Biogeography and Migration Routes of Large Mammal Faunas in South±East Asia During the Late Middle Pleistocene: Focus on the Fossil and Extant Faunas from Thailand

Palaeogeography, Palaeoclimatology, Palaeoecology 168 72001) 337±358 www.elsevier.nl/locate/palaeo

Biogeography and migration routes of large mammal faunas in South±East Asia during the Late Middle Pleistocene: focus on the fossil and extant faunas from Thailand

C. Tougard*

Laboratoire de PaleÂontologie, Institut des Sciences de l'Evolution, UniversiteÂ Montpellier II, Place E. Bataillon, C.C. 064, 34095 Montpellier cedex 05, France Received 10 November 1999; received in revised form 29 September 2000; accepted for publication 5 October 2000

Abstract Thailand has long held a key position in South±East Asia because of its location at the boundary of the Indochinese and Sundaic provinces, the major biogeographical regions of South±East Asia. These provinces are distinct climatically, ¯oristi- cally and faunistically. The present-day limit between them is located at the Kra Isthmus, in peninsular Thailand. Previous studies of the Javanese large mammal fossil faunas and the recent study of fossil large mammal faunas from Thailand strengthen the hypothesis of a ªcontinentalº migration route 7in contrast with the ªinsularº hypothesis via Taiwan and the Philippines) during the Late Middle/Late Pleistocene period. Thailand was even part of this migration route. During the glacial periods, the faunal exchanges were favored by the emersion of a huge continental shelf called Sundaland 7South±East Asian continental area connected to Borneo and Indonesia islands by land bridges), when the sea level was low. No geological, biogeographical or paleobiogeographical evidence supports the hypothesis of a migration route via Taiwan and the Philippines. Analysis of the extant and Late Middle Pleistocene large mammal faunas 7Carnivora, Primates, Proboscidea and Ungulata) points out the antiquity of the Indochinese and Sundaic provinces. This idea is also supported by the fact that mainland faunas already displayed modern character, whereas the Javanese faunas were mainly composed of endemic forms. However, the occurrence of the extinct species, Elephas namadicus found in a Malaysian site 7Tambun) indicates that the limit between the two provinces, if not close to the present place, was located more southward than today, possibly in peninsular Malaysia. It is also con®rmed by the presence of northern species, such as Ailuropoda melanoleuca baconi and Crocuta crocuta ultima, in Late Middle Pleistocene sites of Thailand. In fact, because of climatic cooling that occurred in the northern hemisphere during the Pleistocene, the northern faunas had to move southward. To our knowledge, the hypothesis of the antiquity of the South±East Asian provinces with a boundary south of the present-day one is mentioned here for the ®rst time. Previous biogeographical studies were based mainly on insular data from the Philippine Archipelago 7geological and mainly present-day zoogeographical data) but also from Indonesia and Borneo 7paleontological data). However, the recent discovery of Thai large mammal faunas from Late Middle Pleistocene allows to bridge the gap in the fossil record between the northern faunas of South±East Asia and the southern ones. These Thai faunas are characterized mainly by extant forms but some are today absent from the Thai territory 7Ailuropoda melanoleuca, Crocuta crocuta, Pongo pygmaeus, Sus cf. barbatus, Rhinoceros cf. unicornis, Cervus eldii). q 2001 Elsevier Science B.V. All rights reserved.

Keywords: Biogeography; Faunal exchanges; Large mammal; Pleistocene; South±East Asia

*Tel.: 133-0-4-67-14-38-90; fax: 133-0-4-67-14-36-10. E-mail address: [email protected] 7C. Tougard). 0031-0182/01/$ - see front matter q 2001 Elsevier Science B.V. All rights reserved. PII: S0031-0182700)00243-1 338 C. Tougard / Palaeogeography, Palaeoclimatology, Palaeoecology 168 32001) 337±358 1. Introduction: present biogeographical context of to the northern part of Thailand, the former province South±East Asia also includes South China, Myanmar 7ex-Burma), Vietnam, Laos and Cambodia. The Sundaic province The world is today divided into six biogeographical includes the southern part of Thailand, Malaysia, regions: Palearctic, Oriental, Ethiopian, Australian, Sumatra, Java and Borneo 7Lekagul and McNeely, Neoarctic and Neotropical 7Watts, 1984). Within 1988). The boundary between the two provinces is these regions, distinct subregions or provinces have presently located at the Kra Isthmus, in peninsular been proposed. For example, the Oriental region clus- Thailand, north to Tavoy on the Myanmar side ters the Indochinese, Sundaic, Indian and Wallacean 7148N) and the Chumpon-Prachuap border 7118N) on provinces 7Lekagul and McNeely, 1988). the Thai side 7Whitmore, 1984; Lekagul and Thailand 720±78N) straddles two distinct biogeo- McNeely, 1988). graphical areas: the Indochinese province 7northern), The Indochinese and Sundaic provinces display and the Sundaic province 7southern) 7Lekagul and important climatic differences. Indeed, the Indochi- McNeely, 1988; Gray et al., 1994) 7Fig. 1). In addition nese province presents a stronger seasonality with a

Fig. 1. Map of South±East Asia showing the boundaries of Indochinese and Sundaic provinces 7after Lekagul and McNeely, 1988). C. Tougard / Palaeogeography, Palaeoclimatology, Palaeoecology 168 32001) 337±358 339

Table 1 List of richest large mammal fossil sites taken into consideration in the present study

Country Faunasa Reference

Middle Pleistocene Philippines Luzon, Mindanao von Koenigswald, 1956; Bautista, 1991 early Late Middle Pleistoceneb South China Koloshan, Hoshangtung, Hsingan Kahlke, 1961 Myanmar Mogok Colbert, 1943 Vietnam Tham Om, Hang Hum, Keo Leng Olsen and Ciochon, 1990 Laos Tam Hang Fromaget, 1936 late Late Middle Pleistocene Thailand Thum Wiman Nakin, Thum Phra Khai Phet Tougard et al., 1996; Tougard, 1998 Cambodia Phnom Loang Beden and GueÂrin, 1973 Malaysia Tambun Hooijer, 1962; Medway, 1973 Java Ngandong Oppenoorth, 1932; Medway, 1973; van den Bergh et al., 1996; FalgueÁres et al., 1998; de Vos, pers. comm. Late Pleistocene Vietnam Lang Trang Vu et al., 1996 Sumatra Sibrambang, Lida Ajer de Vos, 1983 Java Punung Badoux, 1959; van den Bergh et al., 1996 Borneo 7Sarawak) Niah Medway, 1964, 1988; Harrison, 1996

a The faunas are grouped by period 7for details of the age, see their bibliographic references), except for the Philippines since the sites are not very well known. b The terms used to designate the different period follow Harland et al. 71989); ªearlyº and ªlateº 7without capital letter) de®ned the beginning and the end of the Late Middle Pleistocene, respectively. lower rainfall 7Whitmore, 1984; Gray et al., 1994). species, from Corbet and Hill 71992) and Nowak However, the boundary between both provinces is 71999), are given in Appendix A and those of fossil especially valid for plants 7limit between rain forest species in Appendix B 7see also Table 1). and monsoon forest) 7Whitmore, 1984). It is also true From a paleontological point of view, the Late for insects, amphibians, birds and mammals 7Whit- Middle Pleistocene biogeography of large mammal more, 1984; Lekagul and Round, 1991; Corbet and faunas is less well known than for the extant faunas. Hill, 1992; Gray et al., 1994). The Pleistocene fossil record is more documented in However, has Thailand already been a biogeogra- South China, Vietnam and Java, whereas few large phical privileged place as today? In order to answer this mammal fossils were discovered in the other countries question, we will use data from the Pleistocene geolo- of South±East Asia, except in Thailand. Indeed, until gical framework as well as data from the analysis of the the 1980s, no Pleistocene large mammal fauna was extant and fossil large mammal faunas. The present known in Thailand. So, there was an important gap study is only based on the large mammal faunas from between the northern and southern fossil faunas of South±East Asia and the Philippines, because compar- South±East Asia, and the Thai fossil faunas should isons are possible only with large mammal fossil link them. faunas. Few small mammal fossil faunas are available in South±East Asia, except in South China 7Huang, 1991; Zheng, 1993) and Thailand 7Chaimanee, 2. Geological context 1998). It is also important to note that the marine mammals are not taken into consideration. The faunal 2.1. Tectonic history composition of the Indochinese and Sundaic provinces is compared. Moreover, complete faunal lists of extant An extensive literature 7Molnar and Tapponnier, 340 C. Tougard / Palaeogeography, Palaeoclimatology, Palaeoecology 168 32001) 337±358

Fig. 2. Bathymetric map of South±East Asia showing the 50, 100 and 200 m isobaths 7from van den Bergh et al., 1996).

1975; Tapponnier et al., 1982; Hutchison, 1989; as the Red River 7Yunnan province), Sagaing 7Myan- Tapponnier et al., 1990; Dheeradilok, 1992) deals mar), Mae Ping, Three Pagodas 7Thailand) and Suma- with the geological evolution of South±East Asia in tran faults for the most important 7Hutchison, 1989; some details, so I will just be focused on some points. Polachan et al., 1991). The southern part of the Sunda- The South±East Asian region is characterized by land lithospheric continental plate 7Malaysia, Suma- two major lithospheric continental plates, that of tra, Java and Borneo) and then the northern part Sundaland 7South±East Asia landmass including 7Southern China, Myanmar, Indochina and Thailand) Sumatra, Java and Borneo) representing a south-east- were pushed away 7Tapponnier et al., 1982; Hutchi- wards extension of the Eurasian plate, and the north- son, 1989; Dheeradilok et al., 1992). The second colli- wards extension of the Australian platform, into the sion was that between South±East Asia and Australia south and eastern Indonesian region 7Hutchison, about 15 myr ago, leading to the formation of the 1989). Between Sundaland and the Australian plat- Lesser Sunda islands and parts of Sulawesi and the form, lie a set of small continental lithospheric frag- Philippine Archipelago 7Tapponnier et al., 1982; ments mixed with island arcs such as the Philippines Hutchison, 1989). During the Pleistocene, the 7Vondra et al., 1981; Hutchison, 1989). South±East Asian volcanic and tectonic events lay The geological history of South±East Asia is in the framework of the collisions of the continental mainly linked to two major plate collisions. The ®rst Eurasian and oceanic Indo-Australian tectonic plates. one was the collision between India, separated from These events had mainly effects in Indonesia located Eastern Gondwanaland 7Australia), and Eurasia about in the southern limit of the Sundaland, and in the 65±55 myr 7Courtillot et al., 1986; Beck et al., 1995). Philippine Archipelago located in the area of four This event led to the formation of several faults such major subduction zones 7Heaney, 1985; Hutchison, C. Tougard / Palaeogeography, Palaeoclimatology, Palaeoecology 168 32001) 337±358 341 1989). Especially eastern Indonesian and Philippine these latter islands is over 1000 m 7Heaney, 1985; islands are subject to uplifts. South±East Asia is Hutchison, 1989). currently an aseismic region except the Cenozoic fault zones 7Hutchison, 1989). 3. Zoogeography of extant large mammals in South±East Asia and Philippines 2.2. Glacio-eustatic events 3.1. Features of the large mammal faunas One of the most important Pleistocene geological events in South±East Asia has been a series of glacio- The extant faunas of South±East Asia include 129 eustatic sea level ¯uctuations 7Fig. 2). Before 0.8 myr, species of large mammals 7Primates, Carnivora, sea level ¯uctuations were moderate with a mean of Proboscidea, Perissodactyla and Artiodactyla) 7for 70 m and lowest sea levels at around 100 m below 18 details, see Appendix A). Among them, 53 species present-day sea level after the planktonic d O of large mammals 741%) are of Indochinese af®nity 7Prentice and Denton, 1988). At 0.8 myr, there was whereas 44 species 734%) are of Sundaic af®nity. The a dramatic change with minimum sea levels down to remaining species are of both af®nities 716 species of 170 m and an average sea level of around 90 m below large mammals; 12.5%) or of other 7Palearctic, present-day sea level. This ¯uctuation mode contin- Indian, ¼) af®nities 716 species of large mammals; ued up to recent times. In details, several less impor- 12.5%). The large mammal faunas of the Indochinese tant drops of the sea level seem to have occurred and Sundaic provinces display a greater faunal simi- during the Late Middle and Late Pleistocene. It was larity to each other than to that of the Philippine due to cooler climatic conditions around 0.28 and Archipelago 7Wallacean province). Despite their af®- 0.24 myr and then around 0.18 and 0.125 myr, accord- nities, both provinces have 48 species in common. On ing to palynological data 7van der Kaars, 1991; Zheng the other hand, the whole Philippine Archipelago and Lei, 1999). The sea levels were respectively shares ®ve large mammal species with the Indochi- between 50 and 130 m for the former coolings and nese faunas and nine with the Sundaic ones 7see around 150 m for the latter ones 7Prentice and Denton, Appendix A). Among the nine species in common, 1988). three species are only found in Palawan. This latter During the glacial periods of the Pleistocene, low island has 58% of its whole mammalian fauna in sea levels led to the emersion of a huge continental common with that of Borneo 7Heaney, 1985). shelf extending until the marine areas of 200 m depth 7Hutchison, 1989). This shelf was also considered as 3.2. Endemism rate Sundaland. The Philippine islands were out of this contour line except Palawan, since the depth of The number of large mammal endemic species water is about 145 m between Borneo and Palawan 7from islands or small continental area) is 23. 7Heaney, 1985). In a modern bathymetric setting, a Among them, 19 species are found in the Sundaic sea level lowering of 50 m would be enough to province, especially Indonesia 7Sumatra, Java) and connect Sumatra, Java and Borneo with Asian main- Borneo, and only four are present in the Indochinese land 7van den Bergh et al., 1996). However, Java province. For the larger Indonesian islands, the large would still remain quite isolated as a large elongated mammal endemism rate is around 9% in Sumatra, bay would have separated Java from Borneo to the 22% in Java and 20% in Borneo. However, if the north 7van den Bergh et al., 1996). Moreover, the whole mammalian fauna is considered, this rate is minimum sea depth between Borneo and Luzon much lower except for Borneo 724%), whereas it is 7Philippine Archipelago) via Palawan is 485 m, and around 3% in peninsular Malaysia, 6.5% on Sumatra between Borneo and Mindanao via the Sulu archipe- and 12% on Java 7Heaney, 1985). Moreover, the lago 7other Philippine islands) is 290 m 7Heaney, faunas of the Indonesia and Borneo are balanced 1985). It is worth noting also that there is a small faunas, contrary to that of the Philippine Archipelago. island arc extending north from Luzon toward An unbalanced fauna is an endemic fauna with very Taiwan. However, the minimum sea depth between few taxa 7de Vos et al., 1999). On the other hand, the 342 C. Tougard / Palaeogeography, Palaeoclimatology, Palaeoecology 168 32001) 337±358

Fig. 3. Dendrogram based on the occurrence of large mammal species found in South±East Asian countries showing the af®nities of the extant Thai large mammal fauna with those of the Indochinese province. The fauna of Thailand is here considered as a unit. The informative species 7species found at least in two faunas) are considred to elaborate a matrix of appearance/disappearance, coded in 1/0, respectively. With the program ªS. A. H. N. Clusteringº 7Sequencial Agglomerative, Hierarchical and Nested Clustering, version 1.80; Rolf, 1993) from the software NTSYS-pc, this matrix was transformed in a distance matrix. The dendrogram is obtained with the method U. P. G. M. A. 7Unweighted Pair Group Method, Arithmetic average; Sneath and Sokal, 1973). This analysis was based on the species listed in Appendix A without distinction between the Thai large mammal faunas found north and south of the Kra Isthmus, cross being replaced by the number 1. endemism rate is 85% for the whole mammalian fauna north and south of the Kra Isthmus, the dendrogram of the Philippine islands: 42% on Palawan and 7Fig. 4) shows that the northern Thai fauna is related between 55 and 87% on the other Philippine islands to the Indochinese fauna and the southern Thai fauna 7Heaney, 1985; van Oosterzee, 1997). is closer to the Sundaic one. In fact, 31% of the Thai large mammals 720 of 64 large mammal species) are 3.3. Zoogeographical af®nities of the Thai faunas of Indochinese af®nity and 20% 713 of 64 species) are of Sundaic af®nity. It is worth noting that, in both Concerning the Thai large mammal fauna, it dendrograms, Java and the whole Philippine Archi- appears that this fauna, considered as a unit, is pelago seem to be closely related. It is maybe due to more similar to the Indochinese province fauna as the UPGMA method 7distances considered of equal shown by the dendrogram 7Fig. 3) based on the length) or to high endemism rate on the Philippine occurrence of the species found in each country of Archipelago and to the impoverishment of the Java- South±East Asia. On the other hand, if the Thai nese fauna since the island isolation more than a fauna is decomposed in large mammal fauna found strong faunal af®nity.

Fig. 4. Dendrogram based on the occurrence of South±East Asian large mammal species showing the division of the Thai extant faunas into two groups: one with Indochinese similarities and the other with Sundaic similarities. This dendrogram was elaborated like that of Fig. 3. However, the Thai fauna was decomposed in large mammal faunas found north and south of the Kra Isthmus 7see Appendix A for the detailed north and south Thai large mammals). C. Tougard / Palaeogeography, Palaeoclimatology, Palaeoecology 168 32001) 337±358 343

Fig. 5. Location map of the late Late Middle and Late Pleistocene richest large mammal sites in South±East Asia 7for references, see Table 1).

4. Late Middle and Late Pleistocene fossils is worse for the Philippine fossil record. Few sites have been found 7Luzon, Mindanao) and poorly studied 7von 4.1. General context Koenigswald, 1956; Shutler and Mathisen, 1979; Bautista, 1991). For that study, sites speci®cally diversi- In South±East Asia, the fossil record is unequally ®ed and found in the Indochinese and Sundaic provinces documentedfortheMiddleandLatePleistoceneperiods. are taken into account 7Fig. 5, Table 1, Appendix B). Numerous localities are known for the Middle Pleisto- Studies of fossil faunas from South China, Laos, Myan- cene of the Indochinese province but few, related to this mar, Cambodia, Malaysia and Philippines, are old and period, have been found in the Sundaic one. As for the often limited to the genus determination, whereas recent Late Pleistocene, several fossiliferous sites are known in taxonomic reviews were done for the fossil faunas found the Sundaic province, contrary to the Indochinese one. It in Vietnam, Sumatra, Java and Borneo. 344 C. Tougard / Palaeogeography, Palaeoclimatology, Palaeoecology 168 32001) 337±358 4.2. Temporal comparison 3Late Middle Pleistocene/ due to a body mass adaptation in order to better resist Late Pleistocene) to cooler climatic conditions to avoid important body heat wastes 7Bergmann, 1847; James, 1968; Fooden If the Late Middle Pleistocene faunas are compared and Albrecht, 1993), as recorded during the late Late with those of the Late Pleistocene, a faunal turnover Middle Pleistocene 7van der Kaars, 1991; Zheng and can be observed. It is more marked on Indonesian Lei, 1999). The Late Pleistocene fauna is composed of islands, mainly on Java 7de Vos et al., 1999; Sondaar extant species, except S. orientalis and E. namadicus, et al., 1999). Indeed, the Punung fauna, also called even if some species 7P. pygmaeus, A. melanoleuca, Pongo-Homo sapiens fauna, is completely different T. indicus and Cervus unicolor) are not living today in from the archaic Ngandong fauna, also called Stego- Vietnam 7Vu et al., 1996). don-Homo erectus fauna 7Aziz et al., 1999). In the Punung fauna, genera and species are still living on 4.3. Geographical comparison 3mainland/islands) the Asian continent or in Indonesia. Eight out of 11 According to Vu et al. 71996) and de Vos et al. taxa de®ned at the species level were and are still 71999), the Lang Trang fauna is similar to the Indo- widely widespread in South±East Asia. One species nesian Late Pleistocene faunas, especially those of 7Elephas maximus) of the Punung fauna is of Indochi- Sumatra 7Lida Ajer and Sibrambang), Java 7Punung) nese af®nity. Three species, Pongo pygmaeus, Tapirus and Borneo 7Niah). However, the extinct S. orientalis indicus and Sus barbatus, were found ®rst in Middle as well as Ursus thibetanus and A. melanoleuca are Pleistocene sites of the Indochinese province. Later, absent from the Indonesian sites. On the other hand, S. when the land bridges were submerged by the sea barbatus and Bos javanicus do not occur in the Lang during interglacial periods, some specimens of these Trang fauna. For the late Late Middle Pleistocene species were probably trapped on islands, whereas faunas, the difference of faunal composition between they became extinct on the mainland. Probably all the Indochinese and Sundaic provinces is strongly new Javanese faunal elements are the result of a marked. Indeed, in the mainland, the faunas have a novel migration wave 7Aziz et al., 1999). On the recent character since all species found in the Thai and mainland, the early Late Middle Pleistocene was char- Cambodian sites are extant. The Tambun 7peninsular acterized by the Ailuropoda-Stegodon fauna, mainly Malaysia) and Ngandong 7Java) sites still have de®ned by the occurrence of Ailuropoda melanoleuca archaic forms 7Stegodon trigonocephalus, Hexaproto- baconi and Stegodon orientalis but also Ursus thibe- don sivalensis, S. macrognathus, Homo erectus, E. tanus kokeni, Arctonyx collaris rostratus, Elephas namadicus, Duboisia santeng and others; see Appen- namadicus, Megatapirus augustus and P. pygmaeus dix B for the complete faunal lists). These forms, 7Kahlke, 1961; Aigner, 1978; Huang, 1979; Li, 1981). related to the Siwalik faunas, are from an earlier However, archaic forms such as S. orientalis, E. migration route 7Early to Middle Pleistocene), the namadicus, M. augustus or Rhinoceros sinensis were so-called ªSiva-Malayanº route 7from the continent not found in the late Late Middle Pleistocene local- to the Sunda shelf; von Koenigswald, 1935). ities from Thailand and Cambodia 7Beden and GueÂrin, 1973; Tougard et al., 1996; Tougard, 1998), whereas 4.4. Zoogeographical af®nities of the fossil faunas S. orientalis, and perhaps, E. namadicus were discovered in the Late Pleistocene site of Lang Trang, Studies of the Late Middle and Late Pleistocene Vietnam 7Vu et al., 1996). The Thai and Cambodian South±East Asian faunas reveal that some large faunas are modern in their composition because they mammals which today characterize either the Indochi- do not contain extinct species. On the other hand, nese province or the Sundaic province have been some extinct subspecies occurred 7Crocuta crocuta discovered in several localities 7Table 1, Appendix B). ultima, A. collaris rostratus, Rhinoceros sondaicus In fact some living species such as Macaca assamensis, guthi and others; for more details, see Appendix B). U. thibetanus, A. melanoleuca, but also Elaphodus In general, subspecies are larger than their homolo- cephalophus were present in fossil faunas of South gous Holocene or recent species 7Hooijer, 1949; China, Vietnam, Myanmar and/or Thailand, whereas Tougard, 1998; de Vos et al., 1999). It was probably Hylobates syndactylus or S. barbatus have only been C. Tougard / Palaeogeography, Palaeoclimatology, Palaeoecology 168 32001) 337±358 345 found in Sumatra, Java and/or Borneo. These species bution is today restricted to small continental areas, occur today exclusively in the same provinces in which islands or other world region. The spotted hyena, they occurred during the Late Middle and Late Pleisto- Crocuta crocuta, is only found in Africa, south of cene. The number of species is probably not exhaustive, the Sahara 7Nowak, 1999). The habitat of the giant but all the present-day large mammal species have not panda, A. melanoleuca, is restricted to some South been found in fossiliferous deposits. Some were and are China mountains, whereas the orang-utan lives only also too widespread in South±East Asia 7for more in Sumatra and Borneo 7Nowak, 1999). Three situa- details about the geographical af®nity of species, see tions are thus presented. According to Kurten 71956), Appendices A and B). C. crocuta ultima took over from Hyaena brevirostris Except these extant species, some other extinct or sinensis ®rst in South China. Indeed, this latter species missing species are also known exclusively in the became extinct at the same time a cooling event northern part of South±East Asia or in the southern occurred. On the other hand, the cooling caused an part. It is notably the case for S. orientalis, E. nama- altitudinal drop of the mountain vegetation zones, as dicus and M. augustus in the northern part, and S. those of the bamboo species 7Zheng and Lei, 1999). In trigonocephalus and D. santeng in the southern one this way, this kind of plants could have covered some 7for more details about the province af®nity of each lowland areas perhaps in contact with each other. This fossil species, see Appendix B). It is dif®cult to could have allowed the southward migration of A. consider some species as endemic, for example melanoleuca baconi, that subsists on this plant Szechuanopithecus yangtsensis or Parameles simpli- 7Tougard et al., 1996). According to Heaney 71985), cidens, because it may be a taphonomic bias or a most of the species that are endemic to a single island wrong determination of a taxa due to the preservation are restricted to mountain habitats. However, because state or the paucity of the fossil material. of the altitudinal depression of mountain forests As for the faunas of the Philippine Archipelago during the glacial periods, most mountain mammal 7Luzon and Mindanao), they were characterized species were widespread in appropriate habitats. mainly by endemic species 7von Koenigswald, 1956; This indicates that immigration across the Sundaland Bautista, 1991; Appendix B). Only one species, S. cf. took place in the recent past. Concerning P. trigonocephalus, was found in common with the Ngan- pygmaeus, fossil remains from the Late Middle Pleis- dong fauna 7Late Middle Pleistocene, Java). However, tocene and the Late Pleistocene were found in South elephants are generally considered as good swimmers China, Vietnam, Laos, Cambodia and Thailand. On 7Heaney, 1985; van Oosterzee, 1997). This form can not the other hand, some fossil remains were also discov- strongly testify to a land connection between Java and ered in Late Pleistocene sites of Indonesia and Luzon, probably via Borneo. Borneo. The migration of Pongo from mainland to Indonesia and Borneo was probably due to the sea 4.5. Zoogeographical af®nities of the Thai fossil faunas level lowering 740±65 m below present-day sea level) before 70,000 yr, but after the glacial maximum The recent study of the Late Middle Pleistocene at about 135,000 yr 7van den Bergh et al., 1996). fauna from northeastern Thailand 7Tougard, 1998) These three examples show how the late Late Middle shows that this fauna is characterized by a more Pleistocene cooling might have had an in¯uence on important number of Indochinese large mammal the southward extension of the northern faunas. species 740% of the 27 Thai large mammal de®ned Unfortunately, it is currently not possible to know at the species level) compared to the Sundaic ones whether the extension of the Indochinese species, 715%). Moreover, this fauna is characterized notably except P. pygmaeus, was limited southward to Thai- by the occurrence of the hyena C. crocuta ultima, the land or to peninsular Malaysia, because of the nearly panda A. melanoleuca baconi and the orang-utan P. complete lack of Pleistocene fossiliferous sites in the pygmaeus. These species were widespread during the latter area. However, the discovery of the extinct Late Middle Pleistocene in the northern part of species, D. santeng and especially E. namadicus,in South±East Asia. On the other hand, they do not the Tambun site 7Hooijer, 1962; Medway, 1973) is a live in Thailand any more. Their geographical distri- valuable evidence. Indeed, E. namadicus was found 346 C. Tougard / Palaeogeography, Palaeoclimatology, Palaeoecology 168 32001) 337±358

Fig. 6. Map of South±East Asia showing the Sundaland boundary during the Late Middle Pleistocene 7adapted from Hutchison, 1989; Heaney, 1991). The migration route hypothesis 7by continental or insular ways) are illustrated by the arrows. The Early and Middle Pleistocene Siva-Malayan route 7a) and the Late Pleistocene Sino-Malayan one 7b) are considered here as part of the continental way. mainly in South China and Vietnam 7Kahlke, 1961; 5. Discussion Olsen and Ciochon, 1990). Therefore, it was a characteristic of the Indochinese province. Concerning D. 5.1. Late Middle and Late Pleistocene migration santeng, this species was also found in Lower Pleis- routes tocene localities of Java. It seems that it was characteristic of the Sundaic province. However, the 5.1.1. Historical review of previous works attribution of the fossil remains to this species has to In South±East Asia, numerous important faunal be considered carefully. The remains, especially the exchanges occurred, notably between the Late Middle teeth, could belong to Boselaphus. Pleistocene and the Late Pleistocene. This is testi®ed C. Tougard / Palaeogeography, Palaeoclimatology, Palaeoecology 168 32001) 337±358 347 by the large mammal turnover in the Punung site 5.1.3. Evidences from extant large mammals and 7Java) of Late Pleistocene age. However, the way other vertebrates used by faunas to migrate from mainland to Indone- The analysis of the present-day Indochinese and sian islands and inversely was long debated. Two Sundaic large mammal faunas compared to those of migration routes were considered: a ªcontinentalº the Philippine Archipelago 7Wallacean province) way via Myanmar, Thailand or Indochina 7Laos, Viet- 7Corbet and Hill, 1992) can provide information on nam and Cambodia) and then peninsular Malaysia, or the most plausible migration route. The Indochinese an ªinsularº way via Taiwan and the Philippine Archi- and Sundaic faunas are more similar to each other pelago 7Fig. 6). than to that of the Philippines. After the dendrograms von Koenigswald 71935, 1939) supposed that the 7Figs. 3 and 4), the large mammal fauna from all the mammals migrated during the Pleistocene to Java Philippine islands displays greater af®nities with via the Philippines on the base of elements identi®ed faunas from the Sundaic province 7than with those as of Chinese af®nities in Javanese Pleistocene faunas, of the Indochinese one; nine and ®ve large mammal the occurrence of a Pleistocene fauna in the Philip- species in common, respectively). If the Philippine pines, and the lack of fossil evidence from the Sunda- islands had been directly connected with the Asian land area. This author called this way, the ªSino- mainland and a possible passage way between north Malayanº migration route. On the other hand, de and south of South±East Asia, the Philippine fauna Terra 71943) and Colbert 71943) suggested that Java- should also include elements from the Indochinese nese faunas could have originated from Myanmar as province 7Heaney, 1985; Corbet and Hill, 1992), the Javanese faunas of Lower Pleistocene age seems which is not the case. This fact is re¯ected by both to have. These authors are followed by Shutler and large and small mammal faunas 7Rodentia, Insecti- Braches 71985), who considered the island status of vora, Scandentia, Dermoptera, Pholidota) except Luzon as re¯ected in the fossil fauna from Pleisto- bats 7Cynopterus brachyotis, Eonycteris spelaea, cene. De Vos et al. 71999) agreed with a continental Myotis formosus, Tylonycteris pachypus, Murina migration route from China, Vietnam, and Cambodia cyclotis and others; Corbet and Hill, 1992). However, to Indonesia. the Palawan mammal fauna displays 58% of similar- ity with that of Borneo 7Heaney, 1985; van Oosterzee, 1997). 5.1.2. Geological evidences There are also data pertaining to freshwater ®shes In South±East Asia, the Pleistocene geological 7Korn®eld and Carpenter, 1984). Rivers and lakes of events indicate that there have been important Philippines currently support depauperate ®sh faunas. glacio-eustatic sea level ¯uctuations, especially Cyprinid ®shes are the most widespread, and current during the Middle and Late Pleistocene. A sea level evidence indicates that they are descendant of a single lowering between 50 and 150 m could have occurred species that dispersed from Borneo to Mindanao several times 7Prentice and Denton, 1988). Land probably during the Pliocene. The cyprinids of bridges within the mainland and Indonesian islands Palawan also originated on Borneo and could have such as Sumatra, Java and Borneo could exist because arrived probably prior to the Middle Pleistocene of the emersion of a huge continental shelf called 7Korn®eld and Carpenter, 1984). The results of Sundaland 7Heaney, 1985; Lekagul and McNeely, studies about amphibians and reptiles closely parallel 1988). Palawan could have been the only Philippine those of the studies on mammals 7Brown and Alcala, island connected to Asian mainland via Borneo. 1970). Palawan supports a signi®cant number of ende- Indeed, the sea is about 145 m deep between Palawan mic species, but its faunal relationships lie strongly and Borneo, whereas it is much deeper compared to with Borneo. the other Philippine islands 7Heaney, 1985; van Oosterzee, 1997). On the other hand, there is no 5.1.4. Evidences from fossil large mammal faunas proof of land bridge connection between mainland The analysis of the fossil large mammal faunas and the Philippine Archipelago via Taiwan because shows that faunas with modern character were of the sea depth. already present in mainland in the late Late 348 C. Tougard / Palaeogeography, Palaeoclimatology, Palaeoecology 168 32001) 337±358 Middle Pleistocene, whereas the Javanese faunas connection had existed between the Asian mainland still contained archaic forms. As shown by three and the Indonesian islands in the recent past 7Heaney, forms 7P. pygmaeus, A. melanoleuca baconi and 1985; de Vos et al., 1999). On the contrary, a high C. crocuta ultima) from the Thai localities, the endemism rate and an unbalanced fauna as the extant late Late Middle Pleistocene cooling has certainly and fossil faunas testify to a signi®cant isolation as for modi®ed the geographical distribution of the the Philippines, except Palawan 7Heaney, 1985; de South±EastAsianfaunas,notablyinshifting Vos et al., 1999). northern faunas even further southwards, i.e. to From the geological point of view as well as the Indonesian islands. This hypothesis is the extant and fossil large mammal fauna studies, supported by the occurrence of P. pygmaeus, S. it clearly appears that the emersion of the Sunda- barbatus, Naemorhedus sumatraensis, E. maximus land allowed faunal exchanges from continental or Ursus malayanus in the Late Pleistocene South±East Asia to the Indonesian islands, and Punung 7Java) site. These species were ®rst probably inversely, between the late Late Middle found in Late Middle Pleistocene continental Pleistocene and the Late Pleistocene. It was the localities, and then in Late Pleistocene sites from consequence of land bridge connections between Indonesia. According to Aziz et al. 71999), a landmasses when the sea level was low, because marked faunal turnover happened in Indonesia of the continental glacier formation during glacial andmainlyinJavawiththearrivalofnewimmi- periods. grants from the Asian mainland. Indeed, these new The recent discovery of Late Middle Pleistocene immigrants were mainly of Indochinese af®nities, faunas in Thailand brings new information about the or widespread through South±East Asian main- migration routes, strengthening the continental land. It is worth noting that S. barbatus, T. indicus hypothesis. This route could be also called the Sino- and P. pygmaeus have currently only habitats in Malayan route, though it referred ®rst to a migration Indonesia and Borneo, whereas they were found in route from mainland to Java via the Philippines 7von mainland sites older than those of Indonesia Koenigswald, 1935, 1939). Whereas Sundaland 7Punung, Lida Ajer, Sibrambang) and Borneo offered more possibilities to move for the mammalian 7Niah). faunas because of its large surface, Thailand was even Moreover, the Late Middle Pleistocene faunas from part of this migration route. The fossil large mammal Vietnam 7Olsen and Ciochon, 1990; Vu et al., 1996), Thai faunas allow to link and compare those found Cambodia 7Beden and GueÂrin, 1973) and mainly from from South China to Indonesia. The study of the fossil Thailand 7Tougard et al., 1996; Tougard, 1998) share large mammal faunas from the Philippines 7Luzon numerous species with countries located at the north- and Mindanao) do not give strong information to ern and southern ends of the South±East Asian deny this idea. provinces, namely South China, Sumatra, Java, and Borneo. Unfortunately, no evidence of faunal exchanges 5.2. Late Middle Pleistocene biogeography of large from Indonesia to the South±East Asian mainland mammal faunas was found, except the possible presence of D. santeng in the Tambun site, peninsular Malaysia. South±East Asia is currently divided into two However, this species could have migrated via the distinct biogeographical provinces: the Indochinese Siva-Malayan migration route, probably by province and the Sundaic province. The boundary 0.8 myr during the dramatic sea level lowering, between these provinces is presently located at the because it was found only in Lower Pleistocene Kra Isthmus, in peninsular Thailand 7Whitmore, Javanese sites. 1984; Lekagul and McNeely, 1988). As seen earlier, faunal exchanges happened on the Sundaland between 5.1.5. Conclusion the Late Middle Pleistocene and the Late Pleistocene. A low endemism rate and a balanced fauna as the It is also con®rmed by the presence of extant species extant and fossil Indonesian faunas indicate that a characteristic of the Sundaic province in Indochinese C. Tougard / Palaeogeography, Palaeoclimatology, Palaeoecology 168 32001) 337±358 349 faunas and inversely 7Appendix A). However, the the northern part of peninsular Malaysia in the late boundary between the northern and southern faunas Late Middle Pleistocene. of South±East Asia had to moved because several climatic changes occurred during the Pleistocene 6. Conclusions succession of glacial and interglacial periods 7van der Kaars, 1991; Zheng and Lei, 1999). This event With its privileged geographical situation, Thai- is clearly illustrated by the Thai occurrence of the land is located at the crossroads of the South±East three species mentioned earlier 7P. pygmaeus, C. Asian northern 7the Indochinese province) and crocuta ultima and A. melanoleuca baconi), but also southern 7the Sundaic province) parts. These by the northward shift of the panda 7A. melanoleuca) provinces are distinct from the climatic 7seasonality distribution to some mountains of South China during and rainfall differences) as well as from the ¯oristic the Late Pleistocene and the Holocene 7Tougard et al., and faunistic points of view. Their origin seems old 1996). in regard to the large mammal faunas 7carnivores, As seen previously, the continental faunas primates, proboscideans and ungulates) from presented already a modern character, whereas the South±East Asia, notably those of the Late Middle Javanese ones were mainly characterized by archaic Pleistocene. And because a cooling occurred during forms. This fact allows to suppose that two distinct this period, the biogeographical boundary between faunas were present in South±East Asia in the late these two provinces seemed to have been located Late Middle Pleistocene. Moreover, 40% 7of 27 further south than the present one, i.e. south of the taxa de®ned at the species level) of the fossil Kra Isthmus in peninsular Thailand and probably in Thai faunas and 75% 7of eight species) of the fossil northern peninsular Malaysia. Cambodian fauna are of Indochinese af®nity, Moreover, Thailand was certainly the witness of whereas 60% of the fossil Ngandong faunas faunal exchanges between the both South±East 7Java) was of Sundaic af®nity. The Tambun fauna Asian provinces. It was part of the Sino-Malayan is not considered here because three species only migration route between the continental parts of were determined. Among them, Rhinoceros sondai- South±East Asia and the neighboring Indonesian cus was widespread in South±East Asia. E. nama- islands, notably for the human colonization. It is espe- dicus was characteristic of the northern part of cially true since a tooth attributed to Homo sp. was South±East Asia, whereas D. santeng was only found in a late Late Middle Pleistocene site of Thai- present in the southern part. So, it seems that there land 7Tougard et al., 1998). It is worth noting that this was a northern fauna biogeographically distinct from is the only human fossil remain found for this period, the southern one in South±East Asia during the Late hence its importance. Middle Pleistocene. Unfortunately, it is currently dif®cult to specify where the boundary between the northern and south- Acknowledgements ern fossil faunas could be located, because of the nearly complete lack of Middle Pleistocene sites in This work has been completed in the frame of Indonesia and peninsular Malaysia. The lack of data the Thai±French Paleontological Mission of the relative to the peninsular Malaysia Pleistocene fauna French Ministry of Foreign Affairs. Thanks are 7Tambun) and of those relative to Sumatra 7Sibram- expressed to Yaowalak Chaimanee, Varavudh bang and Lida Ajer) and Borneo 7Niah) does not allow Suteethorn and Somchai Triamwichanon, Depart- to know if the species found in Java were distributed ment of Mineral Resources, Bangkok and to Jean- throughout the Indonesian islands and the mainland or Jacques Jaeger, SteÂphane Ducrocq, Jean Pierre if they were endemic to Java. Only the presence of E. Aguilar, Bernard Marandat and Laurent Marivaux, namadicus in the Tambun site is a valuable clue. It ISEM, Montpellier. We are also grateful to J. de allows to suppose that the boundary between the Indo- Vos, C.P. Groves, P.Y. Sondaar and an anonymous chinese and Sundaic provinces, if not close to the reviewer. ISEM Publication No. 2000-089, UMR present place, was located further south, possibly in CNRS 5554. 350 C. Tougard / Palaeogeography, Palaeoclimatology, Palaeoecology 168 32001) 337±358 Appendix A

Faunal lists of the large mammal species found in South China 7SC), Myanmar 7MY), Laos 7LA), Cambodia 7CA), Vietnam 7VI), Thailand 7with a distinction between the faunas found north and south of the Kra Isthmus, NK and SK, respectively), peninsular Malaysia 7MA), Sumatra 7SU), Java 7JA) and Borneo 7BO). The species in common with the Philippines 7PH) of the Wallacean province 7WP) are also indicated, especially those found in Palawan 7P). For each species, the biogeographical af®nity is given in parentheses: Indochinese 7I), Sundaic 7S) and other 7O) af®nities 7Palearctic, Indian, ¼) or widespread 7W) species. The asterisk 7p) indicates that the species has currently a Sundaic habitat but it was exclusively found in the Indochinese province before the Late Pleistocene. Faunal lists and geographical distribution are from Corbet and Hill 71992) and Nowak 71999). Taxonomic names follow Wilson and Reeder 71993).

Indochinese province Thailand Sundaic province WP

SC MY LA CA VI NK SK MA SU JA BO PH

PRIMATES Nycticebus coucang 7I, S) ££ ££££££ £££ Nycticebus pygmaeus 7I) ££££ Tarsius bancanus 7S) ££ Macaca arctoides 7I) ££ ££££ £ Macaca assamensis 7I, O) ££ £ ££ Macaca fascicularis 7S) £££££££££££ Macaca mulatta 7I, O) ££ £ £ £ Macaca nemestrina 7I, S) ££££££££ £ Macaca thibetana 7I) £ Presbytis comata 7S) £ Presbytis femoralis 7S) £££££ Presbytis frontana 7S) £ Presbytis hosei 7S) £ Presbytis melalophos 7S) £ Presbytis potenziani 7S) £ Presbytis rubicunda 7S) £ Presbytis thomasi 7S) £ Trachypithecus auratus 7S) £ Trachypithecus cristatus 7S) ££££££££ £ Trachypithecus francoisi 7I) £££ Trachypithecus obscurus 7S) ££££ Trachypithecus phayrei 7I) ££ £ ££ Trachypithecus pileatus 7I) £ Pygathrix avunculus 7I) £ Pygathrix bieti 7I) £ Pygathrix brelichi 7I) £ Pygathrix nemaeus 7I) £££ Pygathrix roxellana 7I) £ Nasalis larvatus 7S) £ Nasalis concolor 7S) £ Hylobates agilis 7S) ££££ Hylobates concolor 7I) £££ Hylobates gabriellae 7I) ££ Hylobates hoolock 7I) ££ Hylobates klossi 7S) £ Hylobates lar 7I, S) ££ £ ££ £ Hylobates leucogenys 7I) £££ Hylobates moloch 7S) £ Hylobates muelleri 7S) £ Hylobates pileatus 7I) ££ £ Hylobates syndactylus 7S) ££ Pongo pygmaeus 7Sp) ££ C. Tougard / Palaeogeography, Palaeoclimatology, Palaeoecology 168 32001) 337±358 351

7continued)

Indochinese province Thailand Sundaic province WP

SC MY LA CA VI NK SK MA SU JA BO PH

CARNIVORA Canis aureus 7I, O) ££ £ Canis lupus 7W) £ Vulpes vulpes 7I, O) ££ Nyctereutes procyonoides 7I, O) ££ Cuon alpinus 7W) ££ ££££££ ££ Ursus thibetanus 7I) ££ £ ££ Helarctos malayanus 7I, S) ££ ££££££ £ £ Ailuropoda melanoleuca 7I) £ Ailurus fulgens 7I) ££ Mustela kathiah 7I, O) ££ £ £ Mustela lutreolina 7S) ££ Mustela nivalis 7I, O) ££ Mustela nudipes 7S) ££ £ £ Mustela sibirica 7I, O) ££ £ ££ Mustela strigidorsa 7I) ££ £ ££ Martes ¯avigula 7I, S, O) ££ ££££££ £££ Meles meles 7I, O) £££ Arctonyx collaris 7I, O) ££ £££££ £ Melogale everetti 7S) £ Melogale moschata 7I) ££ £ Melogale orientalis 7S) £ Melogale personata 7I) £££ Mydaus javanensis 7S) £££ Lutra lutra 7W) ££ ££££ £ Lutra sumatrana 7S) ££ ££ £ £ Lutrogale perspicillata 7W) ££ ££££££ £££ Amblonyx cinereus 7W) ££ ££££££ ££££7P) Viverra megaspila 7I) ££ ££££££ Viverra tangalunga 7S) ££ ££ Viverra zibetha 7I) ££ ££££££ Viverricula indica 7W) ££ ££££££ ££ Prionodon linsang 7S) £ £ ££ £££ Prionodon pardicolor 7I) ££ £ ££ Paradoxurus hermaphroditus 7W) ££ ££££££ ££££ Paguma larvata 7I, S, O) ££ ££££££ £ £ Arctictis binturong 7I, S) ££ ££££££ ££££7P) Arctogalidia trivirgata 7I, S) ££ ££££££ £££ Hemigalus derbyanus 7S) £££££ Chrotogale owstoni 7I) £££ Diplogale hosei 7S) £ Cynogale bennettii 7S) ££ £ Herpestes brachyurus 7S) ££ ££7P) Herpestes javanicus 7I, S, O) ££ ££££££ £ Herpestes semitorquatus 7S) ££ Herpestes urva 7I) ££ ££££££ Felis chaus 7W) ££ ££££ Prionailurus bengalensis 7W) ££ ££££££ ££££ Prionailurus planiceps 7S) ££ £ £ Prionailurus viverrinus 7W) ££ ££££ ££ Catopuma badia 7S) £ Catopuma temminckii 7I, S) ££ ££££££ £ 352 C. Tougard / Palaeogeography, Palaeoclimatology, Palaeoecology 168 32001) 337±358

7continued)

Indochinese province Thailand Sundaic province WP

SC MY LA CA VI NK SK MA SU JA BO PH

Pardofelis marmorata 7I, S) ££ £ ££ £ £ Neofelis nebulosa 7I, S) ££ ££££££ £ £ Panthera pardus 7W) ££ ££££££ £ Panthera tigris 7W) ££ ££££££ ££ PROBOSCIDEA Elephas maximus 7I, O) ££ ££££££ £ £ PERISSODACTYLA Tapirus indicus 7Sp) ££ ££££ Rhinoceros sondaicus 7I, S) ££ ££££££ ££ Dicerorhinus sumatrensis 7I, S) ££ ££££££ £ £ ARTIODACTYLA Sus barbatus 7Sp) ££ ££ Sus scrofa 7W) ££ ££££££ ££ Sus verrucosus 7S) £ Tragulus javanicus 7S) ££££££££££ Tragulus napu 7S) ££ £££££ £ Moschus berezovskii 7I) ££ Cervus eldii 7I) ££ Cervus nippon 7I) £ Cervus timorensis 7S, O) £ Cervus unicolor 7W) ££ ££££££ £ £ Axis kuhlii 7S) £ Axis porcinus 7I, O) ££ ££££ Muntiacus atherodes 7S) £ Muntiacus feae 7I) ££ Muntiacus gongshanensis 7I) ££ Muntiacus muntjak 7W) ££ ££££££ £££ Muntiacus reevesi 7I) ££ Muntiacus rooseveltorum 7I) £ Elaphodus cephalophus 7I) ££ Hydropotes inermis 7I) £ Bos frontalis 7I) ££ ££££££ Bos javanicus 7I, S) ££££££ ££ Bos sauveli 7I) ££££ Bubalus bubalis 7W) ££££ £ Budorcas taxicolor 7I) ££ Naemorhedus baileyi 7I) ££ Naemorhedus caudatus 7I) ££ £ Naemorhedus sumatraensis 7I, S, O) ££ ££££££ £ C. Tougard / Palaeogeography, Palaeoclimatology, Palaeoecology 168 32001) 337±358 353 Appendix B

Faunal lists of fossil large mammal species of South China 7Hoshangtung, Koloshan, Hsingan), Laos 7Tam Hang), Vietnam 7Tham Om, Hang Hum, Keo Leng, Lang Trang), Cambodia 7Phnom Loang), Myanmar 7Mogok), Thailand 7Thum Wiman Nakin, Thum Phra Khai Phet), peninsular Malaysia 7Tambun), Sumatra 7Lida Ajer, Sibrambang), Java 7Ngandong, Punung) and Borneo 7Niah) 7for site references, see Table 1). The large mammal faunas from the Philippine sites are listed apart because of their high endemism. The biogeographical af®nity is given in parentheses: Indochinese 7I), Sundaic 7S) and other 7O) af®nities 7Palearctic, Indian, ¼) or widespread 7W) species. The asterisk 7p) indicates that the species has currently a Sundaic habitat but it was exclusively found in the Indochinese province before the Late Pleistocene. Late Middle Pleistocene faunas from South±East Asian sites

Hoshangtung Koloshan Hsingan Tam Tham Hang Keo Phnom Thum Wiman Ngandong Hang Om Hum Leng Loang Nakin

PRIMATES Macaca sp. £££ Macaca assamensis 7I) £ Macaca fascicularis 7S) £ Macaca mulatta 7I) £ Macaca cf. nemestrina 7I) £ Macaca robustus 7I) ££ Semnopithecus sp. £ Trachypithecus sp. £ Hylobates sp. ££ Szechuanopithecus yangtsensis 7I) £ Pongo pygmaeus 7p) £££££££ Homo sp. £ Homo cf. erectus 7W) £ Homo erectus 7W) £ Homo sapiens 7W) ££ £

Holoshan Koloshan Hsingan Mogok Tam Tham Hang Keo Phnom Thum Thum Phra Ngandong Hang Om Hum Leng Loang Wiman Khai Phet Nakin

CARNIVORA Canidae indet. ££ Nyctereutes sp. £ Cuon sp. ££ Cuon alpinus 7W) £ Cuon simplicidens 7I, O) £ Ursus cf. angustidens 7I, O) £ Ursus angustidens 7I, O) £ Ursus thibetanus 7I) £ Ursus thibetanus kokeni 7I) ££ £££ Ailuropoda melanoleuca baconi 7I, O) ££££££££ Ailurus fulgens 7I) £ Mustela sibirica 7I) £ Martes sp. £ Martes ¯avigula 7I, S) £ Martes sinensis 7I, O) £ Parameles simplicidens 7I) £ Arctonyx sp. £ Arctonyx collaris cf. rostratus 7I) £££ Arctonyx collaris rostratus 7I) ££ Viverra cf. zibetha 7I) £ Paradoxurus sp. £ Paradoxurus cf. hermaphroditus 7W) ££ Paradoxurus hermaphroditus 7W) £ Paguma cf. larvata 7I, S) £ Paguma larvata 7I, S) ££ Crocuta crocuta ultima 7I, O) £££ £££ 354 C. Tougard / Palaeogeography, Palaeoclimatology, Palaeoecology 168 32001) 337±358

7continued)

Holoshan Koloshan Hsingan Mogok Tam Tham Hang Keo Phnom Thum Thum Phra Ngandong Hang Om Hum Leng Loang Wiman Khai Phet Nakin

Felis sp. ££ Felis lynx 7W) £ Neofelis nebulosa 7I, S) £ Neofelis nebulosa cf. primigenia 7I, S) £ Panthera cf. pardus 7W) £ Panthera pardus 7W) ££ Panthera cf. tigris 7W) £ Panthera tigris 7W) £££ ££ £ £

Hoshangtung Koloshan Hsingan Mogok Tam Tham Hang Keo Phnom Thum Thum Phra Tambun Ngandong Hang Om Hum Leng Loang Wiman Khai Phet Nakin

PROBOSCIDEA Stegodon sp. £ Stegodon orientalis 7I) ££££££ Stegodon trigonocephalus 7S) £ Palaeoloxodon sp. £ Elephas sp. £ Elephas hysudrindicus 7S) £ Elephas cf. maximus 7I) £ Elephas cf. namadicus 7I) ££ Elephas namadicus 7I) £££££ PERISSODACTYLA Tapirus sp. £ Tapirus indicus 7p) ££ £ Tapirus indicus cf. intermedius 7I) £ Tapirus indicus intermedius 7I) £ Megatapirus augustus 7I) £ £ ££££ Rhinoceros sp. ££ Rhinoceros cf. plicideus 7I) £ Rhinoceros sinensis 7I) ££ £££ Rhinoceros cf. sivalensis 7I) £ Rhinoceros sondaicus 7I, S) ££ £ Rhinoceros sondaicus guthi 7I) £ Rhinoceros cf. unicornis 7I) £ Dicerorhinus sumatrensis 7I, S) ARTIODACTYLA Sus sp. £££££ £ £ Sus cf. barbatus 7p) ££ Sus cf. lydekkeri 7I) £££ Sus macrognathus 7S) £ Sus cf. of®cinalis 7I) £ Sus scrofa 7W) £££ £ Hexaprotodon sp. £ Hexaprotodon sivalensis 7I) £ Cervidae indet. £££££££££££ Cervus eldii 7I) £ Cervus unicolor 7W) £££££ Rusa cf. leptodus 7I) £ Axis porcinus 7I) ££ Pseudaxis cf. grayi 7I) £ Muntiacus sp. ££ ££ Muntiacus muntjak 7I) £££ Muntiacus muntjak margae 7I) £ Muntiacus szechuanensis 7I) £ Elaphodus sp. £ Bovinae indet. £££ £ Bos frontalis 7I) ££ Bos frontalis cf. grangeri 7I) £ Bos javanicus 7I, S) £ C. Tougard / Palaeogeography, Palaeoclimatology, Palaeoecology 168 32001) 337±358 355

7continued)

Hoshangtung Koloshan Hsingan Mogok Tam Tham Hang Keo Phnom Thum Thum Phra Tambun Ngandong Hang Om Hum Leng Loang Wiman Khai Phet Nakin

Bos javanicus palaesondaicus 7S) £ Bos sauveli 7I) ££ Bubalus brevirostris 7I) £ Bubalus cf bubalis 7W) £ Bubalus bubalis 7W) £££ £ Bubalus bubalis palaeokerabau 7S) £ Duboisia santeng 7S) £ Caprinae indet. ££ £ Naemorhedus sp. £ Naemorhedus sumatraensis 7I, S) ££ £ Naemorhedus sumatraensis cf. £ kanjereus 7I)

Late Pleistocene faunas from South±East Asian sites

Lang Trang Sibrambang Lida Ajer Punung Niah

PRIMATES Nycticebus coucang 7I, S) £ Macaca sp. ££ Macaca fascicularis 7S) £ Macaca nemestrina 7I, S) £££ Presbytis sp. ££ Presbytis cf. rubicunda 7S) £ Trachypithecus sp. £ Trachypithecus cristatus 7S) ££ £ Hylobates sp. £££ Hylobates muelleri 7S) £ Hylobates syndactylus 7S) £££ Pongo pygmaeus 7p) £££££ Homo sapiens 7W) £££ CARNIVORA Cuon alpinus 7W) £££ Ursus thibetanus 7I) £ Helarctos malayanus 7I, S) £££££ Ailuropoda melanoleuca baconi 7I) £ Mustela nudipes 7S) Arctonyx collaris 7I) ££ Melogale everetti 7S) Lutra perspicillata 7W) £ Lutra sumatrana 7S) Viverra tangalunga 7S) Paradoxurus hermaphroditus 7W) ££ Arctictis binturong 7I, S) Hemigalus derbyanus 7S) Herpestes sp. Catopuma temminckii 7I, S) ££ Neofelis nebulosa 7I, S) Panthera pardus 7W) ££ Panthera tigris 7W) ££££ PROBOSCIDEA Stegodon orientalis 7I) £ Elephas namadicus 7I) £ 356 C. Tougard / Palaeogeography, Palaeoclimatology, Palaeoecology 168 32001) 337±358

7continued)

Lang Trang Sibrambang Lida Ajer Punung Niah

Elephas maximus 7I) ££££ PERISSODACTYLA Tapirus indicus 7p) £££££ Rhinoceros sondaicus 7I, S) ££ Dicerorhinus sumatrensis 7I, S) £££ £ ARTIODACTYLA Sus barbatus 7p) ££££ Sus scrofa 7W) ££ Sus scrofa vittatus 7S) ££ Tragulus javanicus 7S) £ Tragulus napu 7S) £ Cervus sp. ££ Cervus unicolor 7W) £££ Muntiacus muntjak 7W) £££££ Bos sp. £ Bos javanicus 7I, S) ££ £ Bubalus bubalis 7W) £££ Naemorhedus sumatraensis 7I, S) ££££

Pleistocene fossils fromthe Philippines

Luzon: ± Stegodon cf. trigonocephalus ± S. luzonensis ± Elephas beyeri ± Rhinoceros philippensis ± Antelope sp. ± cf. Bubalus ± cf. Cervus Mindanao: ± Stegodon cf. mindanensis ± S. mindanensis

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