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

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

Reexamination of the immature hominid maxilla from Tangier,Morocco

Reexamination of the immature Upper Pleistocene hominid maxilla from Mugharet el-'Aliya (Tangier), Morocco is undertaken in light of new evidence on the growth and development of Upper Pleistocene hominids. Metric and qualitative comparisons were made with 17 immature Upper Pleistocene maxillae, and with a recent Homo sapiens sapiens sample. No unambiguous criteria for aligning the maxilla with Neandertals were found, although one character, the degree of maxillary flexion on the zygoma, strongly suggests that this child could be a representative of H. s. sapiens. The probable lack of a canine fossa in Mugharet el-'Aliya 1, the primary criterion used previously to align it with Neandertals, cannot be accurately extrapolated to its adult form from this juvenile. The present evidence suggests that it is inappropriate to refer to this fossil as “Neandertal-like” or as a North African “neandertaloid.” Thus, the Tangier maxilla should not be cited as evidence for the presence of Neandertal facial features in North Africa during the Upper Pleistocene. © 1993 Wiley-Liss, Inc.     

Mandibular molar root morphology in Neanderthals and Late Pleistocene and recent Homo sapiens

Neanderthals have a distinctive suite of dental features, including large anterior crown and root dimensions and molars with enlarged pulp cavities. Yet, there is little known about variation in molar root morphology in Neanderthals and other recent and fossil members of Homo. Here, we provide the first comprehensive metric analysis of permanent mandibular molar root morphology in Middle and Late Pleistocene Homo neanderthalensis, and Late Pleistocene (Aterian) and recent Homo sapiens. We specifically address the question of whether root form can be used to distinguish between these groups and assess whether any variation in root form can be related to differences in tooth function. We apply a microtomographic imaging approach to visualise and quantify the external and internal dental morphologies of both isolated molars and molars embedded in the mandible (n = 127). Univariate and multivariate analyses reveal both similarities (root length and pulp volume) and differences (occurrence of pyramidal roots and dental tissue volume proportion) in molar root morphology among penecontemporaneous Neanderthals and Aterian H. sapiens. In contrast, the molars of recent H. sapiens are markedly smaller than both Pleistocene H. sapiens and Neanderthals, but share with the former the dentine volume reduction and a smaller root-to-crown volume compared with Neanderthals. Furthermore, we found the first molar to have the largest average root surface area in recent H. sapiens and Neanderthals, although in the latter the difference between M₁ and M₂ is small. In contrast, Aterian H. sapiens root surface areas peak at M₂. Since root surface area is linked to masticatory function, this suggests a distinct occlusal loading regime in Neanderthals compared with both recent and Pleistocene H. sapiens.     

The Exploitation of Ungulate Bones in Homo neanderthalensis and Homo sapiens

Analysis of ungulate bones recovered from a number of Upper and Middle Palaeolithic sites in southern Italy revealed differences in the presence of anatomical elements. There is a lack of clear evidence of carnivore activities, and differences can be attributed to human activity. Indeed, these differences were probably due to different patterns of skeletal exploitation between Homo neanderthalensis and H. sapiens. Small limb bones (carpals, tarsals, sesamoids, long bone epiphyses and especially phalanges) are rarely found in Middle Palaeolithic deposits, but are abundant in the Upper Palaeolithic. The observation of unidentified bone fragments at these sites indicates that during the middle Palaeolithic, marrow extraction regarded essentially the treatment of long bones. First and second phalanges were not frequently used for this practice, but they were often fragmented by H. sapiens. Lack of these bones among the remains of meals of Neanderthal suggests that these bones were probably destroyed by their utilisation as fuel.     

Was Homo erectus responsible for the hand-axe culture?

This paper reviews the evidence from Africa, Asia and Europe of the cultural associations of Middle Pleistocene hominids, as well as the hominid skeletal associations of hand-axe remains.The author points out that it is possible to make a good argument—from the evidence of Steinheim, Kanjera and Swanscombe—that the hand-axes at these sites were made by Homo sapiens. On the other hand, on the basis of Fontéchevade and Vértesszöllös, it could be claimed that Middle Pleistocene Homo sapiens was responsible for primitive flake and chopper cultures. The evidence from Java is negative while that from China is directly opposed to the view that Homo erectus made hand-axes. Only from Ternifine in Algeria and Olduvai in Tanzania is there suggestive evidence that Homo erectus in those areas might have been responsible for the hand-axe culture. Thus, it is not possible at present to make any categorical statements as to the makers of either the great hand-axe culture or the flake and chopper culture, during Middle Pleistocene times.     

The diploic venous system in Homo neanderthalensis and fossil Homo sapiens: A study using high-resolution computed tomography

Objectives

The diploic venous system has been hypothesized to be related to human brain evolution, though its evolutionary trajectory and physiological functions remain largely unclear. This study examines the characteristics of the diploic venous channels (DCs) in a selection of well-preserved Homo neanderthalensis and Upper Paleolithic Homo sapiens crania, searching for the differences between the two taxa and exploring the associations between brain anatomy and DCs.

Materials and Methods

Five H. neanderthalensis and four H. sapiens fossil specimens from Western Europe were analyzed. Based on Micro-CT scanning and 3D reconstruction, the distribution pattern and draining orifices of the DCs were inspected qualitatively. The size of the DCs was quantified by volume calculation, and the degree of complexity was quantified by fractal analyses.

Results

High-resolution data show the details of the DC structures not documented in previous studies. H. neanderthalensis and H. sapiens specimens share substantial similarities in the DCs. The noticeable differences between the two samples manifest in the connecting points surrounding the frontal sinuses, parietal foramina, and asterional area.

Discussion

This study provides a better understanding of the anatomy of the DCs in H. neanderthalensis and H. sapiens. The connection patterns of the DCs have potential utility in distinguishing between the two taxa and in the phylogenetic and taxonomic discussion of the Neandertal-like specimens with controversial taxonomic status.

     

L'hominidé du Pléistocène supérieur en Corée

Since the middle of 1970s, one rock-shelter, six caves and one open archaeological site have yielded several hominid fossils. Among them, fossils from Yonggok, Mandal, Sangsi and Hungsu cave sites are useful to reconstruct the general shape of the Upper Pleistocene Hominids in Korea. The main study of this paper is to focus on the reconstruction of the general shape of the upper Pleistocene hominids, and to examine the cause of cranial changes from the late upper Paleolithic to Mesolithic period in Korea. Biomechanical principles are also applied to reconstruct the social activity of the upper Paleolithic man in Korea. In addition, it is assumed that main proponent of Paleolithic cultures in Korea might be Homo sapiens rather than any other species. Based on the anatomical characteristics of skull, the axillary border of the scapular and the midshaft of the femur, general appearance of Homo sapiens during the Pleistocene in Korea could be reconstructed.     

Allometry,merism, and tooth shape of the upper deciduous M2 and permanent M1

The aims of this study were to investigate the effect of allometry on the shape of dm² and M¹ crown outlines and to examine whether the trajectory and magnitude of scaling are shared between species. The sample included 160 recent Homo sapiens, 28 Upper Paleolithic H. sapiens, 10 early H. sapiens, and 33 H. neanderthalensis (Neandertal) individuals. Of these, 97 were dm²/M¹ pairs from the same individuals. A two‐block partial least squares analysis of paired individuals revealed a significant correlation in crown shape between dm² and M¹. A principal component analysis confirmed that Neandertal and H. sapiens dm² and M¹ shapes differ significantly and that this difference is primarily related to hypocone size and projection. Allometry accounted for a small but significant proportion of the total morphological variance. We found the magnitude of the allometric effect to be significantly stronger in Neandertals than in H. sapiens. Procrustes distances were significantly different between the two tooth classes in Neandertals, but not among H. sapiens groups. Nevertheless, we could not reject the null hypothesis that the two species share the same allometric trajectory. Although size clearly contributes to the unique shape of the Neandertal dm² and M¹, the largest H. sapiens teeth do not exhibit the most Neandertal‐like morphology. Hence, additional factors must contribute to the differences in dm² and M¹ crown shape between these two species. We suggest an investigation of the role of timing and rate of development on the shapes of the dm² and M¹ may provide further answers. Am J Phys Anthropol 154:104–114, 2014. © 2014 Wiley Periodicals, Inc.     

Trigonid crests expression in Atapuerca-Sima de los Huesos lower molars: Internal and external morphological expression and evolutionary inferences

Trigonid crest patterning in lower molars is distinctive among Late Pleistocene hominins such as Homo neanderthalensis, fossil Homo sapiens and modern humans. In this paper, we present an examination of trigonid crest patterning in the Middle Pleistocene permanent lower molar sample (n = 62) of Homo heidelbergensis from Sima de los Huesos (SH). Crest expression was assessed from 3D models of the enamel and the dentine surfaces that were produced using micro-computed tomography (microCT). The aims of our analysis are to: 1) characterize the pattern of trigonid crest expression at the outer enamel and enamel-dentine junction surfaces (OES and EDJ) of the SH sample, 2) evaluate the concordance of expression between both surfaces, and 3) place trigonid crest variation in the SH sample into a phylogenetic context. Our results reveal a greater variability in the expression of trigonid crests at the EDJ (14 types) compared to the OES (4 types). Despite this variability, in almost all cases the expression of a continuous mid-trigonid or distal crest at the OES corresponds with the expression of a continuous mesial/mid-trigonid or distal trigonid crest, respectively, at the EDJ. Thus, it is possible to predict the type of trigonid crest pattern that would be at the OES in the case of partially worn teeth. Our study points to increased variability in trigonid crest expression in M₃s compared to M₁s and M₂s. Moreover, our analysis reveals that the SH sample matches broadly the trigonid crest patterns displayed by H. neanderthalensis and differs from those exhibited by H. sapiens, particularly in the almost constant expression of a continuous middle trigonid crest at the EDJ. However, SH hominins also exhibit patterns that have not been reported in H. neanderthalensis and H. sapiens samples. Other aspects of the variability of the trigonid crest expression at the dentine are presented and discussed.     

Human taxonomic diversity in the pleistocene: Does Homo erectus represent multiple hominid species?

Recently, nomina such as “Homo heidelbergensis” and “H. ergaster” have been resurrected to refer to fossil hominids that are perceived to be specifically distinct from Homo sapiens and Homo erectus. This results in a later human fossil record that is nearly as speciose as that documenting the earlier history of the family Hominidae. However, it is agreed that there remains only one extant hominid species: H. sapiens. Has human taxonomic diversity been significantly pruned over the last few hundred millennia, or have the number of taxa been seriously overestimated? To answer this question, the following null hypothesis is tested: polytypism was established relatively early and the species H. erectus can accommodate all spatio-temporal variation from ca. 1.7 to 0.5 Ma. A disproof of this hypothesis would suggest that modern human polytypism is a very recent phenomenon and that speciation throughout the course of human evolution was the norm and not the exception. Cranial variation in a taxonomically mixed sample of fossil hominids, and in a modern human sample, is analyzed with regard to the variation present in the fossils attributed to H. erectus. The data are examined using both univariate (coefficient of variation) and multivariate (determinant) analyses. Employing randomization methodology to offset the small size and non-normal distribution of the fossil samples, the CV and determinant results reveal a pattern and degree of variation in H. erectus that most closely approximates that of the single species H. sapiens. It is therefore concluded that the null hypothesis cannot be rejected. © 1993 Wiley-Liss, Inc.     

Florisbad and human population succession in Southern Africa.

The human cranium recovered at Florisbad in 1932 is compared with other Sub-Saharan African hominid remains from Broken Hill, the Omo and Klasies River Mouth. The Florisbad frontal is very broad, but despite this breadth and differences in zygomatic form, there is a definite resemblance to archaic Homo sapiens from Broken Hill. There is also some similarity to both Omo I and Omo II, while fragmentary remains from Klasies River are more lightly built and hence more modern in appearance. These impressions are strengthened by measurement and statistical analysis, which demonstrates that Florisbad and Broken Hill are distant from recent African populations. Even if Florisbad is less archaic than the earlier (Middle Pleistocene?) hominid, it is not noticeably Bushman-like. New dates suggestive of early Upper Pleistocene antiquity also place Florisbad securely in a lineage containing Broken Hill, and there is no evidence to support special ties with any one group of living Africans.     

The capitate of Australopithecus afarensis and A. africanus

The capitates of Australopithecus afarensis (AL 288-lw and AL 333–40) and A. africanus (TM 1526) have the identical combination of modern pongid, modern hominid, and unique characteristics. These traits include the combination of a length that is proximodistally shortened (Homo sapiens-like), a facet for the second metacarpal that is distolaterally facing (unique), the reduced styloid process on the third metacarpal (pongidlike), a dorsally placed trapezoid facet (pongidlike), mediolaterally constricted metacarpal III facet (pongidlike), a prominent palmar beak (pongidlike), a single elongated facet for the second metacarpal (H. sapiens-like), a waisted neck (pongidlike), and a reduced amount of “cupping” in the third metacarpal facet (H. sapiens-like). In overall shape the bones are more like H. sapiens than other extant hominids, although they are uniquely different. The two A. afarensis capitates provide no evidence that there are two postcranial morphotypes at Hadar. Available evidence shows that A. afarensis and A. africanus are strikingly similar postcranially. The morphological differences between the capitate of Australopithecus and H. sapiens may relate to the retention of climbing ability and an absence of certain grip capabilities in these early hominids.     

A geometric morphometric analysis of hominin upper second and third molars, with particular emphasis on European Pleistocene populations

The study of dental morphology by means of geometric morphometric methods allows for a detailed and quantitative comparison of hominin species that is useful for taxonomic assignment and phylogenetic reconstruction. Upper second and third molars have been studied in a comprehensive sample of Plio- and Pleistocene hominins from African, Asian and European sites in order to complete our analysis of the upper postcanine dentition. Intraspecific variation in these two molars is high, but some interspecific trends can be identified. Both molars exhibit a strong reduction of the distal cusps in recent hominin species, namely European Homo heidelbergensis, Homo neanderthalensis and Homo sapiens, but this reduction shows specific patterns and proportions in the three groups. Second molars tend to show four well developed cusps in earlier hominin species and their morphology is only marginally affected by allometric effects. Third molars can be incipiently reduced in earlier species and they evince a significant allometric component, identified both inter- and intraspecifically. European Middle Pleistocene fossils from Sima de los Huesos (SH) show a very strong reduction of these two molars, even more marked than the reduction observed in Neanderthals and in modern human populations. The highly derived shape of SH molars points to an early acquisition of typical Neanderthal dental traits by pre-Neanderthal populations and to a deviation of this population from mean morphologies of other European Middle Pleistocene groups.     

Who Killed the Neanderthals?

Homo neanderthalensis, evolved from the European populations of H. heidelbergensis, and shows some special morphological traits, probably due to an adaptation to particular climatic conditions. It also appears that H. neanderthals had a specialized diet which was mostly carnivorous. Anatomically modern humans of the European Upper Palaeolithic seem to differ from the alimentary behaviour that characterized the Neanderthals; their diet was more varied, with a greater contribution from freshwater alimentary resources (molluscs and fishes). Comparison between the various strategies of subsistence adopted by the two species allows us to propose a hypothesis about the extinction of H. neanderthalensis.     

Calvarial shape variation among Middle Pleistocene hominins: An application of surface scanning in palaeoanthropology

The increasing availability of 3D data and tools offers new analytical perspectives in palaeoanthropology, such as the quantitative testing of opposing phylogenetic scenarios. Using optical surface scan data and geometric morphometric techniques, this study explores calvarial shape variation in the “Middle Pleistocene muddle”. The morphological variability between H. erectus on the one hand and H. sapiens/neanderthalensis on the other has long remained obscure: opposing views have attributed the known specimens to any of the three species and possibly one or two more. A large number of landmarks and semilandmarks was extracted from the braincase and the face, in order to quantify the calvarial shape differences among species and key fossils. The results are incompatible with the hypothesis that H. rhodesiensis is the exclusive ancestor of H. sapiens, and offer only weak support for an exclusively European ancestor of Neandertals.     

Cranial vault shape in fossil hominids: Fourier descriptors in norma lateralis

Two major views of human evolution have elicited considerable controversy. These are: [1] the “out of Africa” hypothesis and [2] the “multiregional” hypothesis. This paper is an attempt to try to reconcile these two scenarios using hominid cranial vault data. Elliptical Fourier functions (EFFs) were used to describe, in visual and numerical terms, the shape of the human cranial vault in norma lateralis.Using jpeg images, contours of the cranial vault of a large sample of hominid specimens were pre-processed in Photoshop CS and rotated in 2D space (positional-orientation) so that a line drawn from nasion to porion was horizontal. The cranial vault image was then digitized with 72 closely-spaced points and submitted to a specially written routine that computed EFFs normalized by scaling (size-standardization). This ensured that the representation was invariant with respect to starting point, size and orientation.Statistically significant differences were found between the H. sapiens sample and both the H. erectus and H. neanderthalensis samples. In contrast, there were no statistically significant differences between the H. erectus and H. neanderthalensis groups, leading to three conclusions: [1] the similarity in cranial vault shape between H. erectus and H. neanderthalensis suggests a single gradually evolving lineage; [2] The taxon H. heidelbergensis can be embedded into the H. erectus → H. neanderthalensis line; and [3] H. sapiens seems to be a separate evolutionary development and is considered here either as a separate species or as a possible example of an allopatric semispecies (Grant, 1977). The results here suggest that human evolution over the last 2 Ma may turn out to be neither totally multiregional or simply out of Africa but rather represents a considerably more complicated picture.     

The absolute dating of Upper Pleistocene subSaharan fossil hominids and their place in human evolution

In the evolution of anatomically modern man and his subspecies most specialists have concentrated on investigating geographical areas other than Africa as the possible area of origin.In this study 20 fossil hominids and associated faunal remains from South and East Africa were dated by microanalysis, radiocarbon, and amino-acid dating in order to see whether modern man appears later, was sympatric, or even predated Neandertal man.These dates indicate that anatomically modern man occurs sympatrically and possibly even predates the Rhodesian group of Neandertals in Africa. Modern man might also be contemporary to and possibly even predate the occurrence of Neandertal in Europe.This would indicate that modern man did not evolve from but possibly gave rise to the Neandertals as off-shoots.Two possibilities for the evolution of modern man are suggested. First, that Homo sapiens capensis evolved about 90,000 to 100,000 years ago from possibly Homo erectus by way of a “basic” Homo sapiens and later gave rise to Homo sapiens rhodesiensis, Homo sapiens afer, and possibly Homo sapiens palestinus around 50,000 years ago with Homo neanderthalensis and Homo sapiens capensis evolving separately from Homo erectus. In this case Homo neanderthalensis would be a different species from Homo sapiens which includes Homo sapiens capensis, Homo sapiens rhodesiensis, Homo sapiens afer, and possibly Homo sapiens palestinus.Secondly, Homo sapiens capensis evolved by way of a “basic” Homo sapiens with Homo sapiens rhodesiensis and Homo sapiens palestinus branching off from Homo sapiens capensis around 50,000 years ago. Before that, around 90,000 to 100,000 years ago Homo sapiens capensis evolved first and was then followed by Homo sapiens neanderthalensis from a “basic” Homo sapiens stock, but diverged. This means, all Neandertals, Homo sapiens capensis, Homo sapiens sapiens and Homo sapiens afer can be considered as subspecies of Homo sapiens.The author favors the first scheme since on relative dating grounds the existence of Neandertal man in Europe before the earliest date of Homo sapiens capensis and a “basic” Homo sapiens seems to be fairly well documented. Irrespective of either one of these possibilities, modern man evolved in Africa and seems to have migrated into Europe and other parts of the world.New absolute dating techniques are mentioned in detail like the new radiocarbon-collagen method and amino acid dating.     

Fossil Homo femur from Berg Aukas,northern Namibia

The proximal half of a hominid femur was recovered from deep within a paleokarst feature at the Berg Aukas mine, northern Namibia. The femur is fully mineralized, but it is not possible to place it in geochrono logical context. It has a very large head, an exceptionally thick diaphyseal cortex, and a very low collodiaphyseal angle, which serve to differentiate it from Holocene homologues. The femur is not attributable to Australopithecus, Paranthropus, or early Homo (i.e., H. habilis sensu lato). Homo erectus femora have a relatively longer and AP flatter neck, and a shaft that exhibits less pilaster than the Berg Aukas specimen. Berg Aukas also differs from early modern femora in several features, including diaphyseal cortical thickness and the degree of subtrochanteric AP flattening. The massive diaphyseal cortex of Berg Aukas finds its closest similarity within archaic H. sapiens (e.g., Castel di Guido) and H. erectus (e.g., KNM-ER 736) samples. It has more cortical bone at midshaft than any other specimen, although relative cortical thickness and the asymmetry of its cross-sectional disposition at this level are comparable with those of other Pleistocene fem ora. The closest morphological comparisons with Berg Aukas are in archaic (i.e., Middle Pleistocene) H. sapiens and Neandertal samples. © 1995 Wiley-Liss, Inc.     

Rethinking the dispersal of Homo sapiens out of Africa

Current fossil, genetic, and archeological data indicate that Homo sapiens originated in Africa in the late Middle Pleistocene. By the end of the Late Pleistocene, our species was distributed across every continent except Antarctica, setting the foundations for the subsequent demographic and cultural changes of the Holocene. The intervening processes remain intensely debated and a key theme in hominin evolutionary studies. We review archeological, fossil, environmental, and genetic data to evaluate the current state of knowledge on the dispersal of Homo sapiens out of Africa. The emerging picture of the dispersal process suggests dynamic behavioral variability, complex interactions between populations, and an intricate genetic and cultural legacy. This evolutionary and historical complexity challenges simple narratives and suggests that hybrid models and the testing of explicit hypotheses are required to understand the expansion of Homo sapiens into Eurasia.