Some African fossil foot bones: a note on the interpolation of fossils into a matrix of extant species

Certain fossil foot bones (a terminal toe phalanx from Olduvai, specimens of tali from Spy, Skhūl, Olduvai, Kromdraai, Songhor and Rusinga) have been investigated by other workers using the multivariate statistical approach. The conclusions of their studies have, in the main, been based upon examination of early canonical variates alone. It is demonstrated here that if the full analyses (generalized distances) are taken into account, then almost exactly opposite conclusions obtain. The terminal toe phalanx from Olduvai is widely different from those of modern men; the Neandertal tali differ more from modern human bones than previously realized; the specimens from Olduvai, Kromdraai, Songhor and Rusinga are all completely dissimilar from both African ape and modern human tali.     

Probit analysis of Upper Pleistocene hominids

Probit analysis allows the testing of the normality of a sample population where curve fitting and differential weighting occur simultaneously. Probit analysis may be utilized with small samples, and is ideal for use in the study of fossil populations. It is assumed that population which are normally distributed are likely to be in genetic equilibrium, so that graphic and computational probit analysis may be used to discern the effects of evolutionary forces on human populations. Probit analysis of cranial length in a sequence of Athenian population is presented in order to verify its sensitivity to these evolutionary forces. Probit analysis and other tests of homogeneity are applied to similar variables of Upper Pleistocene European Neandertal and modern man.Despite significant differences between variable means of Neandertal and modern man, probit analyses of their combined populations showed no less deviation from normality than either group separately. This suggests that the evolutionary changes in the cranium from the Neandertal stage to the modern stage of man were effected without significantly disrupting the genetic equilibrium of these populations, i.e. that rapid changes in these populations are unlikely to have occurred.     

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.     

Virtual reconstruction of the Neandertal lower limbs with an estimation of hamstring muscle moment arms

A major problem of fossil hominid analysis is a lack of complete specimens. Many individual specimens have been damaged by the effects of diagenesis and excavation. Significant advances in the field of three dimensional image processing (3D) have enabled the creation of accurately scaled reconstructions of individual fossil bones using mirrored parts of the same fossil bone or human/fossil hominid equivalents. This study presents, for the first time, a method to reconstruct a 3D virtual model of the lower limb of the Neandertal using different bones from different fossil remains (Spy II, Neandertal 1 and Kebara 2) and integrating them into a single model of the Neandertal lower limb. A biomechanical analysis of the model was performed, including computer graphics visualization of the results, motion displacement graphs and muscle moment arms. The overall method has been implemented into an open-source customized software (lhpFusionBox) developed for the biomechanical study of the musculoskeletal system.     

Neandertal Demise: An Archaeological Analysis of the Modern Human Superiority Complex

Neandertals are the best-studied of all extinct hominins, with a rich fossil record sampling hundreds of individuals, roughly dating from between 350,000 and 40,000 years ago. Their distinct fossil remains have been retrieved from Portugal in the west to the Altai area in central Asia in the east and from below the waters of the North Sea in the north to a series of caves in Israel in the south. Having thrived in Eurasia for more than 300,000 years, Neandertals vanished from the record around 40,000 years ago, when modern humans entered Europe. Modern humans are usually seen as superior in a wide range of domains, including weaponry and subsistence strategies, which would have led to the demise of Neandertals. This systematic review of the archaeological records of Neandertals and their modern human contemporaries finds no support for such interpretations, as the Neandertal archaeological record is not different enough to explain the demise in terms of inferiority in archaeologically visible domains. Instead, current genetic data suggest that complex processes of interbreeding and assimilation may have been responsible for the disappearance of the specific Neandertal morphology from the fossil record.     

Ancient DNA: would the real Neandertal please stand up?

Mitochondrial DNA sequences recovered from eight Neandertal specimens cannot be detected in either early fossil Europeans or in modern populations. This indicates that, if Neandertals made any genetic contribution at all to modern humans, it must have been limited, though the extent of the contribution cannot be resolved at present.     

Auditory ossicles from southwest Asian Mousterian sites

The present study describes and analyzes new Neandertal and early modern human auditory ossicles from the sites of Qafzeh and Amud in southwest Asia. Some methodological issues in the measurement of these bones are considered, and a set of standardized measurement protocols is proposed. Evidence of erosive pathological processes, most likely attributed to otitis media, is present on the ossicles of Qafzeh 12 and Amud 7 but none can be detected in the other Qafzeh specimens. Qafzeh 12 and 15 extend the known range of variation in the fossil H. sapiens sample in some metric variables, but morphologically, the new specimens do not differ in any meaningful way from living humans. In most metric dimensions, the Amud 7 incus falls within our modern human range of variation, but the more closed angle between the short and long processes stands out. Morphologically, all the Neandertal incudi described to date show a very straight long process. Several tentative hypotheses can be suggested regarding the evolution of the ear ossicles in the genus Homo. First, the degree of metric and morphological variation seems greater among the fossil H. sapiens sample than in Neandertals. Second, there is a real difference in the size of the malleus between Neandertals and fossil H. sapiens, with Neandertals showing larger values in most dimensions. Third, the wider malleus head implies a larger articular facet in the Neandertals, and this also appears to be reflected in the larger (taller) incus articular facet. Fourth, there is limited evidence for a potential temporal trend toward reduction of the long process within the Neandertal lineage. Fifth, a combination of features in the malleus, incus, and stapes may indicate a slightly different relative positioning of either the tip of the incus long process or stapes footplate within the tympanic cavity in the Neandertal lineage.     

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

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

Human hyoid bones from the middle Pleistocene site of the Sima de los Huesos (Sierra de Atapuerca, Spain)

This study describes and compares two hyoid bones from the middle Pleistocene site of the Sima de los Huesos in the Sierra de Atapuerca (Spain). The Atapuerca SH hyoids are humanlike in both their morphology and dimensions, and they clearly differ from the hyoid bones of chimpanzees and Australopithecus afarensis. Their comparison with the Neandertal specimens Kebara 2 and SDR-034 makes it possible to begin to approach the question of temporal variation and sexual dimorphism in this bone in fossil humans. The results presented here show that the degree of metric and anatomical variation in the fossil sample was similar in magnitude and kind to living humans. Modern hyoid morphology was present by at least 530 kya and appears to represent a shared derived feature of the modern human and Neandertal evolutionary lineages inherited from their last common ancestor.     

The premaxilla in Neandertal and early modern children: ontogeny and morphology.

This comparative study of maxillae in Neandertals, Qafzeh, and extant children examines two specific traits: the premaxillary suture (sutura incisiva) and the interincisive sinuses, proposing a new hypothesis about some features of the Neandertal mid-face. Morphologic study of the premaxillary suture at its different borders (i.e. the nasal aspect of the frontal process, nasal and palatal aspects of the palatal process of the maxilla) indicates a persistence of the suture among very young Neandertal children in comparison to the condition in extant ones. This suggests a longer independence of some parts of the premaxilla in Neandertals. To further examine this possibility, CT scans of two Neandertal children were analyzed: Roc de Marsal, estimated to be about 3 years, and Engis 2, estimated to be about 5-6 years. The results are quite different between the fossils. In the older, the premaxillary suture is represented only by a deep groove. In the younger it extends deep to the surface of the nasal process reaching the Parinaud's canal. Synostosis of the premaxillary suture was found to occur later in Neandertal children than in modern ones. Moreover, we observed the existence of two interincisive sinuses in the fossil children, whereas this is rare in modern children (present on only 2% of our sample of 0-6 year-old infants, n = 247).Persistence of an open premaxillary suture represents the potential for an extended period of growth of the Neandertal mid-face. Although no trace of the premaxillary suture remains in adult Neandertals, Neandertals present many features classically considered as consequences of this persistence. The two interincisive sinuses could be a consequence of the labio-lingual diameter of the incisors. The results presented here can be further investigated by additional studies on the cranial sutural system and by precise morphologic observations and CT scans of the mid-face of a larger sample of fossil children.     

Evolutionary significance of the mandibular foramen area in Neandertals.

An unusual morphology of the mandibular foramen area is described, and its incidence determined for several fossil and modern hominid skeletal samples. This morphology, designated the horizontal-oval type mandibular foramen, is found in 46.2% of the 26 Neandertal foramina examined and in 23.1% of a European Upper Paleolithic sample of 13 foramina. In a total of 747 foramina from five modern skeletal samples, the highest incidence is 3.72%. Possible explanations for the presence of the H-0 trait and its unusually high incidence in Neandertals are examined. It is concluded that this feature is probably a genetic trait which either (1) might be selected for in Neandertals as a part of a massive masticatory apparatus, or (2) represents a discrete cranial trait without functional significance that simply reflects the high incidence of certain genes in Neandertal gene pools.     

What molars contribute to an emerging understanding of lateral enamel formation in Neandertals vs. modern humans

Two hypotheses, based on previous work on Neandertal anterior and premolar teeth, are investigated here: (1) that estimated molar lateral enamel formation times in Neandertals are likely to fall within the range of modern human population variation, and (2) that perikymata (lateral enamel growth increments) are distributed across cervical and occlusal halves of the crown differently in Neandertals than they are in modern humans. To investigate these hypotheses, total perikymata numbers and the distribution of perikymata across deciles of crown height were compared for Neandertal, northern European, and southern African upper molar mesiobuccal (mb) cusps, lower molar mesiobuccal cusps, and the lower first molar distobuccal (db) cusp. Sample sizes range from five (Neandertal M(1)db) to 29 (southern African M(1)mb). Neandertal mean perikymata numbers were found to differ significantly from those of both modern human samples (with the Neandertal mean higher) only for the M(2)mb. Regression analysis suggests that, with the exception of the M(2)mb, the hypothesis of equivalence between Neandertal and modern human lateral enamel formation time cannot be rejected. For the M(2)mb, regression analysis strongly suggests that this cusp took longer to form in the Neandertal sample than it did in the southern African sample. Plots of perikymata numbers across deciles of crown height demonstrate that Neandertal perikymata are distributed more evenly across the cervical and occlusal halves of molar crowns than they are in the modern human samples. These results are integrated into a discussion of Neandertal and modern human lateral enamel formation across the dentition, with reference to issues of life history and enamel growth processes.     

No evidence of Neandertal mtDNA contribution to early modern humans

The retrieval of mitochondrial DNA (mtDNA) sequences from four Neandertal fossils from Germany, Russia, and Croatia has demonstrated that these individuals carried closely related mtDNAs that are not found among current humans. However, these results do not definitively resolve the question of a possible Neandertal contribution to the gene pool of modern humans since such a contribution might have been erased by genetic drift or by the continuous influx of modern human DNA into the Neandertal gene pool. A further concern is that if some Neandertals carried mtDNA sequences similar to contemporaneous humans, such sequences may be erroneously regarded as modern contaminations when retrieved from fossils. Here we address these issues by the analysis of 24 Neandertal and 40 early modern human remains. The biomolecular preservation of four Neandertals and of five early modern humans was good enough to suggest the preservation of DNA. All four Neandertals yielded mtDNA sequences similar to those previously determined from Neandertal individuals, whereas none of the five early modern humans contained such mtDNA sequences. In combination with current mtDNA data, this excludes any large genetic contribution by Neandertals to early modern humans, but does not rule out the possibility of a smaller contribution.     

Thoracic morphology in Near Eastern Neandertals and early modern humans compared with recent modern humans from high and low altitudes

Paleoanthropologists have long noted the unique "hyper-barrel-shaped" Neandertal thorax as inferred from fragmentary ribs, clavicles, and sterna. Yet scholars disagree whether the Neandertal thorax represents an adaptation to cold climates or elevated activity levels. Given the difficulties of reconstructing overall chest shape from isolated and fragmentary thoracic skeletal elements, it is worthwhile comparing Neandertals and contemporaneous early modern human fossils from the same geographic region to recent modern human skeletons that are known to have enlarged chests. This study compares thoracic skeletal morphology in two Near Eastern Neandertals (Tabūn C1 and Shanidar 3) and two early modern humans from the same region (Skhūl IV and V) with four samples of recent modern human skeletons from the Andes (n=347): two coastal groups and two groups from high altitudes. The two highland groups, similar to their living descendants, exhibit morphological evidence of anteroposteriorly deep and mediolaterally wide chests as part of respiratory adaptations to high-altitude hypoxia. I calculated the percentage of deviation of each Neandertal and early modern human fossil from the means of the four recent modern human samples for clavicle and rib lengths and curvatures. Shanidar 3 and Tabūn C1 exhibit ribs that are slightly larger and less curved than the Andean samples, indicating slightly larger thoracic skeletons than modern humans who are known to have enlarged chests in response to increased respiratory demands. Skhūl IV and V have significantly shorter ribs with greater curvature suggesting especially narrow thoracic skeletons. Comparisons with Andean populations suggest that the enlarged thoraces of Neandertals may reflect high activity levels, although results from this study do not exclude cold adaptation as an explanatory factor.     

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.     

Out of the veil of death rode the one million! Neandertals and their genes

Two recent papers report extensive nuclear DNA sequence from a 38,000-year-old Neandertal fossil, comparing it to modern humans and estimating when it diverged from, and whether it contributed to, our gene pool. Based on 65,250 and over a million base-pairs of sequence across the genome, respectively, the groups arrived at slightly different interpretations. The data are an exciting and interesting new contribution, but are not surprising, and a sense of history and question helps put them in perspective.     

Neandertal dental asymmetry and the probable mutation effect

The purpose of this paper is to show that if Brace's hypothesis concerning the role of random mutations in structural reduction (PME) is accepted for the sake of argument, then it cannot be used to explain hominid tooth size reduction since the Mousterian. Specifically it is shown that Brace's Hypothesis implies a smaller variance in dental asymmetry for Neandertal than modern man. Variance analysis reveals that, to the contrary, Neandertal dentitions are more variable than a large sample of modern man's. Finally it is shown that the sequence of dental asymmetry per tooth class is approximately the same in Neandertal as in modern man.     

Differences between Neandertal and modern human infant and child growth models

Studying the emergence of distinctive human growth patterns is essential to understanding the evolution of our species. The large number of Neandertal fossils makes this species the best candidate for a comparative study of growth patterns in archaic and modern humans. Here, Neandertal height growth during infancy and early childhood is described using a mathematical model. Height growth velocities for individuals five years old or younger are modelled as age functions based on different estimates of height and age for a set of ten Neandertal infants and children. The estimated heights of each Neandertal individual are compared with those of two modern human populations based on longitudinal and cross-sectional data. The model highlights differences in growth velocity during infancy (from the age of five months onward). We find that statural growth in Neandertal infants is much slower than that seen in modern humans, Neandertal growth is similar to modern humans at birth, but decreases around the third or fourth month. The markedly slower growth rates of Neandertal infants may be attributable to ontogenetic constraints or to metabolic stress, and contribute to short achieved adult stature relative to modern humans.     

Evolution of the supraorbital region in Upper Pleistocene fossil hominids from South-Central Europe

South-Central European fossil hominids dated to the Upper Pleistocene exhibit a distinct morphological and metric continuum in supraorbital form from early Neandertal (Krapina), through late Neandertals (Vindija), to early Upper Paleolithic hominids. The supraorbital morphologies pertinent to this continuum are documented, and the alterations in size and morphology are discussed ralative to the function of supraorbital superstructures and their relationship to overall craniofacial form. It is concluded that this continuum most likely reflects localized transition between Neandertals and modern man in this region of Europe.