现代人的出现与扩散——中国的化石证据北大核心CSCDCSSCI

自2002年在周口店附近的田园洞发现大约4万年前的现代人化石以来,相继在湖北郧西黄龙洞、广西崇左智人洞等地点发现了早期现代人化石。这些化石发现证实大约10万年前早期现代人在华南地区已经出现。最近在湖南道县福岩洞发现的人类牙齿化石及相关研究进一步揭示具有完全现代形态的人类8万-12万年前在华南局部地区已经出现;而在这个时间段的华北地区,以许家窑人为代表的人类化石形态仍较原始,其演化尚未进入早期现代人阶段。这些研究发现提示,在中国地区,华南是现代人形成与扩散的中心区域,早期现代人以及完全现代类型的人类都可能首先在华南地区出现,然后向华北地区扩散。现有的化石形态证据显示,更新世晚期华南地区人类具有较大的演化变异,可能同时生存有几种不同的演化类群。智人洞属于从古老型智人向现代人演化的过渡类型,而道县则代表着演化进入完全现代类型的人类。基于前人研究及本文的分析,作者认为柳江、资阳、丽江、田园洞等更新世晚期人类化石特征比较进步,在演化上属于与道县相似的现代类型人类。值得注意的是,这些研究进展在引起对现代人在东亚地区出现和扩散关注的同时,古人类学界对其中涉及的许多问题还存在争论。本文在回顾分析这些研究进展的基础上,就相关问题进行了讨论。     

现代人的出现与扩散——中国的化石证据

自2002年在周口店附近的田园洞发现大约4万年前的现代人化石以来,相继在湖北郧西黄龙洞、广西崇左智人洞等地点发现了早期现代人化石。这些化石发现证实大约10万年前早期现代人在华南地区已经出现。最近在湖南道县福岩洞发现的人类牙齿化石及相关研究进一步揭示具有完全现代形态的人类8万-12万年前在华南局部地区已经出现;而在这个时间段的华北地区,以许家窑人为代表的人类化石形态仍较原始,其演化尚未进入早期现代人阶段。这些研究发现提示,在中国地区,华南是现代人形成与扩散的中心区域,早期现代人以及完全现代类型的人类都可能首先在华南地区出现,然后向华北地区扩散。现有的化石形态证据显示,更新世晚期华南地区人类具有较大的演化变异,可能同时生存有几种不同的演化类群。智人洞属于从古老型智人向现代人演化的过渡类型,而道县则代表着演化进入完全现代类型的人类。基于前人研究及本文的分析,作者认为柳江、资阳、丽江、田园洞等更新世晚期人类化石特征比较进步,在演化上属于与道县相似的现代类型人类。值得注意的是,这些研究进展在引起对现代人在东亚地区出现和扩散关注的同时,古人类学界对其中涉及的许多问题还存在争论。本文在回顾分析这些研究进展的基础上,就相关问题进行了讨论。     

中国现代人起源与演化的考古学思考

越来越多古人类学和旧石器时代考古学的发现与研究成果极大地改变了我们对现代人类起源、演化和适应理论问题的看法。中国作为东亚一个幅员辽阔的地理区域,已经成为这项研究的热点地带。来自该地区新的人类化石和石器组合对基于非洲和欧亚大陆西部记录构建的“现代人出自非洲说”（Recent Out-of-Africa）提出了挑战。新的古人类学研究结果表明,早期现代人出现于约100 kaBP的中国南方,可能（至少部分）是由那里的土著居民演化而来。一些古老型智人表现出了与早期古人类镶嵌或过渡性的体质特征,并可能与尼安德特人和丹尼索瓦人杂交混合。同时一些遗址出土的打制石器表现出了早期现代人类技术和行为的复杂性。中国北方的小石片石器主工业和南方的砾石石器主工业贯穿于整个更新世,然而从约40 kaBP开始,石叶技术开始出现在中国北部,紧随其后的是这些地区骨制工具和个人装饰品的出现,这表明,更新世晚期西伯利亚和中亚地区与我国北部可能存在着紧密的文化关系,东北亚地区可能存在着由西北向东南的迁移路线。人类化石和考古学证据表明,中国现代人类起源和适应的过程与机制可能与欧亚大陆西部有所不同。本文对有关中国现代人类起源与演化的考古学研究所取得的新发现和进展进行了总体回顾,从考古学角度阐述了对相关重要学术问题的看法,并为未来的研究提出了方向性建议。     

早期现代人在中国的出现与演化

兴起于上世纪80年代的现代人起源研究与争论在近10年来呈现出一些新的趋势或特点, 主要体现在对现代人起源与演化细节过程的关注。这些新的关注点涉及早期现代人的出现与扩散、中更新世晚期-晚更新世早期人类化石特征变异及演化, 以及早期现代人出现与演化过程中的健康与生存适应活动三个方面。围绕这些问题, 中国古人类学界开展了相关研究并获得了一些新的发现和认识。本文对近10年来早期现代人在中国出现与演化领域的研究进行了回顾, 并对相关问题进行了讨论。     

更新世中期中国古人类演化区域连续性与多样性的化石证据

以往,在东亚大陆发现的更新世中期人类化石被分别归入直立人和古老型智人。这种分类的主要依据是化石形态特征以及年代。魏敦瑞对周口店第一地点人类化石研究描述的一些头骨、下颌骨和牙齿特征通常被作为判定直立人的标准。根据这些化石的年代分布,一般将30万年前的中更新世晚期作为划分直立人与古老型智人的大致年代界限。近20年来,在非洲、欧洲和东亚新发现了一些更新世中期人类化石,目前古人类学界对中国更新世中期人类化石特征及演化有了与以往不同的认识。最近对大荔、许家窑、盘县大洞、许昌、华龙洞等人类化石的研究显示,近30万年以来东亚大陆人类演化呈现复杂的多样性,将这一时期人类全部归入古老型智人难以准确反映更新世中期中国古人类演化模式及规律。本文结合近年中国更新世中期人类演化研究进展,选择部分具有演化及分类价值的形态特征,分析这些特征在更新世中期中国古人类化石的表现特点。在此基础上,对更新世中期中国古人类演化模式做了尝试性探讨。本研究发现,周口店、和县、沂源、南京等中更新世早期人类化石呈现有较多的区域性特征,形态特征表现相对稳定;而大荔、金牛山、许家窑、许昌、华龙洞、马坝、盘县大洞等中更新世晚期人类在化石形态特征表现复杂多样,变异范围大。此外,在这一时期人类化石上发现较多与生存活动、健康、环境适应有关的证据。根据这些发现,作者认为中国中更新世早期组人类演化以形态连续性为主;进入中更新世晚期,中国古人类演化区域性特征减弱,演化模式以多样性为主。一系列新的化石发现和研究证据提示中更新世晚期东亚大陆可能生存有不同的古人类成员。根据目前掌握的化石形态和年代证据,大约30万年前是中国更新世中期演化变化关键时间节点。     

古老型智人向早期现代人演化过渡的研究取得重要进展

中国科学院古脊椎动物与古人类研究所刘武研究员领导的中外研究小组对贵州盘县大洞中更新世晚期(约13-30万年前)4枚人类牙齿化石进行了详细的对比研究,发现一些可能与早期现代人出现有关的形态特征,表明东亚大陆某些中更新世晚期人类可能已经呈现向早期现代人演化过渡的趋势,认为早期现代人在中国的出现时间可能比目前认为的时间为早.3月4日《人类进化杂志》发表了该项研究成果.     

探究早期现代人的南方扩散路线

本文从扩散时间、扩散路径、主要争论以及考古学证据等方面,对早期现代人南方扩散路线的研究现状进行梳理,对相关问题进行了探究。越来越多的证据显示,使用旧石器中期石器技术的早期现代人在晚更新世早期（MIS 5阶段）已经开始沿南方路线扩散,但学术界对于此次扩散的范围和影响程度仍存在争议。在距今约5万年以后,南方扩散路线上的早期现代人出现诸如使用赭石颜料、制作串珠和骨质工具、创作岩画艺术等行为,同时也独立发展出一些区域性适应行为。石器研究显示,南亚地区在5-3万年前逐渐出现了细石器技术及相关产品（细石叶、修背工具等）,而东南亚和大洋洲地区表现出以生产细小石片为主的权宜性石器技术体系,一些细小石片曾被用来加工有机质工具或作为复合型工具使用。中国南方地区紧邻东南亚和印度半岛,无论是从地理位置还是从生态环境来看,都可以纳入南方扩散路线的研究范围。建议从旧石器中期石制品组合和旧石器晚期细小石制品组合两方面入手,开展跨区域比较研究,为探索中国南方地区早期现代人的出现和演化提供重要考古学证据。     

门齿孔位置在中国古人类化石与现代人群的表现及其演化意义

魏敦瑞在研究周口店北京直立人化石时发现,位于上颌骨硬腭表面的门齿孔位置在周口店标本靠后,而在现代人靠近齿槽。此后,门齿孔位置作为具有演化意义的形态特征被用于古人类学研究。迄今,对门齿孔位置在中国古人类化石表现的专门研究仅有周口店一件标本,而在现代中国人的数据尚属空白。鉴于此,本文对门齿孔位置在中国古人类化石以及现代中国人标本进行了观察、测量和数据统计。在此基础上,结合世界其他地区古人类数据资料,本文对门齿孔位置在中国古人类化石的表现特点及其演化意义进行了探讨。本研究发现,从更新世早期到更新世晚期,门齿孔位置在中国古人类呈现由后向前的总体变化趋势。更新世早期和中期直立人(郧县曲远河口、周口店)的门齿孔位置都比较靠后;中更新世晚期的部分中国古人类(大荔、长阳、华龙洞)的门齿孔位置前移,并与现代人接近,而金牛山和巢县门齿孔位置比较靠后,位于直立人范围;在更新世晚期,所有中国古人类都比较靠前,位于现代人变异范围。本文对现代人标本的观测显示,门齿孔位置在现代中国人比较靠前。现代人门齿孔大小及形态存在较大变异,这种表现特点在一定程度上影响对门齿孔位置及演化意义的判定。几乎全部现代人标本门齿孔前缘呈...     

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。     

河北泥河湾及隆化新发现原始牛化石（英文）

原始牛(Bos primigenius)化石新材料包括泥河湾盆地禾尧庄的1件角心、上沙嘴的1件肱骨和承德隆化的1件肱骨、1件胫骨、1件踝骨、2件跟骨和3件距骨;其中上沙嘴的肱骨化石长达457.3 mm,代表迄今在我国境内发现的最大原始牛个体,也是该种的最大记录之一,推算其所属动物的肩高接近2 m;北京门头沟珠窝的原始牛头骨的眶后宽度及角心周长数据也是该种的最大记录之一;禾尧庄的角心标本稍小,但也比晚更新世多数同类要大;这些巨大的化石与欧洲中更新世晚期同类十分接近,其时代也应当与之相当,为中更新世晚期;过去认为原始牛在我国只出现于晚更新世地层,但现在看来该种在中更新世晚期就已扩散到中国。东欧及北亚的最新化石证据表明,过去认为与原始牛共生的晚更新世化石组合的常见属种早在中更新世晚期或更早时期就已出现,其中包括灰狼、虎、真猛犸象、马、真披毛犀、野猪、诺氏驼、赤鹿、驼鹿及草原野牛等。最新测年数据也表明中国北方有几个含原始牛化石的遗址是形成于中更新世晚期;本文认为华北和西北地区个体巨大的原始牛,代表中更新世晚期,而个体较小的则可能代表晚更新世或全新世;近些年在东北地区发现大量原始牛化石,但其中少见个体巨大者。在我国,原始牛化石分布只局限于北方及淮河过渡区,而在南方及毗邻的东南亚地区均未发现此类化石;因此,有人提出原始牛很可能是在中更新世中、晚期通过中亚走廊(Central Asian Corridor)进入中国。不过,近些年在非洲也发现了一些牛属(Bos)化石,因此有人又提出了牛属非洲起源的"奥杜威牛–牛属演化谱系"(Pelorovis–Bos lineage),但该学说所依据的奥杜威牛化石材料,在形态结构上与牛属相去甚远,难以归入同类;此外,非洲可靠的牛属化石记录均未超过1 Ma。本文作者支持传统的牛属"南亚起源学说",因为在南亚西瓦里克地区发现的早期牛属化石不仅时代更古老(早于2 Ma),并且在形态上与原始牛更为相似。中国北方的原始牛化石主要发现于华北平原、黄土高原和松嫩平原的河湖相堆积层,仅极个别出现于黄土地层;由此推断原始牛可能更喜欢水系发育的林缘和/或沼泽环境。     

Neandertals,competition, and the origin of modern human behavior in the Levant

The East Mediterranean Levant is a small region, but its paleoanthropological record looms large in debates about the origin of modern humans and the fate of the Neandertals. For most of the twentieth century, the Levantine paleoanthropological record supported models of continuity and evolutionary transition between Neandertals and early modern humans. Recent advances in radiometric dating have challenged these models by reversing the chronological relationship between Levantine Neandertals and early modern humans. This revised chronostratigraphy for Levantine Middle Paleolithic human fossils raises interesting questions about the evolutionary relationship between Neandertals and early modern humans. A reconsideration of this relationship moves us closer to understanding the long delay between the origin of morphologically modern‐looking humans during the Middle Paleolithic (>130 Kyr) and the adaptive radiation of modern humans into Eurasia around the time of the transition from the Middle to Upper Paleolithic (50 to 30 Kyr).     

The Big Deal about Blades: Laminar Technologies and Human Evolution

Despite the rapid expansion of archaeological knowledge of the Paleolithic over the past several decades, some generalized interpretive frameworks inherited from previous generations of researchers are remarkably tenacious. One of the most persistent of these is the assumed correlation between blade technologies, Upper Paleolithic industries, and anatomically (and behaviorally) modern humans. In this paper, we review some of the evidence for the production of early blade technologies in Eurasia and Africa dating to the late Lower and the Middle Paleolithic. The basic techniques for blade production appeared thousands of years before the Upper Paleolithic, and there is no justification for linking blades per se to any particular aspect of hominid anatomy or to any major change in the behavioral capacities of hominids. It is true that blades came to dominate the archaeological records of western Eurasia and Africa after 40,000 years ago, perhaps as a consequence of increasing reliance on complex composite tools during the Upper Paleolithic. At the same time, evidence from other regions of the world demonstrates that evolutionary trends in Pleistocene Eurasia were historically contingent and not universal. [Middle Paleolithic, Upper Paleolithic, blade technology, human evolution, hominid behavior and capacities]     

中更新世晚期中国古人类化石的形态多样性及其演化意义

近年对许家窑、许昌、华龙洞、澎湖、夏河、哈尔滨等人类化石开展的系统研究,引发了学界对中更新世晚期人类演化及分类的不同认识。基于对相关中国人类化石形态特征的分析,作者提出这一时期中国人类化石形态特征表现为四种类型：1）以中更新世晚期人类共有特征为主;2）以原始特征为主;3）以现代特征为主;4）独特形态组合。多数化石形态特征表现为前三种类型,而许昌和许家窑这种以硕大的头骨和巨大颅容量构成的独特形态组合在其他同时期化石还没有发现。化石形态的多样性提示,不同类型的中更新世晚期中国古人类对现代人的形成贡献不同。作者认为在该时期的人类化石形态多样性规律还未阐明的情况下,将具有混合或镶嵌特征的相关人类化石归入分类地位不确定的人群较为合适。     

Number of ancestral human species: a molecular perspective

Despite the remarkable developments in molecular biology over the past three decades, anthropological genetics has had only limited impact on systematics in human evolution. Genetics offers the opportunity to objectively test taxonomies based on morphology and may be used to supplement conventional approaches to hominid systematics. Our analyses, examining chromosomes and 46 estimates of genetic distance, indicate there may have been only around 4 species on the direct line to modern humans and 5 species in total. This contrasts with current taxonomies recognising up to 23 species.

The genetic proximity of humans and chimpanzees has been used to suggest these species are congeneric. Our analysis of genetic distances between them is consistent with this proposal. It is time that chimpanzees, living humans and all fossil humans be classified in Homo. The creation of new genera can no longer be a solution to the complexities of fossil morphologies. Published genetic distances between common chimpanzees and bonobos, along with evidence for interbreeding, suggest they should be assigned to a single species.

The short distance between humans and chimpanzees also places a strict limit on the number of possible evolutionary side branches that might be recognised on the human lineage. All fossil taxa were genetically very close to each other and likely to have been below congeneric genetic distances seen for many mammals.

Our estimates of genetic divergence suggest that periods of around 2 million years are required to produce sufficient genetic distance to represent speciation. Therefore, Neanderthals and so-called H. erectus were genetically so close to contemporary H. sapiens they were unlikely to have been separate species. Thus, it is likely there was only one species of human (H. sapiens) for most of the last 2 million years. We estimate the divergence time of H. sapiensfrom 16 genetic distances to be around 1.7 Ma which is consistent with evidence for the earliest migration out of Africa. These findings call into question the mitochondrial «African Eve» hypothesis based on a far more recent origin for H. sapiens and show that humans did not go through a bottleneck in their recent evolutionary history.

Given the large offset in evolutionary rates of molecules and morphology seen in human evolution, Homo species are likely to be characterised by high levels of morphological variation and low levels of genetic variability. Thus, molecular data suggest the limits for intraspecific morphological variation used by many palaeoanthropologists have been set too low. The role of phenotypic plasticity has been greatly underestimated in human evolution.

We call into question the use of mtDNA for studies of human evolution. This DNA is under strong selection, which violates the assumption of selective neutrality. This issue should be addressed by geneticists, including a reassessment of its use for molecular clocks. There is a need for greater cooperation between palaeoanthropologists and anthropological geneticists to better understand human evolution and to bring palaeoanthropology into the mainstream of evolutionary biology.     

Throwing in the Middle and Upper Paleolithic: inferences from an analysis of humeral retroversion

When in evolutionary history did long-range projectile weapons become an important component of hunting toolkits? The archeological evidence for the development of projectile weaponry is complex and generally indirect, and has led to different conclusions about the origin and spread of this technology. Lithic evidence from the Middle Stone Age (MSA) has led some researchers to suggest that true long- range projectile weaponry developed in Africa perhaps as early as 80,000 years ago, and was part of the subsistence toolkit carried by modern humans who expanded out of Africa after 50,000 years ago. Alternatively, temporal patterns in the morphology of pointed lithics has led others to posit an independent, convergent origin of projectile weaponry in Africa, the Near East, and Europe during the interval between 50,000-40,000 years ago. By either scenario, projectile weapons would not have been a component of the hunting arsenal of Neandertals, but may have been in use by European early modern humans and thus, projectile technology may have entered into the competitive dynamics that existed between these two groups. The origins of projectile weapons can be addressed, in part, through analyses of the skeletal remains of the prehistoric humans who made and used them. Habitual behavior patterns—including those related to the production and use of technology—can be imprinted on the skeleton through both genetic and epigenetic pathways. Recent studies in the field of sports medicine indicate that individuals who engage in habitual throwing have increased humeral retroversion angles in their throwing arms and a greater degree of bilateral asymmetry in retroversion angles than do non-throwers. This contribution investigates humeral torsion through analysis of the retroversion angle in samples of Eurasian Neandertals, European early modern humans of the middle and late Upper Paleolithic, and comparative samples of recent humans. This analysis was conducted under the assumption that if throwing-based projectile weaponry was used by early modern Europeans but not Neandertals, Upper Paleolithic samples should be similar to recent human groups engaged in habitual throwing in the degree of humeral retroversion in the dominant limb and in bilateral asymmetry in this feature. Neandertals on the other hand, would not be expected to show marked asymmetry in humeral retroversion. Consistent with other studies, Neandertals exhibit increased retroversion angles (decreased humeral torsion or a more posteriorly oriented humeral head) relative to most modern human samples, although this appears more likely related to body form and overall activity levels than to habitual throwing. Although Neandertals with bilaterally preserved humeri sufficient for measurement are rare (consisting of only two males and one female), levels of bilateral asymmetry in humeral retroversion are low, suggesting a lack of regular throwing. While patterning across fossil and comparative samples in levels of humeral retroversion was not clear cut, males of both the middle and late Upper Paleolithic demonstrate a high level of bilateral asymmetry, comparable to or in excess of that seen in samples of throwing athletes. This may indicate habitual use of throwing-based projectile weaponry by middle Upper Paleolithic times. Small sample sizes and relatively great variance in the fossil samples makes these results, however, suggestive rather than conclusive.     

新证据下的现代人起源模型

解剖结构上的现代人是指具有近圆球形头骨、短而平的面颅、纤细的骨骼等特征的区别于其他古老人类的化石和现今的人群。支持多地区演化模型和支持近期非洲起源模型的学者,在“解剖结构上的现代人”的应用范围方面是不同的,前者以连续演化为基本思想,认为这一名词只包括智人中较进步的类群;而后者以分支系统学思想为基础,认为包括所有智人。分子古生物学研究显示,尼人、丹人和智人在遗传学水平上属于不同的人种。新近的以标本-种群为单元的系统分析,因为不是以属、种等分类学阶元进行的,因此与分类学的阶元划分无关。该系统分析的结果显示智人属于单系类群,哈尔滨人、大荔人等组成其姊妹群。尼人与智人的分异早于1百万年,与基因组水平的谱系分析相符合。多次多向的穿梭扩散是统计学上符合系统关系的模型。     

Early agriculture and crop transmission among Bronze Age mobile pastoralists of Central Eurasia

Archaeological research in Central Eurasia is exposing unprecedented scales of trans-regional interaction and technology transfer between East Asia and southwest Asia deep into the prehistoric past. This article presents a new archaeobotanical analysis from pastoralist campsites in the mountain and desert regions of Central Eurasia that documents the oldest known evidence for domesticated grains and farming among seasonally mobile herders. Carbonized grains from the sites of Tasbas and Begash illustrate the first transmission of southwest Asian and East Asian domesticated grains into the mountains of Inner Asia in the early third millennium BC. By the middle second millennium BC, seasonal camps in the mountains and deserts illustrate that Eurasian herders incorporated the cultivation of millet, wheat, barley and legumes into their subsistence strategy. These findings push back the chronology for domesticated plant use among Central Eurasian pastoralists by approximately 2000 years. Given the geography, chronology and seed morphology of these data, we argue that mobile pastoralists were key agents in the spread of crop repertoires and the transformation of agricultural economies across Asia from the third to the second millennium BC.     

Hominin teeth from the early Late Pleistocene site of Xujiayao,Northern China

It is generally accepted that from the late Middle to the early Late Pleistocene (～340–90 ka BP), Neanderthals were occupying Europe and Western Asia, whereas anatomically modern humans were present in the African continent. In contrast, the paucity of hominin fossil evidence from East Asia from this period impedes a complete evolutionary picture of the genus Homo, as well as assessment of the possible contribution of or interaction with Asian hominins in the evolution of Homo sapiens and Homo neanderthalensis. Here we present a comparative study of a hominin dental sample recovered from the Xujiayao site, in Northern China, attributed to the early Late Pleistocene (MIS 5 to 4). Our dental study reveals a mosaic of primitive and derived dental features for the Xujiayao hominins that can be summarized as follows: i) they are different from archaic and recent modern humans, ii) they present some features that are common but not exclusive to the Neanderthal lineage, and iii) they retain some primitive conformations classically found in East Asian Early and Middle Pleistocene hominins despite their young geological age. Thus, our study evinces the existence in China of a population of unclear taxonomic status with regard to other contemporary populations such as H. sapiens and H. neanderthalensis. The morphological and metric studies of the Xujiayao teeth expand the variability known for early Late Pleistocene hominin fossils and suggest the possibility that a primitive hominin lineage may have survived late into the Late Pleistocene in China. Am J Phys Anthropol 156:224–240, 2015. © 2014 Wiley Periodicals, Inc.