Pre-Glacial Bornean Primate Impoverishment and Wallace's Line
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Bornean primates and Wallaces line 393 Pre-glacial Bornean primate impoverishment and Wallaces line Douglas Brandon-Jones Department of Palaeontology, Natural History Museum, Cromwell Road, London SW7 5BD, UK Key words: Asia, Australasia, Borneo, climate, dispersal barriers, island hopping, Java, Mentawai archipelago, Oriental biogeography, rafting, rainforest refugia, Sumatra, Wallacea Abstract bine monkeys (Nasalis, Pygathrix, Presbytis and Semnopithecus, subgenus Trachypithecus), Leaf monkeys (Semnopithecus, subgenus Trachypithecus) macaques (Macaca), loris (Nycticebus) and and lorises (Loris and Nycticebus) are both geographically tarsiers (Tarsius)* Twelve (or thirteen if the pres- disjunct between southern India and SE Asia, with endemic representatives in eastern Indochina! These parallels appear ence of Hylobates agilis is accepted) primate to result from restriction to, and re-expansion from, rainfor- species occur on the island of Borneo* And yet, est glacial refugia in southern India, northeast Indochina despite the presence of suitable habitats, only and west Java! Sureli (Presbytis) and gibbon (Hylobates) dis- the macaques on Sulawesi and the Lesser Sunda tributions reveal further refugia in north Borneo, north Sumatra and the Mentawai Islands! Modern Sumatran pri- Islands and, to a much lesser extent, the leaf mate distribution was moulded by at least two cold dry gla- monkeys on Lombok (purportedly by human cial periods! The earlier one 190,000 years ago eliminated introduction), have crossed Wallaces line* all Sumatran primate habitats whereas, after recolonization, The effectiveness as a faunal barrier of this the later one 80,000 years ago left a north Sumatran rainfor- est refugium! Not only did the Mentawai Islands provide a most widely-adopted division of the Oriental reservoir for the recolonization of Sumatra, but indirectly for from the Australasian zoogeographic region, is an interglacial invasion of Borneo which, like Sulawesi, had generally attributed to the depth of the sea chan- previously been outside the range of Presbytis and gibbons! nel extending from the Bali-Lombok Strait, be- Bornean primate zoogeography indicates that before the tween Borneo and Sulawesi, to the east of the first arid period there may have been fewer than four pri- mate species on Borneo! Most of the present twelve or thir- Philippines* The deep Makassar Strait remained teen Bornean primate species rafted there interglacially or a sea barrier when the Sunda and Sahul shelves post-glacially from Sumatra! Pre-glacial Bornean primate im- were exposed during glacial sea-level depres- poverishment is primarily attributed to a suspected south sions* Huxleys line coincides approximately coastal dry zone which would have inhibited or precluded colonization! Colonization of islands further east must gen- with the eastern edge of the Sunda shelf, and erally have bypassed Borneo via Java or the Philippines! Lydekkers line with the western edge of the The Bornean climatic barrier presented a more severe im- Sahul shelf* These later variations on the divi- pediment to faunal exchange across Wallaces line than did sion have been regarded as clear-cut faunal the sea depth along its course! Such climatic barriers, whose boundaries enclosing a transitional zone* influences waxed and waned with the glacial cycles, would have affected most SE Asian islands and were the prime However, major recolonization of the vol- inhibitor of faunal and floral exchange between the Oriental canic Krakatau archipelago, 12 km away from and the Australasian regions! the next nearest island, has occurred in only a matter of decades (Smith, 1943; Thornton, 1996)* No convincing explanation has been of- Introduction fered as to how a sea barrier such as the Bali- Lombok Strait, little more than three times as SE Asia has a rich primate fauna, comprising wide, could have inhibited colonization for mil- orang-utans (Pongo), gibbons (Hylobates), colo- lennia* Floating islands capable of transporting a Biogeography and Geological Evolution of SE Asia, pp! 393-404 Edited by Robert Hall and Jeremy D! Holloway © 1998 Backhuys Publishers, Leiden, The Netherlands 394 D) Brandon-Jones viable sample of flora and fauna have been reli- Indochina and southern India (Brandon-Jones, ably reported* Even if these crossed a strait only 1993, 1995, 1996a, b, 1997)* Some of these once a century, their impact on floral and faunal refugia are located at one thousand metres or exchange between the two land masses would more in altitude, but most are areas which have been significant* Conditions favouring would have remained sea-bound, either as such rafting would have been enhanced during promontories or islands, or are coastal areas periods of climatic change (see the discussion which retained a maritime climate during below)* desiccative glacial sea-level depressions* The inference (Brandon-Jones, 1996a) that The dual contraction and re-expansion of the during the glaciations Asian rainforest was rainforest led to the fracturing of the distribution reduced by drought to a few scattered pockets of the pied leaf monkeys from continuity be- of distribution provides a more plausible tween Java, northeast Indochina and southern explanation for these biogeographic barriers* India, to survival in those areas alone (Fig*1; Not only could rainforest communities on such Brandon-Jones, 1995)* The prosimian loris has a islands as the Philippines and Sulawesi, have parallel distribution, with an endemic represent- been entirely eliminated, leaving only severely ative, Nycticebus pygmaeus, in eastern Indochi- degraded forest or no forest at all, but the na (Fig*2)* This suggests a similar history of dis- recession of rainforest from some coastal areas, junction and partial recolonization* In both cas- would have seriously impaired the ability of es recolonization has been northward, undoubt- rainforest to disperse by rafting* Contraction and edly from Java in the case of the leaf monkeys, expansion of rainforest distribution has been the and probably so in the case of the loris* Gibbon prime mediator of present primate species (Hylobates) distribution is similar, but without diversity on Borneo (Brandon-Jones, 1996b), in an outlying population in southern India* Recol- sharp contrast to the probable presence there, onization has been extensive but incomplete* At demonstrated in this paper, of only two primate least four primate genera (Hylobates, Presbytis, species before the penultimate glaciation* If Semnopithecus and Macaca) have added Bor- such a faunal turnover can be established for neo to their pre-glacial distribution, but others Borneo, seemingly in the heart of the Asian (Pongo, Nasalis and Pygathrix) have undergone moist rainforest, extending such analysis to little or no post-glacial dispersal* other Indo-Pacific islands should produce The Mentawai archipelago is the key to inter- further insights into the effects of climate change preting the biogeography of Sumatra and Bor- on floral and faunal migration and diversity* neo* Ancestors of its endemic primates (Hylobates klossii, Nasalis concolor, Presbytis potenziani and Macaca pagensis) must formerly Glacial effects on Asian primate distribution have existed on Sumatra, but no longer occur there* The ebony leaf monkey (Semnopithecus Both the pied leaf monkeys (Semnopithecus auratus) can only have reached Java by way of auratus, S) francoisi, S) hatinhensis, S) laotum, Sumatra, from which it is also now absent* S) delacouri and S) johnii) of Java (Indonesia), Presbytis potenziani is sister-taxon to P) comata northeast Indochina and southern India of north Sumatra, north Borneo and west Java (Brandon-Jones, 1995), and the grizzled surelis (Brandon-Jones, 1993)* An ancestral taxon simi- (Presbytis comata) of Java, north Sumatra and lar to P) potenziani is presumed to have been north Borneo (Brandon-Jones, 1993, 1996a, b) the initial coloniser of SE Asia* Hylobates klossii display a tripartite disjunction* Brandon-Jones and the gibbons of Java and (with some varia- (1996a) inferred that the once continuous sub- tion) north Borneo, are chromatically monomor- continental Indian, Chinese and SE Asian rain- phic, unlike those of the Malay peninsula and forest was fragmented by a glacial drought Sumatra* H) lar' vestitus of north Sumatra is re- 190,000 years ago* It subsequently re-expanded, placed in southern Sumatra by the polymorphic although probably not to its former extent, only H) agilis) The call of the south Bornean gibbon to contract again during a second, less severe (whose specific allocation, like that of H) lar drought 80,000 years ago* Asian colobine mon- vestitus, remains debatable) is virtually identical key zoogeography suggests that these droughts to that of H) agilis (Geissmann, 1995)* This indi- eliminated all but a few small pockets of rainfor- cates a geographic relationship between the est* Such rainforest refugia survived in north Mentawai Islands gibbon and other gibbons Sumatra, the Mentawai Islands (off west analogous to that between P) potenziani and P) Sumatra), north Borneo, west Java, northeast comata) H) klossii today is suggested to be mor- Bornean primates and Wallaces line 395 80°W 100°W 120°W Siwaliks CHINA 20°N INDIA Luzon INDOCHINA PHILIPPINES Vietnam Malay Peninsula Mindanao Sabah Singapore Sarawak Sumatra 0° BORNEO Kalimantan Sulawesi MOLUCCAS Belitung LESSER SUNDA ISLANDS Java Krakatau Flores Bali Lombok ? Timor Fig!1! The glacial refugial distribution of the pied leaf monkeys (Semnopithecus auratus,