Brief communication: Investigation of the semicircular canal variation in the Krapina Neandertals

Previous studies comparing bony labyrinth morphology in geographically‐dispersed samples of Neandertals and modern Homo sapiens (H. sapiens) showed that Neandertals generally have smaller semicircular canals than modern H. sapiens (Hublin et al., 1996 ; Spoor et al., 2003 ; Glantz et al., 2008 ). Here we analyze the morphology of a single group of Neandertal specimens from one locale, the Krapina site, to determine the intraspecific variation in Neandertal semicircular canal sizes. Dimensions of the semicircular canals were collected from computed tomography scans of nine temporal bones. With the rare exception, the dimensions of the semicircular canals in the Krapina sample are similar to those previously reported across a geographically‐dispersed sample of Neandertals, further supporting previous studies that suggest low levels of variation in the semicircular canals for Neandertals. Am J Phys Anthropol 154:302–306, 2014. © 2014 Wiley Periodicals, Inc.     

On the meaning of increased fluctuating dental asymmetry: a cross populational study.

Suarez reports a greater magnitude of fluctuating dental asymmetry for Neandertal sample when compared with a sample of modern Ohio whites. He postulates that this greater antimeric variance could be due to a greater degree of inbreeding in the Neandertal populations. In the present investigation, the magnitude of fluctuating dental asymmetry is evaluated for Eskimo and Pueblo populations. These populations were found to exhibit dental variance of equal magnitude to that of the Neandertal population. As these populations are not highly inbred, a stress related mechanism is suggested to explain these observations and the inbreeding hypothesis is rejected. The implications of this mechanism to Brace's Probable Mutation Effect are discussed.     

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.     

Postcranial skeleton of a Neandertal child from Kiik-Koba, U.S.S.R.

A study of the skeletal remains of a Neandertal child from Kiik-Koba has brought to light some features that are typical of adult Neandertal individuals from Europe. Therefore, the ascertained differences in the proportions of the individual parts of upper and lower limbs, as well as a number of marked morphological features, such as the robusticity of ribs, typical Neandertal features of the scapula, curving of forearm bones, back-to-front curving of the femoral diaphysis, retroversion of the upper epiphysis of the tibia, and robusticity of long bones in the Neandertal individual at the age of infans I, may be regarded as phylogenetically ingrained ones. These may in their turn be considered diagnostic features, of importance in determining the phylogenetic classification of postcranial skeletal remains in the development of man.     

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.     

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.     

The Crocuta crocuta spelaea (Goldfuss 1823) population and its prey from the Late Pleistocene Teufelskammer Cave hyena den besides the famous Paleolithic Neandertal Cave (NRW,NW Germany)

Remains from at least seven individuals of the Late Pleistocene Ice Age spotted hyena Crocuta crocuta spelaea (Goldfuss, 1823) from the Teufelskammer Cave in the Neandertal valley (North Rhine-Westphalia, northwest Germany) are described. The small cave was a well-frequented hyena den of the Early to Middle Late Pleistocene which was only 100 m from the famous small Feldhofer Cave, where the first Neandertal human skeleton was found. The high amount of hyena bone material (37%) and its strongly chewed and incomplete prey remains of the mixed mammoth steppe and boreal forest megafauna prove one more of 11 recently known hyena den caves in the Rhenish Massif. Hyenas and cave bears have used the cave, but Neandertal humans lived possibly not at the same time in the same valley. Although hyenas occupied mainly the smaller caves such as the Teufelskammer Cave, humans preferred large portal cave entrances such as in the Neandertal valley with the Small Feldhofer Cave.     

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.     

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.     

Relative dental development of Upper Pleistocene hominids compared to human population variation

The relative development of permanent teeth in samples of Neandertal/archaic Homo and Early Modern/Upper Paleolithic hominids is compared to the range of variability found in three recent human samples. Both fossil hominid samples are advanced in relative M2 and M3 development compared to white French-Canadians, but only the Neandertal/archaic Homo M3 sample is advanced when compared to black southern Africans. Both fossil hominid samples are delayed in relative I1 and P3 development compared to the recent human samples. Two hypotheses concerning the significance of the advanced M3 and M2 development found in both hominid groups and southern Africans compared to French-Canadians are discussed. The first postulates that the differences in relative molar development are due simply to variation in tooth/jaw size relationships. The second postulates that the relatively advanced M3 and M2 development found in the fossil hominids and southern Africans is a correlate of their potential for advanced skeletal maturation compared to French-Canadians and other European-derived populations. It appears that dental development patterns have continued to evolve from the Upper Pleistocene to present times, and that Neandertals and Early Moderns shared similar patterns of relative dental development. © 1996 Wiley-Liss, Inc.     

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.     

Revisiting dental fluctuating asymmetry in neandertals and modern humans

Previous studies have suggested that Neandertals experienced greater physiological stress and/or were less capable of mitigating stress than most prehistoric modern human populations. The current study compares estimates of dental fluctuating asymmetry (DFA) for prehistoric Inupiat from Point Hope Alaska, the Late Archaic, and Protohistoric periods from Ohio and West Virginia, and a modern sample from Ohio to Neandertals from Europe and Southwest Asia. DFA results from developmental perturbation during crown formation and is thus an indicator of developmental stress, which previous studies have found to be higher in Neandertals than in several modern human populations. Here, we use recent methodological improvements in the analysis of fluctuating asymmetry suggested by Palmer and Strobeck (Annu Rev Ecol Syst 17 ( 1986 ) 391–421, Developmental instability: causes and consequences ( 2003a ) v.1–v.36, Developmental instability: causes and consequences ( 2003b ) 279–319) and compare the fit of Neandertal DFA Index values with those of modern humans. DFA estimates for each of the modern population samples exceeded measurement error, with the Inupiat exhibiting the highest levels of DFA for most tooth positions. All significant Neandertal z‐scores were positive, exceeding the estimates for each of the modern prehistoric groups. Neandertals exhibited the fewest significant differences from the Inupiat (9.2% of values are significant at P < 0.05), while for the other modern prehistoric groups more than 10% of the Neandertal z‐scores are significant at P < 0.05, more than 90% of these significant scores at P < 0.01. These results suggest that the Inupiat experienced greater developmental stress than the other prehistoric population samples, and that Neandertals were under greater developmental stress than all other prehistoric modern human samples. Am J Phys Anthropol 149:193–204, 2012. © 2012 Wiley Periodicals, Inc.     

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.     

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.     

Developmental stress and cranial hypostosis by epigenetic trait occurrence and distribution: an exploratory study on the Italian Neandertals

The occurrence and distribution of 35 cranial epigenetic traits in the Italian Neandertals (Saccopastore 1 and 2, Guattari 1) were examined according to morpho-functional cranial regions and with respect to a distinction between hypostosis (i.e., weak osseous development, arrested morphogenesis, retention of infantile features) and hyperostosis (i.e. excess of ossification, not reaching the pathological condition). The results, expressed ashypostotic scores, showed higher levels of hypostosis in these Neandertal specimens than in recent European samples. The highest expressions of hypostotis were observed in those regions of the Neandertal cranium where disequilibrium between skeletal and cerebral growth factors was expected. These results—interpreted as expression of developmental stress—are consistent with a heterochronic interpretation of the development of the Neandertal cranium; namely, a faster ossification of the cranial vault relative to brain growth rates.     

La Ferrassie 6 and the development of Neandertal pubic morphology

Functional interpretations of the mediolateral elongation and superoinferior flattening of the superior pubic ramus of Neandertals require knowledge of its ontogeny. Metric comparisons between the La Ferrassie 6 Neandertal infant, aged 3-5 years, and a sample of modern infants reveal that the acetabulosymphyseal length of La Ferrassie 6, relative to femoral length and iliac breadth, falls at the limits of the range of variation of the modern infants, while the relative height of its superior pubic ramus is indistinguishable from that of the modern sample. It appears that acetabulosymphyseal elongation of the Neandertal pubis is a feature which is expressed quite early in ontogeny, well before puberty. Superoinferior flattening and ventral margin thinning of the superior ramus probably appear later in Neandertal ontogeny.     

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.     

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.