The postcranial dimensions of the La Chapelle-aux-saints 1 Neandertal

The La Chapelle-aux-Saints 1 Neandertal has figured prominently in considerations of Neandertal body size and proportions. In this context, a reassessment of its major long bones and a reassembly of its principal pelvic elements (sacrum and right ilium) was undertaken. There are secure measurements for its humeral and radial lengths and its femoral head diameter, but the femoral and tibial lengths were almost certainly greater than previous values. The resultant humeral, femoral and tibial lengths are similar to those of other male Neandertals, its femoral head diameter is among the largest known for Middle and Late Pleistocene humans, but its radial length is relatively short. The pelvic assembly provides modest bi-iliac and inlet transverse diameters compared with the few sufficiently complete and undistorted Middle and Late Pleistocene archaic human pelves, but its dimensions are similar to those of large male early modern humans.     

The Xujiayao 14 Mandibular Ramus and Pleistocene Homo Mandibular Variation

The earlier Late Pleistocene mandibular ramus from Xujiayao (northern China) preserves traits that vary distributionally among western Old World Pleistocene Homo samples and between Early/Middle Pleistocene archaic humans and Late Pleistocene modern humans in eastern Eurasia. Xujiayao 14 presents a lateral mandibular notch crest, an open mandibular foramen, a wide ramus, an asymmetrical mandibular notch, an enlarged superior medial pterygoid tubercle, (probably) a retromolar space, and gonial eversion, as well as an unusual depression in the planum triangulare. The first two traits appear ancestral for Later Pleistocene and recent Homo and are dominant among modern humans. The second two traits largely separate Xujiayao 14 and archaic Homo from modern humans. The next two traits are found in the highest frequency among the Neandertals, although gonial eversion contrasts with Late Pleistocene Neandertals. Xujiayao 14, in the context of Pleistocene and recent Homo samples and the other Xujiayao human remains, therefore provides a morphological mosaic, highlighting regional variation through the Pleistocene.     

Neandertal knees: power lifters in the Pleistocene?

It has been proposed (Trinkaus, 1983 a; Miller & Gross, 1998) that the marked thickness of Neandertal patellae and/or the posterior displacement of their tibial condyles increased their relative M. quadriceps femoris moment arms, thereby making their legs powerful in extension. However, it is necessary to compare these reflections of muscle moment arm length to appropriate measures of the body weight moment arm and body mass estimates, both of which are influenced by ecogeographically determined body proportions. Reassessment of tibial condylar displacement and patellar thickness, as well as patellar height, relative to an appropriate measure of the moment arm for the baseline load on the knee (body weight), to that moment arm times estimated body mass, and to that moment arm times a skeletal reflection of body mass (femoral head diameter) rejects the hypothesis that the Neandertals had exceptionally powerful knee extension. Relative tibial condylar displacement remains above that of a modern industrial society sample, but similar to that of the Broken Hill tibia, Late Pleistocene early modern humans and a recent human nonindustrial sample. Relative patellar thickness is similar to that of early modern humans, who have relatively thick patellae compared to the late Holocene human samples. Consequently, once body proportions are taken into account, there is little difference between the Neandertals and other later Pleistocene humans in knee extensor mechanical advantage, and all of these fossil hominids are similar in the more important proximal tibial proportions to those of nonindustrial recent humans.     

The Regourdou 1 Neandertal body size

In order to maximize sample sizes for the assessment of body size and proportions among Late Pleistocene humans, the femoral head diameter of the Regourdou 1 Neandertal was estimated from its ischial acetabular lunate surface, so as to provide a reliable measure for body mass assessment. This estimate was accomplished by fitting a sphere to the 3D surface of the acetabulum and then estimating femoral head diameter using a regression of acetabular sphere diameter to femoral head diameter based on associated recent human femora and ossa coxarum. The resultant mean and range of values for the Regourdou 1 femoral head dimension place it among the smaller of the European and southwest Asian Neandertals, although its humeral length is above average for that sample. Regourdou 1 therefore joins Kebara 2 in having moderately long arms for body core size, and it thereby emphasizes the variation in Neandertal body proportions.     

Body size and postcranial robusticity of European Upper Paleolithic hominins

The robust diaphyses of Pleistocene hominins are said to indicate higher activity levels in these prehistoric humans than among people today. Thus, it could be argued that the prediction of body mass from fossil lower limb diaphyseal cortical area (CA) using recent human regressions might lead to erroneously high body mass estimates. This study uses three body mass prediction formulae based on the following features: reconstructed femoral 80% (subtrochanteric) CA, femoral head diameter (FH), and bi-iliac breadth and stature (BIB-St) among European Early and Late Upper Paleolithic (EUP and LUP) and recent humans from Africa and Europe. All three methods produce similar body mass estimates for all groups studied, including recent humans.Gleaning behavioral differences from these data is more difficult, as no significant differences in CA were found among the fossil and recent Europeans. It has been suggested that the EUP had less robust diaphyses than their LUP counterparts. However, here this result is only obtained when CA is size-standardized to femoral length(3) (Ruff et al., 1993, Am. J. phys. Anthrop.91, 21-53 Trinkaus et al., 1998, in Neandertals and Modern Humans in Western Asia, pp.391-404, New York: Plenum). This should not be interpreted as evidence for lower activity levels in the EUP, but rather as an artefact of standardization, for as Wolpoff (1999), Am. J. phys. Anthrop.109, 416-423 points out, these standardized variables are extremely sensitive to limb length differences, and the EUP have longer limbs than their LUP counterparts. With this in mind, these data do not support a pattern of behavioral differences between EUP and LUP humans, and therefore more sensitive measures than CA may be required to detect such differences.     

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。     

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.     

Neandertal scapular glenoid morphology

Analysis of Neandertal and recent human scapular glenoid fossae reveals that the former had long, narrow, and flat glenoid articular surfaces relative to those of modern humans. Comparison of glenoid length, breadth, and curvature to humeral articular dimensions demonstrates that Neandertal glenoid length and curvature scale to proximal and distal humeral articular dimensions in the same manner as those of modern humans. The remaining contrast is in the relatively greater glenoid fossa width seen in modern humans. This difference in morphology implies differences in the habitual degree of dorsoventral glenohumeral movement between Neandertals and modern humans. This in turn may be related to contrasts in tool use, especially with respect to throwing and projectile use.     

Did the lateral enamel of Neandertal anterior teeth grow differently from that of modern humans?

The formation of lateral enamel in Neandertal anterior teeth has been the subject of recent studies. When compared to the anterior teeth of modern humans from diverse regions (Point Hope, Alaska; Newcastle upon Tyne, England; southern Africa), Neandertal anterior teeth appear to fall within the modern human range of variation for lateral enamel formation time. However, the lateral enamel growth curves of Neandertals are more linear than those of these modern human samples. Other researchers have found that the lateral enamel growth curves of Neandertals are more linear than those of Upper Paleolithic and Mesolithic modern humans as well. The statistical significance of this apparent difference between Neandertal and modern human lateral enamel growth curves is analyzed here. The more linear Neandertal enamel growth curves result from the smaller percentage of total perikymata located in the cervical halves of their teeth. The percentage of total perikymata in the cervical halves of teeth is therefore compared between the Neandertal sample (n=56 teeth) and each modern human population sample: Inuit (n=65 teeth), southern African (n=114 teeth), and northern European (n=115 teeth). There are 18 such comparisons (6 tooth types, Neandertals vs. each of the three modern human populations). Eighteen additional comparisons are made among the modern human population samples. Statistically significant differences are found for 16 of the 18 Neandertal vs. modern human comparisons but for only two of the 18 modern human comparisons. Statistical analyses repeated for subsamples of less worn teeth show a similar pattern. Because surface curvature is thought to affect perikymata spacing, we also conducted measurements to assess surface curvature in thirty teeth. Our analysis shows that surface curvature is not a factor in this lateral enamel growth difference between Neandertals and modern humans.     

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.     

Femoral curvature in Neanderthals and modern humans: a 3D geometric morphometric analysis

Since their discovery, Neanderthals have been described as having a marked degree of anteroposterior curvature of the femoral shaft. Although initially believed to be pathological, subsequent discoveries of Neanderthal remains lead femoral curvature to be considered as a derived Neanderthal feature. A recent study on Neanderthals and middle and early Upper Palaeolithic modern humans found no differences in femoral curvature, but did not consider size-corrected curvature. Therefore, the objectives of this study were to use 3D morphometric landmark and semi-landmark analysis to quantify relative femoral curvature in Neanderthals, Upper Palaeolithic and recent modern humans, and to compare adult bone curvature as part of the overall femoral morphology among these populations.Comparisons among populations were made using geometric morphometrics (3D landmarks) and standard multivariate methods. Comparative material involved all available complete femora from Neanderthal and Upper Palaeolithic modern human, archaeological (Mesolithic, Neolithic, Medieval) and recent human populations representing a wide geographical and lifestyle range. There are significant differences in the anatomy of the femur between Neanderthals and modern humans. Neanderthals have more curved femora than modern humans. Early modern humans are most similar to recent modern humans in their anatomy. Femoral curvature is a good indicator of activity level and habitual loading of the lower limb, indicating higher activity levels in Neanderthals than modern humans. These differences contradict robusticity studies and the archaeological record, and would suggest that femoral morphology, and curvature in particular, in Neanderthals may not be explained by adult behavior alone and could be the result of genetic drift, natural selection or differences in behavior during ontogeny.     

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.     

Morphological affinities of the al’a 1 frontal bone

The human frontal bone from al’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.     

The Shift to a Predominantly Logistical Mobility Strategy can Inhibit Rather than Enhance Forager Interaction

Barton et al. (2011) claim that western Eurasia witnessed a shift in hominin land-use from a predominantly residential mobility strategy to a predominantly logistical mobility strategy during the Late Pleistocene, and that this shift would have led to increased interaction rates between Neandertals and modern humans. But an important assumption of their model—that foragers employing a predominantly logistical mobility strategy cover a greater geographic extent over the course of their lifetimes than foragers employing a predominantly residential mobility strategy—may not be universally true. Here, I reevaluate Barton et al.’s conclusion with a simple foraging-radius model. My results show that a shift to a predominantly logistical mobility strategy can inhibit rather than enhance forager interaction under conditions that presume less of Pleistocene hominin foragers.     

Intrapopulational body size variation and cranial capacity variation in middle pleistocene humans: The Sima de los Huesos sample (Sierra de Atapuerca,Spain)

A sexual dimorphism more marked than in living humans has been claimed for European Middle Pleistocene humans, Neandertals and prehistoric modern humans. In this paper, body size and cranial capacity variation are studied in the Sima de los Huesos Middle Pleistocene sample. This is the largest sample of non-modern humans found to date from one single site, and with all skeletal elements represented. Since the techniques available to estimate the degree of sexual dimorphism in small palaeontological samples are all unsatisfactory, we have used the bootstraping method to asses the magnitude of the variation in the Sima de los Huesos sample compared to modern human intrapopulational variation. We analyze size variation without attempting to sex the specimens a priori. Anatomical regions investigated are scapular glenoid fossa; acetabulum; humeral proximal and distal epiphyses; ulnar proximal epiphysis; radial neck; proximal femur; humeral, femoral, ulnar and tibial shaft; lumbosacral joint; patella; calcaneum; and talar trochlea. In the Sima de los Huesos sample only the humeral midshaft perimeter shows an unusual high variation (only when it is expressed by the maximum ratio, not by the coefficient of variation). In spite of that the cranial capacity range at Sima de los Huesos almost spans the rest of the European and African Middle Pleistocene range. The maximum ratio is in the central part of the distribution of modern human samples. Thus, the hypothesis of a greater sexual dimorphism in Middle Pleistocene populations than in modern populations is not supported by either cranial or postcranial evidence from Sima de los Huesos. Am J Phys Anthropol 106:19–33, 1998. © 1998 Wiley-Liss, Inc.     

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.     

A comparative study of frontal bone morphology among Pleistocene hominin fossil groups

Features of the frontal bone that are conventionally used to distinguish among fossil hominin groups were quantitatively examined. Fifty-five fossil crania dating from the early to the late Pleistocene were analyzed. Using a modified pantograph, outlines of the frontal bone were collected along the midsagittal and two parasagittal planes. The profile from nasion to bregma, as well as two profiles above the medial and lateral sections of the orbit, respectively, extending from the orbital margin to the coronal suture were traced. The outlines were measured using Elliptical Fourier Function Analysis (EFFA), which enabled a quantification of aspects of the frontal bone that have historically been described primarily in nonmetric or linear terms. Four measurements were obtained: 1) overall morphology as expressed in the Fourier harmonic amplitudes; 2) maximum projection of the supraorbital torus at three points along the browridge (glabella and the medial and lateral aspects of the torus above the orbit); 3) maximum distance of the frontal squama from the frontal chord, capturing forehead curvature; and 4) nasion-bregma chord length. The results indicate that the midsagittal profile is significantly different among all Pleistocene groups in analyses that include both size and shape, as well as size-adjusted data. Homo erectus is significantly different from the late Pleistocene groups (Neandertals and early modern H. sapiens) in glabellar projection. Anatomically modern humans are significantly different from all other groups in both raw and size-standardized analyses of all three outlines that captured overall morphology, as well as forehead curvature and lateral supraorbital torus prominence, and middle Pleistocene Homo are significantly different in both medial and lateral overall parasagittal form. However, for the majority of analyses there were no significant differences among the Pleistocene archaic groups in supraorbital torus projection, frontal squama curvature, nasion-bregma chord length, or overall frontal bone morphology.     

Tooth wear, Neanderthal facial morphology and the anterior dental loading hypothesis

The Anterior Dental Loading Hypothesis states that the unique Neanderthal facial and dental anatomy was an adaptive response to the regular application of heavy forces resulting from both the masticatory and cultural use of the anterior teeth. Heavy anterior tooth wear frequently observed in Neanderthal specimens is cited as a main source of evidence for heavy forces being applied to these teeth. From this, it might be predicted that the wear shown on the anterior teeth of Neanderthals would greatly exceed that of the posterior teeth and that this differential would be greater than in other hominins with different facial morphologies.In this paper, a new method of examining tooth wear patterns is used to test these predictions in a large assemblage of Late Pleistocene hominins and a group of recent hunter-gatherers from Igloolik, Canada. The results show that all Late Pleistocene hominins, including Neanderthals, had heavily worn anterior teeth relative to their posterior teeth but, contrary to expectations, this was more pronounced in the modern humans than in the Neanderthals. The Igloolik Inuit showed heavier anterior tooth wear relative to their posterior teeth than any Late Pleistocene hominins. There was, however, a characteristic Neanderthal pattern in which wear was more evenly spread between anterior teeth than in modern humans. Overall, the evidence presented here suggests that all Late Pleistocene hominins habitually applied heavy forces between their anterior teeth and that Neanderthals were not exceptional in this regard. These results therefore does not support the Anterior Dental Loading Hypothesis.     

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.