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The Late Neogene Elephantoid-Bearing Faunas of Indonesia and Their Palaeozoogeographic Implications

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The Late Neogene Elephantoid-Bearing Faunas of Indonesia and Their Palaeozoogeographic Implications PP_003_3.90 15-01-2007 10:54 Pagina 1 The Late Neogene elephantoid-bearing faunas of Indonesia and their palaeozoogeographic implications A study of the terrestrial faunal succession of Sulawesi, Flores and Java, including evidence for early hominid dispersal east of Wallace’s Line Gert D. van den Bergh Bergh, G.D. van den. The Late Neogene elephantoid-bearing faunas of Indonesia and their palaeozoo- geographic implications; a study of the terrestrial faunal succession of Sulawesi, Flores and Java, including evidence for early hominid dispersal east of Wallace’s Line. — Scripta Geol., 117: 1-419, 78 figs., 18 pls; Leiden, November 1999. Gert D. van den Bergh, Marine Chemistry and Geology, Netherlands Institute for Sea Research, P.O. Box 59, 1790 AB Den Burg, Texel, The Netherlands. Keywords: Elephantoidea, palaeozoogeography, terrestrial fauna evolution/turnover, Southeast Asia, Sulawesi, Flores, Java, Pliocene-Pleistocene, islands, prehistory, hominids. The stratigraphic framework of the Neogene fossil vertebrate bearing formations of the Indonesian islands Sulawesi and Flores is established and the sediments are dated by means of marine micropalaeontological and/or palaeomagnetic methods. The results allow comparison of the fauna evolution on these islands with the better known fauna succession of Java. On both Sulawesi and Flo- res remains of large mammals and reptiles were excavated from fossil-rich layers, documenting the composition of large vertebrate faunas fairly well. For Sulawesi this procedure allows distinction of separate fauna units which formerly had been lumped together. Both on South-Sulawesi and Flores 3 successive faunas can be distinguished. The oldest terrestrial faunas have recorded ages of 2.5 Ma and 0.9 Ma, respectively. On both islands intermediate faunas have a Middle Pleistocene age, while the youngest faunas are subrecent to Recent. Elephantoids (Stegodon and Elephas) showing various degrees of dwarfing are important constituents in all non-subrecent/Recent faunas. Their dental, cranial and postcranial remains are described in detail and their taxonomic positions are discussed. A new species (Stegodon sondaari sp. nov.) is described from Flores. Geological/tectonic data and the nature of the impoverished and highly endemic faunas from Sulawesi and Flores both indicate longlasting isolation from the southeast Asian mainland and from Java. On Flores and probably also on South Sulawesi fauna turnovers with extinction and immigration of elephantoids occurred around the Early to Middle Pleistocene transition. On both islands the total disappearance of elephantoids presumably took place before the onset of the Holocene. On Flores the Middle Pleistocene faunal assemblage is associated with a palaeolithic industry, which is tentatively ascribed to Homo erectus. Early hominids appear to have been able to cross water barriers and success- fully colonize Flores as far back as 0.6 Ma ago. They may have been in part responsible for the observed faunal turnover on the island, but alternatively, extinctions may have been caused by a vol- canic catastrophe. The existing biostratigraphic framework of Java is discussed and some additions concerning elephan- toid taxa are given. Following the emergence of Java above sea-level from the Late Pliocene until c. 1.5 Ma, mammalian dispersal to the island was limited. This is illustrated by impoverished faunal assem- blages and the occurrence of various dwarfed elephantoids in this time interval, indicating isolated con- ditions. Further uplift and volcanic activity in combination with episodes of moderate low sea-level led to an increasing accessibility of the Java region until c. 0.8 Ma, though the faunas from this time interval remain somewhat impoverished, suggesting filter dispersal. Stegodon elephantoides is for the first time recorded from Java, originating from a layer correlated with this time interval. Corridor dispersal to Java occurred for the first time at around 0.8-0.7 Ma, and can be tied to marked glacio-eustatic sea-level lowerings starting at 0.8 Ma. This period of low sea-level may also have triggered the faunal turnovers observed on Sulawesi and Flores, by enhancing the chance for a second (overseas) dispersal wave to PP_003_3.90 15-01-2007 10:54 Pagina 2 2 Van den Bergh. The Late Neogene elephantoid-bearing faunas of Indonesia. Scripta Geol., 117 (1999) these islands, and perhaps exchange between Sulawesi and Flores. However, superficial similarities between various island stegodonts must be ascribed to parallel evolution in most cases. Common adap- tive trends that can be recognized are dwarfing and increase in hypsodonty. On Java local evolution is documented by the recognition of three successive subspecies of Stegodon trigonocephalus. On Java the extinction of Stegodon and the first recording of Elephas maximus concurs with environmental changes from an open woodland biotope to a rainforest biotope, which took place after the penultimate glacial maximum at 125 ka. On South Sulawesi the Recent fauna presumably entered after the connection with the remaining part of Sulawesi had been established. De rijke biodiversiteit van het enorme eilandenrijk gelegen tussen het vasteland van zuidoost Azië en het Australische continent, dat onder andere het gehele Indonesische territorium omvat, is in grote mate te danken aan de complexe geologische geschiedenis. Daarbij heeft isolatie op eilanden een grote rol gespeeld. Fossiele landvertebraten-voorkomens van de eilanden Java, Sulawesi en Flores vormen het uitgangspunt van de huidige studie. De diverse fossiele faunas van deze eilanden zijn in een litho- en chronostratigrafisch kader geplaatst, waardoor een bijdrage is geleverd om de complexe zoögeografische evolutie van de regio te reconstrueren. Soorten behorende tot de superfamilie der Elephantoidea, waartoe ook de recente olifanten behoren, zijn zeer goed vertegenwoordigd in de fossiele faunas. Vertegenwoordigers van deze groep hebben zich gedurende de laatste 2,5 miljoen jaar met groot succes kunnen verspreiden binnen het gebied. Fossiele vindplaatsen met één of meerdere Elephantoidea, zijn bekend van de eilanden Sumatra, Java, Borneo, Sulawesi, Sangihe, Flores, Sumba, Timor, Luzon en Mindanao. In een recenter verleden is de grote verscheidenheid aan olifanten grotendeels weer verdwenen, niet alleen regionaal, maar wereld- wijd. Vanwege het relatief veelvuldige voorkomen van fossiele olifanten resten, ook op eilanden waar de meeste andere zoogdieren niet konden komen, is de nadruk in dit proefschrift komen te liggen op de olifanten (in de ruimste betekenis van het woord). In het eerste hoofdstuk worden de achtergrond en doelstellingen van het onderzoek geïntroduceerd. Ook wordt de gevolgde taxonomische classificatie van de Proboscidea uiteengezet, gevolgd door een beschrijving van de opgravingsprocedures en een verklaring van de gebruikte terminologie en meet- methoden. Het tweede hoofdstuk handelt over een aantal nog niet eerder beschreven elephantoïden resten van Java, die voor zover mogelijk in het bestaande biostratigrafische kader worden geplaatst. De landfau- na-veranderingen die zich gedurende het Pleistoceen op Java hebben afgespeeld waren in grote lijnen bekend, en daarom is het biostratigrafisch kader van Java van groot belang als referentie voor de vergelijking met de fauna opeenvolgingen op de eilanden Sulawesi en Flores. Het huidige onderzoek laat zien dat wat de olifanten betreft de evolutionaire opeenvolging op Java complexer is geweest dan aanvankelijk werd aangenomen. Sinds Java gedurende het laat Plioceen ten gevolge van vulkanische en tectonische activiteit boven de zeespiegel is uitgerezen, is er een toename in het totaal aantal soorten grote zoogdieren opgetreden tot zo’n 800.000 jaar geleden. Een van de eerste elephantoïden die Java bereikte was Sinomastodon bumia- juensis. Deze mastodont is geassocieerd met de 2 tot 1,5 miljoen jaar oude Satir Fauna, welke slechts een beperkt aantal soorten omvat, waaronder een nijlpaard, herten en een reuzenschildpad, hetgeen duidt op eiland omstandigheden. Vertegenwoordigers van deze taxa hebben Java waarschijnlijk bereikt door een zeebarrière over te steken (sweepstake route). De ongeveer 1,2 miljoen jaar oude, maar slecht bekende, Ci Saat Fauna was oorspronkelijk gedefiniëerd door het eerste voorkomen op Java van Stegodon. Door de relatieve zeldzaamheid van fossielen uit dit tijdinterval was de overgang tussen de Satir Fauna en de Ci Saat Fauna nauwelijks bekend. Een nog niet eerder beschreven onderkaak met een ouderdom van tussen de 1,2 en 1,3 miljoen jaar, wordt hier aan Stegodon elephan- toides toegeschreven. Deze zeldzame soort was tot nog toe alleen bekend uit Myanmar. Opvallend is dat enkele meters boven de laag met S. elephantoides nog een reuzenschildpad aangetroffen is, hetgeen erop lijkt te wijzen dat de overgang tussen de Satir en Ci Saat Fauna een geleidelijke is geweest. Twee jongere faunas, de 0,9 miljoen jaar oude Trinil Haupt-Knochenschicht Fauna en de 0,8-0,7 miljoen jaar oude Kedung Brubus Fauna, zijn veel beter bekend. De Trinil H.K. Fauna is een enigszins verarmde Zuidoostaziatische vasteland fauna, met als enige elephantoïde Stegodon trigonocephalus, ter- PP_003_3.90 15-01-2007 10:54 Pagina 3 Van den Bergh. The Late Neogene elephantoid-bearing faunas of Indonesia. Scripta Geol., 117 (1999) 3 wijl in de Kedung Brubus Fauna daarnaast ook een echte olifant, Elephas hysudrindicus voorkomt. De Kedung Brubus Fauna is een evenwichtige continentale fauna. De laat-Pleistocene Ngandong
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