Is Central Asia the eastern outpost of the Neandertal range? A reassessment of the Teshik‐Tash child

Since its discovery in southeastern Uzbekistan in 1938, the Teshik-Tash child has been considered a Neandertal. Its affinity is important to studies of Late Pleistocene hominin growth and development as well as interpretations of the Central Asian Middle Paleolithic and the geographic distribution of Neandertals. A close examination of the original Russian monograph reveals the incompleteness of key morphologies associated with the cranial base and face and problems with the reconstruction of the Teshik-Tash cranium, making its Neandertal attribution less certain than previously assumed. This study reassesses the Neandertal status of Teshik-Tash 1 by comparing it to a sample of Neandertal, Middle and Upper Paleolithic modern humans, and recent human sub-adults. Separate examinations of the cranium and mandible are conducted using multinomial logistic regression and discriminant function analysis to assess group membership. Results of the cranial analysis group Teshik-Tash with Upper Paleolithic modern humans when variables are not size-standardized, while results of the mandibular analysis place the specimen with recent modern humans for both raw and size-standardized data. Although these results are influenced by limitations related to the incomplete nature of the comparative sample, they suggest that the morphology of Teshik-Tash 1 as expressed in craniometrics is equivocal. Although, further quantitative studies as well as additional sub-adult fossil finds from this region are needed to ascertain the morphological pattern of this specimen specifically, and Central Asian Middle Paleolithic hominins in general, these results challenge current characterizations of this territory as the eastern boundary of the Neandertal range during the Late Pleistocene.     

The M. obturator internus sulcus on Middle and Late Pleistocene human ischia

Recent human ischia and those of Middle and Late Pleistocene hominids exhibit variation in the cranio-caudal location of the sulcus for the internal obturator muscle as it rounds the ischium through the lesser sciatic notch, from being fully cranial of the ischial tuberosity, to bordering the tuberosity, to crossing the superior tuberosity. Among two recent human samples, all three forms exist, with the cranial position of the sulcus being more common in a 20th century Euroamerican sample whereas the intermediate one predominates in a horticultural late prehistoric Amerindian sample. The available Pleistocene Homo fossil remains exhibit the full range of variation with no one form being dominant in Middle Pleistocene archaic humans and Middle Paleolithic late archaic and early modern humans. It is only within the Upper Paleolithic that the cranial and intermediate locations for the sulcus become predominant. These patterns therefore indicate that it is inappropriate to use this feature for distinguishing later Pleistocene hominid groups. © 1996 Wiley-Liss, Inc.     

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.     

辽宁建平古人类肱骨形态结构分析

1957年,在辽宁省建平县发现了一根古人类肱骨化石,编号PA103。通过同一批龙骨中筛选的哺乳动物化石,吴汝康推断PA103应该为更新世晚期古人类,并对该化石进行了表面形态特征观察和描述。为了对PA103化石的内外结构进行更全面的了解,除了线性测量数据的对比,本文还通过计算机断层扫描技术,结合生物力学和形态示量图分析对建平古人类右侧肱骨化石PA103进行了分析。通过本研究发现,PA103骨干横断面的生物力学粗壮度和力学形状指数明显小于尼安德特人,而与同时期欧亚大陆古人类不利手侧最为接近,这说明建平人右侧肱骨可能不是惯用手,同时,建平人的行为活动应该与同时期同地区的古人类处于同一水平,而小于尼安德特人。整体来看,PA103骨干骨密质厚度和截面惯性矩与近现代人的分布模式较为接近,除局部数值增大外,其整体数值小于近现代人的平均水平,这可能与遗传或行为活动有关,由于缺少古人类化石对比数据,更详细的了解还需后期开展更多相关的研究。     

Is human longevity a consequence of cultural change or modern biology?

Increased longevity, expressed as the number of individuals surviving to older adulthood, represents a key way that Upper Paleolithic Europeans differ from earlier European (Neandertal) populations. Here, we address whether longevity increased as a result of cultural/adaptive change in Upper Paleolithic Europe, or whether it was introduced to Europe as a part of modern human biology. We compare the ratio of older to younger adults (OY ratio) in an early modern human sample associated with the Middle Paleolithic from Western Asia with OY ratios of European Upper Paleolithic moderns and penecontemporary Neandertals from the same region. We also compare these Neandertals to European Neandertals. The difference between the OY ratios of modern humans of the Middle and Upper Paleolithic is large and significant, but there is no significant difference between the Neandertals and early modern humans of Western Asia. Longevity for the West Asian Neandertals is significantly more common than for the European Neandertals. We conclude that the increase in adult survivorship associated with the Upper Paleolithic is not a biological attribute of modern humans, but reflects important cultural adaptations promoting the demographic and material representations of modernity.     

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.     

Middle paleolithic human deciduous incisor from Khudji, Tajikistan

In 1997 a human mandibular second deciduous incisor was discovered during excavations at the central Asian Middle Paleolithic site of Khudji, Tajikistan. The specimen was associated with a late Middle Paleolithic assemblage in a minimally disturbed cultural layer. The specimen is average in size for Late Pleistocene archaic human di(2)s and differs from many late archaic human di(2)s in having minimal marginal ridges and tapering markedly distally. In these features it resembles a minority of specimens from the later Pleistocene.     

Metric dental change in the European upper paleolithic and mesolithic.

Evolutionary trends for dental reduction are presented for European Upper Paleolithic and Mesolithic samples. The analysis demonstrates that the greatest decrease in tooth size occurs between the two divisions of the Upper Paleolithic, while little and insignificant change characterizes the Late Upper Paleolithic/Mesolithic transition. Trends for tooth size over this period indicate that (1) human evolution does not stop with the appearance of "anatomically modern Homo sapiens," (2) changes in tooth size fluctuate with increases in the efficiency and complexity of cultural systems, and (3) the Early Upper Paleolithic sample should be considered transitional between Wurm II European Neanderthals and later Upper Paleolithic and Mesolithic groups.     

Diet of upper paleolithic modern humans: Evidence from microwear texture analysis

This article presents the results of the occlusal molar microwear texture analysis of 32 adult Upper Paleolithic modern humans from a total of 21 European sites dating to marine isotope stages 3 and 2. The occlusal molar microwear textures of these specimens were analyzed with the aim of examining the effects of the climatic, as well as the cultural, changes on the diets of the Upper Paleolithic modern humans. The results of this analysis do not reveal any environmentally driven dietary shifts for the Upper Paleolithic hominins indicating that the climatic and their associated paleoecological changes did not force these humans to significantly alter their diets in order to survive. However, the microwear texture analysis does detect culturally related changes in the Upper Paleolithic humans' diets. Specifically, significant differences in diet were found between the earlier Upper Paleolithic individuals, i.e., those belonging to the Aurignacian and Gravettian contexts, and the later Magdalenian ones, such that the diet of the latter group was more varied and included more abrasive foods compared with those of the former. Am J Phys Anthropol 153:570–581, 2014. © 2014 Wiley Periodicals, Inc.     

Evaluating developmental shape changes in Homo antecessor subadult facial morphology

The fossil ATD6-69 from Atapuerca, Spain, dated to ca. 900 ka (thousands of years ago) has been suggested to mark the earliest appearance of modern human facial features. However, this specimen is a subadult and the interpretation of its morphology remains controversial, because it is unclear how developmental shape changes would affect the features that link ATD6-69 to modern humans. Here we analyze ATD6-69 in an evolutionary and developmental context. Our modern human sample comprises cross-sectional growth series from four populations. The fossil sample covers human specimens from the Pleistocene to the Upper Paleolithic, and includes several subadult Early Pleistocene humans and Neanderthals. We digitized landmarks and semilandmarks on surface and CT scans and analyzed the Procrustes shape coordinates using multivariate statistics. Ontogenetic allometric trajectories and developmental simulations were employed in order to identify growth patterns and to visualize potential adult shapes of ATD6-69. We show that facial differences between modern and archaic humans are not exclusively allometric. We find that while postnatal growth further accentuates the differences in facial features between Neanderthals and modern humans, those features that have been suggested to link ATD6-69's morphology to modern humans would not have been significantly altered in the course of subsequent development. In particular, the infraorbital depression on this specimen would have persisted into adulthood. However, many of the facial features that ATD6-69 shares with modern humans can be considered to be part of a generalized pattern of facial architecture. Our results present a complex picture regarding the polarity of facial features and demonstrate that some modern human-like facial morphology is intermittently present in Middle Pleistocene humans. We suggest that some of the facial features that characterize recent modern humans may have developed multiple times in human evolution.     

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.     

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).     

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。     

Quantitative analysis of Neanderthal temporal bone morphology using three-dimensional geometric morphometrics

The temporal bone is the location of several traits thought to differentiate Neanderthals from modern humans, including some proposed Neanderthal-derived traits. Most of these, however, are difficult to measure and are usually described qualitatively. This study applied the techniques of geometric morphometrics to the complex morphology of the temporal bone, in order to quantify the differences observed between Neanderthal and modern human anatomy. Two hundred and seventy modern human crania were measured, representing 9 populations of 30 individuals each, and spanning the extremes of the modern human geographical range. Twelve Neanderthal specimens, as well as Reilingen, Kabwe, Skhul 5, Qafzeh 9, and 4 Late Paleolithic European specimens, were included in the fossil sample. The data were collected in the form of three-dimensional (3-D) landmark coordinates, and specimen configurations were superimposed using generalized Procrustes analysis. The fitted coordinates were then analyzed by an array of multivariate statistical methods, including principal components analysis, canonical variates analysis, and Mahalanobis D(2). The temporal bone landmark analysis was very successful in separating Neanderthals from modern humans. Neanderthals were separated from modern humans in both the principal components and canonical variates analyses. They were much further in Mahalanobis distances from all modern human populations than any two modern human groups were from each other. Most of the previously described temporal bone traits contributed to this separation.     

Who made the Aurignacian and other early Upper Paleolithic industries?

The Aurignacian is typically taken as a marker of the spread of anatomically modern humans into Europe. However, human remains associated with this industry are frustratingly sparse and often limited to teeth. Some have suggested that Neandertals may, in fact, be responsible for the Aurignacian and the earliest Upper Paleolithic industries. Although dental remains are frequently considered to be taxonomically undiagnostic in this context, recent research shows that Neandertals possess a distinct dental pattern relative to anatomically modern humans. Even so, it is rare to find mandibles or maxillae that preserve all or most of their teeth; and, the probability of correctly identifying individuals represented by only a few teeth or a single tooth is unknown. We present a Bayesian statistical approach to classifying individuals represented exclusively by teeth into two possible groups. The classification is based on dental trait frequencies and sample sizes for ‘known’ samples of 95 Neandertals and 63 Upper Paleolithic modern humans. In a cross validation test of the known samples, 89% of the Neandertals and 89% of the Upper Paleolithic modern humans were classified correctly. We then classified an ‘unknown’ sample of 52 individuals: 34 associated with Aurignacian or other (non-Châtelperronian) early Upper Paleolithic industries, 15 associated with the Châtelperronian, and three unassociated. Of the 34 early Upper Paleolithic-associated individuals, 29 were assigned to modern humans, which is well within the range expected (95% of the time 26-33) with an 11% misclassification rate for an entirely modern human sample. These results provide some of the strongest evidence that anatomically modern humans made the Aurignacian and other (non-Châtelperronian) early Upper Paleolithic industries.     

Nonhuman primate hybridization and the taxonomic status of Neanderthals

The present study examines the taxonomic status of Middle Paleolithic Neanderthals by comparing their observed minimum genetic divergence from Upper Paleolithic modern humans in Europe with that observed between macaque species from Sulawesi that are known to hybridize and fully intergrade in the wild. The genetic divergence, and differentiation between Neanderthals and Upper Paleolithic modern humans, as indicated by pairwise minimum genetic distances and F(ST) values calculated from the estimated minimum genetic relationship (R) matrix derived from craniometric data, are significantly greater than those observed both between hybridizing and noninterbreeding Sulawesi macaque species, suggesting that mate recognition and the possibility of gene flow between Neanderthals and Upper Paleolithic modern humans might have been greatly reduced. These results support a species-level taxonomic distinction for the Neanderthals as suggested by proponents of the replacement model. Furthermore, assumptions regarding the monophyletic origin of modern humans from outside Europe are likely valid.     

Brachial and crural indices of European late Upper Paleolithic and Mesolithic humans

Among recent humans brachial and crural indices are positively correlated with mean annual temperature, such that high indices are found in tropical groups. However, despite inhabiting glacial Europe, the Upper Paleolithic Europeans possessed high indices, prompting Trinkaus (1981) to argue for gene flow from warmer regions associated with modern human emergence in Europe. In contrast, Frayer et al. (1993) point out that Late Upper Paleolithic and Mesolithic Europeans should not exhibit tropically-adapted limb proportions, since, even assuming replacement, their ancestors had experienced cold stress in glacial Europe for at least 12 millennia. This study investigates three questions tied to the brachial and crural indices among Late Pleistocene and recent humans. First, which limb segments (either proximal or distal) are primarily responsible for variation in brachial and crural indices? Second, are these indices reflective of overall limb elongation? And finally, do the Late Upper Paleolithic and Mesolithic Europeans retain relatively and/or absolutely long limbs? Results indicate that in the lower limb, the distal limb segment contributes most of the variability to intralimb proportions, while in the upper limb the proximal and distal limb segments appear to be equally variable. Additionally, brachial and crural indices do not appear to be a good measure of overall limb length, and thus, while the Late Upper Paleolithic and Mesolithic humans have significantly higher (i.e., tropically-adapted) brachial and crural indices than do recent Europeans, they also have shorter (i.e., cold-adapted) limbs. The somewhat paradoxical retention of "tropical" indices in the context of more "cold-adapted" limb length is best explained as evidence for Replacement in the European Late Pleistocene, followed by gradual cold adaptation in glacial Europe.     

The Neanderthal taxonomic position: models of intra- and inter-specific craniofacial variation

The Neanderthal taxonomic position is a matter of wide disagreement among paleoanthropologists. Some workers consider this fossil human group to represent a different species, Homo neanderthalensis, while others see it as a subspecies of Homo sapiens. This study developed two models of morphological variation to be applied to a comparison between Neanderthals and modern humans: modern human populations provided a measure of intra-specific variation, while the species and subspecies of Pan provided measures of both intra- and inter-specific morphological differences. Although such an approach has been advocated strongly, it has not been systematically undertaken until recently. The techniques of geometric morphometrics were used to collect data in the form of three-dimensional coordinates of craniofacial landmarks. The data were processed using generalized procrustes analysis, and analyzed by an array of multivariate statistical methods, including principal components analysis, canonical variates analysis and Mahalanobis D(2). The morphological distances between Neanderthals and modern humans, and between Neanderthals and Late Paleolithic/early anatomically modern specimens, are consistently greater than the distances among recent human populations, and greater than the distances between the two chimpanzee species. Furthermore, no strong morphological similarities were found between Neanderthals and Late Paleolithic Europeans. This study does not find evidence for Neanderthal contribution to the evolution of modern Europeans. Results are consistent with the recognition of Neanderthals as a distinct species.     

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.