The Spy VI child: a newly discovered Neandertal infant

Spy cave (Jemeppe-sur-Sambre, Belgium) is reputed for the two adult Neandertal individuals discovered in situ in 1886. Recent reassessment of the Spy collections has allowed direct radiocarbon dating of these individuals. The sorting of all of the faunal collections has also led to the discovery of the remains of a Neandertal child, Spy VI. This individual is represented by two mandibular corpus fragments. The left fragment is the most complete and both sides preserve the mental foramen. Four deciduous teeth are associated with these mandibular remains: three incisors and one canine. The lower left canine (Spy 645a) conjoins with the corresponding alveolar socket in the left part of the mandible. Following extant standards, the developmental stage of the preserved teeth indicate an age at death of about one and a half years. In addition to performing a classical morphometric comparative study of the mandible and teeth, we have evaluated the dental tissue proportions using high-resolution microtomographic techniques. Our results show that Spy VI generally falls within the Neandertal range of variation. However, this specimen also exhibits particular traits, notably in the dental internal structural organization, which reveals that variation in the immature Neandertal variation is larger than what was variation currently represented by the available fossil record. These observations demonstrate the need for investigating the frequency and expression of immature Neandertal traits in fossil anterior teeth, as well as their temporal and geographic variation. Direct radiocarbon dating of the Spy VI specimen has been conducted in two different laboratories. The results of Spy VI confirm the age previously determined for the two adults, making the Spy Neandertal remains the youngest ever directly dated in northwest Europe.     

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.     

Unique ramus anatomy for Neandertals?

The ramus of Neandertal mandibles is said to show a suite of uniquely Neandertal character states that demonstrate the independent course of Neandertal evolution. This is the latest of numerous attempts to define cranial and mandibular autapomorphies for Neandertals. We examine variation in the four presumably autapomorphic ramal features and show they are neither monomorhic within Neandertals (to the contrary Neandertals are at least as variable as other human samples) nor unique to Neandertals, since they regularly appear in populations predating and postdating them. Neandertals differ from other human populations, both contemporary and recent, but the question of whether this fact reflects a divergent evolutionary trajectory must be addressed by the pattern of differences. In this case, as in the other attempts to establish Neandertal autapomorphies, rather than showing restricted variation and increased specialization, the Neandertal sample shows that the range of human variation in the recent past encompasses, and in some cases exceeds, human variation today, even in the very features claimed to be autapomorphic.     

Sexual dimorphism in the human bony pelvis, with a consideration of the Neandertal pelvis from Kebara Cave, Israel.

Sexual dimorphism of the human pelvis is inferentially related to obstetrics. However, researchers disagree in the identification and obstetric significance of pelvic dimorphisms. This study addresses three issues. First, common patterns in dimorphism are identified by analysis of pelvimetrics from six independent samples (Whites and Blacks of known sex and four Amerindian samples of unknown sex). Second, an hypothesis is tested that the index of pelvic dimorphism (female mean x 100/male mean) is inversely related to pelvic variability. Third, the pelvic dimensions of the Neandertal male from Kebara cave, Israel are compared with those of the males in this study. The results show that the pelvic inlet is the plane of least dimorphism in humans. The reason that reports often differ in the identification of dimorphisms for this pelvic plane is that both the length of the pubis and the shape of the inlet are related to nutrition. The dimensions of the pelvis that are most dimorphic (that is, female larger than male) are the measures of posterior space, angulation of sacrum, biischial breadth, and subpubic angle. Interestingly, these dimensions are also the most variable. The hypothesis that variability and dimorphism are inversely related fails to be supported. The factors that influence pelvic variability are discussed. The Kebara 2 pelvis has a spacious inlet and a confined outlet relative to modern males, though the circumferences of both planes in the Neandertal are within the range of variation of modern males. The inference is that outlet circumference in Neandertal females is also small in size, but within the range of variation of modern females. Arguments that Neandertal newborns were larger in size than those of modern humans necessarily imply that birth was more difficult in Neandertals.     

New Neandertal remains from Cova Negra (Valencia, Spain)

New Neandertal fossils from the Mousterian site of Cova Negra in the Valencia region of Spain are described, and a comprehensive study of the entire human fossil sample is provided. The new specimens significantly augment the sample of human remains from this site and make Cova Negra one of the richest human paleontological sites on the Iberian Peninsula. The new specimens include cranial and postcranial elements from immature individuals and provide an opportunity to study the ontogenetic appearance of adult Neandertal characteristics in this Pleistocene population. Children younger than 10 years of age constitute four of the seven minimum number of individuals in the sample, and this relative abundance of children at Cova Negra is similar that in to other Neandertal sites in Europe and southwest Asia. The recognition of diagnostic Neandertal features in several of the specimens, as well as their western European context and late Pleistocene age, suggests that all the human remains from Cova Negra represent Neandertals. The archaeological evidence from Cova Negra indicates sporadic, short-term occupations of the site, suggesting a high degree of mobility among Neandertals.     

Krapina 1: a juvenile Neandertal from the early late Pleistocene of Croatia

The juvenile A Skull from Krapina, Croatia (Krapina 1) has been the subject of considerable debate since B. Skerlj first suggested that it might not be a Neandertal. Although widely known by its original designation, the Krapina A Skull was recatalogued, along with all of the Krapina hominids, in the 1980's (Radovcic, et al., [1988]. The Krapina Hominids: An Illustrated Catalog of Skeletal Collection. Zagreb; Mladost). It is now catalogued as Krapina 1 in the archives of the Hrvatski Prirodoslovni Muzej, Zagreb, Croatia. We present a detailed, morphometric analysis of this specimen, comparing it to other Krapina specimens, juvenile late Pleistocene hominids (including Neandertals), and subadult recent humans. This analysis demonstrates that Krapina 1 possesses morphological features that are primitive retentions; others that represent derived Neandertal specializations; and still others that are typical for all European late Pleistocene humans. Morphological features associated with the browridges are intermediate between Neandertal and early modern European form. Nevertheless, a thorough analysis of the morphology of this specimen, in ontogenetic and regional contexts, leads to the conclusion that it cannot be excluded from the Neandertal range of variation. We conclude that the most parsimonious explanation for this 130 ka specimen is that it should be regarded as a Neandertal.     

ESR and U-series analyses of enamel and dentine fragments of the Banyoles mandible

The Banyoles mandible presents a puzzle. Its anatomy has been described as pre-Neandertal, but the travertine in which it was found has been dated to 45,000 +/- 4000 years. By this time, any pre-Neandertals had supposedly been absent from the European fossil record for more than 100,000 years. It was therefore proposed that the age of the travertine may represent a minimum age estimate, with the mandible possibly having been reworked from older deposits. We carried out a non-destructive ESR analysis of an enamel fragment removed from a molar and performed a series of in situ laser ablation U-series analyses on dentine fragments adjacent to the enamel piece. The analyses resulted in an apparent combined ESR-U-series age of 66,000 +/- 7000 years. The encasing travertine matrix was also analyzed for U-series isotopes and showed signs of U-mobilization. It cannot be excluded that the travertine matrix is older than the previously determined age. If the mandible was not reworked, then the combined ESR-U-series result on the tooth enamel would give its best age estimate. If, on the other hand, the mandible was reworked from another deposit, the actual ESR-U-series age will depend on the external dose rate from the previous matrix and the depth of its burial, which controls the degree of the attenuation of the cosmic dose rate over time. Considering a range of possible burial histories, the mean age of the mandible would lie somewhere between the combined ESR-U-series age and the previously determined age of the travertine matrix. Regarding the morphology of the mandible, a review of its features in the context of larger Neandertal samples indicates that the anatomy of the specimen is not incompatible with such a young age determination, although it further highlights morphological variation in the late Neandertal sample.     

New Radiometric Ages for the BH-1 Hominin from Balanica (Serbia): Implications for Understanding the Role of the Balkans in Middle Pleistocene Human Evolution

Newly obtained ages, based on electron spin resonance combined with uranium series isotopic analysis, and infrared/post-infrared luminescence dating, provide a minimum age that lies between 397 and 525 ka for the hominin mandible BH-1 from Mala Balanica cave, Serbia. This confirms it as the easternmost hominin specimen in Europe dated to the Middle Pleistocene. Inferences drawn from the morphology of the mandible BH-1 place it outside currently observed variation of European Homo heidelbergensis. The lack of derived Neandertal traits in BH-1 and its contemporary specimens in Southeast Europe, such as Kocabaş, Vasogliano and Ceprano, coupled with Middle Pleistocene synapomorphies, suggests different evolutionary forces acting in the east of the continent where isolation did not play such an important role during glaciations.     

Bite force production capability and efficiency in Neandertals and modern humans

Although there is consensus that Neandertal craniofacial morphology is unique in the genus Homo, debate continues regarding the precise anatomical basis for this uniqueness and the evolutionary mechanism that produced it. In recent years, biomechanical explanations have received the most attention. Some proponents of the "anterior dental loading hypothesis" (ADLH) maintain that Neandertal facial anatomy was an adaptive response to high-magnitude forces resulting from both masticatory and paramasticatory activity. However, while many have argued that Neandertal facial structure was well-adapted to dissipate heavy occlusal loads, few have considered, much less demonstrated, the ability of the Neandertal masticatory system to generate these presumably heavy loads. In fact, the Neandertal masticatory configuration has often been simultaneously interpreted as being disadvantageous for producing large bite forces. With rare exception, analyses that attempted to resolve this conflict were qualitative rather than quantitative. Using a three-dimensional digitizer, we recorded a sequence of points on the cranium and associated mandible of the Amud 1, La Chapelle-aux-Saints, and La Ferrassie 1 Neandertals, and a sample of early and recent modern humans (n = 29), including a subsample with heavy dental wear and documented paramasticatory behavior. From these points, we calculated measures of force-production capability (i.e., magnitudes of muscle force, bite force, and condylar reaction force), measures of force production efficiency (i.e., ratios of force magnitudes and muscle mechanical advantages), and a measure of overall size (i.e., the geometric mean of all linear craniofacial measurements taken). In contrast to the expectations set forth by the ADLH, the primary dichotomy in force-production capability was not between Neandertal and modern specimens, but rather between large (robust) and small (gracile) specimens overall. Our results further suggest that the masticatory system in the genus Homo scales such that a certain level of force-production efficiency is maintained across a considerable range of size and robusticity. Natural selection was probably not acting on Neandertal facial architecture in terms of peak bite force dissipation, but rather on large tooth size to better resist wear and abrasion from submaximal (but more frequent) biting and grinding forces. We conclude that masticatory biomechanical adaptation does not underlie variation in the facial skeleton of later Pleistocene Homo in general, and that continued exploration of alternative explanations for Neandertal facial architecture (e.g., climatic, respiratory, developmental, and/or stochastic mechanisms) seems warranted.     

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 medial pterygoid tubercle in the Atapuerca Early and Middle Pleistocene mandibles: Evolutionary implications

Numerous studies have attempted to identify the presence of uniquely derived (autoapomorphic) Neandertal features. Here, we deal with the medial pterygoid tubercle (MTP), which is usually present on the internal face of the ascending ramus of Neandertal specimens. Our study stems from the identification of a hypertrophied tubercle in ATD6‐96, an Early Pleistocene mandible recovered from the TD6 level of the Atapuerca‐Gran Dolina site and attributed to Homo antecessor. Our review of the literature and study of numerous original fossil specimens and high quality replicas confirm that the MTP occurs at a high frequency in Neandertals (ca. 89%) and is also present in over half (ca. 55%) of the Middle Pleistocene Sima de los Huesos (SH) hominins. In contrast, it is generally absent or minimally developed in other extinct hominins, but can be found in variable frequencies (<ca. 25%) in Pleistocene and recent H. sapiens samples. The presence of this feature in ATD6‐96 joins other traits shared by H. antecessor, the SH hominins and Neandertals. Since the TD6 hominins have been attributed either to MIS 21 or to MIS 25, it seems that a suite of assumed derived Neandertal features appeared in the Early Pleistocene, and they should be interpreted as synapomorphies shared among different taxa. We suggest that H. antecessor, the SH hominins and Neandertals shared a common ancestor in which these features appeared during the Early Pleistocene. The presence of the MTP in taxa other than H. neanderthalensis precludes this feature from being a Neandertal autapomorphy. Am J Phys Anthropol 156:102–109, 2015 © 2014 Wiley Periodicals, Inc.     

Middle Pleistocene human remains from the Bau de l'Aubesier

Excavations in later Middle Pleistocene levels at the Bau de l'Aubesier, Vaucluse, France yielded a maxillary molar (M(1) or M(2); Aubesier 10) and a partial mandible from the left C(1) alveolus to the right condylar base lacking the coronoid process (Aubesier 11). Dentally they are similar to other later Middle Pleistocene Europeans in dental dimensions and variable taurodontism (Aubesier 10 but not Aubesier 11). The small Aubesier 11 mandible exhibits a retreating symphyseal profile with a minimal tuber symphyseos, an anterior marginal tubercle at P(4)/M(1), the mental foramen at P(4)/M(1)-M(1), a modest retromolar space, no lingular bridging of the mandibular foramen, an enlarged superior medial pterygoid tubercle, a modest lateral condylar tubercle, and a mandibular notch crest that intersects the middle third of the condylar margin. All of these features fall within the ranges of variation of later Middle Pleistocene Neandertal lineage humans, and some are characteristic of Middle Pleistocene human mandibles in general. In addition, Aubesier 11 exhibits pervasive ante mortem alveolar resorption with apical abscesses, alveolar bone destruction, universal labial/buccal bone loss, ante mortem tooth loss (for > or =81.8% of preserved alveoli), a lingual alveolar fenestration, and two broken root apices with masticatory attrition. These lesions indicate significantly impaired masticatory function, the oldest specimen currently known with such a reduced degree of dental function but one of several Middle Pleistocene human remains with indications of serious abnormalities.     

Brief communication: A morphometric analysis of the neandertal upper second molar leuca I

The scarcity of Neandertal remains from Southern Europe hampers our understanding of Neandertal variability, and can bias interpretations about Neandertal geographic variation. To address this issue, it is often important to reassess human remains that, while discovered decades ago, remain relatively unknown to the scientific community. In this contribution, we provide a complete state‐of‐the‐art comparative morphometric analysis of Leuca I, an unworn left second upper molar (LM²) discovered in 1958 in Bambino's Cave (near Santa Maria di Leuca, Apulia, Italy) and attributed to Homo neanderthalensis. Our study includes comparisons of standard metric and nonmetric data, a 2D image analysis of the occlusal surface and measurements of both 2D and 3D enamel thickness and dental tissue proportions. Although Leuca I follows the Neandertal M²s trend in some morphometric aspects (i.e., small relative occlusal polygon area), in other cases it falls to the higher end (for 3D average enamel thickness) or even outside (for 3D‐relative enamel thickness) the Neandertal M² variability, thus increasing the known Neandertal range of variation. Am J Phys Anthropol 152:300–305, 2013. © 2013 Wiley Periodicals, Inc.     

Fossil human remains from Bolomor Cave (Valencia, Spain)

Systematic excavations carried out since 1989 at Bolomor Cave have led to the recovery of four Pleistocene human fossil remains, consisting of a fibular fragment, two isolated teeth, and a nearly complete adult parietal bone. All of these specimens date to the late Middle and early Late Pleistocene (MIS 7-5e). The fibular fragment shows thick cortical bone, an archaic feature found in non-modern (i.e. non-Homo sapiens) members of the genus Homo. Among the dental remains, the lack of a midtrigonid crest in the M₁ represents a departure from the morphology reported for the majority of Neandertal specimens, while the large dimensions and pronounced shoveling of the marginal ridges in the C¹ are similar to other European Middle and late Pleistocene fossils. The parietal bone is very thick, with dimensions that generally fall above Neandertal fossils and resemble more closely the Middle Pleistocene Atapuerca (SH) adult specimens. Based on the presence of archaic features, all the fossils from Bolomor are attributed to the Neandertal evolutionary lineage.     

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.     

Neandertal radial tuberosity orientation

Examination of adult and immature Neandertal radii demonstrates that the medial versus anterior orientations of their radial tuberosities fall within recent human ranges of variation, but on the average their radial tuberosities are significantly more medially, as opposed to anteromedially, oriented. This more posterior positioning of their radial tuberosities implies a maintenance of an effective moment arm for M. biceps brachii through the full range of supination, an interpretation which fits with the hypertrophy of and increased moment arms for their forearm pronator muscles. It is an additional indication of the muscular hypertrophy evident elsewhere in Neandertal upper limbs.     

Lower limb entheseal morphology in the Neandertal Krapina population (Croatia, 130,000 BP)

Although the Neandertal locomotor system has been shown to differ from Homo sapiens, characteristics of Neandertal entheses, the skeletal attachments for muscles, tendons, ligaments and joint capsules, have never been specifically investigated. Here, we analyse lower limb entheses of the Krapina Neandertal bones (Croatia, 130,000 BP) with the aim of determining how they compare with modern humans, using a standard developed by our research group for describing modern human entheseal variability. The entheses examined are those of the gluteus maximus, iliopsoas and vastus medialis on the femur, the quadriceps tendon on the patella, and soleus on the tibia. For the entheses showing a different morphological pattern from H. sapiens, we discuss the possibility of recognising genetic versus environmental causes. Our results indicate that only the gluteus maximus enthesis (the gluteal tuberosity), falls out of the modern human range of variation. It displays morphological features that could imply histological differences from modern humans, in particular the presence of fibrocartilage. In both H. sapiens and the Krapina Neandertals, the morphological pattern of this enthesis is the same in adult and immature femurs. These results can be interpreted in light of genetic differences between the two hominins. The possibility of functional adaptations to higher levels of mechanical load during life in the Neandertals seems less likely. The particular morphology and large dimensions of the Krapina enthesis, and perhaps its fibrocartilaginous nature, could have been selected for in association with other pelvic and lower limb characteristics, even if genetic drift cannot be ruled out.     

Estimation of stature from the skeletal reconstruction of an immature Neandertal from Dederiyeh cave, Syria

Skeletal reconstruction of a child Neandertal unearthed at Dederiyeh Cave, Syria in 1993, is undertaken and the acquired stature discussed. Although the skeletal remains were well preserved, the reconstruction required several assumptions to be made because of the immature status of the specimen. The assumptions were mainly concerned with distances between bones in the inter-vertebral spaces and in the joints of the hip, knee, and ankle. These were estimated from X-ray films of modern children and data from previous studies. Stature was directly measured on the reconstruction, and found to be 79.2 cm. After corrections for soft tissue thickness and shrinkage of the casts, the stature became 81.7 cm. This estimate is consistent with estimates based on regression equations of long bone lengths, especially from those of the lower extremity. In comparison with longitudinal data for white American boys, the assessment of stature for Dederiyeh varied according to the estimated age. For a younger estimated age, the stature falls in the lower half of the white American range of variation, but with an older estimated age, it falls below the lower limit of the range of variation. Other immature Neandertals including two European specimens, Roc de Marsal and La Ferrassie 6, fall below the lower limit of the 5th to 95th percentile range based on the estimated statures from their long bone lengths. More comprehensive age assessment covering both fossil and modern humans is required before accurate conclusions in relation to Neandertal growth can be drawn.