RESEARCH ◥ the DNH 134 crania deposited at ~2.04 million RESEARCH ARTICLE SUMMARY years ago. The DNH 134 cranium shares af- finities with H. erectus and predates all known PALEOANTHROPOLOGY specimens in that species. The age range of Drimolen Main Quarry overlaps with that of Contemporaneity of Australopithecus, Paranthropus, Australopithecus sediba fromthenearbysite of Malapa and indicates that Homo, Paran- and early Homo erectus in South Africa thropus, and Australopithecus were contem- poraneous in South Africa between 2.04 million Andy I. R. Herries*†, Jesse M. Martin†, A. B. Leece†, Justin W. Adams†, Giovanni Boschian†, and1.95millionyearsago.Itisthefirsttime Renaud Joannes-Boyau, Tara R. Edwards, Tom Mallett, Jason Massey, Ashleigh Murszewski, ◥ that dating has conclu- Simon Neubauer, Robyn Pickering, David S. Strait, Brian J. Armstrong, Stephanie Baker, ON OUR WEBSITE sively demonstrated that Matthew V. Caruana, Tim Denham, John Hellstrom, Jacopo Moggi-Cecchi, Simon Mokobane, Read the full article these three taxa shared Paul Penzo-Kajewski, Douglass S. Rovinsky, Gary T. Schwartz, Rhiannon C. Stammers, Coen Wilson, at http://dx.doi. the same landscape dur- Jon Woodhead, Colin Menter org/10.1126/ ing the same time range, science.aaw7293 making it less likely that .................................................. a population of A. sediba INTRODUCTION: Drimolen is one of several spin resonance (US-ESR) dating on teeth, and is ancestral to Homo, as has been previous- ancient caves located in the Hominid Caves of palaeomagnetism on sediments was under- ly suggested. Analysis of fauna preserved at South Africa United Nations Educational, Scien- taken to establish the age of the site and its Drimolen documents a period of ecological tific, and Cultural Organization (UNESCO) World early human fossils. change, with earlier South African species Downloaded from Heritage Area in South Africa. Between ~2.3 going extinct and new species moving into million and ~1.8 million years ago, there were RESULTS: The DNH 134 cranium shares clear the region from other parts of Africa, includ- major climactic changes and faunal turnovers affinities with Homo erectus,whereastheDNH ing early representatives of H. erectus. in the region, including the last occurrence of 152 cranium represents P. robustus. Stratigraphic the genus Australopithecus and the first oc- analysis of the Drimolen Main Quarry deposits CONCLUSION: Drimolen is the best dated early Paranthropus Homo currence of and ,aswell indicates that unlike many other South African hominin site in South Africa. DNH 134 is http://science.sciencemag.org/ as the first occurrence of stone the oldest and best preserved and bone tools. However, the ex- Early Pleistocene Homo cra- act nature of these changes has nium from South Africa. The been hard to elucidate because DNH 134 Homo cranium has of past difficulties in dating caves affinities with H. erectus and of this age and their perceived extends the species’ temporal geological complexity. Unlike in range by ~200,000 to 150,000 eastern Africa, where volcanic years. DNH 134 being older material is available for dating, than A. sediba complicates the the South African caves have been likelihood of this species being dated with a variety of evolv- ancestral to Homo in South Af- on April 4, 2020 ing methods that have often rica, as previously suggested. given conflicting age estimates. With the oldest occurrence of This means that South Africa’s H. erectus at the southern tip early human record and its rela- ofAfrica,thisarguesagainst tionship to east African hominin asuggestedAsianoriginfor species have been difficult to H. erectus. DNH 152 represents determine. This is especially the oldest P. robustus cranium problematic given that each in South Africa. The Drimolen record is distinct in terms of stone and bone tools are also hominin species until perhaps the oldest from the region. the origin and early evolution The faunal community from of the genus Homo.Although Drimolen as a whole indicates H. erectus many fragmentary fossil speci- The DNH 134 cranium from South Africa. substantial changes in South mens in South Africa have been African ecosystems, with many attributed to early Homo, there is no consen- sites, there was only one major phase of relatively first and last appearance dates of species sus regarding species attribution. short deposition between ~2.04 million years that are related to the extinction of some ago and ~1.95 million years ago. This age has indigenous South African species and the RATIONALE: Drimolen Main Quarry has yielded been constrained by the identification of the migration of others into the region ~2 million one of the richest records of early human fossils ~1.95-million-year-old magnetic field rever- years ago, likely including Homo erectus.▪ in South Africa, including examples of Homo sal at the base of the Olduvai SubChron within and the most complete female skull (DNH 7) of the sediments and by the direct uranium-lead Paranthropus robustus. Excavations between dating of a flowstone that formed during the 2015 and 2018 yielded the first new hominin reversal. This has been augmented by direct The list of author affiliations is available in the full article online. calvaria (DNH 134 and DNH 152) from the site dating on fossils by means of US-ESR that sug- *Corresponding author. Email: [email protected] †These authors contributed equally to this work. in 20 years. A combination of uranium-lead gests that the DNH 134 and DNH 152 crania Cite this article as A. I. R. Herries et al., Science 368, PHOTO: JESSE MARTIN, REANUDdating JOANNES-BOYAU, ANDY I. R. HERRIES on flowstones, uranium-series electron were deposited just before this reversal, with eaaw7293 (2020). DOI: 10.1126/science.aaw7293 Herries et al., Science 368, 47 (2020) 3 April 2020 1of1 RESEARCH ◥ difficulty in dating palaeokarst owing to a lack RESEARCH ARTICLE of suitable material for radiometric dating (5 , 6). Until recently, most dates for South PALEOANTHROPOLOGY African sites were based on biostratigraphic correlation with the better-dated eastern Afri- Contemporaneity of Australopithecus, Paranthropus, can sites some 3000 to 4000 km away (5, 6). The stratigraphic sequences at most South and early Homo erectus in South Africa African palaeocave sites have been defined according to a lithostratigraphic approach in Andy I. R. Herries1,2*†, Jesse M. Martin1†, A. B. Leece1†, Justin W. Adams3,2†, Giovanni Boschian4,2†, which breccia deposits and siltstone deposits Renaud Joannes-Boyau5,2, Tara R. Edwards1‡, Tom Mallett1, Jason Massey3,6, Ashleigh Murszewski1, have been classified as sequential numbered Mem- Simon Neubauer7, Robyn Pickering8,9, David S. Strait10,2, Brian J. Armstrong2, Stephanie Baker2, bers, thought to represent different temporal Matthew V. Caruana2, Tim Denham11, John Hellstrom12, Jacopo Moggi-Cecchi13, Simon Mokobane2, phases of deposition (11–14). In some cases, in Paul Penzo-Kajewski1, Douglass S. Rovinsky3, Gary T. Schwartz14, Rhiannon C. Stammers1, which stratigraphic contacts can be identified, Coen Wilson1, Jon Woodhead12, Colin Menter13 older lithofacies have clearly collapsed, subsided, or been eroded by secondary cave-formation Understanding the extinction of Australopithecus and origins of Paranthropus and Homo in South processes before later lithofacies infilled the Africa has been hampered by the perceived complex geological context of hominin fossils, poor resulting space (15). This often leads to com- chronological resolution, and a lack of well-preserved early Homo specimens. We describe, date, and plexities such as deposits formed by rework- contextualize the discovery of two hominin crania from Drimolen Main Quarry in South Africa. At ing of older units and thus mixing of fossil ~2.04 million to 1.95 million years old, DNH 152 represents the earliest definitive occurrence of material (5, 15, 16). In other cases, in which Downloaded from Paranthropus robustus, and DNH 134 represents the earliest occurrence of a cranium with clear affinities stratigraphic links are obscured, the relation- to Homo erectus. These crania also show that Homo, Paranthropus, and Australopithecus were ship of different lithologies is less certain, and contemporaneous at ~2 million years ago. This high taxonomic diversity is also reflected in non-hominin the depth that deposits have been accumulated species and provides evidence of endemic evolution and dispersal during a period of climatic variability. within the cave has been used to suggest that this also represents temporal depth, even when a continuous depositional sequence cannot be http://science.sciencemag.org/ emnants of infilled and eroded cave sys- (1–4), where the origins of Homo have been identified (6, 13). However, caves do not always tems (palaeocaves) formed within the suggested to occur much earlier (7). However, conform to the law of superposition, and depo- Malmani dolomite in northeastern South the fragmented nature of most South African sits can be inverted or reworked or contain R Africa have yielded one of the richest early Homo specimens (such as StW 53, SK 15, intrusive events (5, 16–18). At other sites, where early hominin records in the world, in- andSK847)makesthetaxonomyofthisgroup mining or erosion has not obscured strati- cluding Australopithecus africanus, Australo- and its relationship to eastern African Homo graphic relationships, it is clear that different pithecus sediba, Paranthropus robustus,and uncertain, with many fossil specimens assigned