Within-group human variation in the Asian Pleistocene: the three Upper Cave crania

Numerous studies on Pleistocene samples have shown that within-group cranial variation was greater than that seen today. The three anatomically modern Upper Cave crania (UC 101, UC 102, and UC 103) from Zhoukoudian, China provide one of the best samples available for addressing the issue of the antiquity of the modern pattern of within-group cranial variability because archaeological evidence indicates that they are spatially and temporally restricted. Research on the Upper Cave fossils usually only includes UC 101 and UC 103 because of postmortem damage to UC 102's cranial vault. However, the face of UC 102 is undamaged, allowing for most facial measurements to be performed accurately. In this study we use facial dimensions to compare all three Upper Cave specimens, and we evaluate whether the variation seen among them is larger than that observed in extant populations.Using a worldwide sample of modern populations to establish a baseline, the three Upper Cave crania were compared to each other. Since there is disagreement over the sex of UC 102, this specimen is treated alternately as a female and as a male. Results show that the Upper Cave specimens exhibit significantly more variation than do individuals within more recent human populations, especially if UC 102 is considered male. Furthermore, results indicate that the fossils never fall into the same modern human group, and that each specimen is significantly atypical of its nearest modern neighbor in multivariate space.We conclude that the three Upper Cave crania do not represent a family group but are representative of the larger contemporaneous heterogeneous Asian Pleistocene population. Our results support the contention that today's within-group homogeneity is a relatively recent phenomenon, and is likely the result of a Neolithic population expansion and its many effects.     

柳江人头骨形态特征及柳江人演化的一些问题

为深入探讨柳江人化石形态特征的表现特点、进化程度及其与其年代数据的吻合性,本文对柳江、山顶洞、资阳、丽江等主要中国更新世晚期人类头骨化石及1 114例全新世以来不同地区现代中国人头骨进行了对比分析,结果发现:柳江人头骨绝大多数特征的出现情况位于现代中国人的变异范围,只有极个别特征与现代人不同;柳江人头骨具有的低眶等特征也可见于其他中国更新世晚期人类化石,说明柳江人化石上保留有少量常见于更新世晚期人类的原始特征,但与其他中国更新世晚期人类,尤其是山顶洞人头骨相比,柳江人显得要现代的多;柳江人与山顶洞人之间头骨形态特征的差异以体现头骨原始性及粗硕强壮程度上的差别居多,而个别特征差异或许与气候环境适应有关。我们认为:柳江人在形态进化上与现代中国人已经非常接近,他们之间的差别非常小;柳江人与山顶洞人头骨特征表现上的差异主要反映了他们之间在演化程度上的差异,同时也在一定程度上体现了各自的生存环境;现有的形态学证据不大可能为柳江人较早的时代提供支持。     

Into Eurasia: A geometric morphometric re-assessment of the Upper Cave (Zhoukoudian) specimens

Since the discovery of the human remains from the Upper Cave of Zhoukoudian in the 1930s there has been speculation over their affinities. In particular, the degree to which the three adult crania exhibit recent East Asian morphology, as well as their degree of within-group variability, has long been debated. Several early researchers described a resemblance to East Asian populations, but these findings have been for the most part rejected by more recent metric and non-metric analyses. Nevertheless, the Upper Cave specimens have not been classified conclusively into any recent modern human population to which they have been compared, and classification results differ for each cranium. Here, the question of the affinities of Upper Cave 101 and 103, the two better-preserved crania, is examined from the perspective of the Late Pleistocene human fossil record using the methodology of 3-D geometric morphometrics. The degree of morphological variation between the two specimens is also evaluated within the context of recent population variability. Neurocranial and facial morphology are analyzed separately so as to maximize comparative samples. Results show a morphological resemblance of the Upper Cave material to Upper Paleolithic Europeans. It is proposed that the Upper Cave specimens retain important aspects of modern human ancestral morphology, and possibly share a recent common ancestral population with Upper Paleolithic Europeans, in accordance with the Single Origin model of modern human origins.     

山顶洞人与现代华北人头骨非测量性特征比较及中国更新世晚期人类演化的一些问题

山顶洞人化石是迄今在东亚地区发现的数量最为丰富、保存状态最为完好的更新世晚期人类化石。多年来,山顶洞人作为东亚地区更新世晚期人类的代表,在研究东亚地区现代蒙古人种及美洲印第安人起源方面发挥着重要作用。随着对现代人起源与演化研究的深入,学术界对更新世晚期人类演化及现代人群形成过程的许多细节及演化机制予以了越来越多的注意,提出了现代人群出现时间及现代人群分化时间的概念。一些研究对山顶洞人的演化程度及其在东亚地区现代人群形成上的作用也提出了不同看法。为了深入探讨这些问题,本文对12项颅骨非测量特征在山顶洞人与现代中国人的表现情况进行了对比研究。发现有8项特征在山顶洞人与现代中国人之间具有不同的表现,作者认为山顶洞人在颅骨特征表现上较现代中国人具有更多的原始性。在此基础上,作者就更新世晚期人群内部变异、现代人群特征标准等问题进行了讨论。     

山顶洞101号男性头骨的三维颅面复原

山顶洞101号头骨化石是东亚地区保存最为完整的化石之一,是探讨东亚地区现代人起源的重要研究材料。本文依据数据集中现生人的面部软组织平均分布,提出了计算机三维颅面复原方法,实现了101号头骨生前面貌的预测复原。主要包括三个步骤：首先使用CT完成了101号男性头骨和下颌骨仿制模型的三维重建。然后,利用计算机技术将现生人的面部软组织分布作为101号头骨的面部软组织分布,实现了颅面虚拟复原,并采用手工绘画技巧再现了复原面貌的形态特征。最后,提出了一种基于面部软组织分布和面貌统计形状模型的形态分析方法,实现了颅面复原结果的评估。山顶洞101号头骨的复原面貌具有头部较长、额头前倾、眉弓粗壮等特征,与101号头骨的几何形态基本一致。该技术再现了更新世晚期人类的脑颅及面部的形态特征,为古人类颅面复原的研究提供了技术支持和参考资料。     

Comparaison de l'expression de certains caractères crâniens sur les hominidés chinois du Pléistocène récent et de l'Holocène (grotte supérieure de Zhoukoudian, sites de Longxian et de Yanqing)

The human remains recovered from “Grotte supérieure de Zhoukoudian” are the best-preserved Late Pleistocene human fossils in East Asia. For decades, as the representative of the Late Pleistocene human in East Asia, the Upper Cave skulls have been playing important role in the research of origins of modern Mongoloids and American Indians. With the advance of the origin and evolution of modern humans, more attention has been paid to the details and the mechanisms for the late Pleistocene human evolution and the formation of modern human populations. Both the origin and diversification of modern humans have been stressed. Some studies further trigger the debaters on the Upper Cave Man concerning its evolutionary level and its role in the formation of modern human populations in East Asia. To further explore these problems, we examined and compared 12 non-metric features on the 3 Late Pleistocene Upper Cave skulls and 162 Holocene individuals earthed from two archaeological sites of North China (Longxian and Yanqing). Our results indicate that 8 on the 12 features have different expression patterns between Upper Cave Man and recent Chinese leading the authors to believe that more primitive expressions appeared on the Upper Cave Man than on recent Chinese populations. Based on these findings, some problems on the intragroup variation in Late Pleistocene and Holocene populations are discussed.     

广西崇左木榄山智人洞的鼠科化石

最近在广西崇左木榄山智人洞遗址采集到1件具有现代智人解剖特征初始状态的下颌骨和丰富的哺乳动物化石。啮齿类中的鼠科化石有锡金小鼠(Mus pahari)、中华姬鼠(Apodemus draco)、黑线姬鼠(Apodemus agrarius)、似德氏狨鼠(Hapalomys cf.H.delacouri)、笔尾树鼠(Chiropodomys gliroides)、社鼠(Niviventer confucianus)、针毛鼠(Niviventer fulvescens)、爱氏巨鼠(Leopoldamys edwardsi)、印度板齿鼠(Bandicota indica)、褐家鼠(Rattus norvegicus)和黄毛鼠(Rattus losea)共8属11种。这种全部由当地或邻近地区现生种类构成的组合显然比当地早更新世的三合大洞和川黔地区中更新世歌乐山期的鼠类组合进步, 而与广西田东雾云洞的鼠类组合相似, 其动物群的时代为晚更新世早期, 与不平衡铀系法得到的测年结果(距今约11万年)相吻合。智人洞鼠科动物的组合明显具有东洋界热带-亚热带动物群的特点, 其中林灌和草地型所占比例较大表明当时的森林面积可能减少、林灌和草地面积可能增加, 反映出当时的气候相对干旱。     

Recent human evolution in East Asia and Australasia.

In both East Asia and Australasia arguments for evolutionary continuity between middle-late Pleistocene hominid populations and modern Homo sapiens are of long standing. In both regions, however, problems of chronological distribution, dating and preservation of hominid skeletal materials provide an effective barrier to extending regional sequences back to 'archaic' Homo sapiens or Homo erectus. The earliest securely dated modern Homo sapiens in East Asia are currently represented by Zhoukoudian Upper Cave at a minimum of 29 ka BP. In Australia skeletal remains of modern Homo sapiens have been dated to 26 ka BP, with archaeological materials at 38 to 50 ka BP. Late Pleistocene human skeletons from sites like Coobool Creek are morphologically and metrically outside the range of recent Australian Aboriginal populations. Similarly Liujiang and the Upper Cave crania can be distinguished from recent East Asian 'Mongoloids'. Evolutionary change within the Holocene needs to be taken into consideration when the evidence for regional evolutionary continuity is considered.     

贵州兴义猫猫洞出土的人类化石

本文主要是对贵州兴义猫猫洞遗址出土的人化石进行的初步研究,并对哺乳动物化石、地层、堆积物作叙述。该遗址的人类化石包括下颌骨4件,股骨3段,是中国南方洞穴遗址中发现的数量较多,系统分类地位较清楚的晚期智人的化石。人类化石同层的鹿牙作铀系法测年结果为14500±1200 BP,相当于更新世的末期。     

中国晚旧石器时代人类与其南邻(尼阿人及塔邦人)的关系

本文计算了中国旧石器时代晚期人类几个头骨与尼阿洞人之间的歧异系数,结果是柳江人与尼阿洞人的距离小于后者与澳大利亚大约同时期的人类的距离,也小于其与山顶洞人之距离。本文还根据一些形态特征的相似性讨论了尼阿洞人、塔邦洞人与亚洲大陆及澳大利亚古人类的关系。结论认为亚洲大陆对尼阿人有过相当大的影响,塔邦洞人也接受过一些来自北方的影响。     

Modern human origins in Australasia: testing the predictions of competing models.

The evolutionary background to the emergence of modern humans remains controversial. Four models have been proposed to explain this process and each has clearly definable and testable predictions about the geographical origins of early Australians and their possible biological interaction with other Pleistocene populations. The present study considers the phenetic affinities of early Australians from Kow Swamp (KS 1 and KS 5) and Keilor to Pleistocene Africans and Asians from calvarial dimensions. The study includes analyses employing log-transformed and size-corrected (Mosimann variables) data. The strongest signals to emerge are as follows: (1) a phenetic pattern in which Australians are most like each other, (2) all three crania possess a mosaic of archaic and modern features, (3) Kow Swamp crania also show strong affinities to archaic remains, (4) Keilor is more modern than KS 1 and KS 5 and (5) Keilor shows affinities to Pleistocene East Asian modern crania (Liujiang and Upper Cave 101) providing evidence for a broad regional morphology. The results refute the predictions of multi-species replacement models for early Australians but are consistent with single-species models. Combined with published evidence from DNA, the present study indicates that the Assimilation model presently offers the best explanation for the origins of Pleistocene Australians.     

The Lake Ndutu cranium and early Homo sapiens in Africa

The partial cranium from Lake Ndutu, near Olduvai Gorge in northern Tanzania, has generally been viewed as Homo erectus, although points of similarity to H. sapiens have also been recognized. Bones of the vault are in fact quite thick, and the cranium is small. Length and breadth dimensions are comparable to those of earlier H. erectus from Koobi Fora and Ileret, and the Ndutu individual is more similar in size to O.H. 12 than to O.H. 9. Unfortunately, the facial skeleton and frontal bone are very incomplete, and little useful information can be obtained from these parts of the existing reconstruction. The parietals are also damaged, but the left temporal is more satisfactorily preserved, and the occiput is nearly complete. Occipital morphology, mastoid shape, and characteristics of the glenoid cavity and tympanic plate probably provide the best available guide to affinities of the Ndutu hominid. In many of these features the cranium resembles Broken Hill, Elandsfontein, and other African fossils referred to archaic H. sapiens. There are some similarities to modern humans also, but no ties to the Neanderthals of Europe. Allocation of Ndutu to an African subspecies of H. sapiens seems most appropriate, even if the pattern of relationships between such archaic populations and recent humans is still unclear.     

Metrical reconsideration of the Skhul IV and IX and Border Cave 1 crania in the context of modern human origins.

The "out-of-Africa" models for origins of modern Homo sapiens incorporate Skhul as one site documenting that early origination. However, only Skhul V is usually considered in the comparative craniology of the question, neglecting the other substantial crania, Skhul IV and IX. Craniometric comparison demonstrates that IV and IX amplify the picture of continuous gradations of Neandertal-to-modern variations throughout the Levant; much variation is thus represented within this one site, raising serious questions about Neandertals and moderns being discrete and long-separated species. Qafzeh 6 too is craniophenetically closer to Neandertals than to the true anatomically modern people of the European Upper Paleolithic. Proper distance analysis of Border Cave 1 cranium shows it is actually far removed from modern African populations. References to Qafzeh, Skhul, and Border Cave as "fully anatomically modern" require reconsideration.     

贵州毕节发现古人类化石与哺乳动物群

2008年12月在贵州毕节团结乡首次发现麻窝口洞化石点,经2009年、2012年和2013年3次发掘,在麻窝口洞上部砂质黏土地层中共发现了四千余件哺乳动物牙齿化石。2013年7月发现的3枚古人类牙齿,分别为左上犬齿、左上第一臼齿和右上第二臼齿。人类牙齿尺寸偏小,臼齿咬合面沟纹简单,没有复杂的咬合面皱纹和附尖齿带结构,牙根短而不显粗壮,上述特点有别于我国已经发现的直立人和早期智人,可归入解剖学上的现代人。与人类相伴的哺乳动物化石,经初步鉴定共计8目20科43属53种,动物群组合反映出亚热带森林生态环境。根据动物群的时代特点,地貌地层及堆积物的光释光年代测定,指示毕节麻窝口洞古人类的时代可能为中更新世晚期,或者晚更新世早期,毕节古人类牙齿的发现为东亚地区现代人的起源及演化增添了新的证据。     

Cnidarian taphonomy and affinities of the Ediacara biota

Plaster impressions and sand casts of extant medusae, a chondrophoran, and a pennatulid share basic structural characteristics with fossils in the Upper Proterozoic Ediacara assemblage. Impressions of extant medusae and Proterozoic circular impressions show general similarities in arrangement and position of radial and concentric structures and a central raised boss. However, annular rings and radial grooves are more numerous in the Proterozoic fossils and strongly folded or deformed fossils are rare as compared with impressions of modem medusae. Recent pennatulids yield impressions that are more deformed and irregular than the Proterozoic genus Charniodiscus. The greater frequency of deformation of most simulated fossil medusoids relative to Precambrian circular impressions implies that Proterozoic medu-soids were substantially stiffer than many modern taxa of comparable sizes. Many fossils with abundant circular rings have no constructional counterparts among the extant forms studied here and their medusoid affinities should remain in doubt. The structural simplicity of impressions of Ediacara organisms and extant cnidarians suggests that their mutual similarities may be due to convergence. However, there is no compelling morphological reason to reject the claim that some Proterozoic fossils may share affinities with living cnidarians. □ Taphonomy. Ediacara biota, cnidarians, phylogenetic relationships.     

Hominid cranial remains from upper Pleistocene deposits at Aduma, Middle Awash, Ethiopia

The Upper Pleistocene localities of Aduma and Bouri have yielded hominid fossils and extensive Middle Stone Age (MSA) archaeological assemblages. The vertebrate fossils recovered include parts of four hominid crania from Aduma and a complete right parietal from Bouri. Archaeological associations and radiometric techniques suggest an Upper Pleistocene age for these hominids. The more complete cranium from Aduma (ADU-VP-1/3) comprises most of the parietals, the occipital, and part of the frontal. This cranium is compared to late Middle and Upper Pleistocene hominid crania from Africa and the Middle East. The Aduma cranium shows a mosaic of cranial features shared with "premodern" and anatomically modern Homo sapiens. However, the posterior and lateral cranial dimensions, and most of its anatomy, are centered among modern humans and resemble specimens from Omo, Skhul, and Qafzeh. As a result, the Aduma and Bouri Upper Pleistocene hominids are assigned to anatomically modern Homo sapiens.     

U-Series dating of Liujiang hominid site in Guangxi,Southern China

It has been established that modern humans were living in the Levant and Africa ca. 100ka ago. Hitherto, this has contrasted with the situation in China where no unequivocal specimens of this species have been securely dated to more than 30ka. Here we present the results of stratigraphic studies and U-series dating of the Tongtianyan Cave, the discovery site of the Liujiang hominid, which represents one of the few well-preserved fossils of modern Homo sapiens in China. The human fossils are inferred to come from either a refilling breccia or a primarily deposited gravel-bearing sandy clay layer. In the former case, which is better supported, the fossils would date to at least approximately 68ka, but more likely to approximately 111-139ka. Alternatively, they would be older than approximately 153ka. Both scenarios would make the Liujiang hominid one of the earliest modern humans in East Asia, possibly contemporaneous with the earliest known representatives from the Levant and Africa. Parallel studies on other Chinese localities have provided supporting evidence for the redating of Liujiang, which may have important implications for the origin of modern humans.     

福建漳平奇和洞发现的新石器时代早期人类头骨

2011年在福建漳平奇和洞发现的距今1万年左右的新石器时代早期人类遗骸"奇和洞III号",是迄今在福建地区发现的最早、最完整的古人类头骨,为探讨华南更新世晚期向全新世过渡阶段人类的体质特征及现代人群的形成与分化提供了重要的研究材料。本文对这件头骨进行了研究,奇和洞III号为35岁左右的男性个体,牙齿龋病严重,推测当时人类的经济模式主要以农耕为主。通过与更新世晚期柳江、山顶洞101号及14组新石器时代人类头骨的比较,发现奇和洞III号头骨兼有更新世晚期人类及新石器南、北方居民的混合体质特征:奇和洞III号头骨长而脑量大,似更新世晚期人类;其高而狭窄的面部、宽阔而低矮的鼻部,呈现出不同于南、北方人群的特殊体质特征。主成分分析显示,奇和洞III号与对比的新石器时代各组在头骨的测量数据上没有表现为明显的南、北地区间差异,但在头骨的测量指数或形状上存在时代和地区间的不同。本文研究为新旧石器过渡阶段人类体质特征的变异提供了进一步证据。     

The current state of korean paleoanthropology

The hominid fossil and Paleolithic archaeology records from the Korean Peninsula are extensive, but relatively little is known about the Korean human evolutionary record outside this region. The Korean paleoanthropological record is reviewed here in light of major research issues, including the hominid fossil record, relative and chronometric dating, lithic analysis, hominid subsistence, and the presence of bone tools, art and symbolism. Some of the major conclusions drawn from this review include: (1) hominid fossils have been found in nine separate sites on the Korean Peninsula; (2) possible Homo erectus fossils are present in North Korea; (3) Ryonggok Cave, in North Korea, has exposed the remains of at least five archaic Homo sapiens individuals; (4) a possible burial of an anatomically modern Homo sapiens child, discovered in Hungsu Cave in South Korea, has been tentatively dated to roughly 40,000 years ago; (5) handaxes and cleavers have been found at a number of sites near Chongokni and they appear to date to at least 100,000 years ago; and (6) taphonomic studies are necessary for addressing issues related to determining the nature of hominid-carnivore interaction over similar resources (e.g. carcasses and shelter); and the presence/absence of Early Paleolithic bone tools, art, and symbolism in Korea.     

The place of Neandertals in the evolution of hominid patterns of growth and development

This study uses the two developmental fields of dental maturation and femoral growth to determine if the pattern of growth and development in Neandertals (archaic Homo sapiens) was intermediate between that of Homo erectus and recent modern humans. Specimens used in the analysis included Neandertals and Upper Palaeolithic early modern Homo sapiens from Europe and individuals from two recent modern human populations. Ontogenetic data for the H. erectus adolescent KNM-WT 15000 and for Gorilla gorilla were included for comparison. Previous reports have indicated that H. erectus demonstrates a pattern of ontogeny characterized by earlier and more rapid linear growth than in modern humans. Results reported here demonstrate that Upper Paleolithic early modern Homo sapiens display a growth trajectory indistinguishable from that of recent modern humans. The pattern of Neandertal ontogeny is not intermediate between the pattern displayed in H. erectus and the derived pattern seen in the modern reference samples and the early modern H. sapiens sample. The Neandertal growth trajectory is consistent with either slow linear growth or advanced dental development.