A reappraisal of the anatomical basis for speech in Middle Palaeolithic hominids

The recovery of a fossil hominid skeleton with a complete hyoid bone from Mousterian deposits in Kebara Cave, Israel, provides new evidence pertaining to the evolution of speech. Previous studies of speech in the Middle Palaeolithic (most notably those on Neandertals) have focused on the basicranium as an indicator of speech capabilities. This work critiques the use of the basicranium and instead presents the anatomical relations of the hyoid and adjacent structures in living humans as a basis for understanding the form of the vocal tract. The size and morphology of the hyoid from Kebara and its relations to other anatomical components are almost identical to those in modern humans, suggesting that Middle Palaeolithic populations were anatomically capable of fully modern speech.     

A deterministic model of admixture and genetic introgression: the case of Neanderthal and Cro-Magnon

There is an ongoing debate in the field of human evolution about the possible contribution of Neanderthals to the modern human gene pool. To study how the Neanderthal private alleles may have spread over the genes of Homo sapiens, we propose a deterministic model based on recursive equations and ordinary differential equations. If the Neanderthal population was large compared to the Homo sapiens population at the beginning of the contact period, we show that genetic introgression should have been fast and complete meaning that most of the Neanderthal private alleles should be found in the modern human gene pool in case of ancient admixture. In order to test/reject ancient admixture from genome-wide data, we incorporate the model of genetic introgression into a statistical hypothesis-testing framework. We show that the power to reject ancient admixture increases as the ratio, at the time of putative admixture, of the population size of Homo sapiens over that of Neanderthal decreases. We find that the power to reject ancient admixture might be particularly low if the population size of Homo sapiens was comparable to the Neanderthal population size.     

Kebara 2 Neanderthal pelvis: first look at a complete inlet

The renewed excavations at the Kebara Cave revealed a Neanderthal skeleton dated at about 50-55,000 years B.P. The pelvis of this individual is the most intact Neanderthal pelvis yet discovered, presenting for the first time a complete inlet. Although the superior pubic ramus is extremely long, as typically seen in the Neanderthals, the size of the pelvic inlet is comparable to that of modern Homo sapiens. The length of the superior pubic ramus is found to stem from a more externally rotated innominate bone and not, as generally assumed, from the larger pelvic inlet. It is suggested that the uniqueness of the Neanderthal pelvis may be attributable to locomotion and posture-related biomechanics rather than to obstetric requirements.     

Loss of air sacs improved hominin speech abilities

In this paper, the acoustic-perceptual effects of air sacs are investigated. Using an adaptive hearing experiment, it is shown that air sacs reduce the perceptual effect of vowel-like articulations. Air sacs are a feature of the vocal tract of all great apes, except humans. Because the presence or absence of air sacs is correlated with the anatomy of the hyoid bone, a probable minimum and maximum date of the loss of air sacs can be estimated from fossil hyoid bones. Australopithecus afarensis still had air sacs about 3.3 Ma, while Homo heidelbergensis, some 600 000 years ago and Homo neandethalensis some 60 000 years ago, did no longer. The reduced distinctiveness of articulations produced with an air sac is in line with the hypothesis that air sacs were selected against because of the evolution of complex vocal communication. This relation between complex vocal communication and fossil evidence may help to get a firmer estimate of when speech first evolved.     

L’architecture de l’épaule au sein du genre Homo : nouvelles interprétations

The morphology of human clavicles can be estimated by projecting them on two perpendicular planes in order to assess the shapes of their cranial and dorsal primary curvatures. In cranial view no differences in curvature appear within the genus Homo, which means the different species had similar arms elevation capacity, especially in protraction. On the contrary, in dorsal view two clavicles morphologies could be defined. The first one is characterized by two curvatures in dorsal view and is possessed by all Homo species, from Homo habilis to Neanderthal, including Homo ergaster, but not modern human, Upper Paleolithic and anatomically modern human remains, who possess clavicles of the second type, characterized by either one curvature, or two slightly pronounced ones in dorsal view. Clavicles displaying two pronounced curvatures in dorsal view are associated with scapula sitting high on the thorax in regard to modern human. However, shoulder with high scapula on the thorax displays two different kinds of architectures: (i) shoulder with short clavicles associated to scapulas sitting more laterally than those of modern human. This group includes earlier Homo like Homo habilis and Homo ergaster and (ii) shoulder with long clavicles associated to scapulas sitting more dorsally on the thorax, like those of modern human. This group includes Homoantecessor and Neanderthals. In other words, within the genus Homo, three shoulders would have existed. Evolution of the shoulder complex is far more complex than previously thought and the arrival of modern bipedalism was not associated to modern shoulder.     

The language capability of Neanderthal man

Considerable publicity has been given a recent article by Lieberman and Crelin ('71) which argues that Neanderthal man lacked the physical features necessary for speech. This present paper presents statistics on some modern men with normal speech who show physical features in common with Neanderthal man. The brain of Neanderthal man was as large as, or larger than, that of modern man. The brain reflected by the endocranial cast of the La Chapelle-aux-Saints skull resembles that of modern man in an area important for speech and thereby suggests Neanderthal man had the neural development necessary for language.     

Phonetic Ability and Related Anatomy of the Newborn and Adult Human, Neanderthal Man, and the Chimpanzee

The sounds of human speech make human language a rapid medium of communication through a process of speech "encoding." The presence of sounds like the vowels [a], [i], and [u] makes this process possible. The supralaryngeal vocal tracts of newborn Homo sapiens and chimpanzee are similar and resemble the reconstructed vocal tract of the fossil La Chapelle-aux-Saints Neanderthal man. Vocal tract area functions that were directed toward making best possible approximations to the human vowels [a], [i], and [u], as well as certain consonantal configurations, were modeled by means of a computer program. The lack of these vowels in the phonetic repertories of these creatures, who lack a supralaryngeal pharyngeal region like that of adult Homo sapiens, may be concomitant with the absence of speech encoding and a consequently linguistic ability inferior to modern man.     

Tissue proportions and enamel thickness distribution in the early Middle Pleistocene human deciduous molars from Tighenif,Algeria

The present study of three human upper deciduous molars from the early Middle Pleistocene site of Tighenif, Algeria, constitutes the first microtomographic-based endostructural exploration of African fossil teeth likely representative of the Homo heidelbergensis morph. Comparative morphological observations and 2-3D measurements describing subtle tooth organization (crown tissue proportions) and enamel thickness topography (site-specific distribution and global patterning) indicate that their virtual extracted structural signature better fits the modern human, rather than the Neanderthal condition. Accordingly, we predict that the inner structural morphology of the deciduous molars from the Middle Pleistocene western European series better fits the primitive, and not the derived Neanderthal figures.     

Genome digging: insight into the mitochondrial genome of Homo

Background

A fraction of the Neanderthal mitochondrial genome sequence has a similarity with a 5,839-bp nuclear DNA sequence of mitochondrial origin (numt) on the human chromosome 1. This fact has never been interpreted. Although this phenomenon may be attributed to contamination and mosaic assembly of Neanderthal mtDNA from short sequencing reads, we explain the mysterious similarity by integration of this numt (mtAncestor-1) into the nuclear genome of the common ancestor of Neanderthals and modern humans not long before their reproductive split.

Principal Findings

Exploiting bioinformatics, we uncovered an additional numt (mtAncestor-2) with a high similarity to the Neanderthal mtDNA and indicated that both numts represent almost identical replicas of the mtDNA sequences ancestral to the mitochondrial genomes of Neanderthals and modern humans. In the proteins, encoded by mtDNA, the majority of amino acids distinguishing chimpanzees from humans and Neanderthals were acquired by the ancestral hominins. The overall rate of nonsynonymous evolution in Neanderthal mitochondrial protein-coding genes is not higher than in other lineages. The model incorporating the ancestral hominin mtDNA sequences estimates the average divergence age of the mtDNAs of Neanderthals and modern humans to be 450,000–485,000 years. The mtAncestor-1 and mtAncestor-2 sequences were incorporated into the nuclear genome approximately 620,000 years and 2,885,000 years ago, respectively.

Conclusions

This study provides the first insight into the evolution of the mitochondrial DNA in hominins ancestral to Neanderthals and humans. We hypothesize that mtAncestor-1 and mtAncestor-2 are likely to be molecular fossils of the mtDNAs of Homo heidelbergensis and a stem Homo lineage. The d_N/d_S dynamics suggests that the effective population size of extinct hominins was low. However, the hominin lineage ancestral to humans, Neanderthals and H. heidelbergensis, had a larger effective population size and possessed genetic diversity comparable with those of chimpanzee and gorilla.     

Evaluating Neanderthal Genetics and Phylogeny

The retrieval of Neanderthal (Homo neanderthalsensis) mitochondrial DNA is thought to be among the most significant ancient DNA contributions to date, allowing conflicting hypotheses on modern human (Homo sapiens) evolution to be tested directly. Recently, however, both the authenticity of the Neanderthal sequences and their phylogenetic position outside contemporary human diversity have been questioned. Using Bayesian inference and the largest dataset to date, we find strong support for a monophyletic Neanderthal clade outside the diversity of contemporary humans, in agreement with the expectations of the Out-of-Africa replacement model of modern human origin. From average pairwise sequence differences, we obtain support for claims that the first published Neanderthal sequence may include errors due to postmortem damage in the template molecules for PCR. In contrast, we find that recent results implying that the Neanderthal sequences are products of PCR artifacts are not well supported, suffering from inadequate experimental design and a presumably high percentage (>68%) of chimeric sequences due to “jumping PCR” events. Electronic Supplementary Material Electronic Supplementary material is available for this article at and accessible for authorised users. [Reviewing Editor: Dr. Martin Kreitman]     

The Neanderthal face is not cold adapted

Many morphological features of the Pleistocene fossil hominin Homo neanderthalensis, including the reputed large size of its paranasal sinuses, have been interpreted as adaptations to extreme cold, as some Neanderthals lived in Europe during glacial periods. This interpretation of sinus evolution rests on two assumptions: that increased craniofacial pneumatization is an adaptation to lower ambient temperatures, and that Neanderthals have relatively large sinuses. Analysis of humans, other primates, and rodents, however, suggests that the first assumption is suspect; at least the maxillary sinus undergoes a significant reduction in volume in extreme cold, in both wild and laboratory conditions. The second assumption, that Neanderthal sinuses are large, extensive, or even ‘hyperpneumatized,’ has held sway since the first specimen was described and has been interpreted as the causal explanation for some of the distinctive aspects of Neanderthal facial form, but has never been evaluated with respect to scaling. To test the latter assumption, previously published measurements from two-dimensional (2D) X-rays and new three-dimensional (3D) data from computed tomography (CT) of Neanderthals and temperate-climate European Homo sapiens are regressed against cranial size to determine the relative size of their sinuses. The 2D data reveal a degree of craniofacial pneumatization in Neanderthals that is both commensurate with the size of the cranium and comparable in scale with that seen in temperate climate H. sapiens. The 3D analysis of CT data from a smaller sample supports this conclusion. These results suggest that the distinctive Neanderthal face cannot be interpreted as a direct result of increased pneumatization, nor is it likely to be an adaptation to resist cold stress; an alternative explanation is thus required.     

Morphological analysis of the neanderthal calotte from Ochtendung,Germany

In 1997 one of the authors (AvB) found three pieces of a calotte of a Neanderthal in a crater depression within the slag-cone volcanic-group called “Wannenköpfe” near the village of Ochtendung in the state of Rhineland-Palatinate, Germany. Three stone tools of the Mousterian culture were directly associated with the hominid fossil. The individual can be securely stratigraphically and absolutely chronologically placed within the early glacial phase of the second last glaciation, the Saale. A comparative-morphological analysis confirms earlier analyses that the remains were those of an adult male. The relatively old age establishes it as another important find of an early Neanderthal in Europe. In absolute years it dates roughly to the transitional time period of the latest EuropeanHomo erectus to the earliest Neanderthal. Morphological analysis confirms that the individual is close to a typical Neanderthal with also some additional erectoid characters. This observation supports the most widely accepted view that the Neanderthal of Europe evolved from an autochthonousHomo erectus group.     

The Neanderthal lower arm

Neanderthal forearms have been described as being very powerful. Different individual features in the lower arm bones have been described to distinguish Neanderthals from modern humans. In this study, the overall morphology of the radius and ulna is considered, and morphological differences among Neanderthals, Upper Paleolithic Homo sapiens and recent H. sapiens are described.Comparisons among populations were made using a combination of 3D geometric morphometrics and standard multivariate methods. Comparative material included all available complete radii and ulnae from Neanderthals, early H. sapiens and archaeological and recent human populations, representing a wide geographical and lifestyle range.There are few differences among the populations when features are considered individually. Neanderthals and early H. sapiens fell within the range of modern human variation. When the suite of measurements and shapes were analyzed, differences and similarities became apparent. The Neanderthal radius is more laterally curved, has a more medially placed radial tuberosity, a longer radial neck, a more antero-posteriorly ovoid head and a well-developed proximal interosseous crest. The Neanderthal ulna has a more anterior facing trochlear notch, a lower M. brachialis insertion, larger relative mid-shaft size and a more medio-lateral and antero-posterior sinusoidal shaft. The Neanderthal lower arm morphology reflects a strong cold-adapted short forearm. The forearms of H. sapiens are less powerful in pronation and supination. Many differences between Neanderthals and H. sapiens can be explained as a secondary consequence of the hyper-polar body proportions of the Neanderthals, but also as retentions of the primitive condition of other hominoids.     

Are Homo sapiens nonsupranuchal fossa and neanderthal suprainiac fossa convergent traits?

The autapomorphic status of the Neanderthal suprainiac fossa was recently confirmed. This was a result of a detailed analysis of the internal bone composition in the area of the suprainiac depression on Neanderthal and Homo sapiens specimens. However, while anatomical differences between Neanderthal suprainiac fossa and the depression in the inion region of the occipital bone of fossil and recent Homo sapiens have been discussed in detail, the etiology of these structures has not been resolved. In this article, the hypothesis that the Homo sapiens non-supranuchal fossa and the Neanderthal suprainiac fossa both formed to maintain the optimal shape of the occipital plane (to minimize strain on the posterior cranial vault) is tested. First, the variation in the expression of the fossa above inion in the crania of recent Homo sapiens from European, African, and Australian samples was examined, and the degree of structural similarity between these depressions and the Neanderthal suprainiac fossa was assessed. Next, the relationship between the shape of the occipital squama in the midsagittal plane and two particular features (the degree of the occipital torus development and the occurrence of a depression in the inion region that is not the supranuchal fossa) were analyzed. Based on the results, it is suggested that the Homo sapiens non-supranuchal fossa and Neanderthal suprainiac fossa are convergent traits.     

The Vindija Neanderthal scapular glenoid fossa: comparative shape analysis suggests evo-devo changes among Neanderthals

Although the shape of the scapular glenoid fossa (SGF) may be influenced by epigenetic and developmental factors, there appears to be strong genetic control over its overall form, such that variation within and between hominin taxa in SGF shape may contain information about their evolutionary histories. Here we present the results of a geometric morphometric study of the SGF of the Neanderthal Vi-209 from Vindjia Cave (Croatia), relative to samples of Plio-Pleistocene, later Pleistocene, and recent hominins. Variation in overall SGF shape follows a chronological trend from the plesiomorphic condition seen in Australopithecus to modern humans, with pre-modern species of the genus Homo exhibiting intermediate morphologies. Change in body size across this temporal series is not linearly directional, which argues against static allometry as an explanation. However, life history and developmental rates change directionally across the series, suggesting an ontogenetic effect on the observed changes in shape (ontogenetic allometry). Within this framework, the morphospace occupied by the Neanderthals exhibits a discontinuous distribution. The Vindija SGF and those of the later Near Eastern Neanderthals (Kebara and Shanidar) approach the modern condition and are somewhat segregated from both northwestern European (Neandertal and La Ferrassie) and early Mediterranean Neanderthals (Krapina and Tabun). Although more than one scenario may account for the pattern seen in the Neanderthals, the data is consistent with palaeogenetic evidence suggesting low levels of gene flow between Neanderthals and modern humans in the Near East after ca. 120-100 ka (thousands of years ago) (with subsequent introgression of modern human alleles into eastern and central Europe). Thus, in keeping with previous analyses that document some modern human features in the Vindija Neanderthals, the Vindija G₃ sample should not be seen as representative of ‘classic’ - that is, unadmixed, pre-contact - Neanderthal morphology.     

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.     

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.     

The Homo sapiens ‘hemibun’: Its developmental pattern and the problem of homology

The occipital bun is widely considered a Neanderthal feature. Its homology to the ‘hemibun’ observed in some European Upper Palaeolithic anatomically modern humans is a current problem. This study quantitatively evaluates the degree of occipital plane convexity in African and Australian modern human crania to analyse a relationship between this feature and some neurocranial variables. Neanderthal and European Upper Palaeolithic Homo sapiens crania were included in the analysis as well. The results of this study indicated that there is a significant relationship between the degree of occipital plane convexity and the following two features in the examined crania of modern humans: the ratio of the maximum neurocranial height to the maximum width of the vault and the ratio of bregma–lambda chord to bregma–lambda arc. The results also revealed that some H. sapiens crania (modern and fossil) show the Neanderthal shape of the occipital plane and that the neurocranial height and shape of parietal midsagittal profile has an influence on occipital plane convexity in the hominins included in this study. This study suggests that the occurrence of the great convexity of the occipital plane in the Neanderthals and H. sapiens is a “by-product” of the relationship between the same neurocranial features and there is no convincing evidence that the Neanderthal occipital bun and the similar structure in H. sapiens develop during ontogeny in the same way.