Cranial base morphology and temporal bone pneumatization in Asian Homo erectus

The external morphological features of the temporal bone are used frequently to determine taxonomic affinities of fossils of the genus Homo. Temporal bone pneumatization has been widely studied in great apes and in early hominids. However, this feature is rarely examined in the later hominids, particularly in Asian Homo erectus. We provide a comparative morphological and quantitative analysis of Asian Homo erectus from the sites of Ngandong, Sambungmacan, and Zhoukoudian, and of Neandertals and anatomically modern Homo sapiens in order to discuss causes and modalities of temporal bone pneumatization during hominid evolution. The evolution of temporal bone pneumatization in the genus Homo is more complex than previously described. Indeed, the Zhoukoudian fossils have a unique pattern of temporal bone pneumatization, whereas Ngandong and Sambungmacan fossils, as well as the Neandertals, more closely resemble the modern human pattern. Moreover, these Chinese fossils are characterized by a wide midvault and a relatively narrow occipital bone. Our results support the point of view that cell development does not play an active role in determining cranial base morphology. Instead, pneumatization is related to available space and to temporal bone morphology, and its development is related to correlated morphology and the relative disposition of the bones and cerebral lobes. Because variation in pneumatization is extensive within the same species, the phyletic implications of pneumatization are limited in the taxa considered here.     

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.     

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.     

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.     

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.     

Chronological changes in stone tool assemblages from Krapina (Croatia)

This study presents the results of the first recent analysis of stone tool assemblages from Krapina (Croatia). All assemblages are Pleistocene in age and many are associated with human remains, the Krapina Neandertals. The assemblages are described typologically and technologically, and subtle chronological changes in raw material selection and technology of tool blank production are observed. These changes involve increasingly sophisticated and selective use of lithic materials. Changing artefact assemblages are considered in light of variability in the hominids from Krapina, and are interpreted as reflecting behavioral change among Neandertals rather than between Neandertal and modern human populations.     

An interesting rock from Krapina

Symbolic items are seldom associated with Neandertals and, when they are, many paleoanthropologists consider them to be Neandertal imitations from modern Homo sapiens. At the Croatian site of Krapina dated to MIS 5e or about 130,000 years ago, among many items, a split limestone rock was excavated by Dragutin Gorjanović-Kramberger between 1899-1905. This brownish rock reveals many dark dendritic forms in cross- and longitudinal section. Of more than 1000 lithic items at Krapina, none resemble this specimen and we propose it was collected and not further processed by the Neandertals because of its aesthetic attributes. Along with other examples from sites in western and central Europe and the recent discovery of eight modified white-tailed eagle talons from Krapina, this unique item suggests that Neandertals were capable, on their own, of incorporating symbolic objects into their culture.     

Evidence for Neandertal Jewelry: Modified White-Tailed Eagle Claws at Krapina

We describe eight, mostly complete white-tailed eagle (Haliaëtus [Haliaeetus] albicilla) talons from the Krapina Neandertal site in present-day Croatia, dating to approximately 130 kyrs ago. Four talons bear multiple, edge-smoothed cut marks; eight show polishing facets and/or abrasion. Three of the largest talons have small notches at roughly the same place along the plantar surface, interrupting the proximal margin of the talon blade. These features suggest they were part of a jewelry assemblage, --- the manipulations a consequence of mounting the talons in a necklace or bracelet. An associated phalanx articulates with one of the talons and has numerous cut marks, some of which are smoothed. These white-tailed eagle bones, discovered more than 100 years ago, all derive from a single level at Krapina and represent more talons than found in the entire European Mousterian period. Presence of eight talons indicates that the Krapina Neandertals acquired and curated eagle talons for some kind of symbolic purpose. Some have argued that Neandertals lacked symbolic ability or copied this behavior from modern humans. These remains clearly show that the Krapina Neandertals made jewelry well before the appearance of modern humans in Europe, extending ornament production and symbolic activity early into the European Mousterian.     

Prevalence and the duration of linear enamel hypoplasia: a comparative study of Neandertals and Inuit foragers

As a dental indicator of generalized physiological stress, enamel hypoplasia has been the subject of several Neandertal studies. While previous studies generally have found high frequencies of enamel hypoplasia in Neandertals, the significance of this finding varies with frequencies of enamel hypoplasia in comparative samples. The present investigation was undertaken to ascertain if the enamel hypoplasia evidence in Neandertals suggests a high level of physiological stress relative to a modern human foraging group, represented here by an archaeological sample of Inuit from Point Hope, Alaska. Unlike previous studies, this study focused specifically on linear enamel hypoplasia (LEH), emphasizing systemic over localized causes of this defect by considering LEH to be present in an individual only if LEH defects occur on two anterior teeth with overlapping crown formation periods. Moreover, this study is the first to evaluate the average growth disruption duration represented by these defects in Neandertals and a comparative foraging group. In the prevalence analysis, 7/18 Neandertal individuals (from Krapina and southern France) and 21/56 Neandertal anterior teeth were affected by LEH, or 38.9% and 37.5% respectively. These values do not differ significantly from those of the Inuit sample in which 8/21, or 38.1% of individuals, and 32/111, or 28.8% of anterior teeth were affected. For the growth disruption duration analysis, 22 defects representing separate episodes of growth disruption in Neandertals were compared with 22 defects in the Inuit group using three indicators of duration: the number of perikymata (growth increments) in the occlusal walls of LEH defects, the total number of perikymata within them, and defect width. Only one indicator, the total number of perikymata within defects, differed significantly between the Inuit and Neandertal groups (an average of 13.4 vs. 7.3 perikymata), suggesting that if there is any difference between them, the Inuit defects may actually represent longer growth disruptions than the Neandertal defects. Thus, while stress indicators other than linear enamel hypoplasia may eventually show that Neandertal populations were more stressed than those of modern foragers, the evidence from linear enamel hypoplasia does not lend support to this idea.     

Brief communication: paleoanthropology and the population genetics of ancient genes

The Mezmaiskaya cave mtDNA is similar in many ways to the Feldhofer cave Neandertal sequence and the more recently obtained Vindija cave sequence. If we accept the contention that the Mezmaiskaya cave specimen is a Neandertal infant, its mtDNA provides no new information about the fate of the European Neandertals. However, there is reason to believe that the Mezmaiskaya cave infant is not a Neandertal, and this places its importance in another light, because it delimits the possible hypotheses of Neandertal and recent human genetic relationships. One possibility is a that the pattern found in ancient mtDNA results from the replacement of an isolated gene pool (Neandertals) by one of its contemporaries (modern humans). A second possibility is natural selection expressed as the substitution of an advantageous mtDNA variant within a single large species, including both Neandertals and modern humans. The geologic, archaeological, and dating evidence shows the Mezmaiskaya cave infant to be a burial from a level even more recent than the Upper Paleolithic preserved at the site, and its anatomy does not contradict the assessment that the Mezmaiskaya cave infant is not a Neandertal. Therefore, the second pattern can be favored over the first.     

Hominid enamel thickness: I. The Krapina Neandertals

Dental x-rays were taken of isolated and in situ adult molar teeth of the Krapina Neandertal (n = 63) and of recent and contemporary molars (n = 423). The radiographs were digitized at high resolution (1,024 × 1,520 × 8 bits) with a 35 mm solid state scanner. Ratios of enamel cap area to the underlying dentinal-pulpal area were determined and comparisons were made between average ratios for the Neandertal and contemporary molars. Neandertal molars had significantly smaller ratios than did contemporary teeth (P < 0.05). It is suggested that the smaller ratios represent relatively thinner enamel for Neandertals and that the thin enamel may have been caused by a metabolic depression that resulted in reduced enamel quantity (hypoplasia). Alternately, the observed differences may be related to expanded pulps seen in various stages of taurodontism. © 1993 Wiley-Liss, Inc.     

Upper pleistocene human remains from Vindija cave,Croatia, Yugoslavia

Human remains excavated from Vindija cave include a large although fragmentary sample of late Mousterian-associated specimens and a few additional individuals from the overlying early Upper Paleolithic levels. The Mousterian-associated sample is similar to European Neandertals from other regions. Compared with earlier Neandertals from south central Europe, this sample evinces evolutionary trends in the direction of Upper Paleolithic Europeans. Compared with the western European Neandertals, the same trends can be demonstrated, although the magnitude of difference is less, and there is a potential for confusing temporal with regional sources of variation. The early Upper Paleolithic-associated sample cannot be distinguished from the Mousterian-associated hominids. We believe that this site provides support for Hrdli?ka's “Neandertal phase” of human evolution, as it was originally applied in Europe.     

The postcranial dimensions of the La Chapelle-aux-saints 1 Neandertal

The La Chapelle-aux-Saints 1 Neandertal has figured prominently in considerations of Neandertal body size and proportions. In this context, a reassessment of its major long bones and a reassembly of its principal pelvic elements (sacrum and right ilium) was undertaken. There are secure measurements for its humeral and radial lengths and its femoral head diameter, but the femoral and tibial lengths were almost certainly greater than previous values. The resultant humeral, femoral and tibial lengths are similar to those of other male Neandertals, its femoral head diameter is among the largest known for Middle and Late Pleistocene humans, but its radial length is relatively short. The pelvic assembly provides modest bi-iliac and inlet transverse diameters compared with the few sufficiently complete and undistorted Middle and Late Pleistocene archaic human pelves, but its dimensions are similar to those of large male early modern humans.     

Is the late Neandertal mandibular sample from Vindija Cave (Croatia) biased?

The late Neandertal sample from Vindija (Croatia) has been described as transitional between the earlier Central European Neandertals from Krapina (Croatia) and modern humans. However, the morphological differences indicating this transition may rather be the result of different sex and/or age compositions between the samples. This study tests the hypothesis that the metric differences between the Krapina and Vindija mandibular samples are due to sample bias. Mandibles are the focus of this paper because past studies have posited this region as particularly indicative of the Vindija sample's transitional nature. The results indicate that the metric differences between the Krapina and Vindija mandibular samples are not due to sample bias. This conclusion is consistent with an earlier analysis of sample bias for the Vindija supraorbital sample.     

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.     

A reconsideration of the Archi 1 Neandertal mandible

A reassessment of the early last glacial immature Neandertal mandibular corpus from Archi indicates a series of features in which it closely resembles other pre-adolescent Neandertal mandibles and contrasts with those of similarly aged recent humans. These are in the context of a re-aging of the specimen to ca. 3 years on the basis of deciduous dental eruption and attrition and permanent dental calcification. The Archi 1 mandible resembles other immature Neandertals in having a “retreating” symphyseal profile in the context of moderate development of mental trigone features. It is relatively robust in the development of lateral and basilar corpus features and some increased symphyseal and lateral corpus thickness. And it exhibits, along with other Middle Paleolithic immature mandibles, anteriorly wide dental arcades, probably due to large developing anterior permanent tooth crowns.     

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.