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The abundant femoral assemblage of Homo naledi found in the Dinaledi Chamber provides a unique opportunity to test hypotheses regarding the taxonomy, locomotion, and loading patterns of this species. Here we describe neck and shaft cross-sectional structure of all the femoral fossils recovered in the Dinaledi Chamber and compare them to a broad sample of fossil hominins, recent humans, and extant apes. Cross-sectional geometric (CSG) properties from the femoral neck (base of neck and midneck) and diaphysis (subtrochanteric region and midshaft) were obtained through CT scans for H. naledi and through CT scans or from the literature for the comparative sample. The comparison of CSG properties of H. naledi and the comparative samples shows that H. naledi femoral neck is quite derived with low superoinferior cortical thickness ratio and high relative cortical area. The neck appears superoinferiorly elongated because of two bony pilasters on its superior surface. Homo naledi femoral shaft shows a relatively thick cortex compared to the other hominins. The subtrochanteric region of the diaphysis is mediolaterally elongated resembling early hominins while the midshaft is anteroposteriorly elongated, indicating high mobility levels. In term of diaphyseal robusticity, the H. naledi femur is more gracile that other hominins and most apes. Homo naledi shows a unique combination of characteristics in its femur that undoubtedly indicate a species committed to terrestrial bipedalism but with a unique loading pattern of the femur possibly consequence of the unique postcranial anatomy of the species.

The morphology and size of the Neandertal thorax is a subject of growing interest due to its link to general aspects of body size and shape, including physiological aspects related to bioenergetics and activity budgets. However, the number of well-preserved adult Neandertal costal remains is still low. The recent finding of new additional costal remains from the Regourdou 1 (R1) skeleton has rendered this skeleton as one of the most complete Neandertal costal skeletons with a minimum of 18 ribs represented, five of which are complete or virtually complete. Here we describe for the first time all the rib remains from R1 and compare them to a large modern Euroamerican male sample as well as to other published Neandertal individuals. The costal skeleton of this individual shows significant metric and morphological differences from our modern human male comparative sample. The perceived differences include: dorsoventrally large 1st and 2nd ribs, 3rd ribs with a very closed dorsal curvature and large maximum diameters at the posterior angle, a large tubercle-iliocostal line distance in the 4th rib, thick shafts at the dorsal end of its 6th ribs, thick mid-shafts of the 8th ribs, large articular tubercles at the 9th ribs, and thick shafts of the 11th and 12th ribs. Here we also describe a new mesosternal fragment: the left lateral half of sternebral segments 4 and 5. This portion reveals that the mesosternum of R1 had a sternal foramen in its inferiormost preserved sternal segment and supports previous estimation of the total length of this mesosternum. The new costal remains from R1 support the view that Neandertals, when compared with modern humans, show a significantly different thorax, consistent with differences found in other anatomical regions such as the vertebral column and pelvis.

Objectives Humans generally comply with the ecological rule of Allen (1877), with populations from tropical environments exhibiting body proportions in which limb segments are long relative to trunk height compared to temperate groups. This study tests whether ecogeographic differences in intralimb proportions are identifiable among two modern fetal samples of differing ancestry. Materials and methods Data are derived from radiographic measurements of long bone diaphyseal length and crown-heel length (CHL) of contemporary, spontaneously aborted fetuses of African Americans ("black") of assumed African (tropical) ancestry and European Americans ("white") of assumed European (temperate) ancestry (n = 184). Population individual limb elements, brachial, and crural indices are compared via analyses of covariance (ANCOVA). Potential patterns of divergent allometric growth are quantified through principal components analysis (PCA). Results African ancestral distal limb elements were consistently, albeit slightly, longer than those of European ancestry, relative to CHL. None of the ANCOVA interactions with ancestry are statistically significant for limb indices. The radius was the only single element that displayed a statistically significant ancestry effect (p = 0.0435) equating to a 1 mm difference. PCA highlights that upper limbs demonstrate negative allometry and lower limbs demonstrate positive allometry with sample-specific multivariate growth patterns being nearly identical. Differences in growth allometry late in gestation make little contribution to observed differences in adult limb proportions. Discussion No statistically significant ecogeographic patterns were appreciated among intralimb proportions between these groups during the fetal period. This study contributes to a greater appreciation of phenotypic plasticity, ecogeographic variation in ontogeny, and the evolution of modern human diversity.

This paper describes the 108 femoral, patellar, tibial, and fibular elements of a new species of Homo (Homo naledi) discovered in the Dinaledi chamber of the Rising Star cave system in South Africa. Homo naledi possesses a mosaic of primitive, derived, and unique traits functionally indicative of a bipedal hominin adapted for long distance walking and possibly running. Traits shared with australopiths include an anteroposteriorly compressed femoral neck, a mediolaterally compressed tibia, and a relatively circular fibular neck. Traits shared with Homo include a well-marked linea aspera, anteroposteriorly thick patellae, relatively long tibiae, and gracile fibulae with laterally oriented lateral malleoli. Unique features include the presence of two pillars on the superior aspect of the femoral neck and a tubercular distal insertion of the pes anserinus on the tibia. The mosaic morphology of the H. naledi thigh and leg appears most consistent with a species intermediate between Australopithecus spp. and Homo erectus and, accordingly, may offer insight into the nature of the earliest members of genus Homo. These fossils also expand the morphological diversity of the Homo lower limb, perhaps indicative of locomotor diversity in our genus.

International audience; The calcium isotopic composition (δ44/42Ca) of bone and tooth enamel can be used for dietary reconstructions of extant and extinct mammals. In natural conditions, the δ44/42Ca value of bone and teeth varies according to dietary intake with a constant isotopic offset of about −0.6‰. Owing to the poor conservation of collagen, carbon (C), and nitrogen (N) isotopic compositions of the Regourdou Mousterian site (MIS 5, Dordogne, France) previously failed to provide any paleodietary information. Therefore, to reconstruct the trophic chain, we have measured calcium (Ca) isotopes from fossil bone samples of the fauna from the Regourdou site, as well as from three bone samples of the Regourdou 1 Neandertal specimen. The results show a taxon-dependent patterning of the Ca isotopic compositions: herbivores generally have higher δ44/42Ca values than carnivores. All the δ44/42Ca values of Regourdou 1 are low (

Aside from the work of Bonifay (see Bonifay et al. 2007 for one of the more recent papers) and various articles following these earlier works (e.g., Binant 1991, Defleur 1993, Maureille et Vandermeersch 2007, Pettitt 2011, see also May 1986 for a more critical analysis), the in situ position of the remains of Regourdou 1 from layer 4 has never actually been discussed on the basis of available data from the salvage operation carried out in October 1957 by E. Bonifay and G. Laplace-Jauretche, under the administrative authority of Francois Bordes, or from the subsequent, more systematic, excavations directed by E. Bonifay between 1961 and 1964. Via the compilation of available information from a number of unpublished documents (Francois Bordes’ field notes, drawings made during the salvage operation, photographs taken in 1957, 1961 and 1962, as well as databases from the 1961 to 1964 excavations), and also a new inventory of human remains (both previously known and recently discovered), it is now possible to more accurately reconstruct the position of the human remains in a Cartesian system. In this, we assume that the concentration of remains uncovered during the salvage operation was in square G2, according to the preliminary systematic excavations carried out in 1961. They also bring to light that while practically no anatomical connections can be demonstrated with any certainty – and despite significant disruptions (all of the hominin remains are spread over 9 squares : G1 to G3, F1 to F3, E1 and E2, D2) – they are mainly positioned in squares G2 and G3 to some degree with respect to the anatomical logic of the human body. We therefore assume that Regourdou 1 was lying flat, with its head to the west – perhaps upon its trunk – close to the wall of the cavity. This result is different from the fetal position hypothesis proposed in Bonifay et al. (2007). Moreover many post-depositional (albeit Pleistocene) disturbances are also evident. We believe that they were likely the result of the utilization and modification of the cavity by brown bears and lagomorphs.Only new excavations at the site, and a better taphonomic understanding of Bonifay’s (1964) layer 4 (in which Regourdou 1 was found), and the exact role of humans in its formation, i.e., their anthropic impact on the layer, will allow us to discuss in more detail the nature of the deposition of the body, and, hopefully, the absence of the skull.

Body mass changes in the Hominini are examined from an evolutionary perspective. Among the conclusions drawn from this examination are the following: 1.) hominids (sensu lato) are among the largest and most terrestrial primates; 2.) early Homo is on average larger in size than Australopithecus (although the two taxa do show some size overlap); 3.) following its emergence in the early Pleistocene, Homo shows a steady increase in body mass, reaching its maximum near the end of the middle Pleistocene; 4.) Homo subsequently decreases in size throughout the late Pleistocene and into the Holocene, only to increase in size again very recently (the so-called “secular trend”); 5.) body mass variation among humans today tends to follow well-known ecogeographical “rules”, which are empirical observations with multiple potential theoretical explanations.

Regourdou is a well-known Middle Paleolithic site which has yielded the fossil remains of a minimum of two Neandertal individuals. The first individual (Regourdou 1) is represented by a partial skeleton while the second one is represented by a calcaneus. The foot remains of Regourdou 1 have been used in a number of comparative studies, but to date a full description and comparison of all the foot remains from the Regourdou 1 Neandertal, coming from the old excavations and from the recent reanalysis of the faunal remains, does not exist. Here, we describe and comparatively assess the Regourdou 1 tarsals, metatarsals and phalanges. They display traits observed in other Neandertal feet, which are different from some traits of the Sima de los Huesos (Atapuerca) hominins and of Middle Paleolithic, Upper Paleolithic and recent modern humans. These Neandertal features are: a rectangular talar trochlea with a large lateral malleolar facet, a broad talar head, a broad calcaneus with a projecting sustentaculum tali, a wide and wedged navicular with a projecting medial tubercle, large and wide bases of the lateral metatarsals, and mediolaterally expanded and robust phalanges that also show hallux valgus in a strongly built hallux.

17 pages; International audience; The understanding of Neanderthal societies, both with regard to their funerary behaviors and their subsistence activities, is hotly debated. Old excavations and a lack of taphonomic context are often factors that limit our ability to address these questions. To better appreciate the exact nature of what is potentially the oldest burial in Western Europe, Regourdou (Montignac-sur-Vézère, Dordogne), and to better understand the taphonomy of this site excavated more than 50 years ago, we report in this contribution a study of the most abundant animals throughout its stratigraphy: the European rabbit (Oryctolagus cuniculus). In addition to questions surrounding the potential bioturbation of the site's stratigraphy, analysis of the Regourdou rabbits could provide new information on Neandertal subsistence behavior. The mortality profile, skeletal-part representation, breakage patterns, surface modification, and comparison with modern reference collections supports the hypothesis that the Regourdou rabbit remains were primarily accumulated due to natural (attritional) mortality. Radiocarbon dates performed directly on the rabbit remains give ages ranging within the second half of Marine Isotope Stage 3, notably younger than the regional Mousterian period. We posit that rabbits dug their burrows within Regourdou's sedimentological filling, likely inhabiting the site after it was filled. The impact of rabbit activity now brings into question both the reliability of the archaeostratigraphy of the site and the paleoenvironmental reconstructions previously proposed for it, and suggests rabbits may have played a role in the distribution of the Neandertal skeletal remains.

The sequencing of the Neanderthal genome answered once and for all the question of whether these hominins played a role in the origins of modern humans—they did, and a majority of humans alive today retain a small portion of Neanderthal genes. This finding rejects the strictest versions of the Recent African Origin model and has been celebrated by supporters of Multiregional Evolution (MRE). However, we argue that MRE can also be rejected and that other, intermediate, models of modern human origins better represent the means by which modern humans became the only extant human species. We argue this because we reject one of the major tenets of MRE: global gene flow that prevents cladogenesis from occurring. First, using reconstructions of Pleistocene hominin census size, we maintain that populations were neither large nor dense enough to result in such high levels of gene flow across the Old World. Second, we use mammalian divergence and hybridization data to show that the emergence of Homo is recent enoug...

Homo erectus and later humans have enlarged body sizes, reduced sexual dimorphism, elongated lower limbs, and increased encephalization compared to Australopithecus , together suggesting a distinct ecological pattern. The mosaic expression of such features in early Homo , including Homo habilis , Homo rudolfensis , and some early H. erectus , suggests that these traits do not constitute an integrated package. We examined the evidence for body mass, stature, limb proportions, body size and dental size dimorphism, and absolute and relative brain size in Homo naledi as represented in the Dinaledi Chamber sample. H. naledi stature and body mass are low compared to reported values for H. erectus , with the exception of some of the smaller bodied Dmanisi H. erectus specimens, and overlap with larger Australopithecus and early Homo estimates . H. naledi endocranial volumes (465–560 cc) and estimates of encephalization quotient are also similar to Australopithecus and low compared to all Homo specimens, with the exception of Homo floresiensis (LB1) and the smallest Dmanisi H. erectus specimen (D4500). Unlike Australopithecus , but similar to derived members of genus Homo , the Dinaledi assemblage of H. naledi exhibits both low levels of body mass and dental size variation, with an estimated body mass index of sexual dimorphism less than 20%, and appears to have an elongated lower limb. Thus, the H. naledi bauplan combines features not typically seen in Homo species (e.g., small brains and bodies) with those characteristic of H. erectus and more recent Homo species (e.g., reduced mass dimorphism, elongated lower limb).

Since the 1984 discovery of the Nariokotome Homo erectus/Homo ergaster skeleton, it has been almost axiomatic that the emergence of Homo (sensu stricto) was characterized by an increase in body size to the modern human condition and an autapomorphic shift in body proportions to those found today. This was linked to a behavioral shift toward more intensive carnivory and wider ranging in the genus Homo. Recent fossil discoveries and reanalysis of the Nariokotome skeleton suggest a more complex evolutionary pattern. While early Homo tend to be larger than Australopithecus/Paranthropus, they were shorter on average than people today. Reanalysis of the Nariokotome pelvis along with the discovery of additional early and middle Pleistocene pelves indicate that a narrow bi-iliac (pelvic) breadth is an autapomorphy specific to Homo sapiens. Likewise, it appears that at least some early Homo (even those referred to H. ergaster/H. erectus) were characterized by higher humero-femoral indices than the H. sapiens avera...

In humans, a notch marking the posterior attachment of the lateral meniscus is often visible on the posterior, lateral plateau of the tibia, adjacent to the intercondylar eminence. In theory, the presence or absence of this notch in dry bone can be used to differentiate the fossil remains of Australopithecus from those of the genus Homo. In a small-scale study, however, we found examples of modern human tibiae that appear not to have such a notch. In other cases, the morphology of the surrounding bone made it difficult to determine whether or not the notch was present. Although based on a small sample, this study questions: 1) the theoretical postulate that the lateral meniscal notch can be used to differentiate between hominin taxa, and 2) the practical reliability of determining the absence or presence of the notch in fossil remains.

Bivariate femoral length allometry in recent humans, Pan, and Gorilla is investigated with special reference to the diminutive Liang Bua (LB) 1 specimen (the holotype of Homo floresiensis) and six early Pleistocene femora referred to the genus Homo. Relative to predicted body mass, Pan and Gorilla femora show strong negative length allometry while recent human femora evince isometry to positive allometry, depending on sample composition and line-fitting technique employed. The allometric trajectories of Pan and Homo show convergence near the small body size range of LB 1, such that LB 1 manifests a low percentage deviation (dyx of Smith [1980] ) from the Pan allometric trajectory and falls well within the 95% confidence limits around the Pan individuals (but also outside the 95% confidence limits for recent Homo). In contrast, the six early Pleistocene Homo femora, belonging to larger individuals, show much greater dyx values from both Pan and Gorilla and fall well above the 95% confidence limits for these taxa. All but one of these Pleistocene Homo specimens falls within the 95% confidence limits of the recent human sample. Similar results are obtained when femoral length is regressed on femoral head diameter in unlogged bivariate space. Regardless of the ultimate taxonomic status of LB 1, these findings are consistent with a prediction made by us ( Franciscus and Holliday, 1992 ) that hominins in the small body size range of A.L. 288-1 (“Lucy”), including members of the genus Homo, will tend to possess short, ape-like lower limbs as a function of body size scaling.

© Berger et al. Homo naledi is a previously-unknown species of extinct hominin discovered within the Dinaledi Chamber of the Rising Star cave system, Cradle of Humankind, South Africa. This species is characterized by body mass and stature similar to small-bodied human populations but a small endocranial volume similar to australopiths. Cranial morphology of H. naledi is unique, but most similar to early Homo species including Homo erectus, Homo habilis or Homo rudolfensis. While primitive, the dentition is generally small and simple in occlusal morphology. H. naledi has humanlike manipulatory adaptations of the hand and wrist. It also exhibits a humanlike foot and lower limb. These humanlike aspects are contrasted in the postcrania with a more primitive or australopith-like trunk, shoulder, pelvis and proximal femur. Representing at least 15 individuals with most skeletal elements repeated multiple times, this is the largest assemblage of a single species of hominins yet discovered in Africa.

Long-term trends in robusticity of lower limb bones in the genus Homo through the Pleistocene until the present have been proposed, which have been interpreted as a consequence of decreasing levels of mobility and activity patterns, changes in lifestyle, and environmental factors. There has also long been evidence that skeletal strength increases over an individual's lifespan. This increase is caused by continuous bone remodeling that optimizes the structure of a bone to resist mechanical loadings and creates a balance between endosteal resorption and subperiosteal apposition. However, none of the previous studies of temporal trends in robusticity has considered both processes and analyzed how individual age-related robusticity might influence higher-level temporal trends. This paper therefore explores temporal trends in robusticity of lower limb long bones within the genus Homo and considers how individual ages-at-death can confound published evolutionary trends, given the fact that some aspects of relative bone strength tend to increase over individual lifespans. Cross-sectional diaphyseal properties of the midshaft and proximal femur and midshaft tibia of Pleistocene and early Holocene individuals, together with data on age-at-death are used to analyze changes in relative bone strength relative to individuals' ages and evolutionary time. The results show increasing bone strength in adulthood until the fourth decade and then a slight decrease, an observation that conforms to previously published results on recent human populations. However, no significant impact of age-at-death on the trends along an evolutionary trajectory has been detected. The evolutionary trends in femoral and tibial relative strength can be described as fluctuating, probably as a consequence of changing mobility patterns, environmentally and technologically influenced behaviors, and demographic processes. The differences between evolutionary trends published in several studies are explained primarily as a result of different ways of standardizing cross-sectional parameters for size, and differences in sample composition.

Hennig differentiated parent‐offspring, or tokogenetic, relationships among organisms within a population, which are network‐like, or reticulate, in their structure, from phylogenetic relationships between species taxa, which are hierarchical. However, some biologists have recently argued that reticulation may occur across taxa at high taxonomic levels, and Jolly, using a papionin analogy for hominin evolution, argued that fossil hominins represent a group of allotaxa that likely exhibited varying degrees of hybridization in zones of ecological overlap. Such potential reticulation among the Hominini has important implications for phylogenetic reconstruction and is likely a significant source of homoplasy in the hominin fossil record. Hybridization between taxa can lead to the merging of taxa, to the reinforcement of behavioral barriers to mating, or even to the emergence of new, hybrid species. Templeton has noted that many mammalian species appear to be grouped into higher‐level, hybridizing taxa called ...

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.

European Neandertals employed a complex set of physiological cold defenses, homologous to those seen in contemporary humans and nonhuman primates. While Neandertal morphological patterns, such as foreshortened extremities and low relative surface-area, may have explained some of the variance in cold resistance, it is suggested the adaptive package was strongly dependent on a rich array of physiological defenses. A summary of the environmental cold conditions in which the Neandertals lived is presented, and a comparative ethnographic model from Tierra del Fuego is used. Muscle and subcutaneous fat are excellent “passive” insulators. Neandertals were quite muscular, but it is unlikely that they could maintain enough superficial body fat to offer much cold protection. A major, high-energy metabolic adaptation facilitated by modest amounts of highly thermogenic brown adipose tissue (BAT) is proposed. In addition, Neandertals would have been protected by general mammalian cold defenses based on systemic vasoconstriction and intensified by acclimatization, aerobic fitness, and localized cold-induced vasodilation. However, these defenses are energetically expensive. Based on contemporary data from circumpolar peoples, it is estimated that Neandertals required 3,360 to 4,480 kcal per day to support strenuous winter foraging and cold resistance costs. Several specific genetic cold adaptations are also proposed—heat shock protein (actually, stress shock protein), an ACP*1 locus somatic growth factor, and a specialized calcium metabolism not as yet understood. Am. J. Hum. Biol. 14:566–583, 2002. © 2002 Wiley-Liss, Inc.

The Late Pleistocene archaic humans from western Eurasia (the Neandertals) have been described for a century as exhibiting absolutely and relatively long clavicles. This aspect of their body proportions has been used to distinguish them from modern humans, invoked to account for other aspects of their anatomy and genetics, used in assessments of their phylogenetic polarities, and used as evidence for Late Pleistocene population relationships. However, it has been unclear whether the usual scaling of Neandertal clavicular lengths to their associated humeral lengths reflects long clavicles, short humeri, or both. Neandertal clavicle lengths, along with those of early modern humans and latitudinally diverse recent humans, were compared with both humeral lengths and estimated body masses (based on femoral head diameters). The Neandertal do have long clavicles relative their humeri, even though they fall within the ranges of variation of early and recent humans. However, when scaled to body masses, their humeral lengths are relatively short, and their clavicular lengths are indistinguishable from those of Late Pleistocene and recent modern humans. The few sufficiently complete Early Pleistocene Homo clavicles seem to have relative lengths also well within recent human variation. Therefore, appropriately scaled clavicular length seems to have varied little through the genus Homo, and it should not be used to account for other aspects of Neandertal biology or their phylogenetic status.

Analyse critique de l'approche de Groves concernant les affinites morphologiques de l'ulna humaine du niveau Middle Stone Age de Klasies River Mouth. Les AA considerent l'analyse variee canonique (CVA) preferable a l'analyse par composantes principales (PCA)

The low brachial and crural indices of the European Neandertals have long been considered indicative of cold adaptation. Recent work has documented lower limb/trunk ratios and deeper chests (anterior-posterior diameter) in European Neandertals than among their successors. The present study uses variables reflective of limb length, body mass and trunk height, and compares European Neandertals to 15 globally diverse recent human samples (1 “Eskimo,” 3 North African, 4 sub-Saharan African and 7 European). Bivariate plots, as well as principal components analysis plots of log shape-transformed data, indicate that European Neandertals had an overall body shape that falls at the extreme end of modern higher latitude groups' range of variation. Cluster analysis (minimum spanning tree on a principal coordinates plot) indicates that the Neandertals are closest in body shape to modern “Eskimos,” but even in this dendrogram, they are joined to the “Eskimo” via a long branch. In fact, it appears that European Neandertals were “hyperpolar” in body shape, likely due to two factors: 1) the extremely cold temperatures of glacial Europe and 2) less effective cultural buffering against cold stress. Am J Phys Anthropol 104:245–258, 1997. © 1997 Wiley-Liss, Inc.

Body proportions covary with climate, apparently as the result of climatic selection. Ontogenetic research and migrant studies have demonstrated that body proportions are largely genetically controlled and are under low selective rates; thus studies of body form can provide evidence for evolutionarily short-term dispersals and/or gene flow. Following these observations, competing models of modern human origins yield different predictions concerning body proportion shifts in Late Pleistocene Europe. Replacement predicts that the earliest modern Europeans will possess “tropical” body proportions (assuming Africa is the center of origin), while Regional Continuity permits only minor shifts in body shape, due to climatic change and/or improved cultural buffering. This study tests these predictions via analyses of osteometric data reflective of trunk height and breadth, limb proportions and relative body mass for samples of Early Upper Paleolithic (EUP), Late Upper Paleolithic (LUP) and Mesolithic (MES) humans and 13 recent African and European populations. Results reveal a clear tendency for the EUP sample to cluster with recent Africans, while LUP and MES samples cluster with recent Europeans. These results refute the hypothesis of local continuity in Europe, and are consistent with an interpretation of elevated gene flow (and population dispersal?) from Africa, followed by subsequent climatic adaptation to colder conditions. These data do not, however, preclude the possibility of some (albeit small) contribution of genes from Neandertals to succeeding populations, as is postulated in Brauer’s “Afro-European Sapiens” model.

Many dramatic changes in morphology within the genus Homo have occurred over the past 2 million years or more, including large increases in absolute brain size and decreases in postcanine dental size and skeletal robusticity. Body mass, as the 'size' variable against which other morphological features are usually judged, has been important for assessing these changes1–5. Yet past body mass estimates for Pleistocene Homo have varied greatly, sometimes by as much as 50% for the same individuals2,3,6–12. Here we show that two independent methods of body-mass estimation yield concordant results when applied to Pleistocene Homo specimens. On the basis of an analysis of 163 individuals, body mass in Pleistocene Homo averaged significantly (about 10%) larger than a representative sample of living humans. Relative to body mass, brain mass in late archaic H. sapiens (Neanderthals) was slightly smaller than in early 'anatomically modern' humans, but the major increase in encephalization within Homo occurred earlier during the Middle Pleistocene (600–150 thousand years before present (kyr BP)), preceded by a long period of stasis extending through the Early Pleistocene (1,800 kyr BP).

The evolution of the human upper limb involved a change in function from its use for both locomotion and prehension (as in apes) to a predominantly prehensile and manipulative role. Well-preserved forelimb remains of 1.98-million-year-old Australopithecus sediba from Malapa, South Africa, contribute to our understanding of this evolutionary transition. Whereas other aspects of their postcranial anatomy evince mosaic combinations of primitive (australopith-like) and derived ( Homo -like) features, the upper limbs (excluding the hand and wrist) of the Malapa hominins are predominantly primitive and suggest the retention of substantial climbing and suspensory ability. The use of the forelimb primarily for prehension and manipulation appears to arise later, likely with the emergence of Homo erectus .

The pattern of variation and covariation of proximal and distal limb segment lengths was examined within and between 20 geographically diverse skeletal samples of modern humans. Analyses of variance-covariance matrices (VCMs) of logarithmically transformed (ln) variates of humerus, radius, femur, and tibia length were performed to test the following hypotheses: first, within populations, the distal and proximal segments will have equal relative (i.e., size-independent) variability. However, between populations, the tibia is predicted to be more variable than the other segments. Tests of fit of computed VCMs to theoretical matrices by an iterative procedure (Anderson [1973] Ann. Stat. 1:135-141) reject the equal variance hypotheses, rather suggesting that the relative variances of the distal limb segments are greater than are those of the proximal. Males and females differ somewhat in that within females, the distal segments of both limbs have equal variance, while within males, the tibia has greater relative variance than the radius. The second hypothesis, regarding between-group variability, is somewhat supported in that between human populations, one cannot reject that the tibia has greater relative variance than the other limb segments. However, neither can one reject an alternative hypothesis that both distal limb segments (tibia and radius) are more variable than the proximal segments. Differential growth allometry is explored, and likely plays a major role in differences seen both within and between human populations.

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, Frayeret al. (1993) point out that Late Upper Paleolithic and Mesolithic Europeans shouldnotexhibit 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 ofoveralllimb 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 Last Neanderthal: The Rise, Success, and Mysterious Extinction of Our Closest Human Relatives (Revised edition). Ian Tattersall. Boulder: Westview Press, 1999. 208 pp.