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Solving mammalian riddles Chapter 3.

The palaeoenvironmental scene based on data mining

Palaeoenvironments of Borneo Geology and palaeogeography of Borneo Wilson and Moss (1999) provided a detailed overview of the geology of Borneo, and their information was used in the palaeoenvironmental reconstructions in this work, alongside the information provided by Hall (1998). A land connection between southern Borneo and mainland Southeast Asia is inferred to have existed during the Eocene and Oligocene (Pupilli, 1973 in Wilson & Moss 1999). During the Late CretaceousEarly Tertiary, a large river (possibly the ancestral Chao Phraya/Mekong River) ran across the length of Sundaland, from its Eurasian source areas to the central and northern Borneo fan area, with a delta in the Natuna Island region (Moss & Chambers 1999). The central ranges of Borneo were uplifted towards the end of the Oligocene, and the erosion of these areas supplied sediments eastwards towards the Makassar Straits (Moss et al, 1998 in Wilson & Moss 1999).

Figure 3.10. Main geological features of Borneo (after Steinshouer et al. 1997); for legend refer to Fig. 3.5.

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An important aspect of the geology of Borneo was brought to light by the work of Haile et al. (1977), who obtained palaeomagnetic data from Cretaceous igneous rocks in West Kalimantan. They stated that, although Borneo had remained approximately in the same paleolatitude since the late Cretaceous, it had rotated counter-clockwise about 50 acting as a unit with the Malay Peninsula. Similarly, Nishimura & Suparka (1997) stated that Borneo, the Celebes Sea basin and the western arm of Sulawesi rotated 50 counterclockwise during the early Miocene (2017 Mya). This rotation continued until some 10 Mya (Hall 1998), although Lumadyo et al. (1990 in Lee & Lawver 1994) had stated that the stable western part of Kalimantan has not been rotated since the Eocene. The rotation history of Borneo and Western Sulawesi is therefore still highly contentious (for an overview see Wilson & Moss 1999), and data reported by Morley (2000a) suggest that significant rotation of Borneo cannot be accommodated by Tertiary structures onshore and in the Gulf of Thailand. Pieters et al. (1987) provided an overview of the early Tertiary development of Borneo. In Late Eocene times the deposition of terrestrial to shallow marine sediments began over a large part of the island of Borneo around a central highland formed by an emergent orogenic complex (the Kuching Arch). During the Paleogene the Kuching Arch comprised island and shallow water areas, which separated the more rapidly subsiding portions of the Sarawak and Kutai Basins (Rose & Hartono 1978). The continental basement of the Schwaner Mountains in Southwest Kalimantan probably persisted as a highland area, connected by land to SE Sumatra and the Malay Peninsula. The Meratus Mountains (Fig. 3.10) and the Barito region were still submerged (Pieters et al. 1987), and the Paternoster Block, offshore of present-day SE Kalimantan, was only partly emergent during the Early Paleogene, being later transgressed during the Oligocene (Rose & Hartono 1978). In Early Oligocene times a westward transgression of the sea reached the upper Mahakam River and the Barito Shelf. A long and narrow arm of this sea extended westward almost across to the present-day west coast of Kalimantan, confined between the tectonically active emergent orogenic complex to the north and the persisting highland basement to the south. Continuing deformation and uplift along the southern margin of the orogen led to the demise of this central Borneo Basin and by Early Miocene times the basin had disappeared altogether according to Rose and Hartono (1978). In Middle Miocene

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times, the Kutai Basin became separated from the Ketungau-Melawi Basin by a high area near the present-day Mller Mountains, after which both basins slowly filled up (Ott 1987). This resulted in volcanic activity, which formed a high plateau of tuffs and other volcanic material (Molengraaff & Weber 1920, p. 467). This was then also the time when the northern part of Borneo (present-day Sarawak, north East Kalimantan, Sabah, and Brunei) became connected to the southern part (consisting of the Schwaner Mountains). Interestingly, considering that the entire Kapuas and Mahakam River valleys are low-lying (< 100 m a.s.l. even in the upper reaches), it is not impossible that, during extreme sea level highstands, the two river systems became connected again, thereby effectively separating northern Borneo from the rest of Sundaland. Between 17 and 11 Mya, sedimentation rates rose everywhere in SE Asia, spectacularly so in the Sarawak and Sabah Basins, north of Borneo. Rates of deposition of carbonates and shallow-water detrital material changed from 0.17 to 0.4 mm/year. During the Pliocene, this rate increased even more to 0.69 and 0.65 mm/year for the Sarawak and Sabah Basins respectively (Metivier & Gaudemer 1999). It remains unclear whether this increase was due to the climatic optimum during the Middle Miocene, which may have led to higher rainfall and erosion, or whether increased uplift provided the sediments. Considering that the increase continued into the Pliocene, mountain building may be a more likely explanation. Adding up the solid phase volume accumulated in the Sabah and Sarawak sedimentary basins also suggests the occurrence of considerable mountain areas on Borneo. Mtivier et al. (1999) provided a total estimate of 1.3 * 106 km3 of sediment accumulation in the Sabah and Sarawak Basins since 17 Mya. Under the assumption that the Bornean highlands were equally drained in a northern, southern, and eastern direction this would add up to approximately 4 * 106 km3 of upland being eroded in 17 Mya, if no further uplift occurred. The present Bornean uplands are approximately 1 * 106 km3, which would indicate that over the last 17 Mya, a mountain area with a height of several kilometres has been eroded. This was confirmed by R. Hall (pers. comm.), who remarked that Borneo shed vast amounts of sediments in the last 10 Mya, equivalent to the removal of 6 km of crust; as much as the Himalayas now but on a third of the area. Based on research on geological and topographic criteria, Thomas et al. (1999) estimated that in NW Kalimantan the groundsurface was lowered between

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1,200 and 1,500 m since 30 Mya, giving an average denudation rate of 4050 mm/Kya (which is considerably lower than the estimates by Metivier & Gaudemer 1999). East Borneo Zanial and Luki (1984) described the depositional cycles in the Tarakan Basin. From the latest Oligocene to early Middle Miocene sedimentation in the basins occurred in a marine environment. By the end of the Early Miocene the delta front had advanced approximately 200 km eastward of the present-day coastline (Moss et al., 1997 in Friederich et al. 1999), which brought north-east Kalimantan very close to the Sulawesi area. At the end of this period the area was uplifted, but presumably reflooded during the Middle Miocene highstand. A major change in the sedimentation history of north-east Kalimantan occurred in early Middle Miocene, when a deltaic environment developed at the western side of the region. This delta front started to prograde eastwards during the Miocene until all sedimentary processes in this area were terminated by a Late Miocene uplift at approximately 6.6 Mya. The eastern coastline of Borneo in Early Miocene times ran approximately from the south-west corner of Kalimantan, via the area of the upper reaches of the Barito and Mahakam to just north of the Mangkalihat Peninsula (Pieters et al. 1987). During the Miocene, delta systems in the Kutai Basin prograded south-eastward filling the Kutai Basin so that by the Late Miocene, deltaic deposition had generally reached a position beyond that of todays East Kalimantan coastline (Rose & Hartono 1978). By the end of the Miocene, the drainage system within Borneo was similar to the present day (Wilson & Moss 1999), although Smit-Sibinga (1953b) stated that the Mahakam River came into being only 2 Mya, and that this river initially flowed into the very large Kutai Lake (of which the present lakes are only remnants). In the Late Miocene (Middle Miocene according to Ott 1987) and Early Pliocene the Meratus Graben was uplifted and started to shed sediments to the west and to the east (Rose & Hartono 1978; van de Weerd et al. 1987), eventually leading to the rise of the Meratus Mountains. At that time, the Barito Basin became separated from the Kutai Basin by the Adang Flexure/Fault, which resulted from the uplift of the Meratus Mountains (Satyana et al. 1999). Miocene coals on the west and east side of the

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Solving mammalian riddles Chapter 3. The palaeoenvironmental scene based on data mining

Meratus Mountains, suggests that the basins on either side of these mountains were becoming terrestrial and that the climate was warm and wet (Friederich et al. 1999). The uplift of the Meratus Mountains continued into the Pleistocene (Satyana et al. 1999). East of the Meratus Mountains, a large island existed during the Tertiary (van Bemmelen 1970). The elevated area was surrounded by the depositional basins of SE and E Borneo, West Sulawesi, the Kangean-Madura-Rembang belt, and Bawean. Van Bemmelen named this land area the Pulau Laut centre of diastrophism. According to him this Pulau Laut centre was elevated at the end of the Pliocene, but the crest of this dome was rapidly engulfed during the Pleistocene, presumably leaving only the present-day land area of Pulau Laut. Emmet and Bally (1996) claimed that the eastern extension of the Kangean High was emergent in the Late Miocene, but it is unclear whether the Kangean High was part of this Pulau Laut centre. West Borneo Lloyd (1978 in Wilson & Moss 1999) suggested that Borneo lost its connection to the Malay Peninsula during the latest Miocene or Pliocene, which was possibly caused by global sea-level changes and/or plate readjustments (Wilson & Moss 1999). Quaternary sediments on the Sunda Shelf lie directly on pre-Tertiary rocks. However, as Wilson and Moss (1999) pointed out, it is possible that marine sediments deposited during possible regressions of this region may have been removed by later erosion. For further details on the divergence between the Bornean and Malay landmasses see below in the section Palaeoenvironments of the South China Sea. Ter Bruggen (1955) hypothesized on the palaeocourse of the Kapuas River. He suggested that only in the Quaternary the present upper reaches of the Kapuas broke through the Semitau uplands to join the Melawi River near present day Sintang (Fig. 3.11). The part of the Kapuas below Sintang, therefore, is the original continuation of the Melawi. In that scenario, the proto Kapuas would have flowed northward through the upper Kapuas Lakes area and followed the course of the present Batang Lupar River.

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Batang Lupar River Kapuas River

Melawi River

Figure 3.11. Map of the Kapuas River and its hypothesized change from drainage into the Batang Lupar (top) to its merge with the Melawi (bottom).

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A right tributary of the Melawi, now the piece of river between Sintang and Semitau, reached the Kapuas plain by cutting back, and decapitated the Kapuas River, which thus obtained its present course. The Kapuas Lakes area is therefore likely to consists of the former riverbed of the Kapuas. With regard to the course of the Kapuas it is also interesting to note that Smit-Sibinga (1953a) suggested that the Boh River, which is now a tributary of the Mahakam, used to be part of the upper reaches of the Kapuas; stream capture later joined it to the Mahakam River. Sabah Rice-Oxley (1991) and Tan and Lamy (1990) provided a detailed description of the Early MiocenePliocene palaeoenvironments of north-west Sabah. The whole of offshore NW Sabah was a realm of deep marine shale deposition during the Early Miocene to early Middle Miocene. The coast ran approximately in the same area as the present-day coastline and was prograding in a NW direction, while sediments were sourced from the highlands of the Rajang accretionary prism (Tan & Lamy 1990). The Middle Miocene to early Late Miocene saw the uplift of the Crocker Range and a further NW shift of the coastline (Tan & Lamy 1990), while the Kinabalu intrusion was emplaced in the middle Late Miocene (Jacobson, 1970 in Tan & Lamy 1990). Mt. Kinabalu, the highest mountain in the Sundaland area, was uplifted between 4 and 10 Mya, although uplift to its current height is thought to have occurred only within the last 1.5 Mya (Barkman & Simpson 2001), while the mountain reached its present height about 100 Kya (Choi, 1996 in Tanaka et al. 2001) providing suitable habitat for mountainous species. During the Late Miocene, the northern Borneo coastline changed from having a N-S orientation to a SW-NE orientation, similar to today (Wilson & Moss 1999). The middle Middle Miocene to Pliocene stage is characterised by two major depositional cycles, each starting with an initial phase of coastal plain sedimentary upbuilding, followed by rapid transgression (see detailed maps in RiceOxley 1991). Between the Late Miocene and Pliocene, the palaeocoastline in NW Sabah was positioned off-shore the present-day coastline, approximately following the Mangalum and Morris Faults (see Figure 3 in Tan & Lamy 1990). Early Miocene coal finds in the Maliau Basin suggest wide tidal flats in this area and warm and wet environmental conditions (Tjia et al. 1990).

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Sarawak Most of Sarawak consists of deep sedimentary basins although, in the region east of Kuching, basement rocks appear near the surface (Isaacs 1963), which suggests that this area has been emergent for a long time. Agostinelli et al. (1990) produced palaeogeographic maps for Sarawak. Through the Miocene, the palaeocoastline was oriented more or less orthogonally to the present one, suggesting that a considerably part of what is now northern and central Sarawak was below sea-level. The main source of sediment for the coastal Sarawak area appears to have been somewhere north of Kuching, which fits Isaacs (1963) hypothesis of an old emergent land area in that region. Uplifted anticlines on the present Sarawak land area caused the deposition of vast amounts of sediments on a rapidly prograding shelf during the early Late Miocene (ca. 11 Mya) (Mat-Zin & Tucker 1999)), which led to the following palaeogeography: A coastline running parallel to the present one, a broad shelf, and a steep transition to deep water (Agostinelli et al. 1990). Towards the end of the Tertiary, most of west Sarawak was raised above sea-level. A prolonged period of erosion followed in the Late Tertiary and Early Quaternary times, reducing much of the area to a peneplane (Tan 1986), probably levelling down the summits of all mountains to the 1,500 m contour in Late Miocene times (Muller 1971). At various times during the Quaternary, the present-day rivers were able to extend their levees much further into the coastal shelf than at present (Andriesse, 1972 in Tan 1986). Vegetation of Borneo The pollen record for Brunei suggests a strongly seasonal palaeoclimate in Late Oligocene and Early Miocene and to a lesser extent also in the Late Miocene Pliocene, with abundant gymnosperms such as Pinus, Abies, Tsuga, Keteleria and Ephedra (Muller 1966, 1975). These species gradually disappeared during the Late Miocene and Pliocene (Germeraad et al, 1968 in Watanasak 1990), although during the Late Miocene the appearance of the seasonal mangrove taxa Aegialites and Camptostemon in Brunei indicates a return towards a more seasonal climate (Morley 1977). Also, the conifers Phyllocladus and Podocarpus first appear in the Late Pliocene of Borneo, with the former arriving at the Plio-Pleistocene transition, indicating a land connection between Malaya and Borneo at that time, and either cooler climates and/or substantially higher mountains in Borneo than at the present

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(Muller 1971). According to Morley (1977), for the montane taxa to arrive in Borneo a direct mountainous land connection with Southeast Asia had to exist, but later, Morley (1999) revised this idea by suggesting that cooler and drier climatic conditions could have accounted for the southward spread of these species. Because most of these taxa are wind dispersers, a land connection between Borneo and Malaya may not even have been necessary. Pollen and spores finds in Early Miocene coals in the Maliau Basin in Sabah (Tjia et al. 1990) suggest warm everwet climates, and not strong seasonality as suggested above, as they indicate the presence of ombrogenous peat swamps in everwet climates, mangroves, seasonal swamp forests, and possibly other habitats (Morley 1991). This may suggest that there was climatic variation between seasonal and everwet climates within the Miocene. The vegetation of Middle Miocene peat swamps from SE Kalimantan (Demchuck and Moore, 1993 in Morley 2000) and the Tarakan Basin (Morley 1991) can be inferred from palynological analysis of coal. The presence of mangrove and back-mangrove species suggests that the peat accumulated partly under brackish conditions, whereas elsewhere watershed peats occurred; the presence of Meyeripollis naharkotensis in coal suggests everwet climatic conditions. Mangrove peats formed at a time of maximum Miocene global sea-levels and temperatures, and it is suggested that conditions somewhat warmer than today are necessary for the formation of these peats (Morley 2000). Further occurrence of EarlyMiddle and MiddleLate Miocene coals in the Mahakam Delta, as described by Peters et al. (2000) and coals in Miocene and Early Pliocene sequences of the Kutai, Barito, and Asam Asam Basins (Friederich et al. 1999) suggest that overall the MioceneEarly Pliocene climate in north and east Borneo was warm and wet. In the Late MiocenePliocene the climate became cooler and drier. Gramineae maxima in Late Miocene and Pliocene sediments of the Mahakam Delta suggest more open savannah vegetation intermittently replacing the rain forests. Also, the distribution of the mangrove taxa Aegialites and Camptostemon in Sarawak and the Mahakam Delta may relate to phases of dry climate, coinciding with periods of low sea-level (Morley 2000). East and SE Borneo are still much drier than the other parts

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of the island, and increased seasonality or reduced rainfall would have first translated into vegetation changes on that part of the island. I was unable to find palaeoenvironmental data for the earlier part of the Pleistocene, a period of considerably importance to the evolution of present-day mammal species of Borneo. However, there are several sources that suggest considerably colder temperatures and reduced rainfall during the Late Pleistocene. Thomas (1987) found geomorphological features in West Kalimantan that suggested that 40 Kya ago this area underwent a 50% reduction in rainfall, and that a monsoon climate existed accompanied by a tree-savannah vegetation. These findings were corroborated by Thorp et al. (1990) who interpreted extensive braided and fan-like alluvial landforms to be the results of dry, savanna-like palaeohydrological conditions during oxygen isotope 3 (post-60 Kya). These alluvia were dissected, their surface sediments podzolized and their incised valleys re-alluviated during the last 15 Kyr (Thorp et al. 1990). Jirin (1993) described Late Pleistocene vegetation changes in five palynological zones from Sabah. Pollen in the oldest zone, probably representing the penultimate glacial period, suggest a cold and dry climate, which led to the expansion of montane vegetation. Lowland cover contracted as precipitation was reduced, while sea-level was low, which led to the reduction of mangrove vegetation. A sea-level high represented in the next zone caused the mangrove vegetation to expand. The climate was warm and wet, and montane vegetation was reduced, while lowland vegetation expanded. The next zone represents an extensive sea-level fall during the LGM. The cooler and possibly drier climate caused montane forest to expand to lower altitudes. Expansion of lowland vegetation at the end of this period indicates climatic amelioration, and after the Pleistocene-Holocene boundary mangrove cover expanded, and montane vegetation retreated to its present altitudinal range (Jirin 1993) (note that there is no mention of grasslands). Another pollen diagram from East Borneo shows an increase in savanna at ca. 20 Kya, which might serve as an indicator of a drier climate than at present (Flenley 1998). Dated charcoal finds caused Goldammer and Siebert (1989) to believe that the coastal area of East and South Kalimantan was more continental and drier than at present and they assumed that forest formations at that time were more seasonal and had probably a temporary fire climax character. Majid (1982) suggested that at the height of the LGM, the Niah area in Sarawak was covered

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by deciduous monsoon forest, while Caratini et al. (1988) suggested that, during the LGM, the hinterland of the Mahakam Delta probably consisted of grassland or savannah. Not everywhere on Borneo did the cooler and drier conditions lead to more open vegetation types as shown by the findings of Anshari et al. (2000) and Kershaw et al. (2001). Their data from West Kalimantan indicate that during the LGM the upper Kapuas area was covered in rainforest, very similar to that found during the midHolocene. Also, in the Sebangau area of Central Kalimantan, peat datings showed that peat had started to develop at 18.3 Kya 0.05, and had continued to do so until c. 7 Kya, after which no peat was formed until it started again at 1.3 Kya (Page et al. 1999a); again this indicates that the climate remained relatively warm and wet during the LGM, although peats may only have started to develop after the height of the glacial period. Palaeoenvironments of the Malay Peninsula and Malacca Strait Geology and palaeogeography of the Malay Peninsula and Malacca Strait Hutchison (1990) described the palaeogeography of the Malay Peninsula (Fig. 3.12) during the Paleogene. The Paleogene drainage from Sundaland flowed southwards on the tilted basement surface through the NS Bengkalis Graben, the fault zone that forms the eastern coast of Sumatra, the Sunda and Asri Basins offshore west Java, and the grabens of Northern Sumatra (Hutchison 1996). The Muar River in Peninsular Malaysia, which was still connected to the Pembeling River, is a relict of this palaeogeography. This river system flowed south down the regional slope, probably reaching the Indian Ocean on the South Sumatran coast. Hutchison (1990) stated that the Paleogene Chao Phraya-Mekong River flowed southward along a regional slope as well, a direct analogue of the Tembeling-Muar, but because the Chao Phraya-Mekong River probably debouched near the Natuna Islands it is unlikely that the two river systems were connected. East of the Malay Peninsula lies the Malay Basin, which is of Oligocene to recent age. The basin is about 350 km long and 250 km wide (Madon & Watts 1998), and it is closely associated with the Thai Basin in the north and the Penyu and West Natuna

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Basins in the south (Fig. 3.12). It is bounded at the north-eastern flanks by the Khorat Swell and in the south-west by the Tenggol Arch (Ramli 1988).

Figure 3.12. Main geological features of the Malay Peninsula (after Steinhouser et al. 1997); for legend see Figure 3.5.

Armitage and Viotti (1977) described the depositional environments of this basin, on a location some 300 km east of Kuala Trengganu, which lies on the present east coast of Peninsular Malaysia. From Early to Middle Miocene a fluvial plane existed, grading to a coastal plain, which changed towards a brackish coastal plain with more marine influences, while from Pliocene to recent time a marine to neretic environment existed (Armitage & Viotti 1977). Madon and Watts (1998) also mentioned that during the Early to Middle Miocene the basin was characterised by non-marine and brackish

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Solving mammalian riddles Chapter 3. The palaeoenvironmental scene based on data mining

depositional environments, while Mtivier et al. (1999) reported that this cycle is composed of successive fluvio-lacustrine, littoral and deltaic or swamp deposits. According to Ramli (1988), the southern part of the Malay Basin (at latitude 400 N) started to become marine in the early Early Miocene, and by the start of the Pliocene the entire Malay Basin was predominantly covered by holomarine inner neritic sediments. The abundance of coal, however, suggests that the basin was at or near sealevel during most of the Miocene and was influenced by only minor fluctuations in sea-level; although this was questioned by Higgs (1999) who suggested that coals in the Malay Basin were allochthonous, and that the basin was much deeper. As coals only form in maritime climates with > 3 m of rainfall and no marked dry season (Cole 1987), we do get an idea of the prevalent climatic conditions in the Malay Basin area during the Miocene. A similar structural history has been described in the Penyu Basin, which lies between the Malay and West Natuna Basins (for the latter see the section on Palaeoenvironments of the South China Sea). In Late Miocene to Pliocene times, both basins developed a shallow marine environment, where water depths probably never exceeded 200 m (Azmi et al., 1994 in Madon & Watts 1998). By the Early Miocene, the Malay Peninsula had almost reached its present-day position, after having moved with the Indochina block in a southerly direction, but the adjacent North Sumatra Basin and central Thailand basins were still undergoing extension (McCabe et al., 1998 in Lee & Lawver 1995). Still, the area was affected both by vertical movements and changing sea-levels. Scrivenor (1931) suggested that Mt. Ophir (named Gunung Ledang on more recent maps) and Mt. Kedah were once islands, and that a line from Alor Setar (in Kedah State) to Songkhla (in Thailand) was the coast-line of land that was once an island, or group of islands, but had since become the southern part of the Thai-Malay Peninsula. Scrivenor provided further tentative evidence of former high sea-levels on the Malay Peninsula. He described marine sponge-spicules of relatively recent age (Pliocene or Pleistocene) at an altitude of at least 70 m in the upper Perak River area (or rather its tributary the Plus River). Tjia (1973) also mentioned raised Quaternary shorelines in Perak, Selangor, and the Kinta Valley which were found at altitudes of 6075 m above present day sea-level. He tentatively correlated these high sea-levels to the Milazzian, an Early-Middle

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Pleistocene interglacial. Possibly, these can be correlated to the high sea-levels at about 21.7 Mya, as seen in sea-level curves presented by Hutchison (1989, p. 70). These high sea-levels are correlated with the Malayan boulder beds and Old Alluvium, which Tjia, and in the same book also Stauffer (1973), give an EarlyMiddle Pleistocene age, but which others have dated as much younger (see discussion below). The fact that the boulder beds are deformed structurally, with dips of 60 locally, and a thickness of up to 300 m strongly suggest that considerable time has passed since their deposition, making a Late Pleistocene age unlikely (Stauffer 1973). Burton (1964) thought that the Older Alluvium on terraces of at most 85 m a.s.l. were deposited during an Early Pleistocene highstand. Interestingly, Burton distinguished between two types of Older Alluvium, i.e. wide-spread terraces with a maximum altitude of 70 m a.s.l., and older, coarser textured sediments at an altitude of between 90 and 140 m a.s.l. In the last 5 Myr, sea levels highstands over 50 m a.s.l., have probably only occurred 5 times, as judged from sea level curves presented by Haq et al. (1987) and Mitchum et al. (1993): 1. During the EarlyMiddle Pliocene; 2. at ca. 4.0 Mya; 3. at ca. 3.4 Mya; 4. at ca. 1.05 Mya; and at ca. 0.3 Mya. Only the EarlyMiddle Pliocene highstand seems to have been close to or over 80 m a.s.l. making it a likely candidate for the higher terraces of Sundaland. Gupta et al. (1987), in their description of the Old Alluvium deposits, suggested that these were associated with seasonality of water flow. Such a seasonal component may have resulted from a dry southwest monsoon due to the increased size and relief of Sumatra. Up until recently, the age of these deposits remained a matter of much debate (see Batchelor 1993; Thorp & Thomas 1993). Batchelor (1993) considered the Old Alluvium of Peninsular Malaysia and Singapore, which he correlated with the Older Sedimentary Cover offshore Peninsular Malaysia, to be of Late Pliocene/Early Pleistocene to Middle Pleistocene age (at least 700 Kya). Thorp and Thomas (1993), on the other hand, disagreed with this dating and suggested that the Old Alluvium was of much younger age (Late Pleistocene). They based this on what they considered reliable datings by thermoluminescense. Furthermore, they were convinced that this Old Alluvium correlated with the Late Pleistocene alluvial bodies that they found in West Kalimantan. These latter deposits were built during a period of increased erosion and sedimentation during the isotope 3 stage of the last glacial cycle, which was

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characterized by rainfall of 23 m/year over a period of probably more than 30 Kya. Part of this debate has recently been decided in favour of Thorp and Thomas by Teeuw et al. (1999), Kamaludin and Azmi (1997), and Kamaludin et al. (1993). The former dated west Bornean sediments at 753 Kya, which is largely in agreement with the Late Pleistocene dating by Thorp and Thomas. Similarly, Kamaludin and Azmi (1997) and Kamaludin et al. (1993) dated the upper strata of the Old Alluvium deposits in western Malaysia at between 67 and 29 Kya, which again agrees with Thorp and Thomas. Still, the fact that fossils of typical Middle Pleistocene species, such as Palaeoloxodon namadicus were found in the Old Alluvium north of Kinta Valley (Peninsular Malaysia) indicates that at least parts of that stratum are of genuinely Middle Pleistocene age (Stauffer 1973) (unless the species survived into the Late Pleistocene), and it probably needs to divided into different units. In a study of the geology of the Malacca Strait, Emmel and Curray (1982) found the existence of a rugged basement in the southern Strait (south of 3N). This basement, which they considered to be of pre- or Early Pleistocene age, consisted of low peaks, 40 m higher than the valleys. Emmel and Curray (1982) considered this basement indicative of strong erosion during lowered sea-level. Overlying this rugged topography were several types of sediments. Firstly, the old sea floor, like the present one, represents an abrasion surface, formed initially during a low-level sea stand and followed by a transgression. During the low-level phase, this surface was subaerial, as evidenced by the small erosional valleys cut into it. Emmel and Curray correlate the old sea floor with the low sea-level and the transgressive phase preceding the Sangamon (Eem, or Riss-Wrm) interglacial, although this is speculative. Further sediments were described by Kudrass and Schlter (1994). Firstly, they discussed the sequence I sedimentary cover that was found directly on top of sedimentary bedrock in both the northern and southern Malacca Straits. They ascribed a tentative age to this sequence, i.e. Tertiary to Early Pleistocene, and suggested it to be an exclusively terrestrial deposit, which could possibly suggest that Sumatra and the Malay Peninsula were then connected. On top of sequence I, Kudrass and Schlter found an unconformity that was probably caused by rejuvenated fluviatile erosion during an extended period or several periods of lowered sea-level. They tentatively

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dated the termination of this unconformity at 0.9 or 0.6 Mya, when the increased build-up of glacial ice-sheets enhanced the amplitude of sea-level changes (Berger et al., 1993 in Kudrass & Schluter 1994). In contrast to sequence I, sequences II and III were deposited in an environment which was influenced by frequent shifts of the base level of erosion. The small-scale fluctuations may be correlated to the transitional periods from the interglacial to the glacial period, when sea-level shifted several times across the 30 m depth contour (Shackleton, 1987 in Kudrass & Schluter 1994). Kudrass and Schlter (1994) found two major unconformities at the boundaries of sequences II/III and III/IV, which they ascribed to prolonged periods of widespread erosion caused by long periods of low sea-level. These periods were tentatively correlated to the two last high glacial periods, when sea-level was lowered more than 50 m from 190125 Kya and from 7010 Kya. The final sequence IV was deposited after the LGM. During the transgression, the Strait of Malacca was an elongated shallow bay where a great supply of fine-grained terrigenous sediment partly compensated the rapid sea-level rise. Up to 30 m thick Holocene coastal mudmangrove-peat accumulated in a short period (Kudrass & Schluter 1994) and the final opening of the Strait between the Indian Ocean and the South China Sea may only have occurred as recently as 5 Kya (Geyh et al., 1979 in Kudrass & Schluter 1994). Vegetation of the Malay Peninsula and Malacca Strait The flora in the Early to Middle Miocene Malay Basin area included species that indicated a swamp or mangrove setting (Morley 2000), while Gramineae were also found. Especially during the drier climate phases of the Miocene and Pliocene, which were possibly correlated to the periods of low sea-level, Gramineae maxima in the Malay Basin suggest more open savannah vegetation intermittently replacing the rain forests. These Gramineae maxima are more common in Late Miocene and Pliocene sediments than in the latter part of the Middle Miocene and Early Miocene (Morley 1999). Morley (1999) reported on a very distinctive occurrence of Dacrydium, which is a good indicator of heath forests, within the Pliocene of the Malay and Thai Basins. Stauffer (1973) and Hing and Leong (1990) described plant remains in Late Tertiary coal beds in basins of the Malay Peninsula. Coal at Batu Arang near Kuala Lumpur contained a forest flora indicating a drier climate than now, or a partly upland source for the transported material (Stauffer 1973). More recent dating, however, suggested

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that these coals are probably of Eocene to Oligocene age (Ahmad Munif, 1993 in Hasiah & Abolins 1998). The coal composition of PlioceneEarly Pleistocene (?) deposits in the Kepong and Kluang-Niyor Basins (Stauffer 1973), and the Merapoh Basin (Hing & Leong 1990) indicates that these were derived mainly from woody and herbaceous plants as well as from spore-bearing plants, as would have been found in a tropical forest dominated by angiosperms and to a lesser extent gymnosperms (Hing & Leong 1990). Morley (1999) suggested that Pinus savannah was probably widespread on the Malay Peninsula at 660 Kya, 480 Kya, 200 Kya, and 22 Kya, while Heaney (1991) mentioned that pine-grasslands were found near Kuala Lumpur at 160 Kya. All of these, apart from the 200 Kya time, are periods of low sea-level and presumably drier, slightly cooler conditions2. During the interstadials the climate in the lowlands of Peninsular Malaysia was probably as that prevailing today, which is suggested by pollen found at interglacial deposits dated at ca. 80 Kya and 55 Kya (Kamaludin & Azmi 1997). Also, finds of peat, wood, laterite, and oxidized iron described by Stauffer (1973) suggests that during the Pleistocene there were at least several periods in which perhumid conditions existed, although these finds were not dated and can therefore not be reliable correlated with any particular glacial or interglacial periods. Elsewhere, Ayob (1970) provided carbon-dated peat and wood samples from deposits containing pollen indicating perhumid vegetation; these samples were dated at ca. 36.4 Kya and 41.2+ Kya, indicating that before the LGM an evergreen vegetation type existed in this area close to present-day Kuala Lumpur. Price et al. (1997) suggested that bauxite formation on the Malay Peninsula took place continuously from 115 Kya to the present, which would suggest mostly warm and wet conditions. Geomorphological research by De Dapper (1985) on landforms in the Malay Peninsular uplands suggested a shift from dry climatic conditions with a fairly open vegetation (tree or grass savannah), rather unprotected slopes and braided river systems (T2-terrace and P2 pediment) to much wetter conditions with slopes well protected by a dense forest cover (T1-terrace). The T2 surface was locally covered by

According to Hantoro (1995), the 480 Kya glacial is part of tge same glacial period and sea-level low as the 450 Kya sea level low mentioned in Table 3.1.

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ashes, which X-ray microanalysis ascribed to the Toba volcano, while the ashes never occur on T1 surfaces and are even locally covered by the T1 alluvia. The Toba ashes could either be from explosions at 74 Kya or 30 Kya (see section on Geology and Palaoegeography of North Sumatra), and the climatic shift probably refers to conditions during the 80 Kya glacial and the interglacial conditions following it, although it cannot be excluded that the terraces refer to LGM and post-LGM conditions. A combined coastal and offshore survey in the Strait of Malacca between Port Dickson and Singapore yielded evidence of per-humid climatic conditions in this region before the sea-level rise following the LGM. Between 50 and 10 Kya, dry land conditions with peats and mangroves prevailed in most of the southern Malacca Strait, at times in association with freshwater lakes, as indicated by the presence of diatom ooze (Geyh et al. 1979). North of 5N, Emmel and Curray (1982) found evidence of deltaic progradation. The upper Pleistocene deposits here consisted of silty clay in which peat was regularly found, suggesting deposition in calm waters, probably into vegetated waters or lagoons. Emmel and Curray suggested abundant vegetation in the emergent Malacca Strait, probably resembling the lowland vegetation of tropical regions, with mangroves in the low-lying areas and Nipah palm along the banks of muddy creeks. Tropical rain forest would have covered the higher drier parts of this area. These conclusions are in contrast with those by de Dapper (1987) who reviewed arguments against and in favour of drier conditions and more open vegetation types during the LGM in the Malay Peninsula. Based on geomorphic evidence in the uplands of the Malay Peninsula he concluded that during the LGM vegetation in these areas was much more open, and only reverted to tropical rainforest in the Holocene. Finally, Taylor et al. (2001) investigated cores of sediment from Nee Soon, a peat swamp in the perimarine zone of Singapore, which yielded a record of environmental change comprising the LGM and Holocene periods. The evidence indicated the occurrence of swamp conditions at Nee Soon during the late glacial and early Holocene and taxa presently associated with highland areas in dryland forest at low altitude. This would suggest either temperatures substantially lower that those at

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present and, possibly, humid conditions, or cold, seasonally dry climates and reduced levels of atmospheric CO2. In summary, on and around the Malay Peninsula, there appears to be evidence of alternating phases of open, grassy vegetation types during glacials to closed evergreen forest during interglacials. Palaeoenvironments of the South China Sea Geology and palaeogeography of the South China Sea The South China Sea, in very broad terms, is that part of the Pacific Ocean generally west of the Philippines and north and west of Borneo. During the Palaeocene to Middle Miocene, the South China Sea opened, leading to a southward migration of mainland Asian continental crust of about 700 kilometres in 25 Mya. This spreading of the South China Sea carried the present-day areas of Palawan and Mindoro from the Asian mainland to their present position in the Philippine archipelago (Holloway 1982). Sundaland, or the Sunda Shelf Plate, is considered to be composed of a mosaic of continental and oceanic microplates accreted and sutured together in the Late Triassic (Pulunggono 1985; Cole & Crittenden 1997). Since the Early Tertiary, the Sunda Shelf plate has generally tilted southward and has been subsiding (Ponto et al. 1988). This resulted in the development of a large basin in the Gulf of Thailand and the Sunda Shelf (White & Wing 1978), between the Natuna Ridge, an extension of the Sunda landmass and the Khorat Con Son Platform, which was part of the Asian mainland. The eastern part of the basin extends from offshore Vietnam, across Indonesian and Malaysian waters into Sarawak and Brunei (White & Wing 1978). The Natuna-Khorat Ridge appears to have had enough local topographic relief to prevent a marine environment from penetrating the West Natuna Basin until the Middle Miocene when this sill was breached and a more marine environment developed in the West Natuna Basin (White & Wing 1978). By the early Middle Miocene, the entire oceanic crust of the South China Sea appears to have been subducted beneath the Borneo accreted continental crust, and the continental crust of the South China Sea Platform collided with Borneo. The more rigid and buoyant continental crust does not

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bend as much and this resulted in a period of regional uplift and erosion throughout early Middle Miocene times in the South China Sea area (Tan & Lamy 1990). The East Natuna Basin and most of the West Natuna Basin indicate Miocene deposition in deltaic or littoral environments (Wongsosantiko & Wirojudo 1984). The area west of the Natuna Islands was uplifted during the Miocene and became temporarily emergent in Late Miocene to Early Pliocene (Holloway 1982). Similarly, the East Natuna area emerged in the Late MioceneEarly Pliocene, probably as a result of a sea level lowstand (Martono 2000). Later in the Pliocene open marine muds once more buried the Natuna Basins (White & Wing 1978). Hall (2002) suggested uplift of this part of the Sunda Shelf (the area between Borneo, the South Malay block, and Sumatra, i.e. the Singapore Platform) from about 15 Mya because of the presence of thick Neogene sediments filling the offshore NW Java basins. This appears to fit van Bemmelens (1970) theory of an emergent Singapore Platform during the Miocene and Pliocene that supplied sediments to northern Java. Also, Tjia and Liew (1996 in Hanebuth et al. 2000) claimed that the Singapore Platform was tectonically stable during the Pliocene and Pleistocene, and may have been emergent during most of that time. Ben-Avraham and Emery (1973) similarly suggested that after the Early Miocene, the Singapore Platform, Lampung High, and Karimunjawa Arch were the main high areas supplying sediments to the surrounding basins. Furthermore, considering the age of the AnambasNatuna Arc and the Riau-Bangka Arc, these must have been much higher and wider in earlier times (Inger & Voris 2001), possibly providing mountainous corridors. During Pleistocene low sea-level stands, these arcs protruded 7501,000 m above the surrounding flatlands, trapping moisture on one side and creating a rain shadow on the other. Hall (in litt., 20 March 2000 and 12 November 2001; 2002) remarked that on the Sunda Shelf generally marine conditions come in the Late Miocene (later to the NW and earlier to the NE) and may be as late as Pliocene, while at the same time global sea-level was falling. Preceding this marine transgression, there was probably a tectonic subsidence event, since there is a major unconformity in this area, not seen further north in the Gulf of Thailand. The result of this event was to remove about

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2,000 m of strata in the Malay Basin, and further south, the ridge connecting Borneo and the Malay Peninsula has only Quaternary strata resting directly on basement. Isaacs (1963) reported on a minor sedimentary basin about 80 km west of the Tambelan Islands (west of Borneo), which reached a depth of ca. 700 m below sealevel. This area is probably where Borneo and the Malay Penisula were first separated; considering the amount of sediment, it may have remained submerged during most of the Quaternary. This basin which has become filled with sediments can tentatively be followed further south towards the islands of Tujuh, Karimata, and Bangka. Aleva et al. (1973) reported on a MiocenePliocene sedimentary cover, with a thickness of over 100 m in some areas; on top of this they found a sequence of sedimentary layers and erosional surfaces. Only one of these erosional surfaces, the Red Clay Formation, clearly developed when the area became emergent, which, according to Aleva et al., happened during the LGM. An older terrestrial deposit also existed, the Alluvial Complex, which may have been deposited during the Late Pliocene sea-level lowstand (but see discussion on the age of the Alluvial Complex in the section on Palaeoenvironments of the Malay Peninsula). It consisted of valleys and depressions, which deeply incise the Older Sedimentary Cover (which Aleva and colleagues considered to be of Late Tertiary age). The occurrence of peat layers and the alternation of deep incisions of valleys with thinly stratified sedimentation indicate frequent vertical oscillation in relation to the sea-level (Aleva et al. 1973). Aleva et al. mentioned that the pollen content of the Alluvial Complex and the Older Sedimentary Cover are rather similar, and that the latter had been dated as MiocenePliocene. The above information suggests that the opening-up of the land between Borneo and the Malay Peninsula started in the north, possible in relation to the deepening of the Malay Basin. A sea inlet developed west of the Tambelan Islands, which moved south towards the Karimata, Bangka, and Belitung Islands. This presumably happened during the Pliocene. The data suggest that this sea inlet reached the Java Sea at the time when the Older Sedimentary Cover was deposited (MiocenePliocene according to Aleva et al., but possibly younger). This first sea connection between the South China and Java Seas probably closed again during a major sea-level lowstand, possibly the one at 2.4 Mya, as evidenced by the Alluvial Complex. The area once

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more became inundated, presumably separating Borneo from Malaya and Sumatra, and then re-emerged during the LGM. Molengraaff and Weber (1920) and Verstappen (1975) hypothesized that during Pleistocene low sea-levels the rivers of west Borneo and SE Sumatra formed part of one river system, the North Sunda River. Two other rivers emptied into the present day South China Sea: the Mekong, and the Chao Phraya and tributaries collected south to the tip of peninsular Malaysia. The drainage of the present day Java Sea was eastward and a ridge along the islands of Bangka-Belitung-Karimata separated this system from those to the north (Verstappen 1975). During the Late Pleistocene glacial, Sumatras rivers flowed into a number of major river systems. The flowage between the Malay Peninsula and Sumatra had as tributaries the north-eastern Sumatran rivers, such as the Simpang Kanan, Panai, Rokan and Siak, and also several large rivers from the west of the Malay Peninsula, i.e. the Perak, Bernam, Muar and Lenek Rivers (Voris 2000) (note that according to Scrivenor 1931, the Pahang River flowed until recently through the Tasek Bera area and the present Muar River to debouch into the Malacca Straits). Further south along the Sumatran coast the Indragiri, Hari and Musi joined the river systems from West Borneo to form the North Sunda River System or Molengraaff River (also see Wyrtki 1961). Judging from the 120 m sea-depth contour, the mouth of this river would have been at approximately E 109.60 and N 5.11, between the northern and southern Natuna Island groups. In between these two river systems, a third river system existed according to Voris (2000), which drained the present Kampar River (running through the Singapore Straits) and was joined by the Johore River before flowing into the large river system from the Gulf of Thailand. These rivers all disappeared when the shelf area became inundated once more. Pelejero et al. (1999a) found evidence for the beginning of the inundation of the Sunda Self at 14.9 Kya, although this could also have happened at 13 Kya, as suggested by Broecker et al. (1988). A reconstruction of this inundation suggests that at the beginning of the deglaciation the position of the Molengraaff River mouth remained unchanged during the first small step in sea-level rise, due to the morphology of the shelf break. It is not until 1513.5 Kya that a fundamental change occurred in the oceanographic setting of the southern South China Sea, when the Sundashelf was

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flooded across the threshold depth of 70 m. At this time, the river mouth was rapidly displaced landward by more than 200 m/year. The last stage of the flooding and submergence of Sundaland was accomplished at about 10 Kya, when the 30 m depth of the present Karimata Strait was reached (Pelejero et al. 1999b). This is in line with the findings by Wang et al. (1994) who suggested that the gateway between the Java and South China Seas (sill depth 36 m) opened at about 10.5 Kya. Vegetation of the exposed South China Sea area Land exposure on the Sunda Shelf was often linked to drier and colder climates, and possibly increased seasonality (Verstappen 1975, 1980, 1997; Gupta et al. 1987; Stuijts et al. 1988; Thorp et al. 1990; Thomas et al. 1999). Some authors have therefore suggested that the land connection between Borneo, Sumatra, Java and the mainland was covered in savannah-type vegetation (Muller 1975; Morley 1981; Morley & Flenley 1987; Broecker et al. 1988; Caratini & Tissot 1988; Heaney 1991), although monsoon forest (Morley, pers. comm., in Whitmore 1981; Whitten et al. 1996; Adams & Faure 1997; Taylor et al. 1999) has also been suggested. Adams and Faure (1997) and Chappell and Thom (1977) suggested that, at least in the final glacial stages of the Pleistocene, rainforest colonisation rates may have been insufficient for rainforest to recolonize the exposed land between Java, Sumatra, and Borneo, because of the brackish soils left behind as the sea retreated. Here, I investigate the evidence for drier, more open vegetation types on the exposed Sunda Shelf. Based on pollen data Morley (1978, in Morley & Flenley 1987) suggested that in the Early Miocene, Late Miocene and Pliocene, much of the South China Sea (SCS) experienced a strongly seasonal climate. Earliest Miocene palynomorph assemblages from the Natuna and Malay basins are thought to reflect drier climates than those in the Late Oligocene, due to the common occurrence of Gramineae, and the low representation of fern spores and spores characterizing peat swamps. Interestingly, pollen, comparable to that of Shorea and Hopea, was common, which suggests that low diversity Dipterocarp monsoon forests must have been widespread at this time. When sea-levels rose and land became submerged at approximately 20 Mya, this association disappeared (Morley 2000). The common occurrence of temperate conifers, such as Abies, Picea and Tsuga in north Borneo, the Natuna and Malay

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basins and Indochina, probably relates to cooler climatic conditions in the low latitudes in Late Oligocene and Early Miocene (Morley 2000). Further north, on the northern Sunda Shelf, the Pliocene is characterised by the presence of abundant pollen of the conifer Dacrydium and pollen of the Polystachyus type, which indicates either montane environments, or heath forest, especially where there is impeded drainage, with or without peat formation (Morley 1977, 1999). No further data were found on the PlioceneMiddle Pleistocene environments of the southern SCS. Pelejero et al. (1999a) modelled the MiddleLate Pleistocene climate of the SCS area by looking at a wide range of biomarkers. They found that sea surface temperatures in this shallow enclosed system showed much more variation between glacial and interglacial times than open ocean areas at the same latitude. For instance, the difference in SCS temperature between present-day times and the LGM is 2.8 C, whereas elsewhere this difference varies between 1.3 and 1.8 C. During the LGM, sea surface temperatures in the SCS were down to 25 C. Further climatic events suggested by the results of Pelejero et als (1999a) research include the following: 1. at 19.5 Kya a period of extreme precipitation and river run off; 2. a very warm period between 127 and 116 Kya, with sea surface temperatures between 28.7 and 29.9 C (as opposed to present-day 28 C) and sea-levels at 6 m higher than present-day levels (also see below); and 3. two periods dated at 2928 Kya and 37.7 43.2 Kya that were characterised by extreme tropical precipitation and enhanced summer monsoon activity. Pollen spectra from the southern SCS indicate that during the last glaciation, the lowland was covered by tropical lowland rainforest, and mangroves grew along the river mouths and along the coasts. The periodic expansion of montane gymnosperms implies falling temperature, at least on the mountains of surrounding islands, but no desiccation was found during the glacial period (Sun et al. 2000). There is further support for the maintenance of high rainfall and rainforest in the southern part of the SCS. Pollen data from peat sediments taken from the Pulau Tujuh area, southeastern Sumatra, strongly suggests rainforest or peat swamp forest vegetation during the LGM. Pandanus is common, while values for Cyperaceae are low and Poaceae occurs at a low percentage. In 4 samples, there are significant percentages of Rhizophoraceae

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suggesting increased proximity of mangroves (van der Kaars, unpubl. data in Kershaw et al. 2001). Also Kawamura (1998) found Late Pleistocene to Holocene pollen on the Sunda Shelf suggesting the presence of mangroves (with Sonneratia and Rhizophora), while also Pinus pollen was detected. Unfortunately, the exact location and age of the sediments cannot be determined from the papers abstract. In the northern South China Sea, high levels of the herb Artemisia may indicate that during the LGM at least parts of the exposed continental shelf of the northern South China Sea was occupied by grassland, dominated by this species. This would indicate an annual precipitation of between 300 and 500 mm and an average July temperature of around 1524 C (Sun & Li 1999; Sun et al. 2000). Interestingly these data suggest that in the northern part of the South China Sea frequent changes occurred in the character of the exposed shelf vegetation during glacial times (Sun & Li 1999; Sun et al. 2000). These changes were mostly from a relatively cool and humid climate to more drier and/or temperate conditions. During the drier and temperate conditions the shelf was covered by Artemisia, while other species indicate the presence of swamps or wetlands scattered on the shelf. Similar cycles were detected by Li (1997, in Sun & Li 1999) in the southern part of the South China Sea (at 610N, 11213E), where the pollen diagram shows alternative predominance of upper montane rain forest and of lowland rain forest with mangrove. Meijaard (2003, also see Appendix 1) provided further information on Late Pleistocene environments of the SCS area.

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Palaeoenvironments of southern mainland Asia Geology and palaeogeography of southern mainland Asia Thailand is located on what is considered to be a stable land area compared to adjacent regions in SE Asia (Dheeradilok 1995). The western coast of the Thai Peninsula has, however, been affected by crustal movement. The distinctive coastal landform with steep cliffs and short but steep-gradient river courses suggest uplift. This uplift resulted in the emergence of thick Tertiary coal beds and associated terrestrial fossils (Dheeradilok 1995). During the Oligocene and Miocene the ca. 30 intermontane basins of mainland Thailand and the Gulf of Thailand were occupied for long periods by lakes, as indicated by extensive lacustrine (or fluvio-lacustrine) deposits. By the end of the Miocene, many of these lakes were replaced by a fluviatile, erosional regime similar to that of the present time (Roberts & Jumnongthai 1999).

Figure 3.13. Main geological features of Indochina (after Steinshouer et al. 1997); for legend see Fig. 3.5.

The Quaternary deposits of the Lower Central Plain of Central Thailand represent a complex sequence of alluvial, fluvial, and deltaic sediments. About 2,000 m of

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Pleistocene and Holocene sediments were deposited in the basin, of which only the uppermost 300 are known (Sinsakul 2000). In the Late Pleistocene, the sea transgressed over the Lower Central Plain beyond Uthai Thani, and subsequently receded during the last glacial period (Dheeradilok 1995). There seems little doubt that high sea-levels occurred in Thailand during the Mid Holocene (ca 6 Kya) and that it reached as far land inward as 70 km north of Bangkok (Dheeradilok 1995; Woodroffe 2000). South of the Lower Central Plain, the Gulf of Thailand is a shallow epicontinental sea (maximum depth 86 m) that separates the Thai-Malay Peninsula to the west from the Indochina massif to the east. It is composed of a series of north-south trending ridges, which separate 12 major basins (Fig 3.13). These basins are divided by the Kro Kra Ridge into the Westerns Graben Area that contains 10 basins, and the Eastern Graben Area that contains the Pattani and Malay Basins (Watcharanantakul & Morley 2000). The Pattani Basin was probably temporarily emergent at 14 Mya due to low sealevels; tectonic uplift at 5 Mya, however, did not lead to its emergence (Pigott & Sattayarak 1993). Sedimentary units in the Thai basins are identical to those found in the Western Malay Basin (Metivier & Gaudemer 1999). In OligoceneEarly Miocene times, depositional sequences in the Pattani Basin are dominated by continental lacustrine, fluvial, and delta plain deposits. After this, marginal marine and fluvial deposits show increasing marine influence towards the Malay Basin, although according to Martens et al. (2000) fluviatile sedimentation in the Pattani Basin continued into the Pliocene. Tertiary sedimentary fill in the Gulf of Thailand (predominantly Neogene) varies in thickness from about 8,000 m in the deepest parts of the basins to less than 300 m over some of the basement highs (Highton et al. 1997). The Middle Miocene unconformity represents the onset of a regional marine transgression in the Malay Basin and a return to the delta plain environments during the Late Miocene in the Pattani Basin (Watcharanantakul & Morley 2000). Another unconformity at 10 Mya may have some tectonic significance, as it seems to mark the end of a major inversion in the Malay and W. Natuna Basins. However, it also coincided with a major sea-level low stand. Several million years may be absent in this unconformity (Morley et al. 2001). The Gulf of Thailand environment from the Late Miocene to recent was described as flood plains with more mangrove swamp

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and marine deposits in the upper part (Lian and Bradley, 1986 in Watcharanantakul & Morley 2000). During the Pliocene and Pleistocene, in the northern part of the Gulf near Bangkok, there were at least three major breaks in deposits, probably associated with sea-level changes. The first of these was deposited when the sea transgressed in the Pliocene (Dheeradilok & Kaewyana 1986). The thin sedimentary cover on the basement highs of the Gulf of Thailand (as little as 300 m) (Highton et al. 1997) suggests that these structures would have been emergent thoughout most of the Pleistocene and possibly Pliocene. If the Kro Kra Ridge was one of these structures, it would effectively divide the Gulf of Thailand into two seas. Also, the Ko Phangan Ridge (see Pigott & Sattayarak 1993), would add to the watershed division in the Gulf of Thailand. Between these two ridges lies the 250 km long and 50 km wide Kra Basin, which received more than 2.5 km of predominantly lacustrine Tertiary sediments (Pigott & Sattayarak 1993). These distinct valleys are also mentioned by Sawamura and Laming (1974), who recognized three sea floor valleys in the northern part of the Gulf of Thailand. My investigation of bathymetric maps (Hydrographic Department of Thailand 1978) showed that the most eastern valley, which appears to connect with the present Chao Phraya River, closely follows the eastern shore of the Gulf of Thailand. The western valley is less clearly defined, but seems to closely follow the western shore of the Gulf, thereby diverging from the eastern valley. The two seem to be separated by a shallower area, which could coincide with the Kro Kra Ridge as described by Watcharanantakul and Morley (2000) and Highton et al. (1997). It would be interesting to know whether the western river at an earlier time could have flowed west of the Kro Kra Ridge, across the low part of the Malay Peninsula near Krabi into the Andaman Sea. The ridge can be followed to N 1120 E 11045. The question is whether the two valleys and associated rivers would have merged downstream or whether they stayed separated during periods of low sea-level. In the latter case it becomes conceivable that the western river system would have crossed the present-day Malay/Thai Peninsula and flowed into the Indian Ocean. There is some support for this model. Garson et al. (1975) describe the Tertiary Krabi Series in the lowland areas north of Krabi. These deposits appear to fill shallow marine or lacustrine basins in a Tertiary landscape, and primarily lie unconformably on and between limestone sediments. The Krabi Series

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seem to consist primarily of deposits, which were tentatively dated at Middle-Tertiary or later. Ducrocq et al. (1995) estimated mammal fossil in these deposits to be of Late Eocene age, but considering that such fossils would most likely be found in lacustrine environments, it could well be that later marine deposits occur in the same Tertiary series. Interestingly, Garson and colleagues stated that the land area of the present-day Phangnga Bay was submerged in the Late Tertiary or Quaternary due to a relative sealevel rise, while also on Phuket Island small areas of marine sediments suggest that high eustatic sea-levels once inundated a much larger area than today. Overall, it therefore seems likely that this whole land area between Surat Thani and Krabi was low-lying during the Late Tertiary and Quaternary and that during periods of high sealevels at least parts became submerged. Whether this led to the development of a complete sea strait cutting the Thai-Malay Peninsula into two halves remains unclear, while there is also further need for supporting evidence regarding a river that flowed from central Thailand, west of the Koh Kra Ridge, past Surat Thani and into the Indian Ocean near Krabi. Of great importance to the biogeography of non-volant, terrestrial species in Indochina and China are the three main rivers, the Salween, Mekong, and Yangtze. The changes in these river courses and their effect on the regions biogeography were described by Meijaard and Groves (in press-b) (see Appendix 6). This river flow model is supported by river sedimentation calculations by Mtivier and Gaudemer (1999). They found that sedimentation in all SE Asian rivers, apart from the Mekong, Red River, and Chao Phraya has remained constant during the Quaternary. In the case of the Mekong River, the present-day load is much larger than the average filling rate over the last 2 Myr. Mtivier and Gaudemer suggest that the Mekong River discharged into the Gulf of Thailand or the Malay Basin before shifting its course to the Mekong Basin. There does, however, not seem to be much support for a direct connection between the Salween and Chao Phraya Rivers, as suggested by Attwood and Johnston (see above). For further discussion of this see Appendix 6. Vegetation of southern mainland Asia Toward the end of the Early Miocene, rising global temperatures and sea-levels corresponded with a change to predominantly moist forests in Indochina, and tropical

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and paratropical rain forests became established beyond the tropics. From this time, Dipterocarpaceae became prominent in Thailand (Watanasak, 1990 in Morley 2000) and alternating wet and dry climates were characteristic of Vietnam (Dzanh, 1994 in Morley 2000). Palynological study of coal deposits in northern Thailand confirms this change from a more temperate climate in the Oligocene to more tropical conditions in the Miocene. Between the Early and Middle Miocene these deposits suggest a mixed environment of lakes and forest swamps, with Pinus and Florschuetzia trilobata possibly indicating some seasonality (although coal formation would suggest mostly everwet conditions) (for detail see Figure 1 in Ratanasthien et al. 1999). Similarly, palynological studies of coals in the Krabi Basin, southern Thailand, suggest wet conditions and also proximity to mangroves (Watanasak et al. 1995). The Middle Miocene vegetation of the Pong area in eastern Thailand suggests a bushy mangrove environment (Vozenin-Serra et al. 1989), which would indicate that the coast was considerably further inland than today. However, Ducrocq et al. (1994) reported that neither the vertebrate, nor invertebrate fossils indicate a mangrove environment during the Middle Miocene in the named area. Instead, they suggested that between 16 and 14 Mya, a stable palaeoenvironment existed, with wet and warm characteristics. Cenograms3 of the Middle Miocene faunas suggest a quite open habitat, or possibly small areas of forest intermixed with grassland. These faunal structures indicate a tropical climate characterized by an alternation of dry and rainy seasons. The cenograms exclude the possibility of environments dominated by either closed forest or steppe (Ducrocq et al. 1994). The Miocene palaeoenvironments, further south in the Pattani Basin consisted of low-lying swamps (in a subsiding basin) dissected by rivers (Jardine 1997). Based on an ecological shift from C3 to C4-dominated vegetation types in central Asia, Quade et al. (1989) described major climatic changes that occurred towards the end of the Miocene (C3 plants include all trees, shrubs and herbs, and grasses favoured by a cool growing season, whereas C4 plants include grasses favoured by a warm growing season). From at least 18 until 7.47.0 Mya, the northern Pakistan floodplain was dominated by closed canopy forest, with or without an understory of C3 shrubs and

3 Cenograms are graphs of body weight frequencies, which, for mammals, may give some indication of the kind of vegetation type in which a community occurred.

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herbs, while early Middle Miocene (16 Myr) Siwalik deposits in Himachal Pradesh clearly indicate the presence of evergreen tropical rainforest of Malesian affinity (Morley 2000). Towards the end of the Miocene, evergreen forest still existed in Nepal, but after that time they were gradually replaced by deciduous forests (Morley 2000). Similarly, coal-bearing sediments in northern Vietnam considered to be of Late Miocene age are indicative of a humid, subtropical climate (Covert et al. 2001). Between 7.47.0 Myr and 5 Myr, a vegetation mosaic of grasslands and forest was probably present in the Himalayan foothills (Quade et al. 1989; Awasthi, 1992 in R.J. Morley 2000). After 5.0 Mya, up until 0.4 Mya, grasses by far dominated the vegetation, possibly interspersed with riparian habitats in which some C3 trees and shrubs grew. Drier Pliocene climates in Rajasthan are shown by the presence of fossil woods, which indicate deciduous forest at that time (Guleria, 1992 in Morley 2000). Elsewhere, palaeobotanical data from India and from Myanmar indicate that a rich tropical to subtropical vegetation covered the region ca. 5 Mya under a prevailing warm humid climate (several authors in Poole & Davies 2001). However, during the Pliocene, the increasingly arid climate, engendered by the rising Himalayas, caused a consequent change in the vegetation of this region (Poole & Davies 2001). During the Pliocene, an alpine flora would have occupied the higher ranges of the Himalayas and the plateaux of Yunnan and North Burma, while during the cold stages of the Pleistocene, these areas were covered by glaciers (Kingdon Ward 1939). Quade et al. (1989) speculated on the cause of the vegetation change around 7.4 Mya, as it remains unclear whether this happened due to climatic changes or because of the evolutionary first appearance of C4 plants. They do, however, suggest that a strong intensification of the monsoon system at 7.47.0 Mya played an important role. The vegetation changes are also reflected in faunal turnovers: browsers are replaced by grazers; rodents show considerable turn-over; and Sivapithecus (a possible ancestor of the Orang-utan) disappears from the record (Barry et al. 1985; Quade et al. 1989; 1991; 2002). These palaeoenvironmental changes are also reflected in north Thailand, where the Li Basin contains Oligocene and Early Miocene coal (C.K. Morley et al. 2000), indicating a wet and warm, fluvial or lacustrine environment. Coals in Middle Miocene sediments still indicate a peat swamp, fluvial or lacustrine environment, but in the Late Miocene, Pliocene and Quaternary sediments no coal was found (C.K.

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Morley et al. 2000), possibly indicating a climate that was either too cool or dry for coal formation. Mammalian fossil assemblages in Yunnan Province (also see Chapter 4.1) indicate that between Early and Middle Pliocene a mixed forest, and open bush-grassland vegetation existed, with several carnivores typical of forested areas, and ungulate species suggesting more open, grassy vegetation (Pan 1993). Slightly wetter conditions seem to have prevailed further east and southeast of Yunnan. Reconstructions of the climate and vegetation of this area in the Middle Pliocene (3 Mya) indicate that there was a considerable expansion of evergreen forest in southern China, whereas Indochina was mostly covered by rainforest, possibly with patches of deciduous forest in the area of present-day Burma (Dowsett et al. 1994). Based on rodent fossils distributions, Chaimanee (1998) reconstructed Late Pliocene Early Pleistocene palaeoenvironments for several locations in Thailand. Khao Samngam (9942 E; 1327 N) probably had an environment with some mixed vegetation with grasslands in the floodplain and forests on the surrounding limestone hills. At 1.961.79 Myr, Longgupo, in South China, 15 latitude north of Khao Samngam had a forest community, but of different vegetation composition than that in Thailand. Early Pleistoceneearly Middle Pleistocene fossil communities indicate a typical forest faunal assemblage, probably of a dry evergreen type with some open patches. At this time, the boundary between the Indochinese and Sundaic faunistic subregions may have been about 500 km south of the present Kra boundary [Note that this line further south would approximately have been in the same location as the Kangar-Pattani floristic boundary (Whitmore 1984), which separates Indochina from Malesia]. This situation occurs also in some more recent, probably Middle Pleistocene, localities from Peninsular Thailand, when several Sundaic endemic taxa moved north, with some typical Indochinese forms moving south. Chaimanee (1998) suggested that these localities (at 827 N) are indicative of dense evergreen forest with no indication of grassland, and they probably correspond to an interglacial stage of the earliest part of the Middle Pleistocene. The Snake Cave deposits in northeast Thailand (1630 N; 10149 E), which were dated as middlelate Middle Pleistocene (minimum age 169 Kyr), suggest cooler climatic conditions than today, probably with

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dry evergreen forest with some clearings with grasses or bamboo. The present-day vegetation in this area is dense evergreen forest (F. Blasco, pers. comm. in Chaimanee 1998). According to Dongsheng and Menglin (1984), Indochina and southern China were still covered by humid rainforest at the Pliocene-Pleistocene boundary, although this rainforest zone was progressively pushed south during the Pleistocene until no rainforest remained in China. By the Middle Pleistocene, the subtropical and tropical zones had shifted south- and eastward, and by the Late Pleistocene, these zones had migrated even farther southward and were considerable reduced in area (Jablonski & Whitfort 1999). Evidence for grasslands is very limited through most of the Pliocene, but subsequently shows a marked increase, together with charred grass cuticle, during the Early Pleistocene, indicating the expansion of savannah vegetation, which was subject to burning (Morley 1999). Thick laterites of PlioceneEarly Pleistocene age in the Lower Central Plain of Thailand (Thiramongkol 1986) may also indicate seasonality in a generally humid tropical climate (Whittow 1984). In Yunnan Province, towards the Late Pliocene, the fossil assemblages indicate a forest and woodland environment, while also appearing to reflect a period of rapid faunal and environmental change. In the Early Pleistocene, the fauna indicates a more open, grassland-bush environment, whilst pollen analysis suggests a fairly cool subtropical climate. Finally, towards the end of the Pleistocene most likely a grassland-dominated environment existed (Pan 1993). The MiddleLate Pleistocene in the eastern parts of SE Asia led to some considerable changes in vegetation distribution. For instance, Zheng and Lei (1999) provide palaeoenvironmental data for the Leizhou Peninsula, north of Hainan Island, for the last 400 Kyr. Between 400 and 340 Kya, pollen data indicate slightly drier and cooler conditions compared to today with an increase of montane forest formation. Between 340 and 280 Kya this was replaced by a wetter and warmer climate with predominant evergreen oak forest. Between 280 and 240 Kya, mean annual temperatures were more than 4C lower than today, which led to a substantial increase in typical montane conifer forest elements. Between 240 and 180 Kya, significant warming occurred and montane elements were replaced by fagaceous evergreen forest, while tropical

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lowland rainforest taxa increased significantly. Between 180 and 125 Kya, vegetation changes in tropical China were less drastic and only the coldest glacial stadials affected the local forest. The inter-glacial stage between 125 and 65 Kya, was accompanied by a warmer climate and dense forest, which, between 65 and 29 Kya, was followed by a drop in temperature of between 5 and 6C. The period between 29 and 15 Kya was the first during the whole study period in which the Pleistocene dense forests were transformed to grassland or savanna vegetation. Furthermore, the dominance of Poaceae and Artemisia implies not only a cooler, but also a much drier climate. After 15 Kya the pollen assemblage shows the resurgence of montane forest, and the disappearance of savanna formation (Zheng & Lei 1999). Zheng and Lei (1999) also found that a fundamental change from densely forested formation to savanna did not occur until the LGM. The replacement by grassland in southeastern China during the LGM could be a result of extremely dry conditions caused by the increased continentality in southern China and SE Asia, the reduction of the South China Sea, as well as the lowering of sea surface temperatures. As Zheng and Lei showed that precipitation during earlier glacial-interglacial cycles may have been very stable, and that even during the cooler period wet conditions prevailed, it can be hypothesized that the South China Sea area did not become subaerial in any other periods during the last 400 Kyr. Only, during the LGM did the exposure of the South China Sea area lead to both dry and cold conditions, which caused forest formations to change to savanna. Ferguson (1993) reviewed the Late Pleistocene environmental changes for south and southwest China. Judging from the vegetational changes in the Late Pleistocene cold phases, the mean annual temperature in south and southwest China was normally no more than 23C lower than at present (Luo, 1991c; Tong and Shao, 1991; Wu, 1991, all in Ferguson 1993). Probably the only exception was the extremely cold and arid phase at the end of the Pleistocene (2015 Kyr). At that time, the vegetation in the extreme south of China (2123 N) resembled that now growing in the Changjiang region, indicating a fall in the mean temperature by some 6C. In the interstadials, warm temperate taxa were replaced by tropical elements. In Jiangxi Province, centralsoutheast China, the Late Pleistocene climate (between 12.8 and 10.5 Kya) was similarly drier and cooler than todays, and the subtropical, mixed deciduous-

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evergreen broad-leaved forest, which is now in the area, was reduced and herbaceous cover expanded (Jiang & Piperno 1999). Data by Liu et al. (1986) suggested that Yunnan, at a time approaching or including the LGM (ca. 3620 Kyr), experienced greater winter humidity and rainfall, while mean annual temperatures were only slightly, if at all, lower than present. Yunnan may therefore have represented a Chinese tropical refugium. Other such extensions of the tropical belt in China during the LGM were suggested for Guangxi and Guangdong, although this remains contested (see Ferguson 1993). Overall, mainland SE Asia appears to have been cooler and more seasonal than today during much of the Late Pleistocene. For instance, between 62 and 28 Kya and between 28 and 19 Kya, the reworking and redeposition of aeolian sands along the southeastern Vietnam coast points to reduced vegetation cover and landscape instability in this area (Murray-Wallace et al. 2002). Also, Nutalaya et al. (1986) described loess deposits in the Khon Kaen district near the Mun River and also in North Thailand. Again, these indicate dry climates and devegetation, which possibly occurred during the LGM and/or the penultimate glaciation at 130 Kya. Further to the north in north-east Thailand (17 N, 103 E), the pollen assemblages in the Late Pleistocene/Early Holocene (1610 Kya) suggest xeric, species poor, and strongly seasonal vegetation, with seasonally inundated floodplains or stream margins (Kealhofer & Penny 1998). Follow-up work by Penny (2001) described palaeoenvironments from the same region in north-east Thailand from 40 Kya. Between 40 and 10 Kya the vegetation in this area, consisting of FagaceousConiferous forest, is remarkably stable, with clear evidence of lower temperatures and possible geomorphic evidence of significant drying in that region. At the start of the Holocene, at ca. 12 Kya, a rapid transition occurred from the pine/oak forests to tropical broad-leaf taxa. Finally, after the LGM, climatic conditions rapidly improved in much of SE Asia. In Thailand, along the eastern coast of the peninsula, peat deposits and lateritic layers that were dated as post-LGM, but before 912 Kya, suggest a warm and wet climate in that period (Dheeradilok 1995). Meanwhile, in eastern Cambodia, the climate was still relatively cool and dry at 9.3 Kya with vegetation being at least partly semi-evergreen,

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but of a drier type than present. Also, open forest with grasses was common at this time. At ca. 8.4 Kya grasses disappeared, and other signs indicate an abrupt transition to a warmer, more humid climate (Maxwell 2001). This resembles patterns described for Sichuan and, possibly, Yunnan (Sun et al., 1986; Li and Liu, 1988; Jarvis, 1993, all in Maxwell 2001), although the dry late glacial period may have lasted longer in Indochina than in southwest China. A possible reason for this may have been that the source of Indochinas monsoon precipitation lies in the Gulf of Thailand, which even by 8 Kya was still 20 m lower than today, and large areas were therefore still dry. At ca. 5.3 Kya, changes in pollen indicate a climatic shift to drier conditions, which lasted until ca. 2.5 Kya with a recovery of the monsoonal strength. The Palaeogeography of Sulawesi, Nusa Tenggara, and the Philippines Zoogeographic evolution in island SE Asia cannot be investigated without observing the environmental and geological changes that have occurred in Sulawesi, the lesser Sundas, and the Philippines. Many mammalian taxa now found in the latter areas are closely related to Sundaland forms, although it still remains largely unclear how they migrated between those areas. The Borneo-Palawan-Calamian-Mindoro link seems to have been one migration route, while another one may have followed the Sulu Archipelago, connecting Borneo and Mindanao. It remains unclear whether mammals also migrated from north Sulawesi, across the Sangihe Islands to Mindanao. The biogeography of these areas is outside the scope of this thesis, but some aspects will be highlighted below, as they are important to the discussion of some species groups studied here, e.g. the Sus barbatus group (Appendix 8), mouse-deer (Tragulus sp., Appendix 5), and Cervinae (Appendix 7). One of the key questions regarding mammalian biogeography in Sulawesi is whether at any time in the Late Tertiary or Quaternary there was a land connection between this island and Sundaland. Groves (1976) suggested that Sulawesi and Java were once connected, as did Hooijer (1975) who postulated the existence of what he named Stegoland. A similar idea was proposed by P. and F. Sarasin (in Weber 1902) who followed and measured a ridge from Madura to the Kangean and Sabalana Islands, and on to the southern-western tip of Sulawesi, and suggested that this once provided a land link between Java and Sulawesi. More recent data, however, suggest that

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Sulawesi has had no land connections with either Java or Borneo since the Eocene. It is indeed now generally accepted that in the Middle Eocene, the southwest arm of Sulawesi formed part of a low-lying swampy area along the south-eastern margin of Sundaland. Part of this arm became submerged from the Late Eocene to Middle Miocene, but the presence of a continuous pollen record in clastic sediments indicates that some areas remained emergent when the arm drifted away from Sundaland, during the opening of the Makassar Straits (Morley 2000). The north arm of Sulawesi was nearly in its present-day orientation by the end of the Early Miocene (Lee & Lawver 1995), while East and West Sulawesi collided sometime during the Middle Miocene (Audley-Charles et al., 1988 in Lee & Lawver 1995). Wilson and Moss (1999) suggestion that western Sulawesi was connected to Borneo and emergent up until the Eocene cannot explain the presence of recent Sulawesi faunas. They do, however, suggest that island hopping routes may have existed along volcanic arcs, such as the long-lived arc along the north arm of Sulawesi and the Cagayan and Sangihe arcs, and later along the younger Sulu arc. Furthermore, they concluded that the uplift and subsequent erosion of Borneo from the Late Oligocene to the present day and the emergence of more extensive land areas in Sulawesi led to the progressive reduction in width of the Makassar Straits. This may have facilitated interchange of biota in the Late Tertiary, especially during times of low sea-level. Data provided by Guntoro (1999) suggested that the Straits were at their shallowest during the Early Miocene, and between mid-Miocene and recent times as then shallow marine limestones were deposited. Also, Pulau Laut was once much larger (see Borneo section) providing another stepping stone between Borneo and Sulawesi. The age of the volcanic islands in the inner Banda Arc (consisting of the islands in a chain along Bali, Lombok, Sumbawa, Flores and Banda) is not known accurately, but Burrett et al. (in Schmitt et al. 1995) indicated that they are post-Miocene, and some (e.g. Bali) may be only about 3 Myr old. Similarly, Lombok and Sumbawa were elevated only in the relatively recent past (Hall 2002). Komodo is the exception among the mainly Tertiary volcanic islands, with a history dating back to the Late Mesozoic (Auffenberg, 1980 in Monk et al. 1997). At one time, Flores was also thought to be a geologically old island that existed before the Miocene, but it is now

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thought to have its origins in the Mio-Pliocene (Monk et al. 1997). Table 3.2 shows the time since several of the islands of Nusa Tenggara have been last emergent, although it should be realized that many of these islands have complex histories and may have been emergent and subsequently submerged before their latest emergence. They could therefore have provided stepping-stones for dispersing species (Monk et al. 1997). The exact geographical arrangements of the land masses during the Pleistocene glacial maxima are uncertain, partly because the available bathymetric maps lack detail (Schmitt et al. 1995). Heaney (1991, in Schmitt et al. 1995) estimated that, during periods of low sea-level, the present islands formed six islands: Greater Sumbawa (Lombok, Sumbawa and Moyo); Sangeang; Greater Flores (Komodo, Flores, Adonara, and Lembata); Pantar; Alor; and Sumba. There may, however, have been additional ridges connecting some of these islands (Schmitt et al. 1995). Island Sumbawa Atauro Sumba Timor Tanimbar Kai Buru Seram Age or epoch of emergence Plio-Pleistocene Pliocene 1 Mya Quaternary Quaternary Quaternary Plio-Pleistocene Pliocene

Table 3.2. Nusa Tenggara islands and the time when they became last emergent (after Monk et al. 1997).

The Philippine archipelago first appeared as dry land during the late Eocene or early Oligocene (Peterson & Heaney 1993). At about 25 Mya, the Philippines were part of an arc that connected them with Sulawesi and Halmahera. This arc created a discontinuous land connection between these island areas (Hall 1998). Between 15 and 5 Mya, the deep marginal basin of the South China Sea was eliminated and in the Early Pliocene an area of shallow sea run from the northern end of the Philippines to the south coast of present-day China (Hall 1998). Morley (2000) suggested that the Palawan microcontinent that rafted from south China to its present position may have acted as a dry-land raft for mainland Asian species. It should be noted, however, that sediments on Palawan appear to indicate that Palawan was submerged for much of its

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rafting time (Hall 1996). Still, Holloway (1982) described late Middle Miocene unconformities in Palawan and in the neighbouring areas of the Sulu Sea and offshore west and north-west Palawan. These, possibly correlated, unconformities indicate a phase of uplift at that time, resulting in emergence and non-deposition. None of the other parts of the Philippines had any land connection to Sundaland. Also, volcanic arc activity occurred along the Cagayan and Sulu ridges during the Miocene and PlioPleistocene, respectively, and probably resulted in emergent chains of islands (Rangin and Silver, 1991 in Wilson & Moss 1999); these may have been times when migration from Borneo to Mindanao via these arcs was possible, although a continuous land connection probably never existed. Most of the growth of the Philippine islands has occurred since the beginning of the Miocene, and much since the Pliocene. Because the islands are on a shallow platform, they were heavily influenced by repeated Pleistocene changes in sea-level (Heaney 1985). Heaney (1985) reviewed the distribution of many animal species in the Philippines. Based on this, he concluded that there has been no land bridge to the Philippines from Asia during the Late or Middle Pleistocene, with the exception of a Middle Pleistocene connection from Borneo to Palawan. Reis et al. (2001), however, suggested that this Pleistocene land connection did not exist, but a narrow water gap instead, which prevented some species from colonizing. According to others (e.g., Earl of Cranbrook 2000), Palawan was connected to northern Borneo during the LGM and only became separated at about 14 Kya. Heaney (1985) found some evidence from the primary fresh-water fish (those that are never found outside freshwater habitats) for the existence of a land-bridge from Palawan to Mindanao during the Early Pleistocene or (more likely) the Pliocene, although this remains somewhat speculative. During Late Pleistocene climatic fluctuations, the islands of Luzon and Catanduanes merged to form Greater Luzon, while Leyte and Biliran merged to form Greater Mindanao (Peterson & Heaney 1993). In a paper on the stratigraphy of Palawan, Wolfart et al. (1986) provided evidence for the Pliocene inundation of parts of the southern end of Palawan. The Pusok Conglomerate found in the eastern side of southern Palawan is thought to be of Pliocene age, has a thickness of some 100 m, and overlies pre-Tertiary rocks

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transgressively, suggesting submergence during a sea level highstand. This is further supported by Pliocene limestones that were found at the southern edge of Palawan, southeast of Canipaan. These deposits are unconformably overlaying Miocene sediments in much of southern and central Palawan, which suggests that, in the Late Miocene or Early Pliocene, this area was emergent, possibly providing dispersal opportunities for terrestrial mammals. Southern and central Palawanat least their coastal areasbecame emergent on the Pliocene-Pleistocene boundary (Wolfart et al. 1986). Northern Palawan, on the other hand, probably became emergent in the Early Miocene and was not or only to a slight extent submerged during any of the later high stands. South-west of Palawan, parts of Balabac Island also have Pliocene sediments, while raised beaches and drowned valleys suggest that during high sea levels parts of the island became submerged (John 1963). Summary of palaeoenvironmental and palaeogeographical reconstructions Above, I have provided detailed information on the changes in land/sea distribution and the most important vegetation types in island SE Asia since the Early Miocene. In the next section, I have mapped these changes and these maps will be used to see whether mammalian evolution can be linked to the most important palaeoenvironmental and palaeogeographical changes. For that purpose I will provide each map with a code which will be used later to refer phylogenetic events to these coded palaeogeographies: 1. Early Miocene (MIO 1) 2. Middle Miocene (MIO 2) 3. Middle to early Late Miocene (MIO 3) 4. Late Miocene (MIO 4) 5. Early Pliocene lowstand (PLIO 1) 6. Early to Middle Pliocene highstand (PLIO 2) 7. Middle Pliocene (PLIO 3) 8. Late Pliocene (PLIO 4) 9. Early Pleistocene (PLEI 1)

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10. Early to Middle Pleistocene (PLEI 2) 11. Early to Middle Pleistocene highstand (PLEI 3) 12. Middle Pleistocene lowstand (PLEI 4) 13. Late Pleistocene (PLEI 5) In summary, during the Early Miocene, Sundaland existed as an extension of the Asian mainland, and primarily covered the western part of Borneo, the Sunda Shelf and the Malay Peninsula; at that time a warm and wet climate existed. During the Miocene, the land areas of Java and Sumatra slowly started to shape up, initially as a chain of small volcanic islands. Much of the land area was flooded during the Middle Miocene highstand. Towards the end of the Miocene, when global climatic conditions had cooled and become drier, more land was exposed, leading to connections of parts of Sumatra and Java to the rest of the Sundaic land mass. Vegetation became increasingly open, and grasses started to become more common. During the Pliocene, the sea between the Malay Peninsula and Borneo started to open, eventually leading to the latter becoming an island. At that time the climate was becoming increasingly dry and environmental fluctuations increased. It appears that an Early-Middle Pliocene sea level high caused a break-up across the Malay/Thai Peninsula which could have lasted for as long as 1 Myr. During the Pleistocene, Java and Sumatra started to take on their present-day shape. The considerable fluctuations in sea level at that time led to alternating connections and disconnections between the different islands of this region, but the exact sequence of this remains unclear. It appears that during a Middle Pleistocene glacial Java was directly connected to the Malay Peninsula by a land bridge that followed the Riau and Lingga area, Bangka and Belitung, and reached Java via the Karimun Jawa area. During the Pleistocene glacials, tropical wet evergreen rainforest probably became restricted to refugia, as suggested by the vegetation patterns that existed during the LGM.

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3.5 PALAEOGEOGRAPHICAL RECONSTRUCTIONS

Early Miocene (MIO 1) During the Early Miocene, the islands of Sumatra and Java did not exist as we know them today. Instead several smaller islands were in place, none of which probably connected to Sundaland. At this time, it appears that northern Borneo with the Kuching High was separated from the rest of Sundaland. The location of upland areas (see Fig. 3.14) outside the area of present-day Borneo is largely hypothetical. The presence of marine influences in the Malay and Thai Basins suggested that the eastern coastline of Sundaland was located considerably further west than suggested by Hall (1998). This was also suggested in the palaeoenvironmental maps by Jinmin (1994). Pollen findings suggest a markedly seasonal climate in Borneo and Sumatra. The key biogeographic issues are that Borneo was part of Sundaland, while parts of Sumatra were possibly connected. Most of Java had probably not yet emerged.

Figure 3.14. Palaeoenvironmental reconstruction for the Early Miocene. The grey areas show the land area at this time in relation to the present-day shape of the region; black areas are uplands, and the dotted lines are rivers.

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Middle Miocene (MIO 2) The EarlyMiddle Miocene saw the re-expansion of tropical and sub-tropical forest zones as the climate became warmer, and wet forest types such as peat swamps developed in Sundaland. This was also a time of generally high sea-levels. Sundaland is still one landmass and connected to the Asian mainland. Sumatra probably consisted of an island arc, while Java was still mostly submerged.

Figure 3.15. Palaeoenvironmental map for the early Middle Miocene (for legend see Fig. 3.14)

An important period of low sea-levels occurred between 16.5 and 16.2 Mya, which may have led to a brief reconnection of Sumatra with Sundaland. However, the further sea-level rise during the EarlyMiddle Miocene resulted in inundation of most of northern Sumatra, which suggests that no terrestrial mammals survived in that area until later in the Miocene. At 16 Mya, however, a faunal turn-over occurred in southern Asia (Barry et al. 1985), and this may have resulted in an influx of new mammal taxa into Sundaland. Figure 3.15 shows the palaeoenvironmental map at the height of the early Middle Miocene climatic optimum.

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Middle to early Late Miocene (MIO 3) Compared to the palaeoenvironmental reconstructions by Hall (1996; 1998), the data in this research suggest less emergent land in Java and on Sumatra, which are here shown as a chain of several islands rather than the two larger land masses of protoJava and proto-Sumatra shown by Hall. In Fig. 3.16 I hypothesize that the Lampung High in southern Sumatra provided a link between the Sumatran land mass and Sundaland. The climate would have been mostly hot and humid and a tropical evergreen vegetation would probably have covered most of Sundaland. The river courses are largely hypothetical, although it could be that the Mae Khlong and Chao Phraya/Mekong Rivers were separated by the upland area of the the Koh Kra Ridge. A large island, in the present-day Pulau Laut region, may have existed south-east of Borneo.

Figure 3.16. Palaeoenvironmental reconstruction for the late Middle to early Late Miocene (for legend see Fig. 3.14).

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Late Miocene (MIO 4) Similarly to the preceding reconstruction, it was still found that only small parts of Java had emerged in this period, as opposed to the complete emergence of Java suggested in Halls (1996; 1998) reconstruction. West Java was probably a small land area now, connected to the Lampung High, while in the central Java area some small islands may have occurred that were however flooded again later in the Pliocene and did therefore not play a role in Tertiary mammal evolution. Towards the Late Miocene, the climate in mainland Asia was becoming drier and cooler, although it is unclear whether that significantly changed the vegetation of Sundaland. Judging from the vegetation for Indochina, it is likely that Sundaland remained largely covered in tropical evergreen forest.

Figure 3.17. Palaeoenvironmental reconstruction for the Late Miocene (for legend see Fig. 3.14).

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Early Pliocene lowstand (PLIO 1) In the Early Pliocene, the prograding southern Sunda Shelf edge ran quite close to the Java area, but it is unclear whether the Sundaland area actually connected with the exposed parts of Java. In Fig. 3.18 I have still drawn a connection between Sundaland. southern Sumatra, and west Java, but this is also the time when the Sunda Dome collapsed which probably led to the break-up of this connection. The further deepening of the Malay and Thai Basins probably also led to the opening up of the sea between Borneo and the Malay Peninsula, although the timing of this is poorly known. Similarly, I have drawn a connection between north Sumatra and the Malay Peninsula via the Asahan Arch, but again the timing of the submergence of the arch is unclear.

Figure 3.18. Palaeoenvironmental reconstruction for the Early Pliocene lowstand (for legend see Fig. 3.14).

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Early to Middle Pliocene highstand (PLIO 2) The EarlyMiddle Pliocene sea level highstand (ca. 4.53.5 Mya) (e.g. Haq et al. 1987; McNeill et al. 1996; McNeill et al. 1998), which Haq et al. estimated at ca. 100 m above present-day sea levels, could have significantly changed the geography of the region (see Woodruff 2003). It is possible that it cut the Malay/Thai Peninsula in two, while it is also possible that Borneo was separated, or almost separated into two parts when the valleys of the Kapuas and Mahakam Rivers became sea arms within close reach of each other. It was probably also at this stage that the land connection between Borneo and the Asian mainland was first broken, or at least narrowed down to a small land corridor. This is likely to have happened somewhere along the Riau/Lingga/Belitung/Bangka ridge. The high sea levels suggest warm and wet conditions, although no detailed vegetation information is available for this period.

Figure 3.19. Palaeoenvironmental reconstruction for the EarlyMiddle Pliocene highstand (for legend see Fig. 3.14).

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Middle Pliocene (PLIO 3) During the Middle Pliocene, sea levels remained relatively high and I hypothesize that at that time the connection between Borneo and the Malay Peninsula was severed (but exact information on this is lacking). This would have been the first time that Borneo became an island (unless it happened earlier during the Pliocene), which would have led to the divergence of Bornean species. Also the Sumatra and Javan islands were isolated.

Figure 3.20. Palaeoenvironmental reconstruction for the Middle Pliocene (for legend see Fig. 3.14).

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Late Pliocene (PLIO 4) The Late Pliocene glacial led to very low sea levels and cool and dry conditions. I think that at that time Borneo became reconnected to the rest of Sundaland (as sea levels were at a similarly low level as during the LGM). I have also drawn connections between Sumatra and the Malay Peninsula. At this time it is likely that evergreen rainforest became restricted to refugia although their locations remain

unknown.
Figure 3.21. Palaeoenvironmental reconstruction for the Late Pliocene (for legend see Fig. 3.14).

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Early Pleistocene (PLEI 1) During the Early Pleistocene, sea levels were generally high which presumably led to the fragmentation of the large area that resulted from the lowstand between 2.8 and 2.4 Mya. Data for this period are tentative, however.

Figure 3.22. Palaeoenvironmental reconstruction for the Early Pleistocene (for legend see Fig. 3.14).

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EarlyMiddle Pleistocene (PLEI 2) The arrival on Java of the Ci Saat Fauna at this time suggests that a land bridge or an arc of small islands connected central Java to the Asian mainland. This could either have been a direct connection between Bangka, Belitung, and Karimun Jawa, or one via the southwest of Borneo (see Figs. 5.8 and 5.9).

Figure 3.23. Palaeoenvironmental reconstruction for the EarlyMiddle Pleistocene (for legend see Fig. 3.14); see Fig. 5.8 for an alternative scenario.

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EarlyMiddle Pleistocene highstand (PLEI 3) The landbridge described on the previous page (via the Karimun Jawa Islands) was probably low-lying and would have been affected by sea level changes and the general subsidence of the Java Sea area. Intermittently it would have provided dispersal opportunities for species from the Asian mainland to Java and vice versa.

Figure 3.24. Palaeoenvironmental reconstruction for the EarlyMiddle Pleistocene highstand (for legend see Fig. 3.14).

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Middle Pleistocene lowstand (PLEI 4) The Middle Pleistocene lowstand occurred during one of the more severe glacials of the Pleistocene. It led to the dispersal of many new species to Java, which at that time was presumably connected to the Asian mainland. Java had quite an open environment at that time, which was probably also found in south-eastern Borneo, and possibly on the central Sunda Shelf.

Figure 3.25. Palaeoenvironmental reconstruction for the Middle Pleistocene lowstand (for legend see Fig. 3.14).

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Late Pleistocene (PLEI 5) The palaeogeography of the LGM is reasonably well known. At this time, Sundaland was connected into one landmass, probably the largest it had ever been. This area was dissected by some very large rivers of which the courses are known with considerable accuracy. Tropical evergreen rainforest did probably not occur anywhere on Java (apart maybe from some mountain areas) and may also have been absent from southeastern Borneo, the central Sunda Shelf and from parts of the Malay Peninsula. In these areas, the evergreen forest was probably replaced by more open deciduous forest types or wooded grasslands. Tropical evergreen forest remained on Sumatras west coast and the Mentawai Islands, around the Natuna area, and around the Bornean mountains.

Figure 3.26. Palaeoenvironmental reconstruction for the Late Pleistocene lowstand (for legend see Fig. 3.14).

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