A Mitochondrial Stratigraphy for Island Southeast Asia
2006; Elsevier BV; Volume: 80; Issue: 1 Linguagem: Inglês
10.1086/510412
ISSN1537-6605
AutoresCatherine Hill, Pedro Soares, Maru Mormina, Vincent Macaulay, Douglas J. Clarke, Petya B. Blumbach, Matthieu Vizuete-Forster, Peter Forster, David Bulbeck, Stephen Oppenheimer, Martin Richards,
Tópico(s)Forensic Anthropology and Bioarchaeology Studies
ResumoIsland Southeast Asia (ISEA) was first colonized by modern humans at least 45,000 years ago, but the extent to which the modern inhabitants trace their ancestry to the first settlers is a matter of debate. It is widely held, in both archaeology and linguistics, that they are largely descended from a second wave of dispersal, proto-Austronesian–speaking agriculturalists who originated in China and spread to Taiwan ∼5,500 years ago. From there, they are thought to have dispersed into ISEA ∼4,000 years ago, assimilating the indigenous populations. Here, we demonstrate that mitochondrial DNA diversity in the region is extremely high and includes a large number of indigenous clades. Only a fraction of these date back to the time of first settlement, and the majority appear to mark dispersals in the late-Pleistocene or early-Holocene epoch most likely triggered by postglacial flooding. There are much closer genetic links to Taiwan than to the mainland, but most of these probably predated the mid-Holocene “Out of Taiwan” event as traditionally envisioned. Only ∼20% at most of modern mitochondrial DNAs in ISEA could be linked to such an event, suggesting that, if an agriculturalist migration did take place, it was demographically minor, at least with regard to the involvement of women. Island Southeast Asia (ISEA) was first colonized by modern humans at least 45,000 years ago, but the extent to which the modern inhabitants trace their ancestry to the first settlers is a matter of debate. It is widely held, in both archaeology and linguistics, that they are largely descended from a second wave of dispersal, proto-Austronesian–speaking agriculturalists who originated in China and spread to Taiwan ∼5,500 years ago. From there, they are thought to have dispersed into ISEA ∼4,000 years ago, assimilating the indigenous populations. Here, we demonstrate that mitochondrial DNA diversity in the region is extremely high and includes a large number of indigenous clades. Only a fraction of these date back to the time of first settlement, and the majority appear to mark dispersals in the late-Pleistocene or early-Holocene epoch most likely triggered by postglacial flooding. There are much closer genetic links to Taiwan than to the mainland, but most of these probably predated the mid-Holocene “Out of Taiwan” event as traditionally envisioned. Only ∼20% at most of modern mitochondrial DNAs in ISEA could be linked to such an event, suggesting that, if an agriculturalist migration did take place, it was demographically minor, at least with regard to the involvement of women. Island Southeast Asia (ISEA), the area encompassed by modern Indonesia, East Malaysia, and the Philippines, was colonized by modern humans at least 45,000 years ago1Barker G Barton H Beavitt P Bird M Daly P Doherty C Gilbertson D Hunt C Krigbaum J Lewis H et al.Prehistoric foragers and farmers in southeast Asia: renewed investigations at Niah Cave, Sarawak.Proc Prehist Soc. 2002; 68: 147-164Google Scholar and possibly >50,000 years ago.2Détroit F Dizon E Falguères C Hameau S Ronquillo W Sémah F Upper Pleistocene Homo sapiens from the Tabon cave (Palawan, The Philippines): description and dating of new discoveries.C R Palevol. 2004; 3: 705-712Crossref Scopus (75) Google Scholar At that time, the region was split between the Pleistocene continent of Sunda, which stretched from Sumatra to Bali and Palawan, and Wallacea, which included the islands east of Wallace’s line (fig. 1). The Sunda shelf was flooded when sea levels rose in the early-Holocene epoch, spurring the development of maritime exchange between populations on the remnant Sunda islands (especially Borneo and Palawan) and populations in Wallacea.4Bellwood P Prehistory of the Indo-Malaysian archipelago. University of Hawaii Press, Hawaii1997Google Scholar, 5Oppenheimer S Eden in the East. Wiedenfeld and Nicholson, London1998Google Scholar, 6Solheim WG Archaeology and culture in southeast Asia: unraveling the Nusantao. University of Philippines Press, Quezon City2006Google Scholar However, despite this evidence for a dynamic population history in early ISEA, paleoanthropologists tend to classify all early human remains in the region as “Australo-Melanesian” (i.e., related to the indigenous people of Australia and New Guinea) and argue for a mid-Holocene immigration of the ancestors of most of the present-day inhabitants.4Bellwood P Prehistory of the Indo-Malaysian archipelago. University of Hawaii Press, Hawaii1997Google Scholar The primary justification for a two-tier population history in ISEA is historical linguistics. Almost all indigenous populations in the region speak languages belonging to the Malayo-Polynesian branch of Austronesian, also found in the Pacific and Madagascar, whereas the other nine primary branches of Austronesian (the Formosan languages) are spoken only by aboriginal Taiwanese.7Blust R Subgrouping, circularity and extinction: some issues in comparative Austronesian linguistics.Symp Ser Inst Linguis Academica Sinica. 1999; 1: 31-94Google Scholar This is interpreted as implying that the Austronesian languages must have developed in Taiwan and spread, by sea, from there to across their current distribution, with the various branches of Malayo-Polynesian separating along the voyage,8Blust R Austronesian culture history: the window of language.in: Goodenough WH Prehistoric settlement of the Pacific. American Philosophical Society, Philadelphia1996: 28-35Google Scholar a model known as “Out of Taiwan.” In the 1970s, archaeologists began to draw a direct relation between the dispersal of early Austronesian languages and the onset of the Neolithic period in ISEA (and in the Pacific outside of New Guinea). Documentation of the prehistory of ISEA was (and, to an extent, remains) meager,4Bellwood P Prehistory of the Indo-Malaysian archipelago. University of Hawaii Press, Hawaii1997Google Scholar, 9Spriggs M Chronology of the Neolithic transition in Island Southeast Asia and the western Pacific: a view from 2003.Rev Archaeol. 2003; 24: 57-80Google Scholar but several sites have yielded preceramic sequences followed by the introduction of pottery at a mid-Holocene date. The appearance of pottery was used as a proxy for the arrival of the Taiwanese “Austronesians,” whose agricultural subsistence allowed them to supplant or assimilate the indigenous Australo-Melanesian foragers.10Schutler R Marck JC On the dispersal of Austronesian horticulturalists.Archaeol Phys Anthropol Oceania. 1975; 10: 81-113Google Scholar Subsequent archaeological work in China, however, demonstrated the establishment of large settlements dependent on rice agriculture during the early Holocene. Archaeological comparisons suggested that the earliest Neolithic cultures in Taiwan resemble earlier cultures found in South China and also have parallels to Neolithic sites found in the Philippines. According to the model most popular among ISEA archaeologists today, rice agriculturalists in South China dispersed into Taiwan ∼5,500 years ago, where they developed the Austronesian languages before expanding again into the Philippines and the rest of ISEA ∼4,000 years ago.4Bellwood P Prehistory of the Indo-Malaysian archipelago. University of Hawaii Press, Hawaii1997Google Scholar, 11Bellwood P First farmers, the origins of agricultural societies. Blackwell Publishing, Oxford2005Google Scholar Vigorous debate continues among archaeologists, however, as to how the evidence should be interpreted. Although the model allows for the assimilation of indigenous populations, the extent of any assimilation cannot be assessed from the archaeological record.12Pawley A Austronesian dispersal: languages, technologies and people.in: Bellwood P Renfrew C Examining the farming/language dispersal hypothesis. McDonald Institute for Archaeological Research, Cambridge2002: 251-273Google Scholar Moreover, the archaeobotanic evidence in ISEA seems not to fit the simple rice-fueled Out of Taiwan model, placing the motive force for the dispersal in question.13Paz V Island Southeast Asia—spread or friction zone?.in: Bellwood P Renfrew C Examining the farming/language dispersal hypothesis. MacDonald Institute for Archaeological Research, Cambridge2002: 275-285Google Scholar, 14Hunt CO Rushworth G Cultivation and human impact at 6000 cal yr B.P. in tropical lowland forest at Niah, Sarawak, Malaysian Borneo.Quaternary Res. 2005; 64: 460-468Crossref Scopus (43) Google Scholar, 15Anderson A Crossing the Luzon Strait: archaeological chronology in the Batanes Islands, Philippines and the regional sequence of Neolithic dispersal.J Austronesian Stud. 2005; 1: 25-44Google Scholar South of northern Luzon, the only possible evidence for Neolithic (pre–Iron Age) rice in ISEA is restricted to western Borneo,16Bellwood P Gillespie R Thompson GB Vogel GS Ardika JW Datan I New dates for prehistoric Asian rice.Asian Perspect. 1992; 31: 161-170Google Scholar where it predates the supposed Austronesian arrival. Generally speaking, the Neolithic of ISEA is characterized by tremendous local diversity, rather than a uniform dispersal package,17Szabó K O’Connor S Migration and complexity in Holocene Island Southeast Asia.World Archaeol. 2004; 36: 621-628Crossref Scopus (18) Google Scholar and it has even been suggested that Taiwan received some aspects of its Neolithic technology from the mainland and some from ISEA.18Meacham W On the improbability of Austronesian origins in South China.Asian Perspect. 1984-5; 26: 89-106Google Scholar The model, therefore, relies heavily on the linguistic argument—but this is not quite as powerful as often assumed. The Austronesian languages may indeed be most diverse in Taiwan, but assigning a root to the language tree is still not straightforward. Since the reconstructed tree is starlike, with nine Formosan branches and one Malayo-Polynesian branch,7Blust R Subgrouping, circularity and extinction: some issues in comparative Austronesian linguistics.Symp Ser Inst Linguis Academica Sinica. 1999; 1: 31-94Google Scholar the root could be anywhere within Taiwan or ISEA. Indeed, some linguists cluster the Philippine languages with those of Taiwan,19Wolff J The position of the Austronesian languages of Taiwan within the Austronesian group.in: Li P Tsang J-K Ho D Tseng C Austronesian studies relating to Taiwan: Institute of History and Philology Symposium Series 3. Academica Sinica, Taipei1995Google Scholar and recent attempts to apply phylogenetic methodologies from biology to the linguistic data have only suggested a root somewhere within the general area of Taiwan, the Philippines, and Borneo.20Greenhill S Gray RD Testing dispersal hypotheses: Pacific settlement, phylogenetic trees and Austronesian languages.in: Mace R Holden C Shennan S The evolution of cultural diversity: phylogenetic approaches. UCL Press, London2005: 31-52Google Scholar A root in ISEA would mesh better with alternative views of the archaeological evidence that place ISEA at the center of population dispersals and Taiwan at the periphery—with only sporadic mainland influence—and with the languages most likely emerging during the Holocene within “Austronesia”18Meacham W On the improbability of Austronesian origins in South China.Asian Perspect. 1984-5; 26: 89-106Google Scholar itself. This would imply an early offshoot to Taiwan and subsequent leveling of language diversity across the Malayo-Polynesian area as a result of the formation of extensive socioeconomic networks.5Oppenheimer S Eden in the East. Wiedenfeld and Nicholson, London1998Google Scholar, 6Solheim WG Archaeology and culture in southeast Asia: unraveling the Nusantao. University of Philippines Press, Quezon City2006Google Scholar, 18Meacham W On the improbability of Austronesian origins in South China.Asian Perspect. 1984-5; 26: 89-106Google Scholar The debates reviewed above can be tested with genetic data. The Out of Taiwan model would predict that at least some ISEA lineages should trace back to ancestral populations in Taiwan and, ultimately, South China, with the age of those ISEA-specific lineage clusters being no more than ∼4,000 years ago. Earlier dispersals, perhaps stimulated by climate change and sea-level rise, as suggested in several alternative models,5Oppenheimer S Eden in the East. Wiedenfeld and Nicholson, London1998Google Scholar, 6Solheim WG Archaeology and culture in southeast Asia: unraveling the Nusantao. University of Philippines Press, Quezon City2006Google Scholar, 18Meacham W On the improbability of Austronesian origins in South China.Asian Perspect. 1984-5; 26: 89-106Google Scholar would, in contrast, predict dispersals centered on ISEA and dating to the late Pleistocene or early Holocene. If several distinct dispersal processes shaped the genetic variation in the region, their signatures should each be evident in the genetic record. A number of genetic studies have already attempted to address the question of a putative Austronesian expansion, but most mtDNA studies to date have focused on Pacific islanders, with little work on the potential source populations. Because of this and because most studies only examined the fast-evolving first hypervariable segment of the control region (HVS-I), firm conclusions have been elusive. 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However, different studies with similar data sets have failed to agree on how the data should be partitioned. This is a consequence of poor phylogenetic resolution—insufficient markers to allow reconstruction of the main branches of the genealogical history. This, in turn, results in poor phylogeographic resolution, since branches of the tree that may be distributed differently in space cannot be distinguished. The use of more markers will be necessary before the Y-chromosome contribution to the debate can be clarified. In the meantime, we have addressed the poor quality of previous mtDNA data, both by sampling almost 1,000 individuals from locations throughout ISEA and by analyzing the samples at a higher resolution than done previously, by including coding-region as well as control-region variants gleaned from complete sequence data. This dramatically improved data set substantiates recent suggestions that major rethinking is needed with regard to the prehistory of the region. Sampling locations are shown in figure 1. A total of 929 anonymous, unlinked DNA samples from across ISEA and Taiwan were analyzed and comprised 180 Sumatrans (42 from Medan, 24 from Padang, 52 from Pekanbaru, 28 from Palembang, and 34 from Bangka—previously published by Hill et al.35Hill C Soares P Mormina M Macaulay V Meehan W Blackburn J Clarke D Raja JM Ismail P Bulbeck D et al.Phylogeography and ethnogenesis of aboriginal Southeast Asians.Mol Biol Evol. 2006; 23: 2480-2491Crossref PubMed Scopus (113) Google Scholar), 46 Javanese (36 Tengger, 1 from Yogyakarta, 1 from Banjumas, 1 from Garut, 1 from Jakarta, 1 from Probolinggo, 1 from Semarang, 3 from Solo, and 1 from Wonogiri), 157 individuals from Borneo (68 from Kota Kinabalu and 89 from Banjarmasin), 2 individuals from Bajawa in Flores, 82 Balinese (including 67 from Denpasar, 3 from Gianyar, 1 from Nusa Dua, 1 from Semarapura, 4 from Singaraja, 2 from Tabanan, and 2 from Ubud), 44 individuals from Mataram in Lombok, 237 individuals from Sulawesi (46 from Ujung Padang, 38 from Palu, 89 from Manado, and 64 Toraja), 50 individuals from Waingapu in Sumba, 43 individuals from Ambon, 45 individuals from Alor, 61 Filipinos, and 78 Taiwanese aboriginals (21 Ami, 18 Atayal, 18 Bunun, and 21 Paiwan—data updated from Sykes et al.24Sykes B Leiboff A Low-Beer J Tetzner S Richards M The origins of the Polynesians—an interpretation from mitochondrial lineage analysis.Am J Hum Genet. 1995; 57: 1463-1475PubMed Google Scholar and Melton et al.36Melton T Clifford S Martinson J Batzer M Stoneking M Genetic evidence for the proto-Austronesian homeland in Asia: mtDNA and nuclear DNA variation in Taiwanese aboriginal tribes.Am J Hum Genet. 1998; 63: 1807-1823Abstract Full Text Full Text PDF PubMed Scopus (98) Google Scholar). 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RFLP screening was used to resolve haplogroup status in a hierarchical fashion, as follows: haplogroups M (+10397 AluI; +10394 DdeI), N (−10397 AluI; −10394 DdeI), M7 (+9824 HinfI), D (−5176 AluI), E (−7598 HhaI), G (+4830 HhaI), P (+15606 AluI), U (+12308 HinfI), and I (+10032 AluI). Haplogroup B affiliation was checked by sc
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