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The Pleistocene Endemic Fauna of the Indonesian Archipelago

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The Pleistocene Endemic Fauna of the Indonesian Archipelago TROPICS Vol. 10 (1): L35-1.43 Issued May 30, 2000 The Pleistocene Endemic Fauna of the Indonesian Archipelago Fachroef Azrz Geological Research and Development centre (GRDC), Jalan Diponegoro No. 57, Bandung,40L22, Indonesia ABSTRACT The Indonesian Archipelago is situated between the Asian and the Australian continents. It is divided into the Sunda and the Sahul shelves and the Wallacea area in between. The Sunda and the Sahul shelves have been connected with the continents several times, as the result of tectonic movements and sea level fluctuations in Pleistocene time. On the other hand, the Wallacea area has never been connected to any continent. Asiatic faunas have migrated to Java, and some ofthem have even been able to cross over Wallace's line by sweepstake dispersal. Thereafter, they adapted to island environments and developed into the unbalanced endemic island faunas. This paper gives an overview of the unbalanced endemic island faunas of the Indonesian archipelago. Key words: Pleistocene / unbalanced / endemic / island fauna / sweepstake dispersal / Indonesian Archipelago The Indonesian Archipelago is a natural bridge connecting the Asian with the Australian continents. Physiographically, it is divided into the Sunda continental shelf to the west and the Sahul continental shelf to the east with the Wallacea province in between (Fig.l). The Sunda shelf is the southeastem edge of the Asian continent, consisting of the Greater Sunda Islands in which the islands of Sumatra, Java, Kalimantan, and the surounding shallow sea areas are included. The Sahul shelf is the northeastern edge of the Australian continent, consisting of the islands of Irian, Aru, Kai and shallow seas surrounding them. The edges of these continents are marked by the 200m isobath. These two continental shelves are separated by an intervening belt of deep sea basins, trenches, channels and island festoons. This area corresponds to the Wallacea province in which the Ipsser Sunda islands (the islands of Lombok, Sumba, Sumbawa, Flores, Selor, Alor and wetar), plus Timor, Tanimbar, Maluku, Halmahera, Karakelong, Sangir and Sulawesi are included. During sea level drops in glacial times, the Sunda shelf become one landmass with the Asian continent, as did the Sahul shelf with the Australian continent. When sea level rose up during the interglacial time, the islands on both continental shelves became separated once again. ZOOGEOGRAPHY It was Wallacp (1863, in Whitmore, 1981) who drew an imaginary red line along the Makassar strait 136 F. Aztz r,tuy\r',p ,rF1- - i c J | \ uluzoN 'dI?Nc! ps,,, ilS^""'LrPPrNEs Fig. 1. Geography oflndonesian Archipelago and adjacent areas. to separate the Indo-Malayan region to the west and the Austro-Malayan region to the east. Wallace's line has prompted controversy among zoogeographers, and many other lines have also been advocated (e.g. Huxley's, Weber's, Lydekker's lines). In general, those later lines were drawn on the basis of limited grcups of fauna like land snails, mammals, birds or bats. The ability of dispersal, especially of crossing sea barriers, varies among faunal groups. Simpson (1977) recommended that the Oriental Faunal Region be bounded by the Huxley's line, coinciding with the Sunda shelf, whereas the Australia Faunal Region should be bounded by Lydekker's line which coincides with the Sahul shelf. The faunal region between the two shelves is considered an intermediate zone. According to Van den Bergh (1997), no real objection attaches to the designation of the islands in between the Sunda and the Sahul shelves as transitional; however, the fauna of each island should be studied separately. MA,.IOR EACTORS FOR MIGRATION OF EAUNA The past migration of land vertebrates from one place (mainland) to the other (island) depends mainly on the paleogeography of the area and the biological aspect of the fauna. Simpson (1965) and Dermitzakis and Sondaar (1979) discussed the relationship between the fauna and geography and give a model for migration as follow: L. Corridor dispersal, dispersal of fauna across a broad land connection, in which faunal interchange is possible in both directions. It is indicated by a balanced fauna, where there are few or no differences between mainlands. 2. Filter dispersal, or dispersal through or across a small barrier, such as a restricted environment or a disconnected landmass. Spread is possible for some fauna but not for others. A somewhat impoverished but balanced mainland fauna develops. 3. Sweepstake dispersal, across a broader barrier/wide sea. Spread is impossible for most and very improbable for some faunas, but does occur accidentally. It is characterized by an unbalanced The Pleistocene endemic fauna of the Indonesian Archipelago r37 endemic fauna. 4. Puddle dispersal across a nalrow sea. A route which is insurmountable for most fauna but easy for others. Biological aspects of the fauna are the most important factors for migration. Some faunal elements, like elephants, hippos and deer are able to swim across open seas. Sondaar (1977) mentioned that the elephants have a digestive system which produces gases making it easy for them to float. In addition, their trunk can be used an excellent snorkel, which enhances their potential to survive for long period in water. Another factor is their nomadic way of life. They live and migrate including swimming in crowds. Under pressure and stress conditions, elephants have the capability to swim as far as 48 km and to survive for three days in the open sea (Johnson, 1978). THE EAUNA OF INDONESIAN ARCHIPEI,AGO JAVA (Fig.l: (D) The Pleistocene faunal biostratigraphy of Java was originally proposed by Von Koenigswald (1934, 1935). Recently, it has been realized that there aro many inconsistencies present in his concept. A new concept was then proposed and discussed (e.g. De Yos et al.,1982;De Vos 1983, 1996; Sondaar, 1984; Irinders et al., 1985; Sudijono, 1986; A2i2,1,996). The new concept ofJavan biostratigraphy, in ascending order (Fig.2), involves recognition of the Satir (1.5 Ma), Cisaat (1.2 Ma), Trinil H. K. (1.0 Ma), Kedungbrubus (0.8 Ma), Ngandong (0.4? Ma), punung (0.0s Ma) and the wajak faunas (0.01 Ma). After the emergence of Java about 1.5 Ma ago, some Asiatic faunal elements from Siwalik, Narbada and Burma via the Siva-Malayan route got to Java by sweepstake dispersal, and these developed into an endemic unbalanced fauna. This is reflected in the Satir fauna of the Bumiayu area in Central Java. The faunal composition is very limited in number of species, of which Sinomastodon bumiayuensis, Hexaprotodon simplex, Cervids and Geochelone sp. are the main components. These associated species are called by De Vos (1996) Tetralophodan-Geochelone fauna. A similar fauna is also known from the lower black clay of the Sangiran (Pucangan) Formation in the Sangiratr area, Central Java. On the basis of pollen spectrum data, it seems that the Tetralophodon-Geochelone fauna lived in an open area with partly mangrove forest environments (Polhaupessy, 1990; Semah, 1984). About 1.2 Ma - 1.0 Ma ago, a migration of mainland fauna took place to form the Cisaat-Trinil H. K. faunas which are characterized by the arival of Stegodon trigonocephalns and Homo erectw. These elements reached Java from the Siwalik, Narbada and Burma areas via the Siva-Malayan route by means of filter dispersal. This new and successful arival of mainland faunal elements caused a dramatic faunal turnover in which the endemic ktralophodon-Geochelone fauna was replaced by the so-called Stegodon-Homo erectus fauna. In this event, Sinomastodon bwniayuensis was replaced by Stegodon trigonocephalus, whereas Hexaprotodan simplac was replaced by Hexaprotodon sivalensis. During the course of time, the connection between the Sunda shelf and the Asian continent became stronger. The mainland fauna of the Siwalik and Narbada area reached Java by means of corridor dispersal, in which new arrival of Elephas and Tapirus (Kedungbrubus-Ngandong faunas) took place. The large number of bovids suggest that the Cisaat-Ngandong fauna lived in an open woodland environment. The next episode, during the sea level drop of the Late Pleistocene, permit a full land 138 F. Aztz AGE FAJNAL UNITS DISPERSAL ROUTE CHRACTERZED ENV z ool WAJAK ogz PONGO- 3fr 6d HOMO SAPIENS FAUNAL TURN- PUNUNG oo8 ?8 OVER @RRIDOR = o.40 NGANDOT{G z zo J o.80 KEDUIT3BRUzuS J STEGODON- o o o = HOMO ERECTIJS r.oo TRINIL H. K z= FILTER I lrl (L HAUNAL TURN- r.20 CISAAT o OVER z (9rlJ {- a TETRALOPHODON r.50 SATIR SWEEPSTAKE GEOCHELONE 3p Fig.2. New concept ofthe biostratigraphy in the Indonesian Archipelago' connection between the Sunda shelf and the Asian continent. A migration of mainland fauna to Java (punung fauna) from China, Vietnam, and Cambodia via the Sino-Malayan route by conidor dispersal took place, characterized by the arrival of Pongo pygmaeus and also Homo sapiens. The abundance of pongo pygnaeas in the Punung fauna indicates a humid rain forest environment. A dramatic faunal turnover occurred in which archaic faunal elements and Homo erectu.s became extinct. After the development of the Punung fauna, probably at the beginning of the Holocene, the Sunda shelf broke up again as the consequence of sea level rising during the present interglacial period. At this time, Java was separated from Kalimantan and subsequently from Sumatra. The fauna on each island became impoverished. SULAWESI (Fie.l: @) Sulawesi is the largest island in the Wallacea area, separated from the Sunda Shelf (Kalimantan) by the deep Makassar Strait which now is about 2000 m deep and 250 km wide. The Makassar Strait is the main part of Wallace's line, an effective barrier for the eastward migration of the fauna of Asiatic origin. Nevertheless, a Plio-Pleistocene fauna of Asiatic affinities has been discovered in the Wallacea the older area in the western arm of South Sulawesi.
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