An Australasian Test of the Recent African Origin Theory Using the WLH
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John Hawks An Australasian test of the recent African Department of Anthropology, origin theory using the WLH-50 calvarium University of Utah, Salt Lake City, UT 84112, This analysis investigates the ancestry of a single modern human U.S.A. E-mail: specimen from Australia, WLH-50 (Thorne et al., in preparation; [email protected] Webb, 1989). Evaluating its ancestry is important to our understand- ing of modern human origins in Australasia because the prevailing Stephen Oh models of human origins make different predictions for the ancestry Paleoanthropology Laboratory, of this specimen, and others like it. Some authors believe in the Department of Anthropology, validity of a complete replacement theory and propose that modern University of Michigan, humans in Australasia descended solely from earlier modern human Ann Arbor, MI 48109-1382, populations found in Late Pleistocene Africa and the Levant. These U.S.A. ancestral modern populations are believed to have completely replaced other archaic human populations, including the Ngandong hominids of Indonesia. According to this recent African origin theory, Keith Hunley the archaic humans from Indonesia are classified as Homo erectus,a Paleoanthropology Laboratory, different evolutionary species that could not have contributed to the Department of Anthropology, ancestry of modern Australasians. Therefore this theory of complete University of Michigan, replacement makes clear predictions concerning the ancestry of the Ann Arbor, MI 48109-1382, specimen WLH-50. We tested these predictions using two methods: U.S.A. E-mail: a discriminant analysis of metric data for three samples that are [email protected] potential ancestors of WLH-50 (Ngandong, Late Pleistocene Africans, Levant hominids from Skhul and Qafzeh) and a pairwise difference analysis of nonmetric data for individuals within these Seth Dobson samples. The results of these procedures provide an unambiguous Department of Anthropology, refutation of a model of complete replacement within this region, and Washington University, indicate that the Ngandong hominids or a population like them may St. Louis, MO 63130, have contributed significantly to the ancestry of WLH-50. We there- U.S.A. E-mail: fore contend that Ngandong hominids should be classified within the [email protected] evolutionary species, Homo sapiens. The Multiregional model of human evolution has the expectation that Australasian ancestry is in all three of the potentially ancestral groups and best explains modern Graciela Cabana Australasian origins. Paleoanthropology Laboratory, 2000 Academic Press Department of Anthropology, University of Michigan, Ann Arbor, MI 48109-1382, U.S.A. E-mail: [email protected] Praveen Dayalu Paleoanthropology Laboratory, Department of Anthropology, University of Michigan, Ann Arbor, MI 48109-1382, U.S.A. Milford H. Wolpoff Paleoanthropology Laboratory, Department of Anthropology, University of Michigan, Ann Arbor, MI 48109-1382, U.S.A. E-mail: wolpoff@umich.edu 0047–2484/00/070001+22$35.00/0 2000 Academic Press 2 . ET AL. Received 15 July 1998 Revision received 12 September 1998 and accepted 15 September 1999 Keywords: modern human origins, Australasia, recent African origins, Journal of Human Evolution (2000) 39, 1–22 multiregional evolution, doi:10.1006/jhev.1999.0384 WLH-50, Ngandong. Available online at http://www.idealibrary.com on Introduction hypothesis of modern human origins in Australasia have explained the seemingly Since its discovery in 1982, Willandra Lakes archaic nature of WLH-50 differently, as a Hominid (WLH) 50, as reconstructed by A. consequence of either its size and related Thorne, has figured prominently in discus- robusticity or possibly pathology (Stringer, sions of modern human origins in Australia 1998; Webb, 1990), but not as a result and Indonesia (Thorne, 1984; Webb, 1989; of any significant ancestor–descendant ff Thorne & Wolpo , 1992; Frayer et al., relationship with earlier Australasian 1993, 1994; Stringer, 1998). Dated to some populations. Despite these differences, both 15–13 ka by gamma spectrometric U-series interpretations agree on one fundamental item; analysis (Simpson & Gru¨n, 1998), and to WLH-50 is a modern human (Thorne, 1984; about double that by ESR (on bone, by Wolpoff, 1989, 1999; Stringer, 1998). Caddie et al., 1987), the WLH-50 cal- We therefore begin with the supposition varium appears on inspection to exhibit that earlier Late Pleistocene Africans, many features that closely resemble earlier Levantines (Near Eastern individuals from Indonesian hominids, including the Skhul and Qafzeh), and WLH-50 are mem- Ngandong fossils of Java [Thorne, cited in bers of the same species, Homo sapiens, and Wong (1999); Thorne et al., in preparation; all represent modern humans or their im- ff Wolpo , 1999]. Those authors who support mediate ancestors. There are two different a multiregional model of human evolution predictions about the pattern of ancestry, view the similarities between the Ngandong and it is from these that we can develop a hominids and WLH-50 as phylogenetic, test for the African replacement model for meaning in this case that these specimens modern human origins. The replacement are members of the same evolutionary model asserts: species and share some features by virtue of ( 1) The earlier Late Pleistocene Africans having come from the same region of the and Levantines are likely to be direct world (Frayer et al., 1993, 1994). Ngandong ancestors of WLH-50 by virtue of age is one of the probable ancestors of WLH-50, 1 and geography. in this view, or the two may share a recent ( 2) The Ngandong hominids cannot be common ancestor. However, authors who direct ancestors of WLH-50, by support the recent African replacement virtue of being in a different species, 1Even if the most recent Ngandong date estimate is H. erectus (Rightmire, 1990), whose correct (Swisher et al., 1996), and the oldest WLH-50 habitation on Java may have persisted estimate is correct (Caddie et al., 1987), Ngandong is older than WLH-50, and the Indonesian site may be well into the late Pleistocene (Swisher considerably older (Gru¨n & Thorne, 1997). et al., 1996). -50 3 The complete replacement hypothesis Table 1 Specimens in the comparative samples requires that there is a unique relationship Late Pleistocene between early modern humans, represented Ngandong Africans Levantines by the earlier Late Pleistocene Africans Levantines, and WLH-50, to the exclusion 1 Jebel Irhoud 1 Skhul 5 of more archaic hominids from Ngandong. 4* Jebel Irhoud 2 Skhul 9 We propose to test this complete replace- 5 Laetoli 18 Qafzeh 6 6 Omo 1* Qafzeh 9 ment hypothesis by demonstrating that one 9 Omo 2 or both of the above assertions must be 10 Singa incorrect. A recent African replacement 11 explanation of modern human origins in Australasia is incorrect if it could be shown *Used only in the nonmetric analysis. in a comparison of WLH-50 to earlier Late Pleistocene Africans, Levantines, and the Ngandong hominids from its ancestry. In Ngandong people that the closest relation- this case, replacement would be refuted as a ships are to Ngandong. This is because model for the ancestry of Australasia, and complete replacement predicts unique re- the Ngandong hominids must be accepted lationships between WLH-50 and the late as H. sapiens. Pleistocene Africans and individuals from Skhul/Qafzeh to the exclusion of Ngandong. Materials and methods Multiregional evolution, in contrast, views all three of these groups as potential Our fossil sample consists of the possible ancestors for WLH-50. Pleistocene ancestors for WLH-50 from the We have chosen two procedures to Levant, Africa and Indonesia, as described examine the relationship of WLH-50 to above. Seventeen crania from the earlier the African, Levantine, and Indonesian part of the Late Pleistocene were chosen for samples: comparison with WLH-50 (Table 1). We ( 1) A discriminant analysis of metric data assigned all fossils except for WLH-50 to which differentiates the Africans, one of three groups based on geography. Levantine, and Ngandong hominids These are the most complete crania from into three groups and can be used to this time period in these regions and include assess the group affinity of WLH-50, the specimens that preserve all or most of and the areas that are observable on WLH-50. ( 2) a pairwise difference analysis using Fragmentary specimens and specimens too non-metric data, comparing WLH-50 recent to be potential ancestors of WLH-50 with the individuals in these three were excluded from this study. The metric samples. variables were chosen to maximize the The prediction of the replacement model is number of measurements we could replicate that WLH-50 will sort with the Africans or in all the crania. The incomplete condition Levantines in both tests, since it must be of WLH-50 limited us to 21 variables, uniquely descended from these groups. If mostly standardized chords and arcs, which these procedures find that WLH-50 clusters could be accurately and repeatedly with Ngandong rather than with Africans measured. All of them were present on all or Levantines, then it must follow that crania, and no missing data were allowed for WLH-50 shares no special relationship with the discriminant analysis. Late Pleistocene hominids from Africa Four measurements of WLH-50 required and the Levant that would exclude the a reconstruction of the glabella region, 4 . ET AL. Table 2 Comparison of selected measurements different from Stringer’s, as addressed ff by di erent authors* for WLH-50 below. Stringer Brown All other measurements of fossil crania WLH-50 Measurement (1998) (1998) This were taken on the original specimens by one (in mm) value value study of us (MHW), with a small number of exceptions. A combination of published and Biasterionic breadth 123 127 123 cast measurements were used for Laetoli Biauricular breadth 138 138 Biparietal breadth 142 139 142 Hominid 18 (Magori & Day, 1983), and Bregma-asterion 151 151 150 Omo2(Day & Stringer, 1991).