Dating the Drimolen hominins Fossil hominins from South Africa are enriching the story of early human evolution and dispersal. Herries et al. describe the geological context and dating of the hominin-bearing infilled cave, or palaeocave, at a site called Drimolen in South Africa (see the Perspective by Antón). They focus on the age and context of a recently discovered Homo erectus sensu lato fossil and a Paranthropus robustus fossil, which they dated to ∼2.04 million to 1.95 million years ago. This makes Drimolen one of the best-dated sites in South Africa and establishes these fossils as the oldest definitive specimens of their respective species ever discovered. The age confirms that species of Australopithecus, Paranthropus, and early Homo overlapped in the karst of South Africa ∼2 million years ago. Science, this issue p. eaaw7293; see also p. 34 Multiple hominin genera, including the earliest Homo erectus lineage, were present in South Africa 2 million years ago. INTRODUCTION Drimolen is one of several ancient caves located in the Hominid Caves of South Africa United Nations Educational, Scientific, and Cultural Organization (UNESCO) World Heritage Area in South Africa. Between ~2.3 million and ~1.8 million years ago, there were major climactic changes and faunal turnovers in the region, including the last occurrence of the genus Australopithecus and the first occurrence of Paranthropus and Homo, as well as the first occurrence of stone and bone tools. However, the exact nature of these changes has been hard to elucidate because of past difficulties in dating caves of this age and their perceived geological complexity. Unlike in eastern Africa, where volcanic material is available for dating, the South African caves have been dated with a variety of evolving methods that have often given conflicting age estimates. This means that South Africa’s early human record and its relationship to east African hominin species have been difficult to determine. This is especially problematic given that each record is distinct in terms of hominin species until perhaps the origin and early evolution of the genus Homo. Although many fragmentary fossil specimens in South Africa have been attributed to early Homo, there is no consensus regarding species attribution. RATIONALE Drimolen Main Quarry has yielded one of the richest records of early human fossils in South Africa, including examples of Homo and the most complete female skull (DNH 7) of Paranthropus robustus. Excavations between 2015 and 2018 yielded the first new hominin calvaria (DNH 134 and DNH 152) from the site in 20 years. A combination of uranium-lead dating on flowstones, uranium-series electron spin resonance (US-ESR) dating on teeth, and palaeomagnetism on sediments was undertaken to establish the age of the site and its early human fossils. RESULTS The DNH 134 cranium shares clear affinities with Homo erectus, whereas the DNH 152 cranium represents P. robustus. Stratigraphic analysis of the Drimolen Main Quarry deposits indicates that unlike many other South African sites, there was only one major phase of relatively short deposition between ~2.04 million years ago and ~1.95 million years ago. This age has been constrained by the identification of the ~1.95-million-year-old magnetic field reversal at the base of the Olduvai SubChron within the sediments and by the direct uranium-lead dating of a flowstone that formed during the reversal. This has been augmented by direct dating on fossils by means of US-ESR that suggests that the DNH 134 and DNH 152 crania were deposited just before this reversal, with the DNH 134 crania deposited at ~2.04 million years ago. The DNH 134 cranium shares affinities with H. erectus and predates all known specimens in that species. The age range of Drimolen Main Quarry overlaps with that of Australopithecus sediba from the nearby site of Malapa and indicates that Homo, Paranthropus, and Australopithecus were contemporaneous in South Africa between 2.04 million and 1.95 million years ago. It is the first time that dating has conclusively demonstrated that these three taxa shared the same landscape during the same time range, making it less likely that a population of A. sediba is ancestral to Homo, as has been previously suggested. Analysis of fauna preserved at Drimolen documents a period of ecological change, with earlier South African species going extinct and new species moving into the region from other parts of Africa, including early representatives of H. erectus. CONCLUSION Drimolen is the best dated early hominin site in South Africa. DNH 134 is the oldest and best preserved Early Pleistocene Homo cranium from South Africa. The DNH 134 Homo cranium has affinities with H. erectus and extends the species’ temporal range by ~200,000 to 150,000 years. DNH 134 being older than A. sediba complicates the likelihood of this species being ancestral to Homo in South Africa, as previously suggested. With the oldest occurrence of H. erectus at the southern tip of Africa, this argues against a suggested Asian origin for H. erectus. DNH 152 represents the oldest P. robustus cranium in South Africa. The Drimolen stone and bone tools are also the oldest from the region. The faunal community from Drimolen as a whole indicates substantial changes in South African ecosystems, with many first and last appearance dates of species that are related to the extinction of some indigenous South African species and the migration of others into the region ~2 million years ago, likely including Homo erectus. The DNH 134 H. erectus cranium from South Africa. PHOTO: JESSE MARTIN, REANUD JOANNES-BOYAU, ANDY I. R. HERRIES Understanding the extinction of Australopithecus and origins of Paranthropus and Homo in South Africa has been hampered by the perceived complex geological context of hominin fossils, poor chronological resolution, and a lack of well-preserved early Homo specimens. We describe, date, and contextualize the discovery of two hominin crania from Drimolen Main Quarry in South Africa. At ~2.04 million to 1.95 million years old, DNH 152 represents the earliest definitive occurrence of Paranthropus robustus, and DNH 134 represents the earliest occurrence of a cranium with clear affinities to Homo erectus. These crania also show that Homo, Paranthropus, and Australopithecus were contemporaneous at ~2 million years ago. This high taxonomic diversity is also reflected in non-hominin species and provides evidence of endemic evolution and dispersal during a period of climatic variability.

中文翻译：

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南非南方古猿、副人猿和早期直立人的同期性

约会 Drimolen 古人类 来自南非的古人类化石丰富了早期人类进化和传播的故事。赫里斯等人。描述了在南非一个名为 Drimolen 的地点的古人类填充洞穴或古洞穴的地质背景和年代（参见 Antón 的观点）。他们专注于最近发现的直立人化石和粗体旁人化石的年龄和背景，它们的年代可追溯到约 204 万至 195 万年前。这使得 Drimolen 成为南非年代最久远的遗址之一，并将这些化石确立为迄今为止发现的各自物种中最古老的最终标本。该年龄证实了大约 200 万年前，南方古猿、副人类和早期人类的物种在南非的喀斯特地区重叠。科学，这个问题 p。eaaw7293; 另见第 34 多个人类属，包括最早的直立人谱系，在 200 万年前出现在南非。简介 Drimolen 是位于南非联合国教育、科学和文化组织 (UNESCO) 世界遗产区南非原始人洞穴的几个古老洞穴之一。在大约 230 万至 180 万年前，该地区发生了重大气候变化和动物群更替，包括南方古猿属的最后一次出现和副人类和人的第一次出现，以及石和骨的首次出现工具。然而，这些变化的确切性质一直难以阐明，因为过去很难对这个时代的洞穴进行年代测定，而且它们感知到的地质复杂性。与东非不同，在火山物质可用于测年的地方，南非的洞穴已经用各种不断发展的方法进行了测年，这些方法经常给出相互矛盾的年龄估计。这意味着南非的早期人类记录及其与东非人类物种的关系难以确定。鉴于每个记录在人类物种方面都是不同的，直到可能直到人类属的起源和早期进化，这尤其成问题。尽管南非的许多零碎化石标本都被归于早期的人类，但在物种归属方面还没有达成共识。基本原理 Drimolen 主要采石场产生了南非最丰富的早期人类化石记录之一，包括人类的例子和最完整的女性头骨 (DNH 7) 粗壮副人猿。2015 年至 2018 年间的挖掘工作在 20 年内从该地点发现了第一批新的人类颅骨（DNH 134 和 DNH 152）。结合流石上的铀铅测年、牙齿上的铀系列电子自旋共振 (US-ESR) 测年和沉积物的古地磁确定该遗址及其早期人类化石的年龄。结果 DNH 134 颅骨与直立人有明显的亲缘关系，而 DNH 152 颅骨代表 P.robustus。对 Drimolen Main Quarry 矿床的地层分析表明，与许多其他南非遗址不同，在 ~204 万年前和 ~195 万年前之间只有一个相对较短的主要沉积阶段。这个年龄受到了~1的识别的限制。9500 万年前的磁场在沉积物中的奥杜威亚纪元底部发生逆转，并通过对逆转期间形成的流石进行直接铀铅测年。通过 US-ESR 对化石进行直接测年，这表明 DNH 134 和 DNH 152 颅骨就在此逆转之前沉积，而 DNH 134 颅骨沉积于约 204 万年前。DNH 134 颅骨与直立人有亲缘关系，并且早于该物种中所有已知的标本。Drimolen Main Quarry 的年龄范围与附近马拉帕遗址的 Australopithecus sediba 的年龄范围重叠，表明在 204 万至 195 万年前，在南非同时期出现了人类、副人类和南方古猿。这是第一次确定性地证明这三个分类群在同一时间范围内共享相同的景观，这使得 A. sediba 种群不太可能像之前提出的那样是人类的祖先。对保存在 Drimolen 的动物群的分析记录了一段生态变化，早期的南非物种灭绝，新物种从非洲其他地区进入该地区，包括直立人的早期代表。结论 Drimolen 是南非最好的早期人类遗址。DNH 134 是南非最古老、保存最完好的早更新世人类头盖骨。DNH 134 Homo 颅骨与直立人有亲缘关系，并将该物种的时间范围延长了约 200,000 至 150,000 年。DNH 134 比 A 年长。如前所述， sediba 使该物种成为南非人祖先的可能性复杂化。由于最古老的直立人出现在非洲的南端，这与建议的直立人亚洲起源相矛盾。DNH 152 代表南非最古老的 P.robustus 颅骨。Drimolen 石器和骨器也是该地区最古老的工具。来自 Drimolen 的动物群落作为一个整体表明南非生态系统发生了重大变化，许多物种的首次和最后出现日期与约 200 万年前一些南非本土物种的灭绝和其他物种迁徙到该地区有关，可能包括直立人。来自南非的 DNH 134 H.直立人头盖骨。照片：JESSE MARTIN、REANUD JOANNES-BOYAU、ANDY IR HERRIES 人类化石的复杂地质背景、年代分辨率差以及缺乏保存完好的早期人类标本阻碍了对南方古猿的灭绝以及旁人猿和人在南非的起源的了解。我们描述了在南非 Drimolen Main Quarry 发现的两个古人类颅骨的发现、日期和背景。在大约 204 万到 195 万年的历史中，DNH 152 代表了粗壮副人猿的最早出现，而 DNH 134 代表了与直立人有明确亲缘关系的头盖骨的最早出现。这些颅骨还表明，大约 200 万年前，人类、副人类和南方古猿是同时代的。