Читать книгу Ecology of Sulawesi - Tony Whitten - Страница 12
ОглавлениеChapter One
Physical, Biological and
Human Background
GEOLOGY
Geological History
The geology of an area and its geological history are the major determinants of the soils, plants and animals that occur there. For this reason a brief account is given below concerning the physical conditions and history of Sulawesi.
About 250 Ma1 ago the earth comprised of two great continents: Laurasi-comprising present-day North America, Europe and much of Asia and Gondwanaland-comprising present-day South America, Africa, India, Australia, Antarctica and the remainder of Asia. Until the last few years the once widely accepted view of the geological history of Indonesia and surrounding regions was that the western half (Malay Peninsula, Sumatra, Java, Borneo and western Sulawesi) had been part of Laurasia, separated until recently from the eastern half (eastern Sulawesi, Timor, Seram, Buru, etc.), which had been part of Gondwanaland, by the broad Tethys Ocean.
This picture has had to change in the light of recent geological and palaeontological evidence. The current view, not without its critics however, is that southern Tibet, Burma, Thailand, Peninsular Malaysia and Sumatra were once part of Gondwanaland and rifted from the northern Australia-New Guinea continental margin some 200 Ma ago. This continental fragment then formed a dissected land connection between Australia and Asia and may have carried with it an evolving higher-plant flora (Audley-Charles 1987). Western Sulawesi together with Sumatra, Borneo, and land that would later form the islands of the Banda Arc2 are considered to have separated from Gondwanaland in the middle Jurassic (Audley-Charles 1983). Australia broke away from Antarctica much later, perhaps in the early Cretaceous (90 Ma), and Australia, New Guinea, and east Sulawesi proceeded to travel northwards at about 10 cm per year. At least part of eastern Sulawesi probably separated from New Guinea before its mid-Miocene collision with western Sulawesi after which the eastern half began to emerge as an island (Audley-Charles 1987) (fig. 1.1; table 1.1).
Figure 1.1. Changing locations of components of Southeast Asia since the first rifting from east Gondwanaland, showing schematic ocean spreading ridges, subduction trenches (triangles) and continental margins (dotted lines). Present coastlines shown for reference only. Horizontal lines: 0°, 30° and 60° South.
After Audley-Charles 1987
The most dramatic event in Indonesian geological history occurred in the Miocene when the northward-drifting Australian Plate caused the bending to the west of the eastern part of the Banda Arc. This westward movement, coupled with the westward thrust along the east-west Sorong fault system from western Irian Jaya, modified the two major landmasses that would form the peculiar shape of Sulawesi we recognize today. It has been proposed that this collision occurred 19-13 Ma ago (Sasajima et al. 1980; Audley-Charles 1987). The Banggai-Sula Islands formed the continental platform section of the east Sulawesi fragment. The Talaud Islands and the small islands of Mayu and Tifore between North Sulawesi and Halmahera are probably also part of the collision suture that formed between Sundaland and Gondwanaland (Audley-Charles 1987).
Thus eastern Sulawesi was like a spearhead that hit western Sulawesi and caused the southwest peninsula to rotate anticlockwise by about 35°, thereby opening the Gulf of Bone (Haile 1978) and causing the northern peninsula to pivot around its northern end, rotating clockwise through nearly 90°. This would have caused subduction3 along the North Sulawesi Trench in Gorontalo Bay (Otofuji et al. 1981) and obduction4 of the ultrabasic rocks of east and Southeast Sulawesi over the erosion debris or molasse deposits of younger rocks.
The physical history of eastern Indonesia has made it one of the most geologically complex regions in the world (Audley-Charles 1981). It is this complexity and the strange shape of Sulawesi, described variously as an orchid, a demented spider and a wobbly 'K', which have long attracted the interest of geologists and others (Davis 1976; Otofuji et al. 1981).
Sulawesi comprises three distinct geological 'provinces' brought together by movements of the earth's crust as described above. These are West and East Sulawesi (divided by the north-northwest fault between Palu and the Gulf of Bone-the Palu-Koro fault), and the Banggai-Sula province comprising the Tokala region behind Luwuk on the northeast peninsula, the Banggai Islands, Butung Island and the Sula Islands (actually part of the political province of Maluku) (fig. 1.2).
West Sulawesi is underlain in the south by a basement of schists (metamorphic rocks of continental origin that split easily along their mineral plates) and ultrabasic rocks (derived from the mantle), and in the north by schists and gneiss (banded, coarse, metamorphic rocks that do not split easily). These are overlain by marine sediments including limestone (primarily calcium carbonate from animal shells), sandstones (consolidated sand), cherts (a compact flint-like variety of silica) and shales (thin layers of consolidated mud, clay and silt). Volcanism began in the Eocene but became widespread in the Miocene, and the volcanic arc that ran from the south to the north deformed the existing sedimentary rocks. The ash and dust from the volcanoes mixed with eroded sedimentary rock derived from the uplifting of the eastern part of the Sulawesi collision zone, to form the Celebes molasse, a generally poorly-consolidated conglomerate rock of gravels, sands, silts and muds formed in terrestrial or shallow-water environments. Molten igneous rocks such as granite and diorite have forced their way by intrusion into Miocene and older rocks at a number of locations (Sukamto 1975a, b; Otofuji et al. 1981). East Sulawesi consists mainly of basic and ultrabasic igneous rocks associated with schists in the west and with Mesozoic limestones in the east and the south.
Figure 1.2. Geological map.
After Katili 1978
The Banggai-Sula province comprises a basement of Palaeozoic metamorphic rocks intruded by granites. Triassic and Permian effusive rocks were deposited locally on the basement. These rocks are overlain by Mesozoic shale, sandstone, conglomerate and marl (consolidated mud and calcium carbonate) deposited in both continental and marine shelf environments. Butung Island is included in this province by virtue of distinctive Jurassic shales which it shares with Banggai-Sula but which do not occur in East Sulawesi (Sukamto 1975b).
Figure 1.3. Tectonic features of Sulawesi.
After Katili 1978
It has been suggested that western Sulawesi collided with eastern Borneo in the late Pliocene (3 Ma ago) thereby closing the Makassar Straits which opened again only during the Quaternary (Katili 1978), although there is no great weight of data to support this. Thick sediments in the Straits from before the Miocene indicate that Borneo and Sulawesi have been separated for at least 25 Ma. During periods of low sea-level (p. 16) it is quite likely that islands would have existed particularly in the area west of Majene and around the Doangdoangan shoals (Audley-Charles 1981). In the latter area a drop in sea-level of 100 m would have exposed an expanse of almost continuous land between southeast Borneo and southwest Sulawesi. One interesting observation, however, is that along the northern (and deeper) section of the Makassar Straits the 1,000 m submarine contour of eastern Borneo more or less exactly matches that of western Sulawesi (Katili 1978) so it is possible that the Straits was once narrower.
As can be deduced from the active lateral movement along the Gorontalo, Palu-Koro, Matano and Sorong faults (fig. 1.3), the island of Sulawesi is at present undergoing a process of fragmentation. The end result could be a cluster of islands separated by narrow straits resembling the complex pattern of the Philippine Archipelago in which the original double island-arc structure is no longer recognizable (Katili 1978).
Volcanoes
The most devastating eruption in Sulawesi in recent times was that of Colo volcano on Una-una Island in Tomini Bay. As far as is known this had erupted only once before, in 1898, when a large eruption spewed out 2.2 km3 of ash which was deposited over 303,000 km2, reaching nearly as far as the border between Sarawak and East Kalimantan 800 km away (Umbgrove 1930). For some days after 14 July 1983 a large number of earthquakes—up to 100 per day—shook Una-una and on 18 July a large 'phreatic' eruption occurred; that is, water caught below hot volcanic material was turned into high pressure steam and exploded, generating a column of steam and debris some 500 m high. The first magmatic eruption occurred in the morning of 23 July, and this sent a plume of ash and other material 1,500 m into the air. The violent climax of the activity came that afternoon when most of the island blew apart and most of the vegetation was destroyed by a 'nuée ardente' (Katili and Sudrajat 1984),5 a glowing mass of turbulent, superheated gases and incandescent solid particles (Francis 1978). Ash from the 15,000 m-high cloud reached Pulau Laut, an island 900 km away off southeast Kalimantan, and covered 90% of the remains of Una-una. All the island's inhabitants were evacuated in time, a wonderful achievement, but all houses, crops, animals, coral and inshore fish were destroyed except along a narrow strip on the east of the island (Katili and Sudrajat 1984).
Colo volcano is just one of a number of volcanoes on or near Sulawesi which have greater and lesser effects on the surrounding human population. By far the most destructive of human life has been Mt. Awu on Sangihe Island (fig. 1.4; table 1.2). Apart from the obvious damage to the surrounding land, ash clouds can have serious effects on aircraft. For example, a Qantas jet had to be grounded for major repairs in 1985 after it had flown into an ash cloud from Soputan volcano.
Figure 1.4. Location of active volcanoes. Numbers refer to table 1.2.
Sulawesi has 11 active volcanoes, compared with 17 on Java, 10 on Sumatra and 6 on Halmahera, and numerous old cones, the most beautiful of which is probably Manado Tua in Minahasa. The volcanoes are associated with the subduction zones north of Toli-Toli (in the case of Colo on Una-una), and east of Minahasa and Sangihe (in the case of the remainder). These regions are often shaken by earthquakes (McCaffrey and Sutardjo 1982). The epicenters or positions on the earth's surface where these originated are frequently cited, but less attention is paid to the depths at which the earthquakes originate. For example, the floor of the Sulawesi Sea is moving southwards but instead of buckling and piling up it is forced down at an angle of about 60° under the northern peninsula. The enormous forces and friction involved generate both earthquakes and heat, and the zone where these occur is referred to as the Benioff Zone. The heat can be so intense that the rocks of the descending plate melt, and the molten material or magma forces its way upwards. In most cases this magma never reaches the surface, hundreds of kilometers above, but cools down in pockets in the earth's crust. The magma which does reach the surface, however, is ejected in volcanic eruptions as lava or pyroclastic deposits such as ash and larger rock particles (fig. 1.5).
A - volcano with eruptions in historical time
B - volcano in solfatara and fumarole stage
C - solfatara and fumarole field
After van Bemmelen 1970; Anon. 1979
Figure 1.5. Vertical section through North and Central Sulawesi showing the Benioff Zone below Una-una.
Based on Katili and Sudrajat 1984
Minerals
Petroleum. Petroleum deposits are the remains of microscopic plants and animals which were buried in mud and sand of shallow prehistoric seas, underwent slow decomposition by bacteria and left a residue of hydrocarbon compounds which, under conditions of high temperature and pressure, were converted into oil and gas. Petroleum in Indonesia is usually found in thick Tertiary deposits (fig. 1.6) but there are leakages of natural gas in Tanjung Api Nature Reserve on the northern coast of the northeast peninsula of Sulawesi, and of oil at various inland and coastal locations in the northeast arm. The largest is in a mangrove area in Kolo Bay on the southern shore where the people have used the viscous oil for corking boats. The first oil well on Sulawesi was drilled in 1902 over one such deposit near the mouth of the Lariang River in northwest South Sulawesi. A gas field to the northeast of Lake Tempe has recently been found by British Petroleum and it is possible that an ammonia or urea plant might one day be built nearby. Development has been postponed, however, due to the low international price of liquified natural gas (Anon. 1986). The first commercially-viable source of oil in the Sulawesi region was found in 1985 by Union Texas 25 km south of Kolo Bay on the southern coast of the northeast Peninsula, and more test wells are planned.
Asphalt. Asphalt is a black, sticky mixture of bitumen or tarry hydrocarbons and mineral matter. About 70,000 ha of limestone in the southeast of Butung is impregnated with asphalt to the extent of 10%-40% by weight (fig. 1.7). These hydrocarbon mixtures have migrated upwards along faults above deep deposits into recent, relatively porous rocks and the lighter fractions have evaporated, leaving the viscous asphalt behind. The asphalt deposits with concentrations of 20% - 30% were first exploited in the 1920s, primarily for tarring roads, and remain the only source of natural asphalt in Southeast Asia (van Bemmelen 1970; Anon. 1985a). About 500,000 tons of asphaltic rock are processed each year (Anon. 1984a).
Coal. Coal, the fossilized remains of plants, has never been mined in Sulawesi and occurs only around Pangkajene, Enrekang, Makale and the Karama River, all in the southwest peninsula. None of these young Tertiary deposits is economically viable (van Bemmelen 1970).
Limestone. Just north of Maros at the edge of the karst hills of elevated Miocene coral reefs (p. 468), are the P.T. Semen Tonasa limestone quarry and cement factories from which about 400,000 tons of cement is produced each year, which supplies the entire needs of eastern Indonesia (Anon. 1984a).
Figure 1.6. Distribution of Tertiary deposits (dark shade) and the location of past and present petroleum exploration activities (dashed lines).
After Anon. 1984a
Copper. Copper ores are found primarily in the northern arm and near the nickel areas of Soroako. None has yet been mined but there were plans to do so just east of Gorontalo6 (Lowder and Dow 1978), and possibly in the Latimojong Mountains of the southwest peninsula. An analysis of leaves from herbs collected on Salayar Island, south of the southwest peninsula, revealed high concentrations of copper (80-600 mg/g compared with normal concentrations of <50 mg/g), indicating that rich deposits might also be present there7 (Brooks et al. 1978).
Gold. Gold is generally associated with copper deposits, and the exploration for copper in North Sulawesi has revealed locations of potential large, commercial gold mines. Four mines that were worked early this century have been exhausted for commercial exploitation. New exploration licenses, however, have recently been issued.
Figure 1.7. Distribution of asphalt-impregnated limestone on Butung Island.
After Anon. 1985a
Nickel. Nickel ores are derived from the weathering of ultrabasic rocks and are also found in the molasse deposits derived from these rocks. The ore deposits around Soroako and Lake Matano are mined by P.T. Inco and those around Pomala'a, south of Kolaka, are mined by P.T. Aneka Tambang.
Minor Products. Deposits of sulphur are known only from Soputan and Mahawu volcanoes in Minahasa and these have been exploited since the 1920s, although production is currently not very active. Kaolin or white clay is produced on a small scale in North Sulawesi for the ceramics industry. Salt is produced in coastal salt pans by small-scale operators in South Sulawesi. Quartz sands and silica were mined in South Sulawesi up to 1977 since when it has not been economically viable to do so.
SOILS
The description of soil types is hampered because there is no system of soil classification that has gained universal acceptance in terms of definitions or names. From experience and observation it has been said that "scientists who are otherwise reasonable and unemotional are liable to behave quite differently when discussing this topic" (Mulcahy and Humphries 1967). In many cases, hybrid systems have evolved using names from various systems with the result that those with little knowledge can become extremely confused. For the purposes of this section the FAO system is used because it has wide recognition and because the names are easily pronounceable.
The major soil types on Sulawesi and their approximate distributions are shown below (table 1.3; fig. 1.8).
* Horizons are layers of soil roughly parallel to the surface which have fairly distinctive characteristics.
After Young 1976
Figure 1.8. Approximate distribution of major soil types on Sulawesi.
After Anon. 1976
CLIMATE
Palaeo climate
The palaeoclimate of Sulawesi does not appear to have been studied specifically, but analyses and summaries of palaeoclimate in Southeast Asia are available (Verstappen 1980; Flenley 1980; Whitmore 1981; Walker 1982; Morley and Flenley in press).
The climates of Sulawesi and the rest of Indonesia today, are quite unlike the climates which dominated the region during most of the Quaternary and before. As was shown on page 2, the world's landmasses have moved around, joining and separating, and this has led to changes in climatic regimes. Tropical and subtropical conditions, and the animals and plants associated with them, extended further away from the equator during the Tertiary than they do now and this has influenced the present distribution of animals and plants (p. 63).
During the latter part of the Quaternary, temperatures in the temperate areas of the world repeatedly rose and fell above and below present temperatures. In the cooler periods the ice sheets of the Arctic and Antarctic extended and this took great quantities of water out of the hydrological cycle (fig. 1.9). This in turn caused sea-levels around the world to fall. The maximum fall in Southeast Asia was about 150 m below present levels. This exposed large areas of dry land beyond the present coastlines-indeed, it uncovered three times the present area of the Sunda Shelf and twice the present area of the Sahul Shelf.8 When the sea-level was only 40 m below present levels it would have been possible to walk in a straight line between Banjarmasin and Surabaya, Saigon and Kuching, Singapore and Pontianak, and Merauke and Darwin. The effective area of Sulawesi was also increased but to a much lesser extent (fig. 1.10). Examination of bathymetric contours reveals features that may have been river valleys when the sea-level was lower (fig. 1.11).
Ocean currents which now enter the Indonesian Archipelago through the Torres Straits, the South China Sea, the narrow straits between many of the Lesser Sunda Islands and the straits between Mindanao and the Sangihe/Talaud Islands would have been blocked and their buffering effect on climate would have been lost. The Sulawesi Sea (between the Philippines and Sulawesi) and the Makassar Straits would have been much more enclosed but the currents entering the Molucca Sea (between Halmahera and Sulawesi) towards and from eastern Sulawesi would have been only marginally obstructed. The main Sunda and Sahul landmasses would have experienced a more continental climate (greater diurnal temperature range, lower rainfall and humidity) but this would have been rather less pronounced on Sulawesi. The lowering of sea and land temperatures during these cool periods would also have reduced rainfall and humidity. It has been estimated that the rainfall 11,000 years ago was 30% of present values in the equatorial zone.
Figure 1.9. Hydrological cycles (a) in warm conditions and (b) in cool conditions. Note the fall in sea-level because water in the form of ice is unable to flow to the sea.
Figure 1.10. The area of Sulawesi and neighbouring land-masses exposed when the sea-level was 100 m below present levels.
After van Balgooy in press
The cooling of the whole earth during the glacial periods lowered the lowest altitudes at which ice remained all year and snow fell, and lowered the upper altitude at which montane trees grew. The maximum temperature depression during the Quaternary occurred 18,000 years ago when, in New Guinea, temperatures at 2,500 m were 10°C lower, but at sea-level only 2° or 3°C lower, than at present. The tree line (the altitude at which trees are no longer able to grow) was lowered about 1,500 m in New Guinea but only about 350-500 m in Sumatra. The sea-level changes also had considerable impact on corals (p. 215).
The zone where the curtains of rising air of the northern and southern hemisphere meet is called the Intertropical Convergence Zone (ITCZ), and in the cool, dry glacial periods this was probably south of its present position (that is roughly over the equator). It is a zone of frequent, showery rain, to the north of which is a high pressure belt with relatively low rainfall. Thus, when the ITCZ lay south of the equator, parts of northern Indonesia (including Sulawesi) would have experienced drier, more seasonal climate with lower rainfall and humidity and greater seasonal change in mean daily temperature (Verstappen 1980).
Figure 1.11. Bathymetric map of the Sangkarang Archipelago to show the deep channels that are probably drowned riverbeds.
Temperatures during the warmest parts of the Quaternary were only 1° or 2°C higher than now (at sea-level) and at this time the water released from the polar ice caps caused sea-levels to rise. There is no unequivocal evidence that shorelines have been more than 6 m higher than at present during the warm periods of the Holocene, but sea-levels could have reached up to 25 m above present levels during the Pleistocene. The most recent sea-level maxima detected off the southwest peninsula were 4,500 and 1,600 years ago when sea-level was 5 m and 2.5 m higher respectively (fig. 1.12) (de Klerk 1983). This agrees closely with evidence from elsewhere in Southeast Asia (Tjia 1980; Tjia et al. 1984). Whereas this rise had a marked effect on the long shorelines of low-lying parts of eastern Sumatra and southern Borneo, the most marked effect on Sulawesi would have been the separation of the blocks of land either side of the Tempe depression. Evidence for this has been found in the vegetation record (p. 29) and there are even stories among local people of a time when travellers did not have to sail around the southern tip of South Sulawesi but could instead sail from the Gulf of Bone, through Lake Tempe and emerge in the Makassar Straits (Sartono 1982). With the exception of the Tempe depression and a few other flat plains (such as Malengke), most of Sulawesi's coastline slopes quite sharply and minor rises in sea-level would not have had significant effects. During this period seasonality in rainfall would have been less, rainfall would have been similar to or even greater than now, and mountain zones of climate, vegetation and fauna would have been raised. However, as stated above, this period occupied only a small fraction of the Quaternary. The majority of the period was characterized by lower rainfall and humidity, greater diurnal and seasonal variations, and by more marked rain shadows. Thus, the seasonal areas of Sulawesi and elsewhere would have been more extensive and, conversely the areas subject to more stable, wetter climates would have been reduced in area.
Present Climate
The climate of Sulawesi is best described with reference to rainfall since temperature is relatively constant, and other climatic variables such as wind velocity, evaporation and humidity change within even small areas. Between September and March, cool northwesterly winds pick up moisture while crossing the South China Sea (between East and West Malaysia, Philippines and Vietnam) and arrive in North Sulawesi via the Sulawesi Sea in about November, and in the west coast of South Sulawesi via the Java Sea in late November or early December. The west coast of the central part of Sulawesi is sheltered from the effects of these winds by Borneo.
Figure 1.12. Changes in sea-level over the last 7,000 years determined from a study in the southwest peninsula.
After de Klerk 1933
After this period, variable, humid, southeasterly winds blow towards eastern Sulawesi and rainfall peaks on the southeast coast occur between April and June, and on the northeast coast somewhat later. The southeasterly winds from the dry and wintery Australian landmass become stronger and these dry winds have a significant influence on the southern tips of the southwest and southeast peninsulas. Manado experiences a short dry season from August to October, but Jeneponto in the south of the southwest peninsula is subject to a long dry season between April and November.
Areas on the west coast of Sulawesi therefore tend to have their highest rainfall in December whereas those on the east coast have their wettest month around May. One might expect to find intermediate areas with two dry seasons (a bimodal distribution) and this is indeed the case; Pendolo and Pinrang in the middle of the southwest peninsula are examples.
Where the orientation of a range of mountains is more or less at right angles to the prevailing winds, the rainfall is higher on the windward side because the water in the air rises and cools as it climbs over the mountains, and this moisture is released as rain. Thus Maros receives over 500 mm per month between December and February but towns on the leeward side of the peninsula receive little rain. Valleys orientated in a north-south direction are in a rain shadow for virtually the whole year and the sheltered nature of the central western coast results in the Palu valley being one of the driest areas of Indonesia with less than 100 mm of rain, on average, falling in each month and an annual total of less than 600 mm.
Various authors have mapped the climatic zones of Sulawesi. The map which corresponds closest to the distribution of vegetation is that which uses the ratio between dry and wet periods (fig. 1.13) (Schmidt and Ferguson 1951; Whitmore 1984a, b).
A second map based on suitability criteria for growing rice has also been devised (fig. 1.14) (Oldemann and Darmiyati 1977), in which five major zones are recognized.
| Zone A | - | an area with ten to twelve consecutive wet months and two or less consecutive dry months; |
| Zone B | - | an area with seven to nine consecutive wet months and three or less consecutive dry months; |
| Zone C | - | an area with five or six consecutive wet months and three or less consecutive dry months; |
| Zone D | - | an area with three or four consecutive wet months and two to six consecutive dry months; |
| Zone E | - | an area with zero to two consecutive wet months and up to six consecutive dry months. |
'Wet' and 'dry' are defined as more than 200 mm and less than 100 mm of rain per month respectively.
Sulawesi has a greater percentage of its area in agroclimatic Zone E than have the islands around it, but more in Zones B and C than Borneo (most of which is in Zone A), Nusa Tenggara and Bali, or the Moluccas (table 1.4).
The most recent and complex climatic map of Sulawesi concerns bioclimate (Fontanel and Chantefort 1978) and recognizes many zones determined by three criteria: mean temperature of the coldest month, mean annual rainfall (fig. 1.15), and number of dry months9 (fig. 1.16). One of the major differences between this map and that on agroclimatology is that it takes into account altitude effects, although not in an absolute sense since the degree of exposure to prevailing winds has a significant effect (p. 21).
The maps discussed above are based on long-term averages but uncommon climatic events, particularly periodic drought, can be extremely important in determining the distribution of certain animals and plants. Most crops, for example, experience stress after only about four days without rain. The variation in annual rainfall is quite considerable (fig. 1.17) with the total for one year sometimes being twice the total of another. Meteorological records normally extract maximum rainfall figures but these are not particularly meaningful ecologically because above a certain quantity, rain will simply run off saturated soils to rivers. Lack of water, with its associated cloudlessness, high temperatures and low humidity, is a much more potent factor and an examination of rainfall minima and their distributions reveal that these too are extremely variable (fig. 1.18). Dry seasons can in fact only be defined by probability since, for the stations examined, at least six different months were recorded as 'driest months' in at least one year. A single location (e.g., Mapanget or Watampone) can have minimum monthly rainfalls between years ranging from 0 to 100+ mm.
Figure 1.13. Rainfall types based on dry/wet period ratios.
After Schmidt and Ferguson 1951; Whitmore 1984a, b
Figure 1.14. Agroclimatic zones.
After Oldemann and Darmiyati 1977
* The figures in the text do not add up to 100%.
After Oldemann and Darmiyati 1977; Oldemann et al. 1980
Figure 1.15. Areas with different mean annual rainfall.
After Fontanel and Chantefort 1978
Figure 1.16. Areas with different numbers of dry months.
Alter Fontanel and Chantefort 1978
Figure 1.17. Annual rainfall at Mapanget (Manado airport), Watampone, Tobea Besar (an island between Butung and the mainland) and Palu.
Data obtained from the Directorate of Meteorology, Jakarta
Figure 1.18. Percentage of years' driest months occurring in each month with agroclimatic zone, average annual rainfall in parentheses, and years of complete data in brackets.
From data obtained from the Directorate of the Meteorology, Jakarta
Droughts in Sulawesi seem to occur about every 20-30 years but information on their ecological effects have not been found. The grassy Sook Plain in Sabah, however, was formed when fire burned rainforest and the peaty topsoil on which it grew after an exceptionally dry period in 1915 (Cockburn 1974). In 1972-73 a prolonged dry spell caused changes on Mt. Kinabalu, Sabah, which it has been estimated will take a century to be reversed (Whitmore 1984a). In most areas of the world, but particularly around the Pacific region, 1982-83 was exceptionally warm and dry, and weather patterns were most unusual. For example, Watampone had 5, and Tobea Besar 6, consecutive rainless months at the end of 1982. This was related to the anomalous sea surface temperatures which in parts of the Pacific rose by as much as 8°C above normal. Around Sulawesi, however, the sea was only about 0.2°C warmer than normal. The sea surface temperature anomaly first appeared in the region of the Sulawesi Sea during the middle of 1982 but its major effects were not felt until December 1982-February 1983 (Barber and Chavez 1983; Cane 1983; Gill and Rasmusson 1983; Rasmusson and Wallace 1983; Chavez et al. 1984; Whitmore 1984a). Although 1982 was generally a very dry year on Sulawesi, 1972 seems to have been even drier.
VEGETATION
Palaeovegetation
Impressions of leaves of grassy plants and rattans have been found in rocks in Minahasa (Koorders 1895) but the vast majority of information regarding palaeovegetation is derived from the careful analysis of pollen remains 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 slightly smaller size than modern specimens. The large relative size of many Pleistocene fossils is a commonly observed phenomenon, possibly a result of the cooler temperatures then prevailing. Larger animals have a lower ratio of skin area to body volume and a consequently lower rate of heat loss. As a result, larger animals are better able to survive at low temperatures (Edwards 1967). Dwarfing of animals on islands generally occurs in species that are relatively large, such as elephants and anoas, whereas gigantism on islands generally occurs in small species such as rats.
The Pleistocene sharks are all still found around Sulawesi and occasionally in rivers, but Hemipristis is now represented only by a single rare species in the Red Sea and Zanzibar (Hooijer 1958). The stingray is also common in Indo-Pacific seas and rivers.
The animal remains found to date are much poorer in numbers and species than those from the Trinil fauna deposits in East Java. Whereas the Trinil deposits may be used to some extent to illustrate the fauna present over some of Sundaland, it is not possible to use the Trinil finds as a basis for discussion of the Sulawesi palaeofauna. Two of the four elephants but none of the three pigs in the Trinil deposits are known from Sulawesi (Hooijer 1974), and the pygmy stegodont on Sulawesi is now also recognized as the same species as the fossil stegodont found on Timor and Flores. The Plio-Pleistocene fauna of Sulawesi is thus not as distinctive as was once thought and the only species not known from outside Sulawesi is the giant pig.
* extinct forms
After Hooijer 1948a, b, 1949, 1950, 1954, 1958, 1964, 1967, 1969, 1972; Clason 1976; 1982; Musser 1984
Figure 1.21. Relative sizes of the giant tortoise, 1.65 m man and a pygmy elephant.
The ancestors of the elephant and stegodonts could have reached Sulawesi via Taiwan and the Philippines (stegodont remains have been found on both islands). The 3,000-m deep sea between Java, Flores, Alor, Timor and Sulawesi might be thought to be too great for even water-loving elephants to cross by swimming (Hooijer 1967, 1970, 1972). If it is assumed that the presence of similar pygmy stegodonts on all these islands must be a result of migrations across land and only short water gaps, it is necessary to postulate massive downfaulting of at least 3,000 m in the intervening straits and seas. Before the downfaulting occurred there would have been routes from Flores to Sulawesi along the Kaloatoa ridge and then via Tanahjampea and Salayar, and from Java to Sulawesi via Madura, the Kangean Islands and the Doangdoangan Islands (Audley-Charles and Hooijer 1973) or perhaps more easily from the Kangean Islands via the Sabalana and Postilyon Islands. The routes from Java may in fact have been viable even without downward faulting during the glacial periods of the Pleistocene during which sea-level dropped 100-130 m below present levels (p. 16).
A recent examination of the swimming powers of elephants has suggested, however, that such massive downfaulting need not have occurred. Elephants have been recorded swimming across lakes and seas to islands in Africa and Asia and the distance record is held by an unfortunate elephant that swam ashore after being washed overboard from a boat 48 km off the South Carolina coast in 1856. Aspects of elephant morphology such as the spongy skull bones and absence of a pleural cavity, the need to bathe regularly and their ancestry (dugongs, or sea cows, are among their closest relatives), suggest that their ancestors may have lived in a semi-aquatic habitat. Some of the distances prehistoric elephants would have had to have swim are in the upper limits of known ability and so the crossing of elephants to and from Sulawesi and other islands does not necessarily require that the sea bed was much higher, and exposed land more extensive, in the recent past (Johnson 1980).
Giant tortoises are said to be able to float and survive long periods in seawater. Thus Geochelone atlas could have drifted to Sulawesi from another landmass (Hooijer 1982), and in fact remains of the same species are found in deposits from Java, Timor and India.
Present Fauna
The fauna of Sulawesi is one of the most distinctive in all Indonesia particularly among the mammals (table 1.8). Of the 127 indigenous mammal species,14 79 (62%) are endemic15 and the percentage rises to 98% if the bats are excluded (table 1.9). The mammal fauna is also characterized by its relatively primitive characters (Musser in press). New species of mammals continue to be found (e.g., Musser 1981, 1982; Bergmans and Rozen-daal 1982; Hill 1983; Boeadi pers. comm.) and revisions of old and new specimens combined with fieldwork help to clarify the exact number and identity of species (e.g., Musser 1971a, b, 1973, 1977; Musser et al. 1982; Groves 1980a, b; Takenaka 1982).
One of the earliest descriptions by a European of a Sulawesi animal was of the curly-tusked babirusa Babyrousa babyrussa by Piso in 1658. It was kept and even bred by early rulers, perhaps as a gift for visiting diplomats, and it is conceivable that some were taken to Bali by Buginese seafarers. It is possible that knowledge of the babirusa influenced the way in which 'Raksasa', a Balinese demonic man-beast, was first drawn, for it has a curved tusk piercing each cheek (Groves 1980a).
The babirusa are enigmatic animals as their name, which means 'pig-deer', implies. They are variable in hair covering (from nearly naked to densely hairy), hair colour (from off-white to brown), and eye colour (from whitish, through grey to brown) (Wemmer and Watling 1982) and none of these characteristics appear to be related to sex. There may be geographical trends in these characters, however, but too few specimens with detailed location data are available. The babirusa is generally grouped with pigs but they have had no common ancestor since the Oligocene (about 30 Ma), and they are barely more similar to pigs than they are to hippos, remains of primitive forms of which have been found in Java (C. Groves pers. comm.).
Two dwarf buffaloes, or anoa, Bubalus depressicornis and B. quarlesi are said to be found in the lowlands and mountains respectively (Groves 1969) but it has been suggested that the differences in horn-shape, a major means of distinguishing the species,16 may simply be a function of age (Wind and Amir 1978). Also the 'mountain' anoa is sometimes found at sea-level and the 'lowland' anoa is sometimes found on high mountains (Thornback 1983). The diurnal bear cuscus Ailurops ursinus is quite a large animal, with a head and body 45 cm long and a tail 55 cm long, and is quite frequently seen in lowland forests. It is very distinct morphologically from other cuscus and is in its own genus in recognition of its primitive characters (C. Groves pers. comm.). The smallest cuscus of all, Strigocuscus celebensis (head and body only 34 cm), is nocturnal, and consequently rarely seen.
The composition of the Sulawesi mammal fauna is very different from that of Borneo or Irian Jaya with many fewer families represented (fig. 1.22). The rats and bats are major constituents of the fauna as they are in the insular regions of the Moluccas and Lesser Sundas.
Based on Anon 1982b
Figure 1.22. The faunas of Borneo, Sulawesi, Flores, Maluku and Irian Jaya (Vogelkop) compared.
There are 332 species of birds known from Sulawesi of which 92 (27%) are endemic, and 81 (25%) are migratory (p. 145; table 1.10) (White 1974, 1976, 1977; White and Bruce 1986). New records of species not previously known from Sulawesi are still being made (Escott and Holmes 1980; Watling 1983). Among the resident birds, 17 genera are endemic to Sulawesi and its surrounding islands including a large number of spectacular endemic birds such as the dark-green bee-eater Meropogon forsteni, the large brightly-coloured hornbill Rhyticeros cassidix, the crowned myna Basilornis celebensis, and the finch-billed starling Scissirostrum dubium which nests in huge numbers in holes bored out of tall dead trees (fig. 1.23). Sulawesi's best-known bird is the maleo Macrocephalon maleo which incubates its eggs in pits that the adult birds dig (p. 155).
Species and genus names in bold indicate Sulawesi endemics, (i) = introduced, (n) = first found by EoS teams, (p) = found on Peleng Island, (s) = found on Salayar Island.
After Groves 1976; Musser 1977, 1981 in press, pers. comm.; Jenkins and Hill 1981; Hill 1983 pers. comm.: Bergmans and Rozendaal in press;
Figure 1.23. Four of Sulawesi's endemic birds. a - Sulawesi crowned myna Basilornis celebensis, b - White-necked myna Streptocitta albicollis, c - Sulawesi dwarf hornbill Penelopides exarhatus, d - Piping crow Corvus typicus.
After Meyer and Wigglesworth 1898; Goodwin 1976
Whereas 100 species of amphibians have been collected on Borneo, only 29 are so far known from Sulawesi.17 Of this total, four frequent habitats associated with man and have probably been accidentally introduced. Of the 25 indigenous species, 19 (76%) are endemic (table 1.11). The Talaud Islands have a tree frog Litoria infrafrenata not found on the mainland but which is common to the east as far as Queensland (van Kampen 1923). The apparently poor amphibian fauna may be an artefact of undercollection since most of the species were found as a result of collections made in the 1870s-1890s and only three small collections have been made this century, each of which has included new species. There may, therefore, be as many species to be discovered as are already known (J. Dring pers. comm.).
After Holmes and Wood 1979; White and Bruce 1986; K.D. Bishop pers. comm., N. Collar pers. comm.
There are 40 species of lizards known from the Sulawesi mainland, 13 of which are endemic, but the group is poorly known and there are certainly new species awaiting discovery. One of the most distinctive of the reptiles, is the large sailfin lizard Hydrosaurus amboinensis, which is usually found near water (p. 301). Better known among the reptiles are the snakes and 64 have been collected from the mainland and its coastal waters (table 1.12) compared with 136 in Peninsular Malaysia, 150 in Sumatra, 110 in Java and 166 in Borneo (Medway 1981). There are 15 endemic species and one endemic monotypic genus Rabdion (den Bosch 1985). Strangely, the small island of Tanahjampea south of Salayar Island has only two species of snakes and both these are endemic: the Jampea ilyssid Cylindrophis isolepis and the Jampea pit viper Trimeresurus fasciatus (de Rooij 1917). Both species were collected by a team from the Bogor Museum that visited Jampea in 1984 (Boeadi pers. comm.). Sulawesi has the distinction of being the locality of the world's longest recorded snake, a reticulated python Python reticulatus that measured 9.97 m in length (McWhirter 1985). These snakes are the only Sulawesi land animals that present any real threat to man.
All of the fish indigenous to Sulawesi are brackish-water species tolerant of freshwater. Some of these appear to be restricted to lakes, while the eels migrate between the lakes and the sea. Many species have been introduced deliberately or accidentally (such as the air-breathing snakehead Channe18 striata and climbing perch Anabas testudineus) and it is these fish that dominate the Sulawesi freshwater fisheries (p. 330).
Until recently the invertebrates of Sulawesi were very poorly known but three expeditions have greatly increased the knowledge of what is present:
• Project Wallace (1985) organized by the Indonesian Institute of Sciences and the Royal Entomological Society of London, based in the Bogani Nani Wartabone National Park in Bolaang Mongondow, North Sulawesi;
• Operation Drake (1980) organised by the London-based Scientific Exploration Society, based in Morowali National Park; and
• a series of medical expedition teams (1970s) organized by the National Institute of Medical Research and the United States Naval Medical Research Unit, based near Lake Lindu.
A flood of papers describing new Sulawesi invertebrates appeared after Project Wallace (e.g., Hoogstraal and Wassef 1977; Bedford-Russel 1981, 1984; Hadi and Tenorio 1982; Hayes 1983; Goff et al. 1986). As an example of the previous lack of information, only one species (endemic) of spring-tail or Collembola was known from Sulawesi before 1985. In a few weeks of collecting in and around Bogani Nani Wartabone National Park no less than 120 species from about 70 genera had been added to the list. Specimens were even collected of two genera that were previously known from single species in North America (P. Greenslade pers. comm.). These small animals are very important in decomposition and mineral cycling (p. 365).
1 Species of western Indonesian forests and clearings.
2 Species found in the Moluccas or New Guinea.
3 Species of cultivation, secondary growth and degraded habitats, common in
western Indonesia and probably introduced by man. Other species are endemic.
From J. Dring pers. comm.
There are 38 species of the large and usually striking swallowtail butterflies on Sulawesi and 11 (29%) of these are endemic. One is Atrophaneura palu, a large black and white swallowtail (forewing 70 mm long) known from only a few specimens collected from close to what is now Lore Lindu National Park (Haugum et al. 1980; Collins and Morris 1985). Another seemingly rare, recently-described butterfly is the 'paper handkerchief or wood nymph butterfly Idea tambusisiana from Mt. Tambusisi in Morowali National Park (Bedford-Russell 1981). Further specimens have since been found by Japanese scientists in North Sulawesi (R. Vane-Wright pers. comm.).
Endemic genus and species in bold italic.
* = freshwater habitats,
# = estuarine and marine habitats.
After Regenass and Kramer 1981; in den Bosch 1985; C. McCarthy pers. comm.
Occasionally insects can be seen in such unusually great numbers that the observation is worthy of record. One example is the stream of pierid white butterflies seen crossing Kalaotoa Island from dawn until dusk one day in 1936 (Doctors van Leeuwen 1937). Such migrations of pierids are well documented in Europe and it has been found that these butterflies travel in more or less straight lines, searching for suitable habitats. By so doing, they avoid returning to a place that they have just left. They feed and breed in different habitats and so, for a chort of butterflies that emerged at more or less the same time, the best place to be is somewhere else. These butterflies sometimes seem to follow rivers but it is possible that they are simply more visible there because experiments have shown that they have a sense of direction rather than a sense of location (Baker 1982).
Endangered Species
The names and status of the world's rare and endangered animals are compiled and monitored respectively by the Conservation Monitoring Centre of the International Union for the Conservation of Nature (IUCN) and Natural Resources based in Cambridge, England. 'Red Data Books' are produced detailing what is known of the ecology and threats facing each of these species, and what conservation measures, if any, are currently in force to protect the animals. Examination of the Red Data Books reveals that 16 species are considered to be at risk of extinction19 on Sulawesi (table 1.13).
During the course of writing this book it has become clear that the above animals are by no means the animals most at risk and indeed some animals may have become extinct unnoticed.
The attractive Caerulean paradise-flycatcher Eutrichomyias rowleyi was discovered in 1873 by a hunter working for the German ornithologist A.B. Meyer (Meyer 1878). The label on the first specimen states that it was male but there was no verification of this (Meyer 1878), and in fact the colour is more typical of a female flycatcher than of a male (S.V. Nash pers. comm.).
That first specimen is in fact the only specimen ever collected of this distinctive bird. One may have been sighted in 1981 (White and Bruce 1986), but intensive ornithological surveys of Sangihe Island by experienced ornithologists in 1985 and 1986 failed to find any. Virtually all of Sangihe has been converted to coconut and nutmeg plantations or else is covered in patches of secondary forest from abandoned gardens. A small patch of montane forest exists on the top of Mt. Sahendaruman in the south of the island and it was felt that this was the only possible habitat for the species (Whitten et al. 1986). Surveys by Action Sampiri in 1998-99 revealed that this bird was clinging to survival in those forests but the numbers are extremely low.
The status of other animals endemic to Sangihe and the Talaud Islands, should now be an immediate cause for concern (see the Introduction). The blue-and-red lory Eos histrio (Sangihe and Talaud20), Sangihe hanging-parrot Loriculus catamene and Elegant sunbird Aethopyga duyvenbodei (Sangihe), and Talaud kingfisher Halcyon enigma have all been seen recently although none of them is at all common. It is possible that the blue-and-red lory is already extinct on Sangihe but it still appears to be common on Karakelang, the main Talaud Island (Whitten et al. in press). The black birdwing butterfly Troides dohertyi (forewing length 73 mm male, 82 mm female) is known only from Sangihe and the Talaud Islands where it lives in lowland forests in which the caterpillars probably feed on the leaves of Aristolochia tagala, a climbing shrub that grows in forests and thickets up to 800 m above sea-level (Ding Hou 1984; Collins and Morris 1985). This butterfly must be considered as being under considerable threat given the greatly reduced area of lowland forest on the islands, for it is not known whether it can adapt to secondary vegetation (Collins and Morris 1985). It is probably already extinct on Sangihe but there is a hunting reserve on Karakelang, which includes hill and mountain forests and efforts should be made to determine whether the birdwing is present. The boundaries of this reserve are under pressure from illegal farmers.
E - endangered, V - vulnerable, R - rare, I - insufficiently known.
After Miller 1977; Thornback 1978; King 1979; Groombridge 1982; Wells et al. 1983; Collins and Morris 1985
The endemic fish of lakes Poso and Lindu, and of lakes Towuti, Matano, Wawantoa and Mahalona (table 4.10) are threatened by the introduction of fish to increase fisheries production. Among the endemic species are four duck-billed fish-Adrianichthys kruyti, about 11 cm long from Lake Poso (fig. 1.24), Xenopoecilus poptae and X. oophorus, about 10-20 cm long, also from Lake Poso, and X. sarasinorum, about 7 cm long, from Lake Lindu. These species used to be thought to comprise the entire family Adrianichthyidae, but recent taxonomic analyses have now placed several more species in this family, some of which are also endemic to Sulawesi, and some of which are found from India to Japan (Rosen and Parenti 1981). There are early reports that X. poptae does not seem to lay eggs as most fishes do, but rather voids eggs which hatch on contact with water. The young fry then swim along with their mother. The broken egg membranes, known locally as momosonya21, rise to the lake surface and used to cover considerable areas of the lake (Weber and de Beaufort 1922). This has not been confirmed and the 'momoso' may be pollen or seeds. It is known that the newly discovered X. oophorus carries its eggs below its body until they hatch (Kottelate 1990) (see Introduction). Similar behaviour in X. poptae may have given rise to the other story.
Teams that visited Lake Poso in 1976 and 1983 both found the two endemic fish, but a survey of fishermen and an examination of fish catches in 1986 could not confirm the continued existence of either species. Some fishermen claimed that some fish had disappeared when Colo volcano erupted in 1983 (p. 7). This seems untenable as a reason since satellite photographs of the ash plume show how this was blown to the west rather than the south (Katili and Sudrajat 1984). At the start of the century it is said that great shoals of X. poptae formed at 12 -15 m between November and January, and were caught by fishermen using hooks (Weber and de Beaufort 1922).
A likely cause of the reduction in the populations of these endemic fish, and also the possible extinction of some of the snails and mussels endemic to Lakes Poso and Lindu (p. 297) (Carney et al. 1980) is the unthinking introduction of exotic fish species, particularly of the tilapia, carp and catfish, to increase fisheries production (Whitten et al. 1986). The extinction of fish species as a result of the uninformed introduction of commercial species has been reported from elsewhere, particularly the enormous lakes of the East African rift valley to which many species of cichlid fish are (were) endemic. Indeed, 60%-80% of the world's freshwater fisheries are based on introduced species. Not all fish introductions damage the indigenous fauna, however, and success stories have been reported from other African lakes. Major agencies now do not, as policy, advocate the introduction of new species of fish into lakes except with the most extreme caution. To avoid any possible extinction of endemic fish on Sulawesi, fisheries staff must first be aware that any introduction may result in the loss of endemic species, and second be able to justify their aims to all parties.
Figure 1.24. Adrianichthys kruyti, one of the three endemic duck-billed fish of Lake Poso.
BIOGEOGRAPHY
Background to Biogeography
Every organism has a spatial distribution related to its ecology, behaviour, physiology, ability to travel long distances, the other organisms living in the same area, and to the geological history and climate of the area in question. A further and important factor in determining distributions is chance. Conditions suitable for an organism in terms of light, humidity, food, etc. do not necessarily occur evenly over an area and this is reflected in that species' distribution. The study of such patterns and the factors causing or limiting them is known as biogeography.
The distribution of an organism is generally bounded by unsuitable habitats, unsuitable climates or the occurrence of a species against which it cannot compete successfully. Alternatively, a species may be actively dispersing and the edge of its range may simply be the furthest point it has reached at that time. Clearly a barrier for one species is not necessarily a barrier for another, and a barrier for one life-stage of an organism is not necessarily a barrier for a different life-stage of the same species. For example, the larvae of many invertebrates and the seeds of plants are much more mobile than the adults.
Distributions can be described on various levels: thus a bird might live in forest; but perhaps only in a few types of forest; at certain altitudes; avoiding forest near rivers; and only using the tops of the taller trees, etc. Such precise micro-habitats are only available in certain locations, and so a species' distribution usually comprises a number of areas inhabited by discrete populations. If these populations are isolated from other populations over a long period, changes may occur which render the two populations reproductively incompatible and new species or subspecies22 may arise.
Organisms disperse from one suitable area to another along one of three routes:
• corridor-a route comprising the same habitats as are present at either end of the route, thereby giving all species a chance to move through it;
• filter-a route comprising only some of the habitats available at either end of the route, thereby preventing certain species that depend on the absent habitats from moving;
• sweepstakes-a route comprising habitats which are absent from the source of species and may represent, in analogy or actuality, a sea separating two areas of land. The chances of a species crossing this 'sea' are very small but, under certain conditions of wind or the stranding of individuals on a raft of floating vegetation drifting down a river and out to sea, a terrestrial species might successfully cross over to another area of suitable habitat. For example, after heavy rains around Manado in 1882, large forest trees were seen floating out to sea, each sufficiently large to provide a temporary home for a few small mammals such as squirrels and invertebrates. Lava can also push trees into the sea. In these two cases one might expect agile animals to jump off the trees, but a few days after the Krakatoa eruption in 1883, a monkey was found, scorched and tired, clinging to a partly-burned tree in the Sunda Straits. The monkey was kept alive for at least a year (Hickson 1889). Even if an animal is washed alive on to a new coast, if there is no member of the opposite sex available then the sweepstakes will have been lost.
So there are three ways in which an organism can disperse:
• by 'jumping' quickly over relatively large expanses of unsuitable habitat;
• by dispersing slowly across habitat which is more or less suitable; and
• by dispersing very slowly and making adaptations on the way allowing the colonizing of areas the environments of which would have been unsuitable for the original stock.
It is well understood by laymen and scientists alike that small islands support fewer species than large islands. After a certain length of time the total number of species on an island will remain more or less constant, and this represents an equilibrium between the colonizing of the island by immigrant species and the extinction of existing species. The rate of colonization is clearly higher when an island is near the mainland because more species are likely to cross the relatively narrow sea gap. Also, the rate of extinction is clearly greater when an island is smaller because the population of any species will be smaller and the chance will be greater of disease and other detrimental events reducing the population to zero or an unviable number. These relationships can be drawn graphically and represent the foundation of the Theory of Island Biogeography (fig. 1.25).
The relationship between island size and number of species is relatively constant for a given group of animals or plants, and in general reducing island area by a factor of ten, halves the number of species (figs. 1.26 and 1.27). Where an island supports fewer species than expected and so falls below the line, the reason may be that:
• the group is not sufficiently well known;
• equilibrium in species number has not yet been reached (where a volcanic island such as Una-una has been destroyed and is being recolonized);
• the island comprises a relatively restricted number of habitats, or habitats which do not support large numbers of species;
• the island is extremely remote and difficult to colonize.
Figure 1.25. The relative number of species on (a) small, distant islands (b) large, distant or small, close islands, and (c) large, close islands.
Where an island supports more species than expected and so falls above the line, the reason may be that:
• more than the equilibrium number are present and some species will in due course be lost;
• the island is peculiarly rich in habitat types; or
• the island is a centre of species radiation in a certain group.
For total species, Sulawesi is below the line of best fit for both plants and birds. For plants this is probably because there has been insufficient collecting (p. 29), but this is unlikely to be the reason for the birds. Sulawesi falls above the line for mammals due, perhaps, to the extraordinary radiation of rat species, and is more or less on the line for snakes. Sulawesi is consistently above the line for the number of endemic species and this reflects its geological history.
These principles are useful in deciding whether an area is well-known biologically. For example, plotting known number of species of milkweed butterflies (Danaidae)23 against island size (by rank) reveals that islands around Southeast Sulawesi and the Togian Islands have probably not been surveyed sufficiently for this group, whereas for the Banggai Islands and islands in South Sulawesi, nearly all the species of this family are probably known (fig. 1.28).
Figure 1.26. Relationship between number of species and island size for revised plants (those with a recent taxonomic review) and three groups of animals.
From Anon. 1982b
When lists of species found on large and small islands are compared, it is generally found that the species absent from the smaller islands are larger than average since these animals generally have large range requirements and low densities. A few species are more abundant and fill a wider niche24 on islands than they do on the neighbouring mainland or larger islands where they have more competitors although this relationship does not always hold (MacArthur et al. 1972).
Figure 1.27. Relationship between number of endemic species and island area for revised plants and mammals, birds and reptiles.
From Anon. 1982b
Figure 1.28. Number of species of milkweed butterflies found on Sulawesi islands.
After Ft. Vane-Wright (pers. comm.)
Wallace's Line
It is an enigma that educated people in Indonesia know about Charles Darwin, even adopting his surname, and yet know nothing about Alfred Wallace, his contemporary and and fellow Englishman. Darwin is usually given credit for formulating the theory of evolution by natural selection, but the primary reason he was encouraged to write down his thoughts was because Wallace had written to him from Indonesia enclosing a manuscript showing that he had reached more or less the same conclusions as Darwin. Both men had been stimulated by what they had seen while travelling, and the interpretation of their observations did not match with the contemporary wisdom concerning the creation of the world. Wallace had been particularly fascinated by a visit to Sulawesi and the first manuscript he sent to Darwin was written in Ternate after leaving Manado. The thoughts of these two men on the distribution and evolution of species turned modern thinking upside-down and Wallace's contribution should not be underestimated.
Whereas Darwin was well-educated and came from a rich family, Wallace left school at 14, in 1837, and eventually earned a living by collecting animals in remote areas of the world to sell to museums. He spent eight years in Sarawak and Indonesia (from Sumatra to Irian Jaya) and the account of his travels totalling about 22,000 km makes splendid reading (Wallace 1869).
In a letter written in 1858, Wallace expressed his view that the Indonesian Archipelago was inhabited by two distinct faunas, one found in the east, one in the west. The following year he defined these two regions, based on the distribution of birds, by placing the boundary between Lombok and Bali and between Borneo and Sulawesi. He was struck that Borneo and Sulawesi should have such different birds and yet be separated by no major physical or climatic barrier. He believed that Borneo, along with Java and Sumatra, had once been part of Asia, and that Timor, the Moluccas, New Guinea and perhaps Sulawesi had once been part of a Pacific-Australian continent. The fauna of Sulawesi seemed so peculiar that he suspected it might have been connected with both the Asian and the Pacific-Australian continents (Wallace 1859). He insisted that an explanation of the origin of the fauna of Sulawesi would have to accept that there had been vast changes in the surface of the earth, a concept which challenged the established view but which we now know to be true (p. 2). The line that Wallace drew east of the Philippines, through the Makassar Straits and between Bali and Lombok (Wallace 1863) came to be known as Wallace's Line. In 1910, three years before he died, Wallace decided that the predominance of Asian forms on Sulawesi should be reflected in the Line being moved east of Sulawesi (Wallace 1910). Many other analyses have been performed on the distribution of animal species resulting in several different Lines (fig. 1.29) (Simpson 1977). Weber's Line attempts to delimit the boundary of faunal balance, that is, where the ratio between Asian and Australian animals is 50:50 (Weber 1904). Weber used molluscs and mammals in his analysis but the exact position of the Line differs from one group of animals to another. For example, Asian reptiles and butterflies penetrate further east then do its birds and snails. Lydekker's Line delimits the western boundary of the strictly Australian fauna in much the same way as Wallace's Line delimits the eastern boundary of the Asian fauna; both these Lines effectively trace the 180-200 m depth contours around the Sahul and Sunda continental shelves respectively. The area between these two Lines has been nominated as a separate region, subregion or transition area called Wallacea (Dickerson 1928). This concept was first suggested by Wallace in 1863, but has been strongly criticized as the area does not comprise a homogenous fauna, and there is no gradual change in species composition across it; instead there are large number of endemic species (Stresemann 1939; Simpson 1977). The name Wallacea should be retained, but to describe the area between the Oriental and Australian regions rather than as the name for a strict biogeographical entity.
Figure 1.29. Biogeographical Lines through insular Southeast Asia.
From Simpson 1977
The concept of Wallace's Line has fascinated biogeographers and it has been found that its validity differs between groups of animals and plants. For example, an early analysis of Sulawesi's flora showed similarities with Borneo, Sumatra and Java, rather than with the Moluccas and New Guinea (Lam 1945). This analysis was based, however, on a limited number (about 700) of species. An analysis at the generic level of the whole Malesian flora25 demonstrated the existence of three provinces in Malesia, of which East Malesia comprises New Guinea, the Moluccas, and Sulawesi (van Steenis 1950). A recent analysis of 4,222 species in 540 genera that have been subject to recent taxonomic revisions revealed that the Sulawesi flora was most closely related to the floras of other relatively dry areas in the Philippines, Moluccas, Lesser Sunda Islands and Java (van Balgooy in press). There is no clear affinity between Sulawesi and the islands east or west of it, but the flora of the lowlands and of the ultrabasic soils show a stronger similarity with that of New Guinea, whereas the montane flora (1,000 m and above) shows a stronger similarity with that of Borneo26 (fig. 1.30). The higher the altitude, the greater the distance between areas of similar altitude, and the less chance there is of receiving plants from similar habitats. This would explain the greater proportion of Bornean plants on Sulawesi mountains, but the greater affinity with New Guinea flora among the lowland plants may be a result of New Guinea having more relatively dry areas than Borneo and therefore being a more suitable source of plant species. Some of these may have been brought to Sulawesi via the 'Sula Spur' (p. 6), whereas others could have island-hopped. An examination of the percentage of taxa that do not cross imaginary lines between or within landmasses in a given direction revealed that the strongest such 'demarcation Line' was for plants of western origin between Borneo and Sulawesi. About 50% of the non-endemic plant species of Borneo do not occur in Sulawesi. This suggests that the Makassar Straits have been a barrier to dispersal for a very long time. Interestingly, however, this Line is very weak when considering non-endemic plants of eastern origin crossing from Sulawesi to Borneo. The easiest routes by which plant species appear to have entered Sulawesi are those via Java and the Lesser Sunda Islands and via the Philippines and Sangihe. The former route argues for the existence of an island chain between Java, Lesser Sundas and Sulawesi in the not too distant past.
When specific groups of animals or plants are examined, many interesting distribution patterns are found. The percentage of a total species list shared between neighbouring islands (fig. 1.31) indicates a generally closer affinity between Sulawesi and islands to the east but this is at least in part an artefact of the relatively impoverished floras and faunas to the east.
The mountain flora of Sulawesi is derived from two sources: those which originated locally (autochthonous) and those for which the centre of origin is outside the area concerned (allochthonous) (van Steenis 1972). The allochthonous flora, although a minority of the total mountain flora, allow hypotheses to be made regarding its origin. This part of the flora belongs to genera whose species are found only in cold climates (i.e., microtherm species), and in the tropics they are generally found only in the subalpine forests on mountains 2,000-2,500 m high. These genera, such as Rhododendron (Eric.) and Gentiana (Gent.), are found in many tropical and subtropical countries yet none can tolerate a hot climate. Soils seem to have little or no influence on the distributions since a single species will be found on soils originating from igneous, sedimentary or recent volcanic parent material. The age of the rocks does seem to be relevant, however, with some species being absent from recent volcanic soils (van Steenis 1972).
Figure 1.30. Floristic affinity of Sulawesi to Borneo (solid line) and New Guinea (dashed line) for each 500 m increase in altitude.
From data in van Balgooy in press
From an analysis of the distribution of about 900 of these cold-adapted mountain species, it has been concluded that there are three tracks by which plants arrived in Sulawesi during some period or periods in the geological past (fig. 1.32) (van Steenis 1972). Continuous ranges of high mountains do not, of course, exist along the entire lengths of these tracks. During the coldest times of the Pleistocene the mean temperature dropped only about 2°C (p. 18) which, with rates of temperature change being about 0.6°C/100 m (not necessarily applicable at that period) is equivalent to a drop in the levels of the forest zones of 350-400 m. The number of suitable mountain tops would obviously have been greater during the cooler periods thereby forming more 'stepping stones' across which plants could disperse.
Palms are a useful group for biogeographical study because their genera, at least, are well known and they represent an ancient group of plants in which several genera had evolved by the Oligocene (30 Ma ago) (Dransfield in press). Only two genera, Gronophyllum and the monotypic Pigafetta, are found in Sulawesi but nowhere further west. This is surprising in the case of Pigafetta because it is a tall fecund palm of secondary growth with small seeds (p. 396). There are, however, 13 genera of palms that are found no further east than Borneo illustrating again that Wallace's Line is most noticeable travelling west to east. Two genera, the spiny Oncosperma and the fan palm Pholidocarpus, cross Wallace's Line from Borneo to Sulawesi but are found no further east, and another sixteen are found in Sulawesi and to the east and west. Of these, the rattans belonging to Calamus, the undergrowth palm Licuala, and the tree palms Cyrtostachys, Areca, and Livistona exhibit a peculiar distribution having species in Sunda-land and New Guinea, but few or none in Sulawesi (Dransfield 1981 in press). Since it is known that Sulawesi was perhaps drier in the Pleistocene than were the continental landmasses (p. 18), it may be that some species became extinct, or that conditions for the evolution of species there were less suitable.
Figure 1.31. Species overlap for plants and three animal groups between Sulawesi and neighbouring islands. Figures are the percentages of total number of species recorded from a pair of areas that are shared between them.
From Anon. 1982b
Figure 1.32. The three tracks by which mountain plants arrived in Sulawesi.
After van Steenis 1936, 1972
A similar distribution is found among the spiny eels (Mastacembel-idae) in which four species are found in Borneo, three in Halmahera, but none in Sulawesi. Fish are in fact a useful group to examine with respect to different opinions concerning the narrowing or closing of the Makassar Straits during the Pliocene. For this it must be understood that freshwater fish can be divided into three ecological groups:
• those confined to freshwater with no tolerance to brackish or salt water;
• those generally encountered in freshwater but showing some tolerance to salt; and
• those with considerable salt tolerance which either migrate between freshwater and marine habitats or are of marine origin and have colonized freshwater habitats (Darlington 1957). This grouping is particularly useful since there are strong correlations between the ecological and systematic groups (Myers 1949; Cranbrook 1981).
The fish fauna of Sundaland has many species from the first two groups whereas Sulawesi has no strictly freshwater fish, only a few with a little tolerance to salt, and the majority are from the third group. The Makassar Straits or Wallace's Line therefore seems to separate two distinct fish faunas and it is therefore unlikely that Sulawesi and Borneo were ever a single landmass.
Freshwater fish may not have been able to reach Sulawesi but all major terrestrial animal groups have. Amphibians, for example, have no indigenous species in common with Borneo but of the endemic species four are related to Sundaland species and two have Papuasian species (p. 47). Two of the three frogs that are found east of Sulawesi may in fact have evolved on Sulawesi and subsequently dispersed (Cranbrook 1981). Twenty-three lizard species found on Sulawesi are also found west of Wallace's Line (C. McCarthy pers. comm.). Similarly, of Sulawesi's 63 snake species, 38 are found both sides of Wallace's Line but only two are found in New Guinea and not in Sundaland (den Bosch 1985).
The tortoise Indotestudo forsteni was thought to be of particular interest because it was the only land tortoise known from both sides of Wallace's Line having been first found in Minahasa and Halmahera. It now appears that it is the same species as the later-described I. travancoria of eastern India, Thailand and Burma, and was brought from there by traders as a food animal (Hoogmoed and Crumly 1984).
The birds of Sulawesi are predominantly western with 67% of the species having affinities with the Sunda region (Mayr 1944). One, the Sulawesi Roller Coracias temmincki, in fact has no relatives in Sundaland and all the other members of its genus are found in Europe, mainland Asia and Africa (Klapste 1982).
Only two of the non-flying mammals on the mainland, the cuscuses, are clearly of Australian/New Guinea affinity;27 the remainder, including the endemics, have their origin in Asia. Wallace's Line delimits the eastern boundary of the distinctive Sundaland fauna comprising moles, flying lemur, tree shrews, lorises, gibbons, pangolins, porcupines, dogs, bears, otters, weasels, cats, elephants, tapir, rhinoceroses and mouse-deer. Thus the fauna of Sulawesi has closer affinities with the Sunda than with the Philippines, Lesser Sundas or Sahul region, but cannot really be said to be part of it.
In general, Wallace's Line is not as clear a demarcation line for invertebrates as it is for vertebrates but there are many fewer genera in Sulawesi than in islands to the west (Gressitt 1961). For example, there are about 1,200 species of butterflies in the Malay Peninsula, 850 in Borneo and New Guinea, but only 450 in Sulawesi (R. Vane-Wright pers. comm.). As with the palms discussed above, the relatively low number of species may be the result of a previously unfavourable climate. Cicadas are poor dispersers and so their distributions show more detectable patterns than do many other groups. The cicadas of Sulawesi are largely of western origin but there are relatively large numbers of endemic species and some endemic genera (Duffels 1983). The same general pattern (major affinity with Borneo, relatively depauperate, but high level of species endemism) can be observed in other groups such as ground beetles, tiger beetles (N. Stork pers. comm.), and pond skaters/water striders (D. Polhemus pers. comm.), but the affinities of dung beetles Scarabaeidae, for example, are by no means clear (J. Krikken and H. Huijbregts pers. comm.). There are now about 200 species of ground beetle known from Bogani Nani Wartabone National Park and although less than half the total known from Borneo, this is still a massive total for such a relatively small area (N. Stork pers. comm.). This needs to be investigated further.
A detailed analysis of butterfly and moth relationships has shown that Sulawesi species are most strongly associated with the fauna, not in Borneo, but in the Philippines, reflecting the ancient connection through the Sangihe Islands (Holloway 1987). As shown above, the plants of Sulawesi also show a greater affinity with the north, east and south than with Borneo, and the difference in affinity of butterflies when compared with other invertebrates, may be a result of the caterpillars being dependent on specific living plants rather than on detritus. The moth families examined in most detail showed virtually no elements from the direction of Australia possibly because this moth fauna had become established by the time the connection through the Sula Islands occurred (Hollaway in press). All these analyses are hampered to some extent, however, by a generally poor knowledge of the Moluccan fauna.
One recent addition to the butterfly fauna of Sulawesi was not through human agency but through natural dispersal. The range of the Monarch butterfly Danaus plexippus used to be restricted to eastern and western North America where it performs remarkable annual migrations between the north and south of its range covering up to 3,000 km in a year travelling up to 125 km per day. This remarkable flying ability and the relatively long life of the adult has meant that when blown off course by strong winds it has reached and successfully colonized new areas. It first reached Hawaii in 1845, Australia in 1870, and Manado in 1873. It has not spread widely within Sulawesi, however, and was not collected by entomologists in or around Bogani Nani Wartabone National Park only 130 km from Manado (R. Vane-Wright pers. comm.).
As might be expected there is no obvious difference in the composition of coastal marine faunas either side of Wallace's Line. Corals, for example, show a great similarity between the species inhabiting reefs throughout the Indo-Pacific region and a general absence of endemic species even in remote areas. Most genera have, in fact, been present for 20-40 million years and some species are probably this old too. This is probably a result of the high-frequency fluctuations in sea-level in the Quaternary (p. 16) which alternately exposed and covered coastal regions. There was probably simply not enough time for populations of long-lived corals to complete many generations before their descendants colonized new habitats, and this may have maximised variation within a species rather than resulted in the appearance of new species. This contrasts with the marked Quaternary speciation observed for more mobile organisms such as fishes and crustaceans (Potts 1983, 1984).
Biogeographical Differences within Sulawesi
The fauna and flora of Sulawesi are far from being homogeneous and evenly distributed, but although the geological history might be expected to have influenced the distribution of species, clear patterns cannot now be seen. The flora of the east peninsula might be expected to have close affinities with the Moluccas, but it is in fact closest to the central and west regions of Sulawesi and to Borneo than to New Guinea. Indeed, all areas of Sulawesi except one are closer to each other in floral composition than to regions outside Sulawesi. The exception is the southwest peninsula whose lowland (<500 m) flora is most closely related to the Lesser Sunda Islands (van Steenis 1972; van Balgooy in press). This could be a result of dispersal from Flores via Kalaotoa, Tanahjampea and Salayar or from Sumbawa via the Sabalana and Postilyon Islands. The tip of the southwest peninsula has a dry climate similar to the Lesser Sunda Islands, bounded to the north by wetter, less seasonal climates (p. 24).
More recent geological history is reflected in the distribution of species within a particular genus. The most striking example of this allopatry (non-overlapping distribution of related species or subspecies) is perhaps the distribution of macaques. The number of species living on Sulawesi is the subject of some debate but four species and seven subspecies seems to be agreed by many (Groves 1980b). These macaques may have evolved from an ancestor of the pig-tailed macaque Macaca nemestrina (found now in Sumatra and Borneo), which crossed to Sulawesi, by rafting or by island hopping from Borneo or Java, possibly in the Middle Pleistocene (Fooden 1969; Takenaka 1982), but their origins and relationships have yet to be determined with certainty (Takenaka 1982; Takenaka and Brotoisworo 1982; Takenaka et al. 1985). It is interesting that M. maura in the southwest has the most primitive characteristics, and M. nigra living furthest away in Minahasa, is the most specialized (Albrecht 1977; Groves 1980b). The original condition was probably a single species but with 'clines' or gradients of character variations existing throughout the range. Disruptions to this continuous distribution would have caused populations to evolve in isolation such that when the disruption was removed the populations were reproductively incompatible (Fooden 1969).
During periods when the sea-level increased by only 4 m (p. 18) the mountainous region south of Lake Tempe would have been cut off by a narrow straits running northwest-southeast, and a narrow isthmus would have been formed between Gorontalo/Limboto and Kwandang. These inundations would help to explain the separation of M. maura and M. tonkeana, and of M. tonkeana and M. nigra. The junction of the two subspecies of M. tonkeana may have arisen from the formation of a narrow isthmus between Tamba and Labuhanbajo (near the prominent Tanjung Manimbaya), and the two subspecies of M. ochreata would have arisen because of the narrow straits between northern Buton and the southeast peninsula. It is not clear, however, how subspeciation arose between the two M. nigra subspecies, or speciation between M. tonkeana and M. ochreata, although unproductive forests on the ultrabasic soils (p. 457) may have formed a biological barrier in the latter case.
Other examples of allopatry are the distribution of subspecies of the large carpenter bee Xylocopa nobilis and of species of the large pond skater/water strider Ptilonera (fig. 1.33) although not enough specimens are known to be able to determine species boundaries.
Another example of closely-related species replacing each other geographically on Sulawesi is sometimes quoted, that concerning the white-eyes of lowland forests: Zosterops anomala in the southwest peninsula, Z. atrifrons in the north peninsula, the central block and Banggai Islands and Z. consobrinorum in the southeast peninsula (Stresemann 1939-41; Lack 1971). This distribution has now been shown to be rather less well-defined than was previously thought (Holmes and Holmes 1985).
Within a well-known group such as the birds, a number of interesting distribution patterns can be seen within those species endemic to the main island. For example, some species are known from only a single peninsula (table 1.14). Half of the 88 endemic birds are found in all regions of Sulawesi and half have partially-restricted distributions. Thus five species are known from only the north peninsula, central area and southeast peninsula, two from only the central area and southwest and southeast peninsulas, two from only the central area and southwest peninsula, etc. As a result the number of endemic species is different between the main areas (table 1.15). Some of these totals may change as more field data become available, particularly from the east peninsula, but they serve to illustrate the apparent paucity of the birds in the southwestern peninsula.
The Sula Islands, to the east of the Banggai Islands, are administratively part of the Moluccas but biogeographically they are part of Sulawesi. Of the birds, there are twice as many species with Sulawesi affinities as with Moluccan affinities (Wallace 1862) and the birds of the Banggai Islands seem to be derived almost equally from Sula and the mainland (Eck 1976). The reptile fauna of the Sula Islands is essentially a poor Sulawesi fauna (Kopstein 1927), and the flying lizards Draco of the Banggai Islands are more similar to those of the Sula Islands than to the mainland (Musters 1983).
From Holmes and Wood 1979; K.D. Bishop pers. comm.
From Holmes and Wood 1979
Figure 1.33. Distributions of the five known species of large pond skater/water strider Ptilonera on Sulawesi. Side views of terminal segments of females' abdomen illustrate the considerable morphological variation. Note the single zone of overlap, in Lore Lindu National Park, a - Ptilonera laelaps, b - P. sumizome, c - P. oribasus, d - P. pamphagus, e - P. dorceus.
After Polhemus and Polhemus 1986
PEOPLE OF SULAWESI
Prehistory
Remains of proto humans Homo erectus from about 500,000 to 1 million years ago have been found in Java but nothing similar has been found in eastern Indonesia. The first traces of modern man Homo sapiens in the latter area have been dated to about 30,000 years B.P.28 These were populations of hunter-gatherer people who were present throughout Indonesia and who successfully crossed the sea between New Guinea and Australia before 35,000 years B.P. They were the direct ancestors of the Australoids, found today in Australia and the New Guinea highlands, and relatives of some of the inland forest tribes of Peninsular Malaysia and the Philippines (Bellwood 1980a).
Between 4000 and 2000 B.C., Austronesian-speaking people on Taiwan and mainland eastern Asia, with an economy based on plant cultivation, began to spread through the Philippines into eastern Indonesia and western Melanesia (the islands of the Bismark, Solomon and New Caledonia groups). This expansion was perhaps initiated by agricultural developments in southern China and Taiwan, and had, by 1500A.D., reached over half the circumference of the globe from Madagascar (colonized from Indonesia) to Easter Island (Bellwood 1980a). The people had taken with them domestic pigs Sus scrofa and dogs, pottery, bows and arrows, a tradition of thatched community houses, fishing and canoe transport with sails. They cultivated taro, bananas, breadfruit, sugarcane,29 sago and possibly coconuts. These people did not replace indigenous inhabitants but rather blended with them (Jacob 1967; Bellwood 1985), and their technological and cultural novelties were adopted. Tracing the migration is extremely difficult because racial history is very complex and some human physical characteristics are easily changed.30
The early setters of South Sulawesi are known from remains excavated in many caves in the southern half of the province (Sarasin and Sarasin 1905; Mulvaney and Soejono 1970; Heekeren 1972) and the most detailed records, indeed among the most detailed in Southeast Asia, come from three caves near Maros (fig. 1.34) (Glover 1976, 1977, 1978, 1979a, b, 1981; Burleigh 1981; Frank 1981; Glover, E. 1981; Mook 1981; Vita-Finzi 1981). The caves, in order of artefact age are Leang Burung 2, Ulu Leang and Leang Burung 1 and represent the period between about 30,000 and 8000 years B.P.. although there is a gap of 10,000 years between Leang Burung 2 and Ulu Leang and overlap between Ulu Leang and Leang Burung 1 (Glover 1977). The culture whose remains are found is sometimes termed 'Toalean' after a tribe of hunter-gatherer and occasionally cave-dwelling people encountered by the Sarasins in the hills half-way between Maros and Watampone. Photos of these people were printed in their book (Sarasin and Sarasin 1905). To-ala means 'forest people' and is a derogatory expression, and the To-ala may simply have been runaway Bugis debt slaves and landless villagers making a poor living in the forest. There is certainly no evidence to link them with the original cave inhabitants and in any case there is a gap of 2,000 years between the most recent, well-dated Toalean finds and the finding of the To-ala people (Mattulada 1979; I. Glover pers. comm.).
Figure 1.34. Caves with known prehistoric remains around Maros. 1 - Leang Burung 2; 2 - Leang Pattae; 3 - Leang Pettae Kere; 4 - Ulu Leang; 5 - Leang Burung 1. (Dark grey shade indicates hills.)
From Glover 1977
Southeast Asia during the mid-Holocene was home to two types of stone tool industry. The Hoabinhian industry was found in northern Sumatra and the Asian mainland, and is characterized by large choppers made from large split river stones. The other industry had its geographical centre in Sulawesi, typified by the 'Toalean' remains and is characterized by the production of numerous relatively small, fine flakes made from chert31 for use as knives, scrapers, etc. These small flakes, known as microliths, are of two main types: backed flakes which appeared about 6000 years B.P. and the specific 'Maros points' which appeared about 4600 years B.P. (fig. 1.35). Similar backed flakes and blades have been found in India and western Asia from about 10,000 years B.P., and in various parts of Australia from about 5000 years B.P. (Glover and Presland 1985) and before. Some of the flakes found in the Maros caves have a gloss resulting from being polished against the materials they were used to cut or scrape. The minute scratches on the glossy surfaces have been analysed and it has been concluded that between 31,000 and 19,000 years B.P. the inhabitants of Leang Burung 2 worked wood and other plant materials with a cutting motion and that the maximum diameter that could have been cut through was only 3 cm or 4 cm. Thus these rude tools could have been used for cutting strips of stems and leaves to make string, mats, baskets and simple weapons such as spears, but would not have been used for whittling or slicing to make more refined objects such as complex wooden points, spear throwers, harpoons, etc. However the tools found in Ulu Leang (dating from 9000 to 3000 years B.P.) could have been used for this finer work. Conclusions from such analyses need to be guarded because it is not possible to be sure that the remains found in the caves represent the whole arsenal or kit of tools used by these prehistoric people (Sinha and Glover 1984). There are groups of forest people today who have taboos about bringing certain tools or weapons into the living area, and this might also have been the case thousands of years ago. Alternatively, it may be just that certain tools were stored close to the areas (outside caves) where they were used.
Animal remains from Ulu Leang indicate that the most important prey were pigs and babirusa, followed by anoa, macaque monkeys, and small animals such as snakes, bats, rodents, cuscus, lizards, tortoises and squirrels. Bird and fish remains are surprisingly rare. Another very abundant food animal which was gathered around the caves was the snail Brotia perfecta. Over 90% of the shells have their dps broken off whereby the small amount of flesh could be sucked out (van Heekeren 1972). These are still found in rivers around the caves which suggests that the environment has not changed dramatically during the period in question.
Plants remains found in Ulu Leang deposits are seeds of sedges, wild grasses Panicum (Gram.), figs Firus (Mora.), Canarium (Burs.) and Bidens (Comp.), a weedy herb used by modern villagers to relieve coughs, toothaches and sore eyes and as a vegetable (Burkill 1966). In addition to the above species, remains of rice husks were found in deposits in a hearth, believed to be 1,500 years old but it is not known for certain whether cultivated forms were important in southern Sulawesi at this time (Glover 1979b, 1985). It is also very likely that the people gathered the large (up to 35 kg) surface tubers of a yam Dioscorea hispida (Dios.) which can still be found in the area and represented an important food for villagers during the Japanese occupation (Burkill 1966; S. C. Chin pers. comm.). The tuber and the foliage contain the poisonous alkaloid dioscorine which must be removed by rasping, pounding, grating and soaking, preferably in salty water. A piece of tuber the size of an apple is enough to kill a man if eaten raw (Burkill 1966).
Figure 1.35. Backed microliths and Maros points.
After Glover and Presland 1985
Two major prehistoric sites have been excavated in North Sulawesi: a coastal cave called Leang Tuwo Mane'e in the north of Talaud's main island Karakelang, and a shell midden (rubbish dump)32 at Paso on the southwest shore of Lake Tondano near some natural hot springs which would obviously have been attractive to early people. The cave deposits contain chert flakes from 4,500 to 6,000 years ago but made in a different style from the chert flakes found in South Sulawesi. In the upper layers, starting about 4,500 years ago, a thin and burnished type of pottery is found; about the same time that it appeared in Ulu Leang. The midden deposits date from about 7,500 years ago and contain scrapers made from obsidian33 flakes (Bellwood 1978). The animal remains found in the midden deposits are similar to these from Ulu Leang but the proportions and quantity of the different animals are markedly different. For example, there are four times as many pigs as anoa in Paso deposits, but fifteen times as many pigs in Ulu Leang as at Paso. There were many rodents and no tortoises at Paso but few rodents and many tortoises at Ulu Leang. At Paso the long bones of animals were broken into recognizable pieces but at Ulu Leang they were chopped up and unrecognizable (Clason 1979). Interestingly, when the Paso midden was being used, the surface of Lake Tondano was higher than it is now (Bellwood 1978), indicating that either some change has since occurred to the riverbed where the water flows out of the lake, or that the rainfall and hence average height of the lake surface used to be higher.
It can be concluded, then, that eastern Indonesia had a variety of stone tool industries worked by isolated communities of people who occupied a wide range of habitats: low swampy areas, steep rocky coasts, high mountains and inland lakes (Glover 1981). The people may have cremated their dead (Boedhisampurno 1982) and buried them in a flexed position. They painted pigs, not babirusa as stated elsewhere (van Heek-eren 1972), geometric designs and made stencils of their hands on the walls and ceilings of their caves. Some of these can still be seen in Leang Pettae and Leang Petta Kere caves in the Prehistoric Park near Maros (Soejono 1978; Anggawati 1985), but they are more or less impossible to date. Red ochre, of the sort used for the paintings, was found in deposits dating from 20-30,000 years ago but this might have been only for decorating the cave-dwellers' bodies. Paintings depicting hunting scenes, boats, and warriors on horseback (fig. 1.36) have been found recently in several rock shelters west of Raha on Muna Island (Kosasih 1983, 1984). These are probably relatively recent but the depiction of horses does not necessarily mark them as particularly recent. For example, horse sacrifices are mentioned by King Mulawarman of East Kalimantan in the 4th century in Indonesia's oldest inscription. Horses and goats would have been brought from western India as items of trade and diplomacy (J. Miksic pers. comm.). Horses are known to have been present in southern Sulawesi in the 16th century (Pelras 1981).
Remains from a Neolithic settlement have been found at Kamasi Hill near Kalumpang, a village 93 km upstream from the mouth of the Karama River in West Toraja (van Stein Callenfels 1951). Plain and decorated pieces of pottery were the most abundant artefact and are similar to remains found in China, Luzon, Vietnam and East Java. Human remains were found, as well as bones of anoa, wild and domestic pigs, and fish. In addition, the excavations revealed highly polished rectangular adzes, small chisels, ground oval axes (similar to those from China and Taiwan), spearheads (similar ones also found in Hong Kong) and stone rings. The minimum estimate for the age of the Kalumpang artefacts is between about 3000 b.c. and 5000 b.c. (Bellwood 1980b, 1985). A nearby site known as Minanga Sipakko is probably older but is less well known (van Heekeren 1972).
Figure 1.36. Hunters or warriors on horseback and man with spear as depicted in paintings on a wall in a rock shelter on Muna Island.
After Kosasih 1983
Shards of perhaps similar smooth, red, unglazed earthenware pots of about 40 cm diameter were found by members of Operation Drake in a cave near Kolonodale but the cave has not yet been properly excavated (Rees n.d.).
The cultural stage after the Neolithic was a form of 'Bronze Age' and Central Sulawesi is rich in artefacts of this period, notably huge stones or megaliths. These megaliths are large worked stones in the form of huge cylindrical vats, large statues, urns and mortars and are found primarily in Central Sulawesi (fig. 1.37) (Sukendar 1976; 1980a, b) but the meaning and creators of these stones are unknown. The decorations on the vats, which were probably multiple burial chambers, are of faces, figures, monkeys and lizards and most are near the village of Besoa, and are very similar to designs found in Laos:34 The statues are mostly larger than life, usually male (only one-quarter are female), legless and armless, and set upright in the ground (Kaudern 1938).
A megalithic culture is still alive (just) in Sa'dang To raja where the people celebrate by erecting large stones in rows or circle (Asmar 1978; Kadir 1980). Megalithic remains also abound in the hills north of Wattang Sopeng as far as Sengkang, and the traditions are still practiced by the Amparita people of Sidenreng (van Heekeren 1958b; MacKnight and Bul-beck 1985). Bronze used to be a highly valued metal and a magnificent, large (115 cm diameter) bronze kettledrum decorated with stylized frogs has been found in Salayar Island.35 It is typical of the Dongson culture, the centre of which was in Vietnam, and may have been made about 2,000 years ago. The presence of this drum may be evidence of a significant settlement in Selayar during the first millenium A.D.,possibly inland from the relatively poor soils of the west coast where most of the population lives today. The houses of the Toraja are almost identical to house motifs used to decorate some Dongson kettledrums. Bronze was regarded as a very special metal and was greatly prized. A description of Minahasan women in 1679 states that the women were seen wearing up to 10 kg of bronze. Bronze was also felt to have magical, particularly protective, powers, and many bronze axes were taken by the Dutch after they had defeated the Gowa army on Buton Island in 1667. In the Luwu and Wotu areas at the top of the Bay of Bone, bronze axes, forged generations earlier, were believed to be teeth of a spirit (van Heekeren 1958b).
Figure 1.37. Locations where megaliths are found in Central Sulawesi.
After Kaudern 1938
Near ports or other centres, rapid cultural changes occurred perhaps 1000 years before they occurred in remote tribal areas. If prehistory is taken as referring to that period before events and thoughts are written down, then there are still groups of people in remote areas of Sulawesi who live in prehistory. The official number of these tribal people in South Sulawesi is 60,000 (e.g., the Sareung, Bentong and Towala), in Central Sulawesi 50,500 (e.g., the Tolare, Towana and Sea-sea), in North Sulawesi 10,000 (e.g., the Gorontalo), and in Southeast Sulawesi 5,800 (e.g., the Tolaki, Tooere and Koro), but only a very small proportion of these are beyond the influence of government institutions. The major trends in dealing with these people are to teach and to develop, and little or no effort is made to learn from them. They hold within their cultures more information about their various environments and about forest products (drugs, rotan, semi-domestic crops, etc.) than could be gleaned in a decade of research, yet this knowledge is being lost to Indonesia and the rest of the world. The only in-depth study of a tribal group in Sulawesi seems to have been on the Towana of the eastern peninsula (Atkinson 1979, 1985), but this concerned their sociology rather than their ecology.
Impacts of Prehistoric Man
The activities of primitive man probably affected populations of animals and plants in four ways (Rambo 1979). First, he exerted direct selection on the species of prey he hunted. In many areas of the world early man has been implicated in the extinction of the giant members of the Pleistocene fauna (these were either absolutely large or large by comparison with living relatives). In Sulawesi, this 'mega-fauna' was represented by at least Heekeren's giant pig, the giant tortoise and the stegodont (p. 34). Both giant tortoises and elephant-like animals have suffered greatly from man and these used to be found in North America, Europe, Africa and Asia. The cause of their extinction is not known but if man were present when these animals roamed Sulawesi's forests (a situation which has yet to be confirmed) no evidence has been found in cave or other deposits. There is no evidence that prehistoric man had any greater effect than causing local extinctions, probably more as a result of forest clearance than overhunting. The only animals known from cave deposits in South Sulawesi which have not been recorded in historic times are two endemic rats, one of which is known from just two specimens collected in Central Sulawesi in the early 1900s. The absence of records from South Sulawesi is probably partly a result of the very small area of lowland forest remaining (p. 98) and partly because there have been no serious studies of small mammals in the province (Musser 1984).
The relative numbers of the different animal species found in the cave deposits cannot be used to judge whether the Toalian hunters were selective or whether they simply caught and ate anything they could catch. This is largely because the excavations made before the war were not done with the rigorous techniques used in recent years and it is likely that only a proportion of the large bones were collected and many of the smaller fragments were overlooked. Many deposits have also since been mined for phosphate fertilizer (p. 553).
The earliest Australoid people would have used spears and cord snares or set traps of camouflaged pits to catch their prey, and the large ground-living species such as pigs, anoa and babirusa were, not surprisingly, the most common prey. Even with the skill and knowledge of a forest-bred hunter it would be scarcely possible to cause anything but local reduction in numbers. They would, however, have produced fear and avoidance responses in their prey. Fear and avoidance of man is a learned trait: the Tasaday, a 28-member tribe in the forests of Mindanao who had had no known contact with other humans before 1962, were able to approach deer, which they did not hunt, and to stroke them (Nance 1975). In the absence of feline predators (such as tigers, clouded leopards and jungle cats), or large eagles, man as a predator would have been a novel component of the ecosystem. The monkeys and pigs may have responded by forming large social groupings because the more eyes and ears a group has the more likely it is that a predator will be detected (van Shaik 1983). Squirrels and birds may have developed more cryptic behaviour.
Second, he dispersed trees by picking fruits in one place and discarding or voiding them in another. An early form of agriculture would have been the accidental or deliberate sowing of tree seeds in the same area, thereby reducing the distance between fruit trees. This phenomenon can frequently be seen at traditional resting places such as ridge tops or along forest paths, where rambutan, durian and other fruit trees can often be found.
Third, when he was able to fell trees, he modified habitats. The long sequences of remains from Talaud and Maros are exceptional, however, in the almost total absence of ground stone axes and adzes. They do appear in the Kalumpang deposits but these date from rather later. So, not only were these tools possibly not part of the tool-kits of the Australoid people but the immigrant Austronesians (or the culture they introduced) apparently did not use them either.36 It is possible to conclude then that the activities of the early horticulturalists in Sulawesi did not involve large-scale forest clearance and cultivation probably took place in small fixed plots next to their dwellings (Bellwood 1980a, 1985). It was probably iron, introduced about 1,500 years ago, rather than stone axes that gave man an efficient means of felling trees and it was probably at about that time that the process of forest clearance truly began. Some of the indigenous wildlife, such as the endemic pig, may have benefitted from the greater areas of relatively succulent secondary growth in the 'edge habitats', but this would also have exposed them to greater hunting pressure. There is evidence of forest clearance from about 9,000 years ago in New Guinea, and about 4,000 to 7,500 years ago in Sumatra.
Fourth, he carried crops, pets/live foods and pests with him on his travels. Among his crops would have been plants whose edible parts were also the regenerative parts, such as yams37 and tubers. Prehistoric man almost undoubtedly introduced the deer Cervus timorensis, common palm civet Paradoxurus hermaphroditus, Malay civet Viverra tangalunga, Prevost's squirrel Callosciurus prevostii, some of the rats, and jungle fowl or wild chicken Callus gallus, as well as the later domesticated animals such as buffaloes, dogs, horses, ducks, etc.
Prehistoric man on Sulawesi had the distinction of having brought into domestication the endemic pig Sus celebensis some time in the early Holocene. This pig accompanied early adventurers or was traded, and morphological analyses of skulls show that Sulawesi pigs are still found in a domestic or feral state in Maluku, Flores, Timor and, amazingly, Simeulue, an island off the west coast of Aceh in northern Sumatra. Unlikely though this may seem, it is of great interest that the Simeulue language is said to be closest to a Bugis-type language! The only other pig brought into domestication was the common wild pig Sus scrofa which was apparently superior to Sus celebensis which it replaced, ousted, or with which it hybridized (Groves 1981).
History
It was in the 13th to 15th centuries that the famous Bugis kingdoms of Bone, Wajo and Soppeng, and the Makassarese kingdom of Gowa arose. During this period the power of the largely Buddhist empire of Srivijaya in Sumatra was waning, and the power of the Hindu Majapahit kingdom of East Java was in ascendency. Around this time the southwest peninsula supported a relatively small population scattered in small settlements centred on resource-rich geographical features such as lakes, rivers and estuaries to supplement the produce of swidden agriculture. Most contacts were with traders from northern Sulawesi, Maluku and Sumbawa rather than from Java (MacKnight 1983). Great quantities of ceramics were imported from China and elsewhere through these trading links, and were used in ritual, two-stage burials (still practiced by some Toraja people today) up to at least the middle of the 17th century (Hadimuljono and Muttalib 1979). Makassar, Luwu, Bantaeng and Salayar were listed as dependencies of Majapahit and their ships are described in a 14th century Javanese poem as harassing trade ships from Malacca. Bugis states were trading with Arab ships during this period. Of these states, Luwu was the first to establish a kingship, and dynasties established later elsewhere were believed to be quasi-divine though still subject to the same forces which influenced their subjects. The 'high-culture' centres of this period in Indonesia tended to be coastal and concerned largely with trade but because of its narrowness, the majority of the southwest arm of Sulawesi was affected. The history of this period is composed mainly of genealogies and was written on leaves of lontar palms Borassus flabellifer (Mattulada 1985) as was the custom in the drier and more seasonal parts of Asia until the Portuguese introduced the techniques of paper manufacture (Whitmore 1977).
At this time there were about 50 kingdoms in the southwest peninsula (although Bone, Wajo and Luwu were clearly dominant), and the kings and their people worshiped images-presumably a mix of the traditional animistic beliefs and traces of Hinduism. These kingdoms, most of them little more than a small town with a feudal chief ruling over a subservient hinterland, had their main settlements near river mouths. These settlements probably comprised 100-200 households and were built backing on to the river (Mattulada 1978).
The name Celebes was first used by the Portuguese historian Tome Pires, whose discerning and comprehensive accounts of eastern Asia in his Suma Oriental were written in India and Malacca between 1512-15. He mentions that the Portuguese sailed to the Spice Islands or Moluccas via Singapore, Tanjung (?)Puting (Central Kalimantan), Buton and sometimes Makassar. Celebes, Banggai and Siau were said to produce foodstuffs and gold for the Moluccas. 'Celebes' applied initially only to the northern point of the north arm (Punta de Celebres) and was first marked on a map in 1524. Only later did it come to apply to the whole mainland (Pires 1944).
There are four hypotheses concerning the derivation of the word Celebes. First, the Bugis word 'selihe' (the 'h' is sometimes pronounced as an 'r') means sea current, so Punta de Celebres would mean Point of Currents (Pires 1944). Second, it derives from 'sula' (island) 'besi' (iron), for the area around Lake Matano in the centre of Sulawesi is one of the richest deposits of iron ore in Southeast Asia. Third, it derives from 'Si-lebih' or the one with more islands (Crawfurd 1856); and fourth, it derives from a corruption of Klabat, the name of the impressive volcano north-east of Manado which dominates the Minahasa landscape (Sarasin and Sarasin 1905; de Leeuw 1931). The modern name Sulawesi clearly derives from Sula-besi ('b' and 'w' are frequently transposed).
The first writer to mention the northern islands was Pigafetta who wrote the journal of the first round-the-world voyage led initially by the Spanish captain Magellan. Magellan himself died in the Philippines in early 1521, but later that year his ships sought a passage from there to the Moluccas. Having left southern Mindanao, they encountered the small Kawio Islands and then sailed to Sangihe Island which was said to be "very beautiful to look at". The island was divided between four kings. The boats then passed the islands of Kalama, Kanakitang, Para, Sanggeluhang and Siau, the last of which was ruled by a king called Raja Ponto. After reaching Ruang, they turned south-east (Pigafetta 1906), never setting eyes on the Sulawesi mainland only 80 km distant.
Gowa was the first dominating power on Sulawesi, due in part to the alliance made with the Bajau or sea nomads (see below) who furnished the kingdom with a wide range of marine produce. Gowa dominated the whole Makassar-speaking region and its capital, Makassar, was a cosmopolitan trading centre. Malay traders began living there in the middle of the 16th century, followed by the Portuguese. The Dutch opened a trading post and factory in 1607, the English in 1613, and the Danes in 1618. Agents from France, the Philippines, India, Arab countries, Aceh and China were also to be found (Reid 1983).
The Dutch displaced the Portuguese from the Moluccas and tried to exercise a monopoly on the spice trade. The importance of Makassar was, however, that people were able to buy spices there without having to tangle with the Dutch. Malay and Gowa ships sailed to the Moluccas with food and foreign goods and traded them for spices which were then sold in Makassar. Makassar became even more popular when it was made a free port.
Catholicism preached by Portuguese missionaries from the 1540s did not make a great impact, and later kings sent envoys to the Malay Peninsula to find Islamic teachers. Islam was accepted as the state religion by the twin kingdom of Goa-Tallo in 1605 and this was the last major Indonesian state to do so.38 Gowa and the states of Bone, Wajo and Sopeng that formed a Bugis alliance were often warring within and between themselves and these last three later accepted Islam at different times after the powerful Gowa. Acceptance of Islam at that time was tantamount to acceptance that they had to be allies (or vassals) of Gowa. Gowa influence spread slowly to south-east Borneo, Buton, Palu, Toli-Toli, Lombok, Flores, Sum-bawa and Timor (Hadimuljono and Muttalib 1979).
Inland, the Toraja people of the central mountains traded with the Bugis but their supposed headhunting traditions tended to minimize the degree of contact. The present-day Aluk-Todolo religion of western Toraja probably resembles the early beliefs of all the inland tribes of that region.
The Minahasans, who are physically most closely related to the Filipinos,39 have never been subject to dynastic rule although it is believed that there was pressure to institute kings in the early 7th century. In response to this, a large open meeting is said to have been held in about 670 A.D.around the stone known today as Watu Pinabetengan,40 and the government of the independent states was discussed (Taulu 1981). In contrast, Bolaang Mongondow and all the islands north of Manado had kings and slaves, and were subject to the rule of the Portuguese-influenced Sultan of Ternate (an important 'spice island' on the west coast of Halmahera, 300 km to the east of Minahasa), as were the Banggai Islands and the mainland area of Luwuk. The first Europeans to live in Minahasa were Spaniards from Magellan's boats escaping from the Portuguese in Ternate in about 1524, although it seems that Portuguese ships had occasionally called before this to buy rice (Jones 1977). Christianity was brought successfully to Minahasa and Sangihe-Talaud in the 1560s by Portuguese missionaries, and the king and people of Siau accepted Christianity in 1568, the same year that Indonesia's third oldest church, the Evangelical Church of Minahasa, was founded. At about the same time most people in Bolaang Mongondow and Gorontalo were accepting Islam, although some had been converted earlier by Bugis traders (Jones 1977). In 1574, Spaniards from Manila sent envoys to Minahasa and in 1619 a pastor was sent out. Eleven years earlier, the Dutch East-India Company built a wooden fort that doubled as a trading post near Manado. In 1643 the Spaniards attempted to impose a half-Spanish king on the Minahasans who were adamant that they wanted to retain their independent status. This spurred the Minahasans to call in the Dutch and ask for a defence and trading alliance (although this was not formally concluded until 1679) (Taulu 1981). The history of this period was written onto a rock face at Watu Pinantik by an ousted Spanish priest, and these writings can still be discerned (Taulu 1981). The trading agreement later turned sour and many states actively opposed the Dutch. This came to a head when Dutch arms and ammunition were seized and taken to Moraya Fort on the shores of Lake Tondano. The Resident began a siege and the will of the Minahasans broke only after he ordered a flock of pigeons with burning palm-fibre tied to their legs to be released; these landed on the thatched roof of the fort and razed it to the ground (Taulu 1981).
The royal court of Makassar was cultured, tolerant, and secure in its success as one of the great entrepots of Southeast Asia (Reid 1983). It was also a thorn in the flesh of the Dutch and as a result its zenith lasted barely 50 years. In 1660 the Dutch destroyed six Portuguese ships in Makassar's deep-water harbour, captured the fort and made an alliance with the Sultan of Gowa. Later they schemed with the Bugis state of Bone, in particular with Arung (Prince) Palakka, against Sultan Hasanuddin of Gowa. They began fighting in 1666 and the Bungaya Treaty was signed a year later forbidding all foreign traders to live in Makassar, and transferring jurisdiction over Bulukumbu-Bira, Maros, Bantaeng and the Makassar fort (Hadimuljono and Muttalib 1979; Andaya 1981). There were several uprisings in Makassar during the following century and the town was totally destroyed at least twice. At this time the Dutch had little or no interest in the land they now ruled, because their main aim had been the suppression of a competitor. The main export during the 18th century had been slaves. After the subjugation of Makassar many Bugis, particularly from Wajo, emigrated and founded royal dynasties in Kutai (East Kalimantan), Johore and Selangor (Peninsular Malaysia). In 1737 the Wajo king of Kutai liberated the Wajo state from Bone, which had become increasingly more powerful, and established the most successful Indonesian maritime commercial operation of the 18th and 19th centuries.
Since most of the written early history of Sulawesi and neighbouring islands was concerned with maritime trade, the life of the traditional nomadic people of East Indonesian seas, the Bajau, is quite well known. The Bajau used to be a group of maritime hunter-gatherers and although now largely settled, they can still be met with around Sulawesi's eastern islands. In contrast to the Bugis who, although seafaring, were based on land or in beach settlements, the Bajau spent more or less their whole lives on or around boats. Another major contrast was the Bugis habit of leaving their family behind when travelling whereas the Bajau always travelled as a family.
The Bajau economy was based primarily on collecting, preparing and selling 'tripang', a collective name for a few types of edible sea cucumbers, a group of echinoderm animals (which also include starfish, brittlestars, and sea urchins) (p. 227). Sea cucumbers are bottom-dwelling (benthic) creatures and are either collected at low tides or dived for in shallow water. The most sought-after tripang were apparently painstakingly prepared by a Bajau group known as the Turijene who were based in the Spermonde Archipelago, particularly around the island of Kuring Aring, 17 km west of Ujung Pandang. All the tripang, together with tortoiseshell, pearls, birds' nests and giant clams, were sold to Bugis seafarers for the China trade in food, tonic and medicine, and this formed the basis of the interdependence of these two groups. Tripang was not eaten by any Malay people even when other food was scarce and, strangely, the Bajau rarely if ever used nets or traps for catching fish, preferring instead to use harpoons and spears.
The narrow entrance to Kendari Bay was discovered in 1839 at which time the east and southeast of Sulawesi were very sparsely populated (fig. 1.38). Inland people in this area have moved south only in the last century or so. A few Bugis and Gowan settlements were established and were dominated alternately by maritime powers such as Luwu and Ternate. Kendari Bay was the only major trading focus in the southeast but this was occasionally abandoned because of the tribal disputes and raids on traders by the inland people. The Bajau did not have a particularly cordial relationship with the Bugis and Gowan traders, but generally had peaceful relations with the inland Tolaki people. This was not always the case, however, and it seems that the Bajau were probably in frequent conflict with, for example, the Toloinang of south Tomori Bay (Sopher 1978).
The Banggai Islands, or more specifically the small islands between Peleng and Banggai, were another traditional centre of Bajau activity; in the 1840s, 100 to 150 boats gathered there at certain times of the year. The Banggai people, of the Towana group, accepted the presence of the Bajau and traded produce from their dry-field agriculture with them. Other centres of Bajau activity were Tomori Bay (west Tolo Bay), Dondo Bay (off Toli-Toli), the Sabalana Islands, and Wowoni Island (Sopher 1978).
One of the earliest dates in the history of Central Sulawesi was 1555 when the Portuguese built a fort at Parigi. In 1602 two Muslim Minangk-abau traders from West Sumatra settled in Palu and Parigi to develop commercial interests and to propagate their faith. In 1680 Palu and Toli-Toli were under the influence of the Sultan of Ternate but the Dutch East India Company had growing interests and established an outpost in Parigi in 1730 to trade in gold (Davis 1976). Meanwhile the people in the hills lived in fortified stockades, and the earthworks that once surrounded these stockades are still visible in villages such as Besoa, Padang Lolo, and Bala (Anon. 1981b). Given the generally low fertility of the hill soils, it is interesting that many old villages and megalithic remains are on or around the beds of long- 'dead' lakes (Davis 1976). The small clans of people were frequently warring and when a peace was settled it was customary to smash plates. This is symbolic and represents the feeling that 'as the shards lay divided and yet together in one place, so shall we all return to our hearths knowing we belong together'. One such place is Poka Pinjang ('broken dish') on the path to the peak of Mt. Rantemario, Enrekang (p. 511), where shards of Chinese porcelain have been found (van Steenis 1937). The shy inland Towana people of the eastern peninsula also paid tribute to the Sultan of Ternate, but later came under the influence of the kings of Bungku to the southwest, and Tojo to the northwest. These rulers did not exert direct rule over this group of Towana, but indirectly through Towana leaders and elders. Tributes of beeswax and small bamboo tubes of uncooked rice were given to the rulers (Atkinson 1979). A succinct description of the culture of the people around Poso and the effects on them of early western influence is available (Kruyt 1929).
Figure 1.38 A sketch map of Sulawesi drawn in 1795 to illustrate just how poorly known the eastern coasts were.
After Woodard 1805
The British took over the Dutch colonial possessions between 1811 and 1816 and it was not until the early years of this century that the Dutch began to exert their rule over all of Sulawesi. In fact, by the late 19th century Minahasa had already received considerable assistance from the Dutch and from Christian mission societies. The degree of support is illustrated by the fact that by 1900 there were 1,200 inhabitants for every school in Minahasa, but 50,000 inhabitants for every school in Java. The Dutch rule of Sulawesi began with much blood being shed, particularly in battles with people in Bone. Until this period remote areas such as Toraja land had withstood or had simply not experienced incursions and assaults into their territory, but now they were brought under an island-wide administration. There then followed 35 years of peace-probably the longest period in Sulawesi's history-during which roads, irrigation schemes and administrative structures and institutions were established.
The Japanese occupied Sulawesi in 1942 but the move towards independence was not entertained until the final five months of the war. Then, Dr. Sam Ratulangi, a Manadonese, and Andi Pangiran, son of the Sultan of Bone, were sent to Jakarta to assist in the preparations for the birth of the infant republic. Ratulangi became the first governor of Sulawesi but was imprisoned after the Australian allied forces handed over the administration to the Dutch in January 1946. By the end of that year a guerrilla operation against the Dutch was well under way and the Dutch responded with their bloodiest campaign anywhere in Indonesia. On 17th August 1950, the Dutch accepted the independence of Indonesia. At this time Sulawesi was a single province. In July 1950, an intermittent rebellion with Islamic and regional goals erupted in and around Luwu and its forces became allied with the Darul Islam movement of West Java. By 1958, when this movement was at its peak, only the cities in South Sulawesi were under government control, and towns such as Soroako, now the centre of an international tin-mining operation, suffered severe privations (Robinson 1983; Kirk 1986). Order was restored in the early 1960s. In the regional rebellions of 1958-61, Minahasa (together with West and North Sumatra) declared their independence from the rest of the Republic, but few lives were lost before national unity was restored.
Present-day People
Despite the relatively small size and population of Sulawesi, the number and make-up of the ethnic groups is extremely complex.41 Early linguists used to ascribe a large number of language groups to Sulawesi. For example, one analysis recognized just one language group in each of Taiwan, the Philippines, Sumatra and Borneo, whereas for Sulawesi nine were recognized. More recent work has lessened the distinctiveness of Sulawesi languages but, even so, two of the nine language families found in Western Malayo-Polynesia are confined to Sulawesi. Linguists classify languages in a stricter fashion than do lay people, and the fifty ways of saying 'no' in Central Sulawesi alone do not necessarily represent different languages (Davis 1976). Rather this has arisen through small kingdoms or states being relatively isolated for long periods, and the development of dialects. The Sulawesi people themselves use region, religion and style of farming as the major criteria for determining ethnic groups. For example, someone from Central Sulawesi may refer to Christian Mamasa speakers of South Sulawesi as 'Toraja', but call Islamic Mamasa speakers 'Bugis' or 'Mandar' (Davis 1976). Similarly, the term 'Bugis' can mean seafaring Makassarese and Mandarese as well as the coastal Bugis (fig. 1.39; table 1.16). About 80% of the population of Sulawesi is Islamic and 20% Christian but there is considerable variations between regions (fig. 1.40).
Figure 1.39. The distribution of the major ethnic groups. Numbers refer to table 1.16.
From Davis 1976
The population is distributed unevenly across Sulawesi, with the area around Ujung Pandang having more than 300 people/km2, much of the rest of the southwest peninsula, Minahasa and Sangihe-Talaud having 100-299 people/km2 and Toli-Toli, Mamuju, the eastern arm and the east of Southeast Sulawesi having less than 30 people/km2 (fig. 1.41).
Figure 1.40. Percentage of population following Islam by county.
Based on Anon. 1981a
1 - Formerly called 'Bare'e'
2 - The group now commonly called 'Toraja'
From Davis 1976
Figure 1.41. Population density of Sulawesi by county.
After Anon. 1981a
PRESENT STATE OF NATURAL ECOSYSTEMS
Man began converting natural forest to other forms of vegetation many-hundreds of years ago but this process has accelerated greatly since the early 1970s when commercial logging, transmigration and estate crop projects began to receive enormous government support. The Sulawesi mainland now comprises an irregular patchwork of natural forest within and between areas of cultivation (fig. 1.42). The forest cover per inhabitant in Sulawesi is more than in Sumatra, Java and Bali, or the Lesser Sunda Islands (table 1.17), but this is at least partly due to the high proportion of land in Sulawesi on slopes which are unsuitable for agricultural development projects (table 1.18). For example, 83% of Sulawesi comprises slopes of 15% or above, compared with only about half of Sumatra, Kalimantan or Irian Jaya. Small islands have lost most of their natural vegetation: for example, Sangihe and other small northern islands were largely deforested by 1920 although Karakelang still has some forest cover (Heringa 1921). Likewise the forests of the small southern islands had all been converted to agricultural uses by 1915 with the exception of small areas of Tanahjampea and Kalaotoa (van Schouwenburg 1915, 1916a, b, c).
Figure 1.42. Extent of cultivated land (black) in 1982.
Adapted from Whitmore 1984b
From Anon. 1982b
Calculating land areas under different forms of land use is best conducted by the interpretation of satellite images. This was done for Sulawesi (Hadisumarno 1978), but the images used were made in 1972 and the usefulness of the information is now somewhat limited to historical comparisons. More recent analyses of land use over relatively small areas have been made by a number of government departments. An excellent series of maps showing land system and suitability, land use and land status was completed in 1987 for the primary use of the Transmigration Department, but these have had great value for others concerned with land use planning and resource management. Meanwhile, different methodologies, definitions and criteria used by different agencies tend to produce somewhat different figures for the land use of Sulawesi and elsewhere (table 1.19). This makes comparisons between tables confusing but does not necessarily affect the comparability of data within tables. Thus it can be seen that Sulawesi had less forest cover than either Sumatra or Kalimantan, and also relatively less land covered with estate crops (p. 487). The percentage of the land under wet rice fields is comparable to the percentage in Sumatra. See the Introduction for more recent data.
A comparison of land area, forest area, rice production and timber exports between the four Sulawesi provinces is instructive (table 1.20). In 1985 North and South Sulawesi were similar, as were Central and Southeast Sulawesi, in their ratio of land to population (about 1 ha/person and 4 ha/person respectively), but the difference in land capability and intensity of use is reflected in the greater rice production in South and North Sulawesi compared with the other two provinces. Southeast Sulawesi generally comprises poor agricultural soils and consequently rice yields are low. Timber ceased being exported out of the natural forests of Southeast Sulawesi in 1979 and out of North Sulawesi in 1981, and it is clear that Central Sulawesi was by far the most important timber producing province. Timber revenues in fact supplied more than 95% of the provincial income in Central Sulawesi. Logging continues in all provinces, of course, to supply local needs.
From Suwardjo et al. 1985
Obtaining definitive data concerning the area of land covered by different categories of forest is as difficult as obtaining accurate data on land use and leads to discrepancies between the figures produced by different departments (tables 1.21 and 1.22). It is well known that some Forestry land no longer has trees growing on it by virtue of inadequate protection, and this is one of the major causes for the discrepancies; there is clearly a difference between Forestry land and forested land. In any case, the percentage of land under some form of legal protection in Sulawesi is less than the percentage on hilly and mountainous slopes which for soil conservation reasons alone should be protected. Estate crops are not allowed on suitable slopes over 25% but some of the steep land has been deforested by second-stage shifting agriculturalists who have, for various reasons, abandoned the traditional or first-stage swidden agriculture practices (p. 570). This land clearly requires legal and enforced protection.
After Suwardjo et al. 1985
Areas with high agricultural potential have clearly been utilized more than areas with low agricultural potential. Thus, nearly all of wet lowland forest on volcanic soils has been felled compared with only 10% of similar forest on ultrabasic soils (table 1.23). Unfortunately none of the remaining wet lowland forest is within either existing or approved nature reserves. This habitat together with wet lowland forest on limestone, dry lowland forest on limestone, freshwater swamp and peatswamp forest are the habitats with the highest priority for conservation on Sulawesi.
The provinces differ strikingly in their geology (p. 5) and so the distribution of habitats between them, and the representation of those habitats within reserves, differ accordingly (table 1.24).
Sulawesi has five National Parks: Bogani Nani Wartabone National Park (formerly Dumoga-Bone) (300,000 ha), Lore Lindu (231,000 ha), Bunaken-Manado Tua Marine National Park, Taka Bone Rate National Park and Rawa Aopa-Watumohae National Park. Lore Lindu has also been chosen as a Biosphere Reserve by UNESCO in recognition of its biological, physical and cultural interest. Biosphere Reserves are areas with a protected core surrounded by utilized buffer zones, managed by a body having institutionalized relationships with the surrounding land and people, centres for management-related research, education and training, and having links with national and international monitoring schemes. Other conservation areas and the areas in Sulawesi under the control of the Directorate-General of Forest Protection and Nature Conservation include Nature Reserves (totalling 322,731 ha on Sulawesi), Wildlife Refuges (144,788 ha), Tourist Parks (97,000 ha), Hunting Parks (22,000 ha), and Marine Parks (none yet declared). Protection Forests (3,867,000 ha) are under the control of the Forestry Department but are also, in theory, a type of conservation forest. The regulations in force or proposed for the different categories of protected areas are shown in table 1.25.
* - areas in million ha; provincial areas given here differ significantly from official government statistics which are quite incompatible with up-to-date provincial maps
From Anon. 1982a. 1984b. 1985a
The conservation areas in the four provinces in 1982 are shown in figures 1.43-46 and are listed with their index of conservation value in table 1.26. Descriptions of these areas, and others which have less or no importance due to exploitation and degradation are described in the Sulawesi volume of the National Conservation Plan (Anon. 1982a).
After Suwardjo et al. 1985
After Anon. 1985c
Based on Anon. 1982a, b
After Anon. 1982
After Anon, 1982b
Figure 1.43. Remaining forest, conservation areas and proposed conservation areas in North Sulawesi. 1 - Tangkoko-Dua Saudara, 2 - Manembo-nembo, 3 - Mt. Soputan, 4 - Mt. Lokon, 5 - Mt. Klabat, 6 - Wiau, 7 - Tamposo-Sinansajang, 8 - Mt. Ambang, 9, 10, 15 - Bogani Nani Wartabone (formerly Dumoga-Bone), 11 - Mt. Simbolang; 12, 13,14 - Marisa Complex, 16 - Mt. Damar, 17 - Labutodoa and Paguyaman Barat, 18 - Karakelang, 19 - Mt. Sahendaruman, 20 - Bunaken-Manado Tua.
From Anon. 1982a
NP - National Park, NR - Nature Reserve, WR - Wildlife Reserve, HR - Hunting Reserve, TP -Tourist Park, PF - Protection Forest, MP - Marine Park, (p) - proposed
After Anon. 1982a
Figure 1.44. Remaining forest, conservation areas, and proposed conservation areas in Southeast Sulawesi. 1 - Napabalano, 2 - Lamedae, 3 - Tanjung Amolenggo, 4 - Mt. Watumohae, 5 - Tirta Rimba, 6 - Buton Utara, 7 - Kayu Kuku, 8 - Rawa Opa, 9 - Polewai, 10 - Tanjung Bati Kolo, 11 - Tanjung Peropa, 12 - Wakouti, 13 - Lasolo Bay, 14 - Wowoni Straits, 15 - Muna Straits, 16 - Lasolo-Sampara, 17 - Tukang Besi Islands.
After Anon. 1982a
Figure 1.45. Remaining forest, conservation areas and proposed conservation areas in South Sulawesi. 1, 2, 7 - Bantimurung and Karanta, 3 - Lampuko-Mampio, 4 - Peruhumpenai Mts., 5 - Lakes Matano/Mahalano, 6 - Lake Towuti, 8 - Lariang, 9 - Mamuju, 10 - Masapu, 11 - Mambuliling, 12 - Rangkong, 13 - Rompi, 14 - Lamikomiko, 15 - Sumarorang, 16 - Samalona Islands, 17 - Lake Tempe, 18 - Torokkapai, 19 - Matanga, 20 - Bulu Saraung, 21 - Palangka, 22 - Bontobahari, 23 - Komara, 24 - Camba River, 25 - Latimojong Mts., 26 - Mt. Lompobatang.
After Anon. 1982a
Figure 1.46. Remaining forest, conservation areas and proposed conservation areas in Central Sulawesi. 1 - Pati-Pati, 2 - Paboya, 3, 6, 12 -Lore Lindu, 4 - Tanjung Api, 5 - Morowali, 7 - Wera waterfall, 8 - Tanjung Matop, 9 - Dolongan Island, 10 - Mt. Dako, 11 - Mt. Sojol, 13 - Peleng waters, 14 - Toli-Toli Mts., 15 - Palu Mts., 16 - Morowali/Balantak Mts., 17 -Bakiriang, 18 - Togan Islands.
After Anon. 1982a
