WLH 50: How Australia Informs the Worldwide Pattern of Pleistocene Human Evolution MILFORD H WOLPOFF Department of Anthropology, University of Michigan, Ann Arbor, MI 48109-1092, USA; [email protected] SANG-HEE LEE Department of Anthropology, University of California at Riverside, Riverside, CA 92521-0418, USA; [email protected] submitted: 15 July 2013; accepted 9 September 2014 MONOGRAPH CONTENTS Abstract 506 Introduction 506 Background 508 Weidenreich 509 New Fossils, New Details 510 Migration versus Sources 511 Was There a Single Source Population? 513 The Simplest Origins Hypothesis 514 WLH 50 514 Goal of the Description and Analysis 514 Pathology or Artificial Deformation as Explanations for the Anatomy of WLH 50 517 Adaptive Explanations 517 Age or Size Related Ruggedness 517 Artificial Deformation of WLH 50—A Red Herring 517 Pathological Explanation for Cranial Thickness 519 Allometry 520 No Compelling Explanations Replace Ancestry 520 Description of WLH 50 and Comparison with Ngandong Sample 520 Condition and Preservation 520 Vault as a Whole 521 Cranial Bone Thickness 528 Frontal Bone 530 Supraorbital Region 530 Internal Surface 534 Parietal Bones 536 Temporal Bones 538 Occipital 540 Zygomatic Bone 541 Statistical Approaches 542 Discriminant Analysis of Metric Data 543 Pairwise Difference Analysis Using Non-Metric Data 545 STET (STandard Error Test) 545 Standard Error of the Regression Slope 547 Does STET “Work”? 547 STET Analysis 548 How Similar is WLH 50 to Ngandong and to the African Sample? 550 Summary of Results From the Statistical Approaches 551 In Some Tests Reported Above, WLH 50 is Most like the Ngandong Sample 551 In Other Tests, the Hypothesis of Multiple Ancestry (Africans and Ngandong) for WLH 50 Cannot be Rejected 551 PaleoAnthropology 2014: 505−564. © 2014 PaleoAnthropology Society. All rights reserved. ISSN 1545-0031 doi:10.4207/PA.2014.ART88 506 • PaleoAnthropology 2014 MONOGRAPH CONTENTS (CONTINUED) Genetic Basis for Mixed Australian Ancestry 551 mtDNA 552 nDNA 553 Who the Denisovans Might Have Been 554 Summary and Conclusions 555 Taxonomic Assessment 555 The Role of WLH 50 in Understanding Human Evolution 555 Acknowledgements 556 Endnotes 556 References 558 MONOGRAPH ABSTRACT WLH 50 is the ~26,000 kyr (16.5–37.4 kyr) year-old calotte of a large (1540cc) young-to-middle-aged male found in dune deposits near Lake Garnpung, one of the Willandra lakes in New South Wales, Australia Its importance stems from the significant similarities it appears to have with the nearby and most probably earlier human sample from Ngandong, Indonesia. Two implications of these similarities are important―the implications for Australian ancestry and the implications for the place of Ngandong in human evolution. We discuss the historic context for is- sues of indigenous Australian ancestry and show how WLH 50 informs these issues We provide both a systematic and comprehensive description of the WLH 50 fossil in a comparative context, and an interpretation of its place in human evolution A comparative analysis of the WLH 50 cranium is important for understanding its morphologi- cal pattern and addressing its ancestry; in particular, whether there are multiple ancestors for WLH 50 and other Australians that include Late Pleistocene Indonesians from Ngandong Besides these anatomical considerations, the issues of indigenous Australian ancestry also are framed by genetic analysis including both paleogenetics and observations of mitochondrial and nuclear DNA from modern populations, and we discuss their implications as well We conclude that WLH 50 is a normal specimen without evidence of pathology or deformation, and that elements of its anatomy are found throughout the Australian fossil record, as it is known today Our compari- sons reveal WLH 50 has substantial similarities to the Ngandong hominids (but not identity with them) and a demonstrable pattern of multiple ancestors that includes Ngandong, as part of the network of human evolution in Australasia “WLH 50, an opalised cranium and partial skeleton, kyr) calotte of a large (1540cc) young-to-middle-aged male found on a lakeshore to the north of Mungo, is much found in dune deposits near one of the Willandra lakes in more robust and archaic than any Australian hominid New South Wales, Australia The overarching issue this found previously It makes the Kow Swamp remains look gracile by comparison This new specimen greatly specimen has raised is about its anatomy and the question reinforces the regional continuity linking Indonesia and of whether and how its anatomy reflects its ancestry. More Australia The morphological sequence found in Aus- than any other Australian fossil, WLH 50 addresses some tralia shows clearly that regional features must be sepa- of the more general issues prominent in studies of Late rated from grade features seen worldwide in order to Pleistocene and Holocene human evolution One of these is understand the differing patterns of Late Pleistocene hu- man evolutionary change.” Alan G. Thorne [1939–2012] the prevalence of evidence for mixture between seemingly (1984). different hominids, now recognized in studies of human genetics (Pääbo 2014). Another is the recognition that these mixing groups had more genetic variation between popu- INTRODUCTION lations than is now evident in Homo sapiens (Hawks 2013). iscussions of the details of Australian prehistory have The consequences of these two facts should be evident in Dcontinued for more than a century, and play a key role anatomical studies as well as genetic ones, and WLH 50 in the ongoing understanding of Late Pleistocene and Ho- has the potential to address them In this regard Australian locene human evolution across the world Questions con- prehistory may be seen to resemble European prehistory, cern how Australians relate to other human populations in that the modern Europeans have a complex ancestry in Do Australians have one or several ancestral sources, and which different human subspecies (Wolpoff 2009) mixed in how can the complex pattern of migrations to Australia the Pleistocene (Wolpoff et al. 2001). be untangled? In this monograph we turn to WLH 50, a Australia is one of the most recent regions of the Old previously undescribed specimen that informs these and World to be inhabited It is widely, perhaps universally, ac- other related issues. WLH 50 is a ~26,000 kyr (16.5–37.4 cepted that the human migration or migrations to Australia WLH 50, Australia, and Pleistocene Human Evolution • 507 Figure 1. The Pleistocene extent of Sahul and Sunda, with the locations of Late Pleistocene fossil sites. Pleistocene Australia, the major part of Sahul, shows the locations of most of the fossils discussed here. At no time during the Pleistocene was it possible to reach Aus- tralia without crossing water (from Pardoe (2006, Figure 1), with permission). were of modern humans with an established, sophisticat- many consider this Australian colonization as migration ed seafaring technology (O’Connell et al. 2010) based on rather than dispersal (Balme 2013). bamboo (Birdsell 1977). Once purposeful, albeit sporadic, The deep prehistory of Australian ancestors cannot be movement to the continent began, there is no reason to ex- found in Australia. It is in this context that comparisons of pect that it was interrupted for long Beginning some 50 kyr the Late Pleistocene/Holocene fossil record of Australia and ago, the ancestors of recent indigenous Australians began the nearby lands are significant. With its peripheral loca- to cross a biogeographic divide (Wallacea) over gaps, some tion, isolation by sea (Figure 1), and the likelihood that gene of which were as much as (depending on the sea level at the flow to the continent was limited to the later Pleistocene time) 70km of water, a distance far enough beyond the ho- and mostly one-way as noted above, Australia has come to rizon that it was not possible to see across to land The pre- play a unique role in understanding the worldwide pattern vailing currents and the absence of suitable, bamboo-like of Pleistocene human evolution, and WLH 50 addresses building materials in Australia assured that the direction of key issues of recent indigenous Australian ancestry Ana- travel was most likely one-way (Allen and O’Connell 2008; tomical comparisons and genetic evidence, discussed be- Westaway and Lambert 2014) Because it was uninhabited, low, clearly reflect what has long been obvious; Australians 508 • PaleoAnthropology 2014 came from East and Southeast Asia. But what parts of Asia, history of the region from genetic analyses and these help and when, have been researched and discussed for more frame the possibilities for WLH 50, Ngandong, and their than a century relationship Throughout the 20th century there was a succession of Paraphrasing the words of Silvana Condemi in intro- scientists who contended the source population of Austra- ducing her monograph on the Saccopastore Neandertal lian ancestors was Indonesian The putative Indonesian an- crania (1992), the goal of this monograph is to provide both cestors were first thought to be the Pithecanthropus remains a description and an interpretation of the WLH 50 fossil In known from Trinil (later including the Sangiran crania), this monograph we ask about the place of WLH 50 in hu- and finally settled to be their descendants, represented by man evolution, and what the pattern of human evolution the more recent Ngandong crania Ngandong is the sample has been that creates such a place for this specimen? of potential ancestors nearest to Australia in space, and clos- est to WLH 50 in time Yet, it would be fair to say that the BACKGROUND contention that some of the ancestors of Australians were Hermann Klaatsch [1863–1916] addressed the pattern of from Ngandong, or from Ngandong-like populations, has ancestry in Australia early in the 20th century, proposing been and continues to be controversial The problem cre- a single lineage interpretation of indigenous Australian ated by a Ngandong ancestry is that the Ngandong crania origins (1908a).