Modern human cranial diversity in the Late Pleistocene of Africa and Eurasia: Evidence from Nazlet Khater,Peştera cu Oase,and Hofmeyr

The origin and evolutionary history of modern humans is of considerable interest to paleoanthropologists and geneticists alike. Paleontological evidence suggests that recent humans originated and expanded from an African lineage that may have undergone demographic crises in the Late Pleistocene according to archaeological and genetic data. This would suggest that extant human populations derive from, and perhaps sample a restricted part of the genetic and morphological variation that was present in the Late Pleistocene. Crania that date to Marine Isotope Stage 3 should yield information pertaining to the level of Late Pleistocene human phenotypic diversity and its evolution in modern humans. The Nazlet Khater (NK) and Hofmeyr (HOF) crania from Egypt and South Africa, together with penecontemporaneous specimens from the Pe?tera cu Oase in Romania, permit preliminary assessment of variation among modern humans from geographically disparate regions at this time. Morphometric and morphological comparisons with other Late Pleistocene modern human specimens, and with 23 recent human population samples, reveal that elevated levels of variation are present throughout the Late Pleistocene. Comparison of Holocene and Late Pleistocene craniometric variation through resampling analyses supports hypotheses derived from genetic data suggesting that present phenotypic variation may represent only a restricted part of Late Pleistocene human diversity. The Nazlet Khater, Hofmeyr, and Oase specimens provide a unique glimpse of that diversity. Am J Phys Anthropol, 2009. © 2009 Wiley‐Liss, Inc.     

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。     

云南丽江古人类股骨的形态结构

1960年,在云南省丽江市发现了三根古人类股骨,通过地层观察,仅PA108可归为更新世晚期。前人对PA108做了初步报导,为了进一步了解丽江人股骨的演化分类地位和东亚早期现代人股骨形态变异,本文对PA108的内外结构进行了详尽的分析。研究发现,PA108具有明显的早期现代人特征,即明显的股骨粗线、骨干中部后侧骨密质最厚和中部横断面轮廓形状偏椭圆。PA108标本也有一定的特殊性,体现在骨干中近端和中部骨密质厚度分布上,这可能与其股骨嵴发育较弱有关,这一特征也导致了PA108与其他东亚早期现代人之间的形态差异,这些形态变异进一步扩大了目前已知的东亚地区早期现代人变异范围。同时,在采用骨密质厚度分布模式进行分类时,建议关注股骨骨干中部骨密质最厚部位。     

现代人出现和演化的化石证据

近年来,新材料的发现、新测年结果的更新以及分子生物学的加入,刷新了关于现代人在各主要区域出现时间、在迁徙扩散中与古老人群的交流模式、晚更新世晚期现代人演化复杂场景等方面的认识。本文梳理了关于现代人出现和演化路径的主要化石证据和研究成果。目前化石证据显示：非洲准现代人的出现时间最早可到MIS 9阶段,非洲现代人在中更新世晚期到晚更新世化石证据连续;欧亚大陆现代人的出现时间可追溯到MIS 6阶段,在MIS 5a-MIS 4阶段经历“瓶颈期”,在MIS 3阶段开始广泛分布。从已有证据来看,MIS 3和MIS 2阶段,欧亚大陆不同区域的现代人演化链条并非单一线性,而是呈现有断点的“网状”演化模式。     

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。     

An Early Pleistocene human pedal phalanx from Swartkrans,SKX 16699, and the antiquity of the human lateral forefoot

In order to assess the antiquity of derived human lateral (lesser) toe morphology, the SKX 16699 Early Pleistocene pedal proximal phalanx from Swartkrans (South Africa) was compared to samples of pedal phalanges attributed to Pliocene/Pleistocene australopithecines, Homo naledi and Late Pleistocene Homo. In contrast to australopith lateral phalanges, the SKX 16699 phalanx exhibits an absolutely (and probably relatively) short length, limited plantar diaphyseal curvature, proximal-to-midshaft and mid-dorsoplantar flexor sheath insertions, and a marked proximodorsal orientation of the metatarsal facet. SKX 16699 is intermediate between the australopith phalanges and later Homo ones in its modest dorsal diaphyseal curvature and mid-dorsoplantar metatarsophalangeal collateral ligament insertion areas. Its diaphyseal robustness is similar to that of Homo phalanges, but overlaps the range of later australopith ones. This combination of features and the close morphological affinities of SKX 16699 to later Homo proximal pedal phalanges suggest the emergence of a distinctly human lateral forefoot by the initial Early Pleistocene.     

Femoral/humeral strength in early African Homo erectus

Lower-to-upper limb-bone proportions give valuable clues to locomotor behavior in fossil taxa. However, to date only external linear dimensions have been included in such analyses of early hominins. In this study, cross-sectional measures of femoral and humeral diaphyseal strength are determined for the two most complete early Homo erectus (or ergaster) associated skeletons--the juvenile KNM-WT 15000 and the adult KNM-ER 1808. Modern comparative samples include an adult human skeletal sample representative of diverse body shapes, a human longitudinal growth series, and an adult chimpanzee sample. When compared to appropriately age-matched samples, both H. erectus specimens fall very close to modern human mean proportions and far from chimpanzee proportions (which do not overlap with those of humans). This implies very similar mechanical load-sharing between the lower and upper limbs, and by implication, similar locomotor behavior in early H. erectus and modern humans. Thus, by the earliest Pleistocene (1.7 Ma), completely modern patterns of bipedal behavior were fully established in at least one early hominin taxon.     

Morphological affinities of the Sal'a 1 frontal bone

The human frontal bone from Sal'a, Slovak Republic, has previously entered into discussions of the morphological patterns of Central European Neandertals and the origins of early modern humans in that region. A morphological reassessment of its supraorbital region and a morphometric analysis of its overall proportions indicate that it falls well within expected ranges of variation of Late Pleistocene Neandertals and is separate from European earlier Upper Paleolithic early modern human crania. It is similar to the Qafzeh-Skhul sample in some metrical and supraorbital robusticity measures, but it contrasts with them in mid-sagittal curvature and supraorbital torus morphology. In the context of its probable oxygen isotope stage 5 age based on inferred biostratigraphic associations, it should not be employed directly for arguments relating to the emergence of modern humans in Central Europe.     

Postcranial robusticity in Homo. II: Humeral bilateral asymmetry and bone plasticity

The analysis of humeral asymmetry in Recent human skeletal samples and an extant tennis-player sample documents minimal asymmetry in bone length, little asymmetry in distal humeral articular breadth, but pronounced and variable asymmetry in mid- and distal diaphyseal crosssectional geometric parameters. More specifically, skeletal samples of normal modern Euroamericans, prehistoric and early historic Amerindians, and prehistoric Japanese show moderate (ca. 5–14%) median asymmetry in diaphyseal cross-sectional areas and polar second moments of area, whereas the tennis-player sample, with pronounced unilateral physical activity, exhibits median asymmetries of 28–57% in the same parameters. A sample of Neandertals with nonpathological upper limbs exhibits similarly low articular asymmetry but pronounced diaphyseal asymmetries, averaging 24–57%. In addition, three Neandertals with actual or possible post-traumatic upper limb alterations have the same low articular asymmetry but extremely high diaphyseal asymmetries, averaging 112–215%. These data support those from experimental work on animals, exercise programs of humans, and human clinical contexts in establishing the high degree of diaphyseal plasticity possible for humans, past and present, under changing biomechanical loading conditions. This lends support to activity-related functional interpretations of changing human diaphyseal morphology and robusticity during the Pleistocene. © 1994 Wiley-Liss, Inc.     

Did a discrete event 200,000-100,000 years ago produce modern humans?

Scenarios for modern human origins are often predicated on the assumption that modern humans arose 200,000-100,000 years ago in Africa. This assumption implies that something ‘special’ happened at this point in time in Africa, such as the speciation that produced Homo sapiens, a severe bottleneck in human population size, or a combination of the two. The common thread is that after the divergence of the modern human and Neandertal evolutionary lineages ∼400,000 years ago, there was another discrete event near in time to the Middle-Late Pleistocene boundary that produced modern humans. Alternatively, modern human origins could have been a lengthy process that lasted from the divergence of the modern human and Neandertal evolutionary lineages to the expansion of modern humans out of Africa, and nothing out of the ordinary happened 200,000-100,000 years ago in Africa.Three pieces of biological (fossil morphology and DNA sequences) evidence are typically cited in support of discrete event models. First, living human mitochondrial DNA haplotypes coalesce ∼200,000 years ago. Second, fossil specimens that are usually classified as ‘anatomically modern’ seem to appear shortly afterward in the African fossil record. Third, it is argued that these anatomically modern fossils are morphologically quite different from the fossils that preceded them.Here I use theory from population and quantitative genetics to show that lengthy process models are also consistent with current biological evidence. That this class of models is a viable option has implications for how modern human origins is conceptualized.     

Cross-sectional geometry and morphology of the mandibular symphysis in Middle and Late Pleistocene Homo

Studies of the evolutionary emergence of the human "chin" have been investigated from a phylogenetic perspective during the later Pleistocene or from a biomechanical perspective across extant primates. Since it was during the Middle and Late Pleistocene that the distinctive human mentum osseum emerged, the relationship between mentum osseum form and resistance to mechanical stress at the mandibular symphysis was examined for forty-two Middle and Late Pleistocene human mandibles. Mentum osseum variation was scored on a five-point ordinal scale (mentum osseum rank). Resistance to bending was represented by second moments of area calculated from symphyseal cross-sections. Relative strength in bending was represented by second moments of area divided by estimated moment arm or beam length. Vertical bending resistance in the coronal plane was maintained across the range of mentum osseum variation within and between later Pleistocene human groups. In contrast, resistance to lateral transverse bending (wishboning) was significantly negatively correlated with the emergence of a protruding mentum osseum. However, Neandertals and early modern humans were equivalent in their abilities to resist this bending regime, while both groups were less resistant in wishboning than earlier archaic humans. In addition, symphyseal inclination, which decreased throughout the later Pleistocene, was highly correlated with mentum osseum rank. Although the overall pattern of differential stasis and change in vertical bending and wishboning resistance at the symphysis is consistent with aspects of the current biomechanical model of the "chin," the decoupling of bending resistance and mentum osseum form in the Late Pleistocene suggests that the evolutionary emergence of the modern human "chin" was at least partly independent of the biomechanical demands placed on the symphysis.     

Late pleistocene human femoral diaphyseal curvature

Anterior femoral curvature is a consistent characteristic of Pleistocene and recent humans, although variation exists in the degree of curvature among individuals and across populations. In particular, one group, the Neandertals, has been characterized for a century as having marked femoral curvature. To evaluate the degree of anterior femoral curvature in both Neandertals and other Late Pleistocene humans, their curvature subtenses and proximodistal positions were evaluated in the context of recent human variation. Recent human comparisons show little relationship between subtense (absolute curvature) and femoral length, suggesting that an index that incorporates subtense relative to the length of the femur is inappropriate for between-group assessments. Neandertals were statistically indistinguishable from Middle or earlier Upper Paleolithic modern humans in the degree of absolute curvature, all of whom had greater curvature on average than all later humans. Additionally, Neandertals and Qafzeh-Skhul early modern humans had a more distal point of maximum curvature than any other group. Curvature was not strongly correlated with functional considerations including body mass estimates, surrogate variables for body size, proximal femoral articular orientation, or knee anteroposterior dimensions. The functional role of femoral anterior curvature is unknown; however, the general decrease in curvature subtense closely parallels the between-group changes in inferred levels of mobility from femoral diaphyseal robusticity and shape, suggesting that femoral curvature may reflect mobility levels and patterns among Late Pleistocene and recent humans.     

Human cranial diversity and evidence for an ancient lineage of modern humans

This study examines the genetic affinities of various modern human groupings using a multivariate analysis of morphometric data. Phylogenetic relationships among these groupings are also explored using neighbor-joining analysis of the metric data. Results indicate that the terminal Pleistocene/early Holocene fossils from Australasia exhibit a close genetic affinity with early modern humans from the Levant. Furthermore, recent human populations and Upper Paleolithic Europeans share a most recent common ancestor not shared with either the early Australasians or the early Levantine humans. This pattern of genetic and phylogenetic relationships suggests that the early modern humans from the Levant either contributed directly to the ancestry of an early lineage of Australasians, or that they share a recent common ancestor with them. The principal findings of the study, therefore, lend support to the notion of an early dispersal from Africa by a more ancient lineage of modern human prior to 50 ka, perhaps as early as OIS 5 times (76-100 ka).     

Patterns of humeral asymmetry among Late Pleistocene humans

Human humeral diaphyseal asymmetry in midshaft and mid-distal rigidity is assessed through the Late Pleistocene in samples of late archaic (Neandertal) and early modern humans. It is considered with respect to directionality (handedness), levels of asymmetry, body size and sexual differences. The overall Late Pleistocene sample indicates a right-handed preference in frequencies (right: 74.8%, left: 15.0%, ambiguous: 10.3%), which are similar to those of recent human samples. Average levels of humeral asymmetry are elevated relative to Holocene samples through all but the small Middle Paleolithic modern human and eastern Eurasian late Upper Paleolithic samples. Humeral asymmetry is especially high among the males relative to the females, and the possibility of a division of labor between uni-manual tasks (mostly male) and bi-manual tasks (mostly female) is considered. At the same time, there is a general pattern of increased asymmetry with larger body size, but it remains unclear to what extent it reflects body size versus sexual effects on bilateral humeral loading. There do not appear to have been substantial changes in humeral asymmetry through time, indicating a continuity of similar manual behavioral patterns through the Late Pleistocene, despite considerable changes in technology through the Late Pleistocene.     

A modern human humerus from the early aurignacian of Vogelherdhöhle (Stetten, Germany)

Implicit in much of the discussion of the cultural and population biological dynamics of modern human origins in Europe is the assumption that the Aurignacian, from its very start, was made by fully modern humans. The veracity of this assumption has been challenged in recent years by the association of Neandertal skeletal remains with a possibly Aurignacian assemblage at Vindija Cave (Croatia) and the association of Neandertals with distinctly Upper Paleolithic (but non-Aurignacian) assemblages at Arcy-sur-Cure and St. C?esaire (France). Ideally we need human fossil material that can be confidently assigned to the early Aurignacian to resolve this issue, yet in reality there is a paucity of well-provenanced human fossils from early Upper Paleolithic contexts. One specimen, a right humerus from the site of Vogelherd (Germany), has been argued, based on its size, robusticity, and muscularity, to possibly represent a Neandertal in an Aurignacian context. The morphological affinities of the Vogelherd humerus were explored by univariate and multivariate comparisons of humeral epiphyseal and diaphyseal shape and strength measures relative to humeri of Neandertals and Early Upper Paleolithic (later Aurignacian and Gravettian) modern humans. On the basis of diaphyseal cross-sectional geometry, deltoid tuberosity morphology, and distal epiphyseal morphology, the specimen falls clearly and consistently with European early modern humans and not with Neandertals. Along with the other Vogelherd human remains, the Vogelherd humerus represents an unequivocal association between the Aurignacian and modern human morphology in Europe.     

Early Modern Humans and Morphological Variation in Southeast Asia: Fossil Evidence from Tam Pa Ling,Laos

Little is known about the timing of modern human emergence and occupation in Eastern Eurasia. However a rapid migration out of Africa into Southeast Asia by at least 60 ka is supported by archaeological, paleogenetic and paleoanthropological data. Recent discoveries in Laos, a modern human cranium (TPL1) from Tam Pa Ling‘s cave, provided the first evidence for the presence of early modern humans in mainland Southeast Asia by 63-46 ka. In the current study, a complete human mandible representing a second individual, TPL 2, is described using discrete traits and geometric morphometrics with an emphasis on determining its population affinity. The TPL2 mandible has a chin and other discrete traits consistent with early modern humans, but it retains a robust lateral corpus and internal corporal morphology typical of archaic humans across the Old World. The mosaic morphology of TPL2 and the fully modern human morphology of TPL1 suggest that a large range of morphological variation was present in early modern human populations residing in the eastern Eurasia by MIS 3.     

史前古人类之间的基因交流及对当今现代人的影响

古DNA实验技术及高通量测序技术的出现和发展,使得直接从古老化石中进行遗传物质的提取及测序成为可能,与古人类相关的基因组学研究因此取得了一系列突破性进展,已灭绝的古老型人类(如:尼安德特人和丹尼索瓦人)与非洲以外现代人之间基因的相互影响已被诸多证据所证实。研究表明,在史前时期,早期现代人向非洲以外地区扩散时,遭遇到了现已灭绝的古老型人类,他们在同一时空内长期共存,并发生了基因交流,有一部分古老型人类基因因此流向了现代人,有些基因一直流传至今,对当今现代人的基因组成产生重大影响;此外,不同古老型人类之间也存在基因交流;而早期现代人也对部分古老型人类的基因组成造成了影响。化石与古DNA信息的证据均表明,史前各种人类之间的基因交流在多个地区发生多次,他们的基因交流共同构建了当今现代人的基因库,并在生理机能、形态和疾病发生率等方面对现代人造成了深远的影响。     

中更新世非洲Bodo人类头骨化石与周口店直立人的比较--中国与非洲人类头骨特征对比之三

发现于埃塞俄比亚MiddleAwash地区Bodo地点距今60万年的人类头骨化石是迄今发现的最为古老和完整的非洲中更新世人类化石。由于Bodo头骨化石在形态特征上兼有直立人与智人的特点,多年来学术界对其分类地位一直存在争议。Rightmire认为Bodo头骨化石与BrokenHill及Petralona等在分类上属于古老型智人的中更新世人类更为接近,是非洲直立人向古老型智人过渡的代表。至少在距今60万年的中更新世早期直立人向古老型智人转变的成种事件在非洲就已经发生。以Bodo头骨为代表的一批更新世中期非洲和欧洲人类化石构成了可能是后期人类祖先的人属海德堡种。这些观点导致了近年学术界对古老型智人在非洲及欧亚出现时间以及更新世中期非洲和欧亚地区古人类相互之间演化关系的关注。基于这样的背景,本文对年代与Bodo化石接近的周口店直立人头骨特征与Bodo头骨的相似及差异表现情况进行了对比研究。结果发现Bodo头骨在一系列特征上与周口店直立人相似,同时在包括颅容量在内的其它一些特征上呈现出后期智人的特点,但总体形态上似乎与直立人更为相似。作者认为尽管这种进化上的镶嵌现象在中国古人类化石记录上也广泛存在,但由于中国人类化石标本在年代上的不确定性,目前还没有可靠的证据说明这种集直立人与智人化石特征为一体的镶嵌性在中国古人类化石出现的时间接近或早于非洲。考虑到中国与非洲直立人生存年代的巨大差异及人类演化的不同步或地区间差异,具有较多后期人类特征表现的人类首先出现在非洲是完全可能的。根据这些研究对比,作者就人类演化的镶嵌现象、更新世中期非洲与亚洲地区人类演化上的差异等问题进行了讨论。     

Chin morphology and sexual dimorphism in the fossil hominid mandible sample from Klasies River Mouth

The site of Klasies River Mouth (KRM) in South Africa has produced a small sample of early Upper Pleistocene hominid remains that have been a focus for discussions of the origins of modern humans. Despite certain primitive characteristics exhibited by these fossils, proponents of a single recent origin have attributed them to early modern humans. Critics of this hypothesis have emphasized the significance of the archaic features evident in this sample, including the absence of pronounced chins among the mandibular specimens. This study compares the size range and chin morphology exhibited by the KRM mandibles with that of Neandertals, Upper Pleistocene humans, and recent humans. The extreme sexual dimorphism documented among the KRM fossils reflects the presence of a very small individual, and previous efforts to classify the KRM sample as archaic on the basis of their robusticity have failed to address the significance of this diminutive hominid. While each KRM fossil falls within the 95% envelope of variability established for chin development in a comparative modern sample, a similarly low frequency of pronounced chins is very unlikely to be found in any recent human population. The morphological pattern of the KRM mandibles is clearly distinct from that of Neandertals and of recent humans. © 1996 Wiley-Liss, Inc.