in deep organic sediments from the bottom of swamps or lakes. The oldest-known Sulawesi pollen is of a relative of Sonneratia mangrove trees from Tertiary rocks in north Central Sulawesi (Sohma 1973).
Attention of palaeobotanists on Sulawesi has been concentrated on two areas: Lake Tempe and in topographic depressions in ultrabasic rocks around Lake Matano. One particularly long core of possibly considerable age has been collected recently from one of the latter sites but results will not be available for some while (Hope 1986).
In the Lake Tempe region cores were taken from the swampy Lake Rawa Lampulung about 4 km east of Sengkang and the pollen shows that at least part of the surrounding area was covered by mangrove vegetation, that is, it was inundated by the sea, from about 7,100 to 2,600 years ago. The rise in sea-level needed to produce this effect is about 5 m which matches the palaeo-climate information (p. 21). A core from the edge of Lake Tempe, 5 km northwest of Sengkang, reveals that at least from 4,400 years ago the area was dominated by freshwater vegetation and that the sea was probably prevented from reaching the lake by a squat molasse ridge to the east.
The history of the vegetation during the Quaternary and late Tertiary is very closely linked with the climatic changes. During the drier periods of the Pleistocene the area of seasonal forest would have extended while the area of rain forest became less. Thus the populations of rain forest species would have been reduced but isolated populations of certain species probably maintained some level of contact or gene flow along riverine areas or wetter soils. In the driest parts of Sulawesi it is quite likely that monsoon or even savannah forest predominated.
Many mountain plants are unable to live successfully in the lowlands, but the lowering of zones on mountains during the cool periods gave opportunities for these plants to spread because the available stepping-stones of suitable habitat increased in area and number.
Present Vegetation
Fewer botanical specimens have been collected on Sulawesi than on any other major island/region in Indonesia. To date only about 23 specimens per 100 km2 have been placed in herbaria whereas over 200 per 100 km2 are known from Java. A density of 100 specimens per 100 km2 would represent an adequately-known flora. Allowing for this, the number of higher plant species10 may be about 5,000. Only seven genera are known to be endemic compared with 17 in Sumatra, 59 in Borneo and 124 in New Guinea (E. de Vogel pers. comm.). In addition, some species are barely known: a forest shrub Thottea celebica (Aris.), for example, has been collected only once, from Lambarese, northeast of Palopo (fig. 1.19) (Ding Hou 1984).
The natural vegetation11 growing in a particular area is dependent on various factors such as soil chemistry, soil water, climate, altitude, distance from the sea, and distance from areas of similar conditions (table 1.5; fig. 1.20).
The coasts of Sulawesi are fringed by coral reefs, mudflats, mangrove forests and rocky or sandy beaches (chapters 2 and 3). The freshwater habitats are generally rather nutrient-poor (although there are striking exceptions) and as a result freshwater vegetation is not well developed (chapter 4). The lowland and hill forests of Sulawesi (chapter 5) have the most tree species of all forest types (table 1.5) but have only seven species of dipterocarp trees, the mainstay of forestry operations in Borneo and Sumatra where there are 267 and 106 species respectively (Ashton 1982). The major trees of commerce are the tall Agathis (Arau.)12 trees with broad, flat leaves; the magnificent yellow-flowered legume Pterocarpus indicus (Legu.) (huge pollarded specimens of which are commonly seen around town squares such as Ujung Pandang) which is deciduous and found most commonly in the more seasonal areas; the gum tree Eucalyptus deglupta (Myrt.), usually found wild in riverine habitats and which is extensively used in reforestation projects; beremban Duabanga mollucana (Sonn.), and gutta percha Palaquium spp. (Sapo.). The hemi-parasitic13 sandalwood Santalum album (Sant.), from the heartwood of which sandal wood oil is extracted, used to be found in the dry Palu valley (chapter 6) but even in the Poboya reserve set up to protect one of the last stands of this tree, only 62 small individuals exist (Sidiyasa and Tantra 1984).
Figure 1.19. Thottea celebica leaves and flower; one of a number of endemic plants known from just one specimen. Scale bars indicate 1 cm.
After Ding Hou 1984
Based on van Steenis 1950; Whitmore 1984a
Figure 1.20. Major natural vegetation types on Sulawesi.
Based on Anon 1982a
Sulawesi has both freshwater and peatswamp forests, though neither is particularly extensive, as well as forests growing on volcanic, limestone and ultrabasic soils (chapter 6). The mountain forests have tree species absent from or rarely found in the lowlands, and the vegetation of the mountain summits includes many brightly-coloured shrubs and herbs (chapter 7).
A large number of areas, particularly in the southwest peninsula, have lost their original vegetation because of the activities of man, and are now made up largely of agricultural landscapes with areas of dry, barren land used to some extent for cattle grazing. The driest areas, in the Palu valley and in the south of the southwest arm, have a generally open appearance with a scrub of prickly pear cactus and other drought-tolerant species. Many grasslands are dominated by alang-alang grass Imperata cylindrica (Gram.) (Steup 1939), but it is not correct to label all grasslands as alang-alang wastelands; some grasslands comprise communities of numerous grasses (not including alang-alang) and small legumes (Steup 1939). Sword grass occurs predominantly along roadsides and around villages. Grassy savannas often have scattered trees of a variety of species such as Morinda tinctoria (Rubi.), Albizia procera (Legu.) and, where burning has been frequent, Fagraea fragrans (Loga.). There are also areas where teak Tectona grandis (Verb.) is common but these are usually the remnants of old plantations (Steup 1939). Teak was introduced to Java and South Sulawesi many centuries ago from India and Burma (Altona 1922; Carthaus 1909).
Based on Anon. 1982a
FAUNA
Palaeofauna
The palaeofauna of Sulawesi is known from just two sets of sites: river sediments near Sompoh, Beru and Celeko in Soppeng district about 100 km northeast of Ujung Pandang, and various limestone caves near Maros. The animals found in the first of these have been called the Cabenge fauna and probably date from the Late Pliocene (more than 1 Ma ago) (Sartono 1979; Hooijer 1982). The animals in the cave sites are known as the Toalian fauna and are of relatively very recent origin, dating from perhaps 30,000 years ago (Hooijer 1950) (table 1.7).
The stegodonts would have looked similar to modern elephants except that the males had huge curving tusks that were so close together that the trunk must have been draped over the sides of the tusks. The pygmy elephant (fig. 1.21), was descended from the prehistoric African elephant Elephas ekorensis (the extant African elephant is Loxodonta africana) and probably left the lineage of the modern Asian elephant E. maximus about 3 million years ago.
The extinct giant pig had peculiarly large tusks in the upper jaw, nearly triangular in cross section, which pointed sideways and extended beyond the lower tusks. The fossil babirusa teeth are larger than those of their living relatives, and the giant tortoise had a carapace nearly two metres long (Hooijer 1948b, 1982), larger than those confined today to the Galapagos Islands in the eastern Pacific and small islands in the western Indian Ocean. The Pleistocene anoa was a similar or possibly
