a putative altitudinal separation (Groves,

1969), with the large form (lowland anoa) inhabiting low-lying areas and the smaller form

(mountain anoa) living at higher elevations. The locality records of available specimens

suggest this altitudinal separation, according to Groves (1969). However, no detailed com-

ment on habitat variation was given to explain this supposed altitudinal boundary between

the ranges of the two species. Furthermore, it has also been suggested that altitude would

cause a clinal variation within one species of anoa, rather than indicate the existence of two

parapatric species (Heller, 1889; Whitten

et al

., 1987).

Weise (1979) attempted a multivariate analysis using 26 measurements from 57 anoa skins

and skulls (a greater number of skulls and measurements than Groves’ study). However,

Weise found insufﬁcient data to show a statistically signiﬁcant clinal range in skull size and

also concluded that there were no biometric differences between lowland and mountain anoas

that could not be explained as a result of allometric transformations because of the smaller

(by 15%) size of the mountain anoa. Careful analysis of Weise’s paper indicated that this is

not a surprising ﬁnding because the criteria he used to classify 36 of the 57 skulls of unknown

locality into mountain or lowland anoas were based on horn and size characters. We feel that

the conclusions of the morphological analysis should be interpreted cautiously because a

priori skull size and horn characteristics were used to identify the anoa species. Weise (1979)

analysed 49 skull characteristics in pairs and in 40 of these correlation cases showed only

size-dependant proportional difference when using multivariate analysis plots: mountain

anoa skulls overlapped well within the lowland anoa grouping. He points to inconsistencies

with characteristics used to identify the forms. He also stated that from the morphological

data and from suggestions that there is contact between the populations and possible admix-

ture in the wild, anoas can only be differentiated to subspecies level.

Others have also reported intermediate forms in the wild and in captive collections, sug-

gesting interbreeding between the two species (Schreiber & Nötzold, 1995; S. Hedges, per-

sonal observations; J. Burton, personal observation). In addition, the different pelage

characters of both species described in Table 1 have been observed in a single animal from

birth to sexual maturity (J. Burton, personal observations).

Nevertheless, recent genetic studies of anoas in zoo collections support the two species

hypothesis. For example, analysis of partial mtDNA cytochrome

gene sequences (285 bp)

revealed clear differences between mountain anoas (from Krefeld Zoo) and lowland anoas

(from Berlin, Leipzig and Antwerp zoos) (Schreiber

et al

., 1999). Of the eight animals sam-

pled, four haplotypes were identiﬁed, within which 11 sites (3.9%) were variable. This level

of nucleotide substitution was far higher than the intraspeciﬁc difference found in three other

species [two dwarf zebus,

Bos (Bos) taurus

; two American bison, Bison bison; and four gaur,

Bos (Bibos) frontalis]; indeed these species showed no variation. Three further species had

0.4% nucleotide substitutions, also lower than in the anoas [four African buffalo, Syncerus

caffer; three European bison, Bison bonanus; and ﬁve banteng, Bos (Bibos) javanicus]. The

interspeciﬁc variation within the anoas (between mountain anoa and lowland anoa) was

Anoa taxonomy, distribution and conservation 29

3.1%. Comparing these data with the interspecies variation between recognized species, the

anoas have greater per cent nucleotide substitution rates than between domestic yak Bos

(Poephagus) grunniens and American bison (2.1%); gaur and banteng (3.2%); and gaur and

yak (3.3%). These results clearly corroborate the two species hypothesis.

Two further studies using complete mtDNA cytochrome b gene sequences (1190 bp) also

support the two species hypothesis (Tanaka et al., 1996; Kikkawa et al., 1997). These studies

were conducted using samples from two and ﬁve captive anoas from Indonesian zoos, respec-

tively. The results of the study by Tanaka et al. (1996) suggest that the mountain anoa and

lowland anoa should be regarded as two species as they found the weighted genetic distance

between lowland and mountain anoas to be 0.0354. This was similar to the weighted genetic

distance between the anoas (mountain and lowland) and water buffalo (both river and swamp

types), clearly distinct species. The weighted genetic distance between anoas was greater than

the intraspeciﬁc variation of domesticated water buffalo (swamp and river buffalo) (0.0256).

Kikkawa et al. (1997) analysed samples from ﬁve anoas. Without clarifying criteria for

describing subspecies, they identiﬁed these animals into three subspecies of anoa, ‘lowland’,

‘mountain’ and ‘quarlesi’. The estimated sequence divergence between their ‘lowland’ and

‘mountain subspecies’ was 3.6%, and between their ‘lowland’ and ‘quarlesi subspecies’ was

3.3%. These sequence divergence differences are closer to those found between ‘good’ species

such as the anoas and water buffalo of the swamp type (3.33%), than those found within

water buffalo (i.e. between river and swamp type buffaloes) (2.67%). Even though the mor-

phological descriptions of the different ‘subspecies’ are unclear, variation between these

animals is equivalent to that seen between other distinct species. Kikkawa et al. (1997) state

differences between anoas and water buffalo were not signiﬁcant at the genus level, but that

the results showed there are two lineages within anoa.

Table 1. Summary of anoa taxonomy

Species

Lowland anoa Mountain anoa

Valid name Bubalus (Anoa) depressicornis (Smith, 1827). Bubalus (Anoa) quarlesi (Ouwens, 1910)

Synonyms Antilope depressicornis, Bos bubalus anoa, Bos

depressicornis fergusoni, Oreas platyceros,

Probubalus celebensis

Anoa quarlesi, Anoa anoa, Bubalus

depressicornis quarlesi

Holotype Nearly complete skull with horns of Antilope

depressicornis Hamilton Smith, 1827; in the

British Museum (Natural History),

London (B.M. 607a)

Three specimens in the Amsterdam

Museum are labelled as syntypes: skin

and skull from a juvenile (ﬁrst molar

present) (a male?) (ZMA 9288); skin

and skull from an older juvenile

(second molar present) (a female?)

(ZMA 9289); the third, an infant skull

(ZMA 9295), is considered by Groves

(1969) to be erroneously labelled as a

type because Ouwens only refers to

two animals in his paper

Type locality Celebes [= Sulawesi, Indonesia] Mountains of central Toradja [= Toraja]

district, Celebes [= Sulawesi,

Indonesia]

Paratype Skull from an unsexed subadult (third molars

erupted but unworn); in the British

Museum (Natural History), London (B.M.

8.12.23.1)

30 J. A. Burton, S. Hedges and A. H. Mustari

Earlier work (Schreiber, Nötzold & Held, 1993), which looked at allozyme distances and

gross body morphology, agreed with, but raised some doubts about the two species hypoth-

esis. The researchers suggested that there was less variation between the anoas in European

zoos than could be found between other closely related species. Electrophoretic distance of

allozymes was used to investigate differences between 25 anoas. Pairwise comparisons of Nei’s

genetic allozyme distances for anoa were D = 0.0206 to D = 0.0505, while for distinct wild

bovid species the values were D = 0.1389 to D = 0.7621 (Hartl et al., 1988). Therefore, these

allozyme distances were more similar to those found between geographical races or subspecies

(Nei, 1987). However, Schreiber and his co-workers cautioned that the taxonomic relevance

of these genetic distances was open to question because of the effect of bottlenecks and

incestuous breeding. In addition, we would argue that these results are also weakened because

of uncertainty over the zoo animals’ origins and lineages. Schreiber et al. (1993) concluded

that the coincidence of differences encountered in karyotypes, polymorphic genes, body size,

hair texture and other phenotypic characters (including horn shape) indicated that the anoas

comprise differentiated taxa, but that the electrophoretic distance variation is less than is

typical for bovine species.

There are at least two possible explanations for the difference in mtDNA and allozyme

distance results discussed above. First, mountain and lowland anoa are well-separated spe-

cies but hybridization in the captive population caused the observed pattern of protein

variation (Schreiber et al., 1993), whereas the maternally inherited mtDNA sequence reﬂects

only the genotype of the founder females (Tanaka et al., 1996; Kikkawa et al., 1997;

Schreiber et al., 1999). Alternatively, the differences in the mtDNA sequences could be a

chance effect caused by the sole female founder of the B. quarlesi species belonging to

another mitochondrial lineage. Such polymorphism would have no taxonomic implications

but would merely indicate the presence of very deep mitochondrial lineages and testify to the

great age of the taxon. Recent studies of well-differentiated haemoglobin sequences are in

accordance with the second possibility (Schreiber & Goltenboth, 1990; Schreiber et al.,

1993). Schreiber et al. (1999) suggest limited gene ﬂow between populations could be sufﬁ-

cient to explain the lack of distinct variation in the wild, but would be insufﬁcient to prevent

differentiation within mtDNA over long periods. This has been shown to occur in other

species (Avise, 1994), including Sulawesi macaques in zones of hybridization (Evans et al.,

2001).

Recently, it was suggested that there might be more than two species of anoa (Sugiri &

Hidayat, 1996). Sugiri and Hidayat took samples from ﬁve wild-caught anoas from the

mountains of Central Sulawesi. Three animals from one area had the same chromosome

number of 2n = 46 as captive mountain anoas, but another of the wild anoas had a chromo-

some number of 2n = 44. The ﬁfth animal’s chromosome number was not reported. Captive

lowland anoas reportedly have a chromosome number of 2n = 48. Sugiri & Hidayat (1996)

suggest that further work is needed, especially as the chromosomes of the individual with

2n = 44 ‘could not be identiﬁed’ as being ‘meta-, sub or acro-centric’. Schreiber et al. (1993)

also argue that the biological meaning of anoa chromosomal variation needs to be investi-

gated, as published chromosome numbers vary considerably, with 2n = 44, 45, 47 and 48 in

European zoos (Koulischer, Tyskens & Mortelsmanns, 1972; Schreiber et al., 1993). In addi-

tion, lowland anoas in Indonesian zoos have been reported to have chromosome numbers of

2n = 36 and 2n = 38 (Amano & Martojo, 1983). More recently, anoas at Ragunan zoo have

been reported to have chromosome numbers of 2n = 48 in lowland anoa, 2n = 46 in mountain

anoa, and 2n = 42 and 2n = 38 in anoas of unidentiﬁed type (Pranadewi, 1998; Marsono et al.,

2001). If this variation exists, then it may show that the taxonomic diversity of anoas has

Anoa taxonomy, distribution and conservation 31

been underestimated in the current classiﬁcation. Other closely related species such as the

water buffalo have chromosome numbers of 2n = 48 and 50 (swamp and Murrah types) and

the tamaraw 2n = 46, less variation than found between anoa species. Clearly this issue

requires further investigation.

A number of studies have suggested the existence of distinct subspecies (Heller, 1889;

Dolan, 1965; Bartikova & Dobroruka, 1973; Foead, 1992; Schreiber & Nötzold, 1995;

Kikkawa et al., 1997). Groves (1969) also discussed the possibility of subspeciation within

the anoas. However, he concluded that while there did appear to be some geographical

variation in size, the number of specimens available was insufﬁcient to assess the signiﬁcance

of this variation.

In conclusion, with the absence of any overwhelming evidence to contradict the systematic

position of the anoas presented by Groves (1969) and adopted by Honacki et al. (1982),

Corbet & Hill (1992) and Wilson & Reeder (1993), the two species, B. depressicornis and B.

quarlesi, are accepted as valid taxa here. However, it does seem likely that revisions will need

to be made within the subgenus Anoa.

MORPHOLOGY

The anoas are the smallest of the extant wild cattle species (Groves, 1969). They are stocky,

short limbed and thick necked. Young anoas are generally reported to have a thick covering

of yellowish-brown woolly hair but there is much variation; newborn calves in zoos have

brown or black pelage. Adults are also variable in colour but are predominantly brown or

black. Males are usually darker than females and both have horns. The anoas are reputed to

have exceptionally thick hides. The following descriptions of the two (putative) species

(Table 2) should be regarded as a guide only, given the apparently high levels of individual

variation shown by these animals and the uncertainty that remains about their taxonomic

status.

Schreiber et al. (1993) concluded that the phenotypic characters conventionally used to

distinguish lowland and mountain anoas including horn cross-section, shoulder height, body

markings and hair characteristics are ambiguous, particularly if only a few animals are

available for comparison. Their conclusions were based on an examination of animals in

zoological collections. Unfortunately, doubts have been raised about these animals’ origins

in Sulawesi. Furthermore, incestuous breeding and possible hybridization between lineages

reduce the value of these conclusions (Schreiber et al., 1993; Nötzold, 1999).

Criteria for identifying anoa species

Morphological criteria were identiﬁed from previous descriptions of anoas (Heller, 1889;

Dolan, 1965; Groves, 1969; Walker et al., 1975; NRC, 1983; Groves, 1985; Grzimek, 1990).

We reviewed all those characters we judged likely to be of utility to select a subset that was

not disputed and that was congruent with our own experiences of examining both live anoas

and skeletal material (Table 2).

These criteria are used in the present review of anoa distribution and status. However, if

an individual showed morphological characteristics from both species, or if no data relating

to these characteristics were available, we consigned the record to the ‘Unknown Bubalus

species’ category. Juvenile B. depressicornis are similar to adult B. quarlesi in some features.

In order to identify a juvenile B. depressicornis, either white leg markings had to be present

or there had to be clear proof that the animal was indeed a juvenile (indicated by the lack of

third molar teeth). Much uncertainty still remains, which is confounded by the overlap in

morphological characteristics (Table 2).

32 J. A. Burton, S. Hedges and A. H. Mustari

Table 2. Morphological characteristics of two anoa species: Bubalus depressicornis and Bubalus quarlesi

Species

Lowland anoa Bubalus depressicornis Mountain anoa Bubalus quarlesi

Body size Shoulder height: 60–100 cm; mass < 300 kg†

Body length 170–188 cm¶

Shoulder height: < 75 cm; mass < 150 kg; body

length 122–153 cm

Horns*† Triangular cross-section ﬂattened, marked transverse ridges &

marked external keel; horn length: lowland male 271–373 mm;

female 183–260 mm*‡

Short, conical and rounded cross-section, no marked

ridges or external keel in juvenile & adult; both sexes:

horn length 146–199 mm*

Overall body pelage colour Black & sparsely (woolly brown in juvenile [M3]); sparse & straight,

often rubbed off with age*

Dark brown to black adult & thick & woolly haired into

adulthood; female coat especially woolly*

White facial/neck markings White facial markings present* White facial markings sometimes present

Often white throat markings*†‡¶ White throat markings never present*

White leg markings (see age

dimorphism)‡§

Forelegs: always white to yellowish-white from knee to hoof (black

strip down front & across pasterns). Hind legs: conspicuous white

spots above hooves. Yellow in juvenile, to white in adult†

For eleg: only whitish-yellow spots above hooves,

sometimes absent§

Tail length (as percentage of

total body length)*‡

19.8–25.8% (9 skins)* 14.6–17.8% (5 skins)*

Groin Light coloured to white* Light coloured but not white*

Age-related dimorphism Juvenile form resembles adult mountain anoa (in following characteristics:

pelage colour, texture, horn shape, body size), BUT lowland anoa always

has white forelegs, conspicuous white spots on hind legs

above hooves (see Dolan, 1965)*

Horn index of least–greatest basal horn breadth

(anterio-posterior to bilateral)*

Skull length Male 293–322 mm; female 290–300 mm* Both sexes skull length 244–290 mm*

*Groves (1969). Species Identiﬁcation Characteristics for the two species.

‡Hedges (1996). Species Identiﬁcation Characteristics. Taken from: Harper (1945); Dolan (1965); Groves (1969); Walker et al. (1975); NRC (1983); Groves (1985); Grzimek

(1990);

†Schreiber et al. (1993). Species Identiﬁcation Characteristics.

§National Research Council (NRC, 1983) states that the entire lower limbs of mountain anoas are creamy white in colour, but all other published descriptions have emphasized

the generally dark-coloured legs of this species.

¶A. H. Mustari, unpublished observation. Note on body mass: all those recorded (n = 8) have been below 150 kg for individuals identiﬁed as B. depressicornis as well as B.

quarlesi.

The features in bold face are those used by the present authors to provisionally identify the species.

Anoa taxonomy, distribution and conservation 33

ECOLOGY AND BEHAVIOUR

Very little is known about the ecology and behaviour of the anoas, although they are

thought to be solitary, forest-dwelling browsers (Whitten et al., 1987; Foead, 1992). In a

recent study, 21 of 40 ﬁeld observations were of a single animal (Mustari, 1995). Anoas

reportedly use many types of forest, from lowland forest, including secondary formations

and mangroves, to montane forest at up to 2300 m above sea level (NRC, 1983; Mustari,

1995). In lowland habitats, anoas have been estimated to occur at densities of 1.3 animals/

km2 (with a 95% conﬁdence interval of 0.5–3.1 animals/km2) in Tanjung Peropa Nature

Reserve (NR) (389 km2) and 1.1 animals/km2 in Tanjung Amolengu Wildlife Reserve (WR)

(5 km2) (A. H. Mustari, unpublished data). These densities were estimated using direct

observation/encounters calculated from line transects, totalling 372 km in Tanjung Peropa

and 124.3 km in Tanjung Amolengo. However, the density estimate for Tanjung Amolengu

WR may have been higher than was sustainable, as animals were known to cross to this

forest from adjacent areas. The typical life span in captivity is reported to be 20–30 years

(Walker et al., 1975; NRC, 1983; Grzimek, 1990). Sexual maturity is attained in the second

to third year (in captivity), with generally only a single calf born per year (NRC, 1983; Jahja,

1987).

DISTRIBUTION

Mohr (1921) provided the ﬁrst distribution map of the anoa but this had little detail. More

recently, Groves (1969) presented a much more detailed map of the anoas’ historical distri-

bution. However, Weise (1979) challenged the taxonomic attribution by Groves of some of

the specimens examined, such as the skulls from the Boro-Boro Mountains area. Since

1990, the present authors have conducted a detailed assessment of anoa distribution, as

well as a review of published information. Criteria for identiﬁcation of anoa species are

described above. Data were collected during forest surveys. Surveyors recorded both direct

and indirect signs (tracks, faeces and feeding traces), although obviously these indirect

signs could not be used to identify anoas to species. Direct observations of anoas are rare,

as indicated from transect data in three areas in South-East Sulawesi: two animals were

seen during 182.1 km of transects in Tanjung Peropa; three anoas during 50 km of

transects in Tanjung Batikolo; and three in 202.7 km of transects in Rawa Aopa Watumo-

hai National Park (NP) (Riley, Hunowu & Soleman, 2001a; Riley et al., 2001b). Further-

more, even the use of automatic cameras (‘camera-traps’) produces few records; for

example, only one individual was photographed during a total camera-trapping time of

4930 hours in Tanjung Peropa (Riley et al., 2001b), while in Rawa Aopa Watumohai NP

none were photographed in 3523 hours of camera-trapping (Riley et al., 2001a). We also

conducted semi-structured questionnaire surveys, and recorded all observations of skeletal

remains found. The majority of information relating to the identiﬁcation of species came

from skulls collected by hunters. These data were used as an indicator of distribution

(presence/absence) of the two species. Questionnaire data were treated with caution

because of the misidentiﬁcation of the different age- and sex-classes of anoas as distinct

species in some local nomenclature.

Survey areas are shown in Fig. 1; the distribution of both anoa species recorded since 1990

is shown in Fig. 2. Further information on anoa distribution and the threats to anoas in each

area can be found below and in Appendix 1. Most mountain anoa records are from the

central region of Sulawesi where much of the forest is above 1000 m asl; the only other

mountain anoa record is from Buton Island, in Buton Utara WR (49) where there are also

mountains of around 1000 m asl (Fig. 2). Lowland anoas records are distributed mainly

34 J. A. Burton, S. Hedges and A. H. Mustari

below 1000 m asl. Thus, the data collected during this study tend to support either the

suggested altitudinal separation of the two anoa species (Groves, 1969) or the altitudinal cline

suggestion (Heller, 1889; Whitten et al., 1987). Nevertheless, the resolution of the map is

rather poor (location of symbols do not exactly represent the location of data points because

of the collection of data from villages and the altitude ranges are broad). So it is worth noting

that MacKinnon (cited in Thornback, 1983) states that in mountainous areas one tends to

ﬁnd mountain anoa, but within this region this species can be found at sea level as well as at

higher elevations. Meanwhile in areas with lowland anoa, this supposedly lowland form

ranges up to at least 1500 m asl (e.g. in the Gunung Tangkoko and Gunung Ambang areas).

Again, further survey work is required.

Figure 1 shows the location of Sulawesi’s protected areas relative to the island’s forest cover.

Signiﬁcant areas of unprotected forest remain between Panua NR/Marisa Complex (6), Gn.

Fig. 1. The location of survey areas and forest cover in Sulawesi. Protected areas are shaded in black. Forested

areas are shaded in light grey according to surveys between 1989 and 1995 (MacKinnon, 1997).

Anoa taxonomy, distribution and conservation 35

Dako Proposed NR (16) and Gn. Tinombala (17) in the north; in the Gn. Lumut/Gn.

Hohoban (27) area in the east; around Pengunungan Tangkeleboke (including Gn. Men-

konga) (53), P. Abuki (54) and P. Matarombea (55); and to the west and south of Lore Lindu

NP (11). It is not known whether most of these unprotected areas contain anoas. Neverthe-

Fig. 2. Distribution of lowland Bubalus depressicornis and mountain anoa Bubalus quarlesi after 1990 and

altitude (m asl). The map symbols represent locations where data have been collected. Each symbol may

represent between 1 and 19 data points from a single location (e.g. numerous skeletal remains, direct

observations or indirect signs on different occasions or locations within the site). See Appendix 1 for key to

survey areas.Key to criteria for anoa distribution: B. depressicornis and B. quarlesi – see Table 1; Unknown

Bubalus species – if an individual showed a morphological characteristic from both species or there was no

data on characteristics, then the individual was included in the data set as ‘Unknown Bubalus species’;

Unconﬁrmed presence – secondary data indicating presence since 1990; Locally extinct? – anoa presumed

extinct during the period since 1990, after extensive ﬁeld surveys or area no longer suitable for anoa; Absent

– anoa never reported from this area; status uncertain – near local extinction.Altitude key: white 0–200 m asl;

light grey 200–1000 m asl; dark grey +1000 m asl. Altitude measured in metres above sea level. Data courtesy

of E. Colijn.

36 J. A. Burton, S. Hedges and A. H. Mustari

less, anoas were still present throughout much of the island of Sulawesi in the 1990s (Fig. 2).

The data compiled and reviewed here and depicted in Fig. 2 are in good agreement with the

distribution map of Groves (1969); for example, both species were present in the south-east

and central regions of Sulawesi. Additional information collected since 1990 has shown that

lowland anoa inhabit the eastern peninsular, where it was previously unreported. Anoas were

thought not to have reached most of Sulawesi’s small neighbouring islands, except for Buton

Island in the south-east. Both species were reported from Buton Island, which is again

noteworthy, as previously only mountain anoa was known to occur. Both species appear to

inhabit Lore Lindu NP and Buton Utara WR. In addition, there are a number of areas where

anoas are known to occur, but they have been incompletely identiﬁed as ‘Unknown Bubalus

species’. This is mainly caused by the lack of morphological descriptions of the anoas present.

More detailed surveys are needed.

The north-eastern and the southern parts of the island are notable for their lack of forest

cover (Fig. 1), and it appears that only small and isolated populations of anoas remain in

these areas (i.e. in Gunung Manembo-nembo WR, Gunung Ambang WR and Bulu Saraung

NR). These populations are at high risk of local extinction. In Tangkoko – Dua Saudara NR,

hunting is thought to have been the main cause of the estimated 90% decline in lowland anoa

numbers between 1978/79 and 1993/94 (O’Brien & Kinnaird, 1996). The need to protect

anoas and their habitat is discussed below.

From our review of the anoas’ distribution and status, it is possible to identify areas that

are likely to be of key importance for the conservation of these animals. Lore Lindu NP in

Central Sulawesi (2310 km2) still has abundant signs of anoa in the high elevation central

areas (Burton, 2002). This park still has much undisturbed montane forest, forming one

continuous area. In 2002, anoas signs were observed in a number of locations within the

park. In Bogani Nani-Wartabone NP (2871 km2) in North Sulawesi and Gorontalo Provinces,

there are reportedly still signiﬁcant numbers of anoa judging by the frequency of indirect

signs seen in the forest (J. Riley, personal communication). Anoas have also been seen and

photographed (using camera-traps) in the area in 2000–2001 (Pangau, 1997; J. Riley, personal

communication). In South-East Sulawesi, camera-trap photographs were taken of anoa in

Tanjung Peropa NR (389 km2) (Riley et al., 2001b). Frequent sightings have also been made

in this area. On Buton Island, observations of lowland anoa have been made in Lambu Sango

WR (250 km2) (Burton, 2001).

CONSERVATION STATUS

Hunting (mainly for meat) and the loss of suitable habitat are the major threats to the anoa,

and recent reports indicate that hunting is by far the more serious of the two (Appendix 1).

In rural areas, there appears to be little awareness of the protected status of anoas, and

villagers readily recounted their hunting experiences to researchers (Schreiber & Nötzold,

1995). Recently, anoas have been extirpated from Tangkoko – Dua Saudara NR (J. Riley,

personal communication), and this is thought to have largely resulted from hunting

(O’Brien & Kinnaird, 1996). In the far larger protected area of Rawa Aopa Watumohai NP

(968 km2), there have been major losses of anoas because of severe anthropogenic effects of

habitat degradation for logging, agriculture and settlements, as well as hunting pressures

(Burton, 2001). In Lore Lindu NP, anoas have apparently declined, and it is suggested that

they are now restricted to core areas that are less encroached upon by humans

(Appendix 1). There have apparently been signiﬁcant declines in distribution within many

other protected areas too (Harper, 1945; Groves, 1982; NRC, 1983; Thornback, 1983; Jahja,

1987).

Anoa taxonomy, distribution and conservation 37

Hunting

Although many hunters are still willing to discuss their activities, this is becoming less

common, making collection of accurate information on hunting pressures more problematic.

However, minimum hunting rates and the reasons behind these are discussed below. Identi-

ﬁcation of skeletal remains and hunting location data gathered from hunters in eight villages

around Lore Lindu NP (2130 km2) showed that about 13 anoas were taken from the park

per year by this small number of villages. In Lambu Sango WR (250 km2), a minimum of 15

animals per year were caught by hunters in 19 villages (J. Burton, personal observations).

The recording of anoa skulls found in villages in Central Sulawesi showed a catch rate of 1.5

anoa/village/year (11 villages, within boundaries of 600 km2). In this area a total of 161 skulls

or skins (from different individuals) were observed, and questionnaires suggested 443 animals

had been collected over a 40-year period (Kasim, 2000). The latter ﬁgure is a catch rate of 1

animal/village/year. A similar survey in Kecamatan Palolo (near Lore Lindu NP) identiﬁed

34 skulls from animals caught in 1998, six skulls in 1999 and three in 2000. A further 153,

22 and 2 anoas having been caught in these years according to questionnaire surveys

(Nurlaela, 2001). The actual numbers of anoas caught in each of these study areas is likely

to be much higher, as people forget the number of animals caught or will be frightened to

admit to the full scale of the hunting of these protected species.

In Morowali NR, Central Sulawesi, hunter-gatherers (from the Wana ethnic group) are

thought to be catching anoas at an unsustainable rate (Avalard, 2000). Avalard suggested

that the Wana would require about 700 km2 of forest containing anoa to sustain a population

large enough for them to hunt at current levels, whereas Morowali is only 225 km2. As the

anoas comprise 40% (by weight) of the large game animals harvested by the Wana, it is likely

that anoas will be hunted to near extinction in Morowali and the Wana will then have to

concentrate hunting effort on other species (Avalard, 2000).

Recent ﬁeld surveys in Lore Lindu NP showed that areas close to settlements had a lower

frequency of indirect signs of anoas (tracks and faeces) recorded on transect surveys (Burton,

2002a). An indication of human pressure was given a subjective category of Low, Medium

or High Human Disturbance relating to the number of trails, agricultural ﬁelds, traps and

people observed in the forest. Areas of higher disturbance level had a low frequency of anoa

indirect signs (Burton, 2002).

In North Sulawesi, where hunting is thought to be a more serious problem than in other

parts of Sulawesi, anoas are still being hunted (Pangau, 1997). However, little anoa meat has

been sold in markets since the mid-1990s, apparently because of the local peoples’ fear of

punishment for engaging in these illegal activities (Clayton & Milner-Gulland, 1999). Patrol-

ling with the aim of stopping the trade in illegally hunted meat on the roads and in the

markets of Minahasa, North Sulawesi has shown that there is less anoa meat on sale relative

to that of the other Sulawesi ungulates (Clayton & Milner-Gulland, 1999). However, it is now

thought that people consume anoa meat in local villages and the forest, instead of transport-

ing it to market and risk being caught.

Hunting methods vary between regions, but the most common techniques are snares or

spears and dogs, and less commonly guns (Whitten et al., 1987; Mustari, 1995; Avalard, 1999;

O’Brien & Kinnaird, 1999). Hunters also set ﬁres to encourage new growth and so attract

anoa and make hunting easier (Whitten et al., 1987).

The main reason for the widespread hunting of anoa is to obtain meat for local consump-

tion (Thornback, 1983; Schreiber & Nötzold, 1995). For example, in South-East Sulawesi,

14 hunters were questioned: over the last 5 years the majority hunted occasionally, catching

between one and 10 anoa each. However, one hunter said he caught 10 animals in 1 year.

38 J. A. Burton, S. Hedges and A. H. Mustari

Five of the hunters said they sold the meat, while the others consumed it in the forest or

within the local community. Anoa meat was usually sold for Rp5000–7500/kg (c. £0.50/kg),

but one hunter said he could sell anoa meat for Rp22 500/kg (c. £1.60/kg) (J. Burton, personal

observations).

Anoa skulls and horns do not seem to be particularly valued as trophies by hunters, and

while quantitative data are unavailable, trade in trophies does not appear to pose a serious

threat to the anoas (Schreiber & Nötzold, 1995; S. Hedges, personal observations). There

have been occasional reports of illegal trade in live animals in the past; for example, it was

reported that an animal trader in South-East Asia offered to sell a pair of lowland anoas at

US$3000 each (Anonymous, 1976). However, no signiﬁcant international trade in either live

animals or body parts has been reported in recent years. Trade does exist within Sulawesi,

where live animals are sold for later slaughter or as pets. An animal can be sold for £29.00–

36.00 (Rp400 000–500 000), although prices can reach as much as £71.00–142.00

(Rp1 000 000–2 000 000) in extreme cases (Lee, 1999a; J. Burton, personal observations).

The effect of hunting on the long-term survival chances of populations of anoa in Sulawesi

was estimated during the Anoa Population and Habitat Viability Analysis Workshop

(PHVA). Available data on life history were used with the VORTEX Program to estimate the

population viability of different population sizes under three levels of hunting pressure

(Manansang et al., 1996). Hunting rates were taken from the estimated 15% per year decline

of the Tangkoko – Dua Saudara NR population (O’Brien & Kinnaird, 1996) and lower levels

of 8% and 4%. Population viability estimates were calculated for initial populations of 25,

100, 360, 1500 and 2860 animals during a 100-year simulation, with the carrying capacity set

at twice the original population size. It is currently unlikely any population in a protected

area is larger than the greatest population modelled above. The simulations showed that all

populations became extinct with the 15% off-take hunting level. The 8% hunting level caused

the local extinction of all except the two largest populations (1500 and 2860 individuals)

within 100 years, but even the largest populations were at high risk of extinction soon after

100 years. Even an initial population size of 2860 anoas with 4% hunting loss caused a greater

than 90% decline in 100 years to just 232 animals. These simulations highlight the threat that

even the largest anoa populations face from hunting levels that could now be occurring

throughout Sulawesi.

Loss of suitable habitat

Habitat degradation and loss is the second most signiﬁcant threat to the anoas. Many anoa

populations are becoming isolated as the forest around the protected areas is cleared or

converted to plantations. Total forested area remaining on Sulawesi in 1985 was estimated to

be between 42 and 49% of the land area of the island (Holmes, 2000; Mathews, 2002). The

loss of lowland forest between 1985 and 1997 was estimated to be 89% (Holmes, 2000).

Recently, it has been said that ‘Sulawesi’s lowland forest has already been reduced to statistical

insigniﬁcance’ (Aden et al., 2001).

Although 11.9% of the island is protected, loss of suitable anoa habitat within protected

areas is as much a problem as in unprotected forest (Aden et al., 2001). For example, the

expansion of agricultural and settled areas, logging, mining and ﬁres have encroached into

many protected areas. Large protected forest areas such as the eastern end of Bogani Nani-

Watabone NP and the Besoa and Lindu areas of Lore Lindu NP are being fragmented by

roads (Burton, 2002). The fragmentation of these once contiguous forests will reduce the

natural movement of anoas between forest patches. This may cause the loss of genetic

diversity and increase the chance of population extinctions (Gilpin, 1991; Wang & Caballero,

Anoa taxonomy, distribution and conservation 39

1999). However, assessing the scale of the threat posed to the anoas by shifting agriculture

and ﬁres is hampered by the lack of knowledge of their habitat requirements. Furthermore,

the effects of disturbance resulting from human presence in the forest (e.g. collecting forest

products or traversing anoa areas) on anoa populations are unknown.

In addition to hunting and the loss of suitable habitat, diseases present a potential threat

to the species. The increased presence of domestic or feral cattle, as well as introduced deer

populations, heightens this risk.

Captive breeding

A total of 125 (58, 67) anoas were reported to be in captivity on 31 December 1998. These

animals were held in 35 institutions: 17 in Europe, 10 in North America and 8 in Asia

(Schreiber et al., 1993; Schreiber & Nötzold, 1995; Nötzold, 1999). Of these 125 animals,

seven were listed as founders. These seven animals were imported from zoos in Indonesia or

Malaysia, or obtained from animal dealers. Their places of origin within Sulawesi are

unknown.

Studbook breeding recommendations have, of necessity, been rather general because of the

uncertainty surrounding the taxonomic afﬁliations of the captive animals. All 125 animals

were listed as B. (A.) depressicornis (Nötzold, 1999) although there has been discussion about

their taxonomic status. Biometric and photographic assessment of the founders of the Antw-

erp, Rotterdam and Leipzig lines (lowland anoa) has indicated their phenotypic similarity.

These populations have subsequently been hybridized. The anoas at Krefeld, Berlin and

Decin Zoos, which are the descendants of three animals originally imported by Berlin Zoo,

have been listed as mountain anoa. The small number and close relationship of these animals

raise questions about the long-term value of this population for the captive breeding pro-

gramme.

Suggestions for an anoa conservation programme

Overall objectives

It is clear from the information presented in this paper that our knowledge of the systematics,

distribution, abundance and general biology of the anoas is still rather scant. As effective

conservation strategies require ﬁrm biological foundations, the recommendations made below

emphasize the need to gather biological information, while at the same time attempting to

conserve as much of the remaining anoa diversity as possible. The top priorities are: (i) to

protect anoas from hunting and prevent habitat loss and degradation in all key sites;

(ii) to complete the genetic studies currently underway in order to determine how many anoa

taxa exist; and (iii) determine the status of the remaining anoa populations (i.e. determine

population size or relative abundance, and monitor population trends) in Sulawesi to allow

the design and implementation of an effective anoa management strategy.

The largest protected area in Sulawesi is 2871 km2. In most protected areas anoa numbers

will be below carrying capacity because of hunting pressure. So, if we use the estimated

density of anoas reported by Mustari above of 1.1 and 1.3/km2, we see that the largest

remaining protected anoa populations are likely to be smaller than 3000 animals, probably

half this population size, with a far smaller number of breeding individuals. A large propor-

tion or the total number of anoa will probably exist in small non-viable populations, as has

been reported recently using similar surveys of the Gaur (Choudhury, 2002). Therefore, for

the long-term conservation of these species active management of the genetic resource may

be required. This could take the form of the movement of individuals between populations

to increase gene ﬂow.

40 J. A. Burton, S. Hedges and A. H. Mustari

Protect anoas from hunting

Anoas need to be protected from hunting by enforcing the existing laws of the Indonesia,

particularly the Conservation of Natural Resources and Ecosystems Act (Udang-Udang

Republic of Indonesia (UURI) No. 5). Law enforcement should be combined with an envi-

ronmental education campaign that stresses that the anoas are unique to Sulawesi and in

danger of being lost forever. Successful anti-poaching activities require frequent patrols of

key protected areas, which should result in the arrest and effective prosecution of poachers.

In addition, realistic ﬁnes for people caught selling live anoas, anoa meat, horns or other

parts or for keeping pet anoas need to be imposed (Lee, 1999b).

Complete genetic research aimed at determining the number of anoa taxa

Until we know more about the genetics of these animals, we cannot ensure that our efforts

are being directed towards conserving the greatest possible proportion of the anoa gene pool.

Genetic research is currently being conducted (by J. Burton) to determine whether the two

presently accepted species of anoa, B. depressicornis and B. quarlesi, are in fact sufﬁciently

distinct to warrant speciﬁc status. The study will also allow the identiﬁcation of the distribu-

tion of both species and of Management Units that will allow effective preservation of the

greatest genetic diversity. Management Units are classiﬁed as populations with morphological

or genetic variation caused by isolation or loss of mixing between populations (Moritz, 1994).

The information collected includes photo-documentation, body measurements and DNA

samples, as well as such information as is available about the animals’ likely places of origin

on Sulawesi. Such information will also help overcome some of the difﬁculties (the small

number of founders, interlineage hybridization and incest breeding) that have hampered

conservation breeding of the captive population in Europe (Schreiber & Nötzold, 1995;

Nötzold & Schreiber, 1996). Results of the genetic work will be a key component in the design

of an in situ management plan for the anoas.

Conduct an island-wide status survey

While the work described earlier in this paper has done much to bring up to date our

knowledge of the distribution of anoas on Sulawesi and its offshore islands, we still know

rather little about likely population sizes and population trends. For the majority of sites we

merely know that anoas are present or are absent/likely to be absent. This makes it difﬁcult

to focus conservation efforts effectively. There is therefore still a need for an island-wide

survey to identify major anoa populations. In order to devise the most representative con-

servation strategy, the surveys should aim to locate populations throughout the island in

order to preserve the full biogeographical variation exhibited by these species.

The proposed island-wide status survey has ﬁve components: (i) conﬁrm the presence and

estimate density of anoas in key areas to identify where sufﬁcient numbers are present for

likely population viability in the long term. These should include Lore Lindu NP, Bogani

Nani-Wartabone NP and environs, the Upper Paguyaman/Nantu area, Morowali NR, Tan-

jung Peropa WR, Lambu Sango WR or Buton Utara NR (at a later date, the following areas

should also be assessed: G. Lumut/G. Hohoban, Pengunungan Tangkeleboke, Pengunungan

Abuki, Dako Proposed NR and Gn. Tinombala). It would also be desirable to survey all

those areas where anoa have been reported since 1990 (Fig. 2); (ii) determine whether anoas

are present in blocks of suitable habitat for which only pre-1990 data exist (Pulau Dolongan,

Masupu Proposed Game Reserve, Sumarorang Katena Rompi Protection Forest, Lampoko

Mampie GR, Bulu Saraung NR, Lasolo-Sampara Proposed NR and Padang Mata Osu WR),

or no reports exist [Gn. Kambuna area (Masupu Proposed WR to Lore Lindu NP), Pungu-

Anoa taxonomy, distribution and conservation 41

nungan Verbeek area (the border between South-East, South and Central Sulawesi)]; (iii)

assess the feasibility of methods for increasing anoa gene ﬂow between protected areas; (iv)

identify major threats to the anoa populations in each site; and (v) collect samples (dry skin,

horns, skulls and, wherever possible, DNA samples) for genetic and morphological analysis.

However, DNA samples should only be collected from pets (or other captive animals).

Development of protected areas

On paper Sulawesi boasts an impressive system of protected areas, but unfortunately many

of these areas are poorly managed, understaffed, and lack the basic resources required for

effective conservation. Once the island-wide survey data have been collected, the need for,

and feasibility of creating additional protected areas should be determined. The priority areas

for conservation management should also be assessed; however, in general the largest areas

of habitat are likely to be the top priorities, because these areas are likely to contain the

populations most resilient to demographic and genetic problems and the effects of hunting.

Management planning for conserving the anoa will have to incorporate logistical consider-

ations such as local infrastructure, Forestry Department support and interest, regional devel-

opment plans and local community support (Anonymous, 1991; Collins et al., 1991;

McCarthy, 1991).

Management of domestic and feral animals to reduce threats to anoas and other wildlife

Domestic water buffalo or other livestock are known to enter into protected areas where

anoas occur (UNDP/FAO, 1982; Zwahlen, 1992). This should be prevented to avoid the risk

that diseases and parasites will be transmitted to the anoas. It is appreciated that genuine

conﬂicts of interest over grazing rights may exist between protected areas and the people

living around them. In the event of an outbreak of a serious disease such as rinderpest or

foot and mouth disease, domestic livestock in the areas surrounding nature reserves and

national parks identiﬁed as containing major anoa populations should be vaccinated.

Ecological and behavioural research to facilitate effective conservation

A detailed ﬁeld study should be regarded as a high priority. We know so little about the

ecology and behaviour of these animals that plans drawn up for their conservation inevitably

have to rely on numerous assumptions. The ﬁeld study should concentrate on those aspects

of anoa biology most pertinent to their conservation. For example, the habitat requirements

of the anoas are still little known. We know next to nothing about what constitutes high-

quality habitat for anoas, or about seasonal variations in habitat utilization. Similarly, almost

nothing is known about the effects of varying degrees of habitat modiﬁcation (e.g. selective

logging) or disturbance (cattle grazing, collection of forest products, etc.) on anoa popula-

tions. Additionally, following suggestions that anoas avoid areas frequented by deer and pigs,

their distribution and ranging behaviour with respect to these species should be studied. A

radio-tracking study would provide much valuable data. Furthermore, research should aim

to determine whether opportunities for improving the quality of available habitat exist and

whether such manipulations would be desirable.

In order to help identify the limiting factors affecting anoa populations, it would be very

useful to gather information about anoa population dynamics (especially recruitment and

mortality) in those areas where hunting intensity, degree of habitat modiﬁcation and other

disturbances are being assessed. The impact of hunting may be assessed by studying how

anoa density and behaviour varies with distance from villages, and with respect to the income

structure and occupation proﬁle in those villages.

42 J. A. Burton, S. Hedges and A. H. Mustari

Sociological/anthropological research

It is recommended that a study of the uses local people make of the areas in and around the

most important sites should be made. Conducting Rapid Rural Appraisals around these sites

would be a useful way of identifying major conﬂicts of interest between protected areas and

the surrounding communities.

Education and training

For effective management of wildlife, it is necessary for protected area staff to be adequately

trained in surveying and monitoring techniques; as already discussed above, the lack of

sufﬁcient numbers of well-trained staff has been identiﬁed as a major limiting factor for the

successful implementation of conservation programmes in Indonesia. It is necessary to

explain to people living around protected areas why their activities need to be controlled and

why wildlife, which they might otherwise utilize, should be conserved. The fact that anoas,

along with many other species, do not occur outside Sulawesi should be stressed because

many people on the island are unaware of their heritage (S. Hedges, personal observations).

Recommendations relating to captive breeding of anoas

Until genetic or morphological studies have provided a clearer idea of the number of anoa

species and their distribution, there should be no further movement of animals from in situ

to ex situ locations or vice versa.

ACKNOWLEDGEMENTS

The authors would like to thank: The Indonesian Ministry of Forestry, Jakarta (PHKA),

Sulawesi’s Provincial Forestry Departments (BKSDA); The Indonesian Institute of Science

(LIPI); and the Indonesian sponsor, Ir. Harayanto MS, Bogor Agricultural University (IPB)

for permission and support to conduct this work. For their collaboration in Indonesia thanks

goes to numerous people: Moh. Yasin; Andi Faisal Alwi; Yusran Zainudin and Jabar Lahadji;

Kaharudin Kasim; Vitri and Piton; Yani Mile and Ridwan; Imran, Ading, A’o; La Ode Naﬁu;

Hally Day; Margaretha Pangau; Arfan Polontalo and Jemi Monoarfa; Duncan Neville and

Edward Pollard; Steve and Puji Oliver. This work was supported by the Stichting Dierentuin

Helpen (Consortium of Dutch Zoos), the University of Edinburgh Development Trust, Royal

Geographical Society, London, Royal Zoological Society of Scotland and The Nature Con-

servancy. A.H. Mustari was additionally supported by AusAID. For their support in the UK

and Indonesia as well as their helpful comments on the manuscript, the authors would like

to thank Dr Alastair A. Macdonald, Dr Roy Wiles and Jon Riley. For assistance in producing

the maps we thank Ed Colijn and Colin Warwick. Thanks are also due to Dr R. McDonald

and an anonymous referee for helpful comments on the manuscript.

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Submitted 14 April 2003; returned for revision 6 June 2003; revision accepted 16 December 2003

Editor: RM

Anoa taxonomy, distribution and conservation 47

APPENDIX 1

Summary of anoa distribution, status and threats in major forested areas of Sulawesi

Map

code Province & area name

Status

(area – km2)

Anoa status

(data type)

Comments (reference codes in

[] brackets, see below for details) Threats

1Gunung Klabat P. NR (53) Unconf (LR) Report by local people during questionnaire [20] Hunting, encroachment [1]

2Tangkoko Batuangus –

Dua Saudara

NR (75) Ex? Present in 1994, limited range [11].

Comparisons of 1993/4 [14] and 1979

[MacKinnon & MacKinnon, cited in 2],

showed decline of 90%; repeat Wildlife

Conservation Society surveys showed no signs

in 1999 & 2001 [21, 24]

Agricultural encroachment and heavy

hunting pressure [7, 11, 12, 14,]

3Gunung Manembo-nembo WR (65) Unconf (LR) Listed 1982 [1] and twice reported in 1994 [11].

Report by conservation staff [20]

Encroachment and hunting [1].

Hunting threat more recently [12]

4Gunung Ambang NR (250) Unconf (LR) Anoa killed in 1992. Report by forestry staff [20] Hunting and habitat loss [12]

5Bogani Nani-Wartabone NP (2871) B.d (Skull) Anoa photographed in 1996 [27] skulls seen

of lowland species. Field sightings in 1995 [20]

Illegal gold-miners hunting [7]

6Panua (in Marisa Complex) NR (940) Conf (LR) Rangers saw anoa in 1995, reports in 1990s [20].

Advantage being within Marisa Complex (940 km2)

Hunting [1, 12]

7Upper Paguyaman/

Nantu area

None (315) B.d (Sighting) Present in early 1990s [11]. Two lowland

anoa seen in 7-day survey in 1991 [6].

Reported sightings in 1990s [20]. Skulls of

Lowland form seen & 2 reports observed 5

and 9 anoa since 1992 [27]

Reports of animals snared in 1993, 1994

and 2002 (inc. B. depressicornis skulls)

[27].

Other threats include habitat loss, road

building) [6]

8Buol/Toli-Toli P. NR (225) Unconf (LR) Reported by local people in 1992 [12] Hunting & habitat loss [12]

9Pinjam/Tanjung Matop WR (16) Unconf (LR) Footprints reported (1995) by forest rangers

[12]. Questionnaires [26]

48 J. A. Burton, S. Hedges and A. H. Mustari

10 Poboya NR (1000) Unconf (LR) Questionnaires [26]

11 Lore Lindu NP (2310) B.d, B.q (Skull) Mountain anoa karyotyped (early 1990s) (unusual

sheep-like head) [11, 15]; horns seen in 1994 [29].

Skulls of mountain and lowland anoa seen [23, 24]

Planned hydroelectric development in

Lake Lindu area [7].

Hunting at boundary [29].

Expanding enclaves [23]

12 Tanjung Api NR (42) Unconf Reported in 1980s [1], Questionnaires [26]

13 Pati-Pati P. WR Unconf (LR) Questionnaires [26]

14 Lombuyan/Pagimanan WR (36) Conf (Skull) Locals reported anoa in 1994, skull observed [17] Hunting [12]

15 Morowali NR (2250) Unconf (LR) Reported in 1980, 1995, 1996 and 2001 [7,

citing the 1980 management plan, 19, 26]

Wana people use spears and dogs [19].

Hunting at unsustainable rate [22]

16 Gunung Dako P. NR Unconf (LR) Questionnaires [26]

17 Gunung Tinombala None B.d (Skull) Questionnaires [26]

18 Gunung Sojol NR (500) Unconf (LR) In 1982 contained anoa [1], Questionnaires [26]

19 Pangi Binangga P. NR (60) Unconf (LR) Questionnaires [26]

20 Bakiriang P. WR (10) Unconf (LR) Questionnaires [26]

21 Bada Valley None Unconf (LR) Present in 1989 [29, 10], Questionnaire [26]

22 Pantai Timur None Unconf (LR) Questionnaires [26]

23 Pantai Barat None Unconf (LR) Questionnaires [26]

24 Pengunungan Takolekaju None B.d (LR, Skull) Skulls of lowland anoa seen in 1990 [18].

Questionnaires 2002 [23]

25 Mayoa area None B.q (Skull) Mountain anoas killed by hunters in 1994 [11] Hunting reportedly intense [11].

26 Gunung Pompangeo None B.q (Sighting) 3 specimens [15] karyotyped as mountain anoa Hunting is the most serious threat [12]

Map

code Province & area name

Status

(area – km2)

Anoa status

(data type)

Comments (reference codes in

[] brackets, see below for details) Threats

APPENDIX 1. Continued

Anoa taxonomy, distribution and conservation 49

27 Gunung Lumut, Gunung

Tempu, Gunung Hohoban

None B.d (Skull) Remains observed by Burton [23]

28 Togians Islands (MR) X (LR) Questionnaires [26]

29 Peleng/Banggai Islands None X (LR) Reports from local forest rangers [26]

30 Dolongan Island WR ? (LR) Reported refuge for anoa, no recent data Human disturbance [1]

31 Pengunungan

Faruhumpenai

NR (900) B.q (Skull) Mountain anoa may occur (1994), skull seen [11, 29] Hunting, clearing lowland forest [1, 12]

32 Masupu WR (25) ? (LR) Reported 1982 [1], repeated visits saw no signs [26] Hunting [1]

33 S. Katena/Rompi PF (100) ? (LR) Reported 1982 [1], repeated visits saw no signs [26]

34 Lampoko Mampie WR (20) ? (LR) Reported in early 1980s [1]; in late 1980s anoa not

mentioned [5]; but may still be present [26]

35 Pengunungan Latimojong P. NR (580) Conf (LR) Reported present (1987) [29]. 2 skulls reported [10] Hunting is the most serious threat [12]

36 Bulu Saraung NR (57) ? (LR) Possibly not present now [26] Deforestation, shifting cultivation [1]

37 Makale area None B.q (Sighting) In 1994 a mountain anoa was examined [11] Hunting is the most serious threat [12]

38 Kambuno Katena PF Conf (LR) Anoa reported [16]

39 Sumarorang P. WR Unconf (LR) Present in montane forest in early 1980s [1],

widespread but uncommon, still hunted [26]

Shifting agriculture and hunting [1]

40 Lasolo-Sampara P. NR (450) ? (LR) Reported in forests in Lawe Solo river valley [1] Hunting and logging [1]

41 Rawa Aopa-Watumohai NP (968) B.d (Skull, LR) Lowland anoa were reported in Rawa Aopa area, not

common [4, 7, 9]. Lowland anoa skulls seen [23]

Hunting pressure high, unsuitable

savanna, encroachment [8, 12, 23]

42 Polewali P. WR (80) Conf (Prints) Mustari observed ﬁeld signs in 2000 [27] Hunting, land clearance [1, 12]

43 Tanjung Peropa NR (389) B.d (Sightings, Skull) Skulls collected and ﬁeld signs in 2002, direct

observation by Mustari in 1996 & 2002 [27]

Hunting 1996, 2002 [12, 27]

44 Tanjung Amolengu WR (8.5) B.d (Sightings, Skull) Estimated 8–12 lowland anoa occurred in this very

small reserve in 1995 [13]

Hunting (snares) 10 anoa killed

in 1980s & 1995; small and isolated [13]

45 Tanjung Batikolo WR (55) B.d (Skull) Mustari saw skulls footprints [27] in 1994, 2000 Land clearance, hunting [27] in 2000

Map

code Province & area name

Status

(area – km2)

Anoa status

(data type)

Comments (reference codes in

[] brackets, see below for details) Threats

50 J. A. Burton, S. Hedges and A. H. Mustari

46 Kolaka Utara NR Conf (Sighting) Anoa seen by Mustari in 1994 [13] Habitat loss and hunting [12]

47 Toronipa NR (20?) Conf (Prints) Mustari [13] saw footprints in 1994 Hunting, habitat loss [12]

48 Lamedia NR (5) Unconf (LR) Questionnaires [26]

49 Buton Utara WR (820) B.q; B.d

(Sighting, Skull)

Mountain anoa seen in 1995 by Mustari [13].

Skulls observed and identiﬁed as lowland anoa [23]

Hunting [12]

50 Lambu Sango WR (250) B.d (Skull) Skulls of lowland anoa, footprints, [23] Hunting [1]

51 Kakinawe NR (50) Unconf (LR) Questionnaires [26]

52 Padang Mata Osu WR ? (LR) Questionnaires [26]

53 Pengunungan Tangkeleboke None Unconf (LR) Questionnaires [26]

54 Pengunungan Abuki None Unconf (LR) Questionnaires [26]

55 Pengunungan Matarombea None Unconf (LR) Questionnaires [26]

56 Kabaena Island None X (LR) Questionnaires [26]

57 Muna Island None X (LR) Questionnaires [26]

58 Wowoni Island None X (LR) Mustari questionnaire to local people [27]

Map

code Province & area name

Status

(area – km2)

Anoa status

(data type)

Comments (reference codes in

[] brackets, see below for details) Threats

Abbreviations: 1. Protected areas: NP, National Park; NR, Nature Reserve; WR, Wildlife Reserve; PF, Protection Forest; P., Proposed status. 2. Anoa status: B.d, Bubalus

depressicornis; B.q, Bubalus quarlesi; Conf, Conﬁrmed presence; Unconf, Unconﬁrmed presence; Ex?, Locally extinct; ?, Status uncertain, near local extinction; X, Never

reported. 3. Data recorded: LR, Local secondary report; Sighting, Direct sighting; Skull, Skeletal remains; Prints, Indirect ﬁeld data (prints, dung).

References (from numbers in [] brackets above): 1– UNDP/FAO (1982); 2 – Thornback (1983); 3 – Anonymous (1984); 4 – Petocz (1989); 5 – Scott (1989); 6 – Clayton et al.

(1991); 7– McCarthy (1991); 8 – Zwahlen (1992); 9 – Gunawan (1995); 10 – Melisch (1995); 11 – Schreiber & Nötzold (1995); 12 – Manansang et al. (1996); 13 – Mustari

(1996); 14 – O’Brien & Kinnaird (1996); 15 – Sugiri & Hidayat (1996); 16 – Tikupadang, Gunawan & Sila (1996); 17 – P. Vercammen, personal communication to Hedges

(1996); 18 – N. Sugiri, personal communication to Hedges (1996); 19 – T. O’Brien, personal communication to Hedges (1996); 20 – Pangau (1997); 21 – Lee (1999a); 22 –

Avalard (2000); 23 – J. Burton, personal observation; 24 – J. Burton, personal observation; 25 – J. Burton, personal observation; 26 – Questionnaire data from collaborating

non-governmental organization (NGO) for J. Burton, 2000–2002; 27 – J. Mustari, personal observations; 28 – S. Hedges, personal observation; 29 – Tikupadang, Gunawan &

Sila (1994); Whitten et al. (1987).

APPENDIX 1. Continued

An Extensive Coverage Anoa Distribution Modelling in Sulawesi Using Maximum Entropy

Article

Full-text available

Apr 2023

As an endangered species, Anoa (Bubalus depressicornis and Mountain Bubalus quarlesi) inhabiting the Sulawesi island requires proper conservation both in and out of their native habitat. The study of anoa habitat is mainly conducted through field studies based on firsthand observations of anoa appearance, footprints, or excrement or through social surveys from residents who saw it directly. The studies are carried out specifically in a particular area with a relatively narrow. However, in practice, this method has limitations, such as the research location determined based on the possibility of anoa, limited research area, and inefficient use of resources. Therefore, this study aimed to model the potential habitat of anoa in the whole of Sulawesi island. This study was based on physical and environmental independent variables such as DEM, surface slope, LST, NDVI, and access to inland water, as well as in-situ species distribution retrieved from scientific papers and reports. This study discovered the likely anoa distribution on Sulawesi island, both inside and outside of its native habitat. LST is the most important independent variable in determining habitat suitability, accounting for 80% of the total, followed by water (15.3%), NDVI (2.9%), DEM (1.6%), and slope (0.3%).

Pigs as Pets: Early Human Relations with the Sulawesi Warty Pig (Sus celebensis)

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Dec 2022

Adam Brumm

The Sulawesi warty pig (S. celebensis) is a wild and still-extant suid that is endemic to the Indonesian island of Sulawesi. It has long been theorised that S. celebensis was domesticated and/or deliberately introduced to other islands in Indonesia prior to the advent of the Neolithic farming transition in the region. Thus far, however, there has been no empirical support for this idea, nor have scientists critiqued the argument that S. celebensis was a pre-Neolithic domesticate in detail. Here, it is proposed that early foragers could have formed a relationship with S. celebensis that was similar in essence to the close association between Late Pleistocene foragers in Eurasia and the wild wolf ancestors of domestic dogs. That is, a longstanding practice of hunter-gatherers intensively socialising wild-caught S. celebensis piglets for adoption into human society as companion animals (‘pets’) may have altered the predator–prey dynamic, brought aspects of wild pig behaviour and reproduction under indirect human selection and control, and caused changes that differentiated human-associated pigs from their solely wild-living counterparts.

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Article

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Jan 2024

Haakon S. Krohn

En este artículo se analizan las palabras españolas registradas en el Diccionario de la lengua española (DLE) que originalmente provienen del malayo o que posiblemente tengan ese origen. Para cada vocablo, se explica su etimología, su llegada al español, la importancia cultural de su referente y sus usos actuales tanto en el español como en los idiomas malayos estándares (el indonesio y el malasio). Además, se identifican palabras etiquetadas en el DLE como procedentes del malayo, pero cuyo origen es una lengua distinta.

New evidence regarding the taxonomic status of the wild buffalo (Bubalus sp.) in Sri Lanka

Article

Full-text available

Aug 2023

Several different views have been expressed regarding the origin and taxonomic status of the wild buffalo (Bubalus sp.) in Sri Lanka. Our study aimed at resolving this issue by using craniometric analysis supported by genetic characterization. The study used buffalo specimens from Sri Lanka stored in museums and private collections, and those recently collected by us from two wildlife protected areas. The craniometric analysis was based on cranial and horn measurements of wild buffaloes (n = 26). Domestic water buffalo specimens (n = 5) were also included for comparison. The molecular analysis was conducted using fragments from 11 of the wild buffalo specimens. The results of the craniometric analysis and genetic analysis using the Cyt b gene showed good congruence, and revealed for the first time, that wild buffaloes in Sri Lanka comprised three species, namely Bubalus arnee (the Asian wild buffalo), B. bubalis (the domestic water buffalo), and B. carabanensis (the swamp buffalo). Thus, the study confirms the presence of B. arnee in Sri Lanka, which was previously uncertain, and indicates the presence of B. carabanensis which was not previously recorded in the country. Importantly, B. arnee in Sri Lanka, referred to as B. a. migona, differs both craniometrically and genetically from the three already known subspecies from the Asian mainland. The recently collected specimens from two protected areas were not of B. arnee. Hence, a wider coverage is needed to identify locations that may now harbour populations of B. a. migona, which is endemic to Sri Lanka. This would serve to strengthen its protection within the country.

Urine biochemical profile analysis and transabdominal ultrasonography for pregnancy detection in the endangered lowland anoa (Bubalus depressicornis)

Article

Full-text available

Dec 2022

Bowolaksono A, Mayasari A, Sundari AM, Simamora ATJ, Suryaningsih R, Suryawan A, Simamora ATJ, Abinawanto, Sjahfirdi L, Hasna A, Supriatna J, Kyes RC. 2022. Urine biochemical profile analysis and transabdominal ultrasonography for pregnancy detection in the endangered lowland anoa (Bubalus depressicornis). Biodiversitas 23: 6056-6061. The population of lowland anoas (Bubalus depressicornis) has decreased significantly due to illegal hunting and habitat loss. As such, the urgency in developing successful ex-situ conservation programs cannot be overstated. Ex-situ breeding programs of anoas, however, are hindered by difficulties in early pregnancy detection. The failure to detect pregnancy at an early stage often results in miscarriage due to repeated breeding attempts of unconfirmed pregnant females. Moreover, frequent false heat during 5 to 6 months of pregnancy corroborated the repeated breeding of pregnant females. Therefore, early detection techniques are needed to improve breeding success. The present study evaluated the use of urine biochemical profiles as a minimally invasive approach for early pregnancy detection and transabdominal ultrasonography as a confirmation tool of false heat in four lowland anoas. Statistical analysis showed that glucose and protein level were significantly elevated during pregnancy since the 0 days of post-mating (p<0.05) and consistently higher compared to the non-pregnant period. Meanwhile, transabdominal ultrasonography successfully identified organ structures and development by the 155th, 176th, 180th, and 191st days of pregnancy. The present study suggests that urine biochemical data provide a promising, minimally invasive approach to early pregnancy detection in anoas. Incorporating urine data and ultrasonography can significantly help to maintain pregnancy by validating false heat.

Spatial Analysis of Mountain and Lowland Anoa Habitat Potential Using the Maximum Entropy and Random Forest Algorithm

Article

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The Anoa is a wild animal endemic to Sulawesi that looks like a small cow. Anoa are categorized as vulnerable to extinction on the IUCN red list. There are two species of Anoa, namely Lowland Anoa (Bubalus depressicornis) and Mountain Anoa (Bubalus quarlesi). In this study, a comparison of potential habitat models for Anoa species was conducted using Machine Learning algorithms with the Maximum Entropy (MaxEnt) and Random Forest (RF) methods. This modeling uses eight environmental variables. Where based on the results of Bubalus quarlesi potential habitat modeling, the RF 75:25 model is the best algorithm with the highest variable contribution, namely humidity of 82.444% and a potential area of 5% of Sulawesi Island, with an Area Under Curve (AUC) of 0.987. Meanwhile, the best Bubalus depressicornis habitat potential model is the RF 70:30 algorithm, with the highest variable contribution, namely population of 88.891% and potential area of 36% of Sulawesi Island, with AUC 0.967. This indicates that Anoa extinction is very sensitive to the presence of humidity and human population levels.

Bibliography

Chapter

Feb 2023

Matthew Richardson

Habitat loss and degradation are currently the main anthropogenic causes of species extinctions. The root cause is human overpopulation. This unique volume provides, for the very first time, a comprehensive overview of all threatened and recently extinct mammals, birds, reptiles, amphibians, and fishes within the context of their locations and habitats. The approach takes a systematic examination of each biogeographic realm and region of the world, both terrestrial and marine, but with a particular emphasis on geographic features such as mountains, islands, and coral reefs. It reveals patterns useful in biodiversity conservation, helps to put it all into perspective, and ultimately serves as both a baseline from which to compare subsequent developments as well as a standardization of the way threatened species are studied.

The Effect of Human Disturbance on the Spatial Distribution of Anoa (Bubalus Sp.) Using MaxEnt Modeling

Article

Full-text available

Jan 2023

Anoa (Bubalus sp.) is one of the endemic species of Sulawesi Island, which is protected because it is threatened with extinction. This study aims to determine the possible distribution of anoa habitat based on environmental parameters by considering the influence of human disturbance. Human disturbance to Anoa is in the form of settlement and transportation infrastructure. The purpose of this paper is to observe whether human disturbance has an effect or not on Anoa spatial distribution. Using 98 encounter species data points, the model discovered that settlement and transportation infrastructure contributed only 4.3% and 1.4%, respectively. Meanwhile, elevation, temperature, and water have a dominant contribution that exceeds 90%. This data indicated that the anoa habitat is less sensitive to human disturbance.

Coalescent-based analysis of demography:: applications to biogeography on Sulawesi

Chapter

Jul 2012

Ben J. Evans

The flora and fauna of Southeast Asia are exceptionally diverse. The region includes several terrestrial biodiversity hotspots and is the principal global hotspot for marine diversity, but it also faces the most intense challenges of the current global biodiversity crisis. Providing reviews, syntheses and results of the latest research into Southeast Asian earth and organismal history, this book investigates the history, present and future of the fauna and flora of this bio- and geodiverse region. Leading authorities in the field explore key topics including palaeogeography, palaeoclimatology, biogeography, population genetics and conservation biology, illustrating research approaches and themes with spatially, taxonomically and methodologically focused case studies. The volume also presents methodological advances in population genetics and historical biogeography. Exploring the fascinating environmental and biotic histories of Southeast Asia, this is an ideal resource for graduate students and researchers as well as environmental NGOs.

Wallace, Darwin and Southeast Asia:: the real field site of evolution

Chapter

Jul 2012

John van Wyhe

POPULATION OF LOWLAND ANOA (Bubalus depressicornis Smith) IN TANJUNG AMOLENGU WILDLIFE RESERVE, SOUTH-EAST SULAWESI

Article

Full-text available

Apr 1996

Abdul Haris Mustari

Pengamatan populasi anoa dataran rendah dilakukan di Suakamargasatwa Tanjung Amolengu dari tanggal 20 Agustus 1994 sampai dengan tanggal 8 Maret 1995. Luas suaka margasatwa tersebut 500 ha terdiri dari hutan primer, hutan sekunder, hutan peralihan (transitional forest), dan hutan mangrove. Delapan ekor anoa dicatat berdasarkan metode konsentrasi, 11 dan 12 ekor anoa dicatat berdasarkan metode jalur dan 10 ekor anoa dicatat berdasarkan metode jejak kaki (footprint). Sex ratio 1: 1 perbandingan anak: setengah dewasa: dewasa adalah 2: 1: 5. Kepadatan populasi 1, 6 ekor anoa/km2.

The haemoglobins of wild cattle (Bovini Simpson, 1945)

Article

Jun 1990

Arnd Schreiber

The haemoglobins of nine bovines (anoa, Cape buffalo, European and American bisons, banteng, gaur, domestic yak, dwarf zebu, and taurine domestic cattle) were investigated by isoelectric focusing (and starch gel electrophoresis). The focusing patterns obtained were characteristic for each wild cattle species, with the exception of the two forms of the genus Bison, the haemoglobin of which could not be differentiated. Despite very small sample sizes, intraspecific Variation of the red cell pigments were found in gaur, anoa and dwarf zebu. The genetic nature of this variability could be rendered likely by finding heterozygous individuals, and by densitometry which suggested dosage effects of genetic expression.

The Conservation Atlas of Tropical Forests Asia and the Pacific

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Jan 1991

The first of a series designed to cover all tropical rain forests in the world. This is a visual portfolio of detailed maps of Asia, accompanied by a text which seeks to analyze the extent and causes of deforestation and to point a way towards sustainable forest development.

Molecular Markers, Natural History and Evolution

Book

Jan 1994

John C. Avise

Preface. Part I: Background: 1. Introduction. Why Employ Molecular Genetic Markers? Why Not Employ Molecular Genetic Markers? 2. History of Molecular Phylogenetics. Debates and Diversions from Molecular Systematics. Molecular Phylogenetics. 3. Molecular Tools. Protein Assays. DNA Assays. References to Laboratory Protocols. 4. Interpretative Tools. Categorical Subdivisions of Molecular Genetic Data. Molecular Clocks. Procedures for Phylogeny Reconstruction. Gene Trees versus Species Trees. Part II: Applications: 5. Individuality and Parentage. Genetic Identity versus Non-Identity. Parentage. 6. Kinship and Intraspecific Phylogeny. Close Kinship and Family Structure. Geographic Population Structure and Gene Flow. Phylogeography. Microtemporal Phylogeny. 7. Speciation and Hybridization. The Speciation Process. Hybridization and Introgression. 8. Species Phylogenies and Macroevolution. Rationales for Phylogeny Estimation. Special Approaches to Phylogeny Estimation. Prospectus for a Global Phylogeny. Special Topics in Molecular Phylogenetics. 9. Conservation Genetics. Issues of Heterozygosity. Issues of Phylogeny. Literature Cited. Index to Taxonomic Genera. General Index.

The impact of traditional subsistence hunting and trapping on prey populations: Data from Wana horticulturalists of upland central Sulawesi, Indonesia

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