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.     

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.     

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.     

The Gravettian occipital bone from the site of Malladetes (Barx,Valencia, Spain)

The juvenile occipital bone from the site of Malladetes in Valencia (Spain) is described and compared with other European Pleistocene representatives of the genus Homo. This specimen derives from a Gravettian cultural context and has been AMS radiocarbon-dated to 25,120 +/- 240 years BP. As such, it provides evidence on early modern human anatomy from the Central Mediterranean region of the Iberian peninsula. The clear evidence for a late survival of Neandertals in southern Iberia, has led to considerable debate surrounding the biological and cultural interactions between these Pleistocene humans and their early modern human successors, and it is within this context that the Malladetes specimen represents an important contribution to the discussion. The recently discovered Upper Paleolithic infant from the site of Lagar Velho in Portugal is said to show a mosaic of Neandertal and early modern human characteristics throughout the skeleton and is argued to represent the strongest evidence yet recovered in favor of hybridization between these two Pleistocene populations. Our analysis of the Malladetes occipital, however, reveals no evidence of Neandertal genetic influence.     

A fossil hominid frontal from Velika Pećina (Croatia) and a consideration of Upper Pleistocene hominids from Yugoslavia

Fossil hominid remains dating to the Upper Pleistocene in Yugoslavia are reviewed. Particular emphasis is placed on the assessment of a hominid frontal from the site of Velika Pe?ina in northwestern Croatia. This specimen represents the earliest absolutely-dated hominid associated with the Upper Paleolithic in Europe. Also the hominid material from the site of Veternica is discussed, and data are presented on the new remains from ?andalja. It is concluded that no Neandertal remains, except for Krapina, have been found in Yugoslavia.     

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.     

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.     

Size variation in early human mandibles and molars from Klasies River,South Africa: Comparison with other middle and late Pleistocene assemblages and with modern humans

Previous studies of the Middle Stone Age human remains from Klasies River have concluded that they exhibited more sexual dimorphism than extant populations, but these claims have not been assessed statistically. We evaluate these claims by comparing size variation in the best‐represented elements at the site, namely the mandibular corpora and M₂s, to that in samples from three recent human populations using resampling methods. We also examine size variation in these same elements from seven additional middle and late Pleistocene sites: Skhūl, Dolní Věstonice, Sima de los Huesos, Arago, Krapina, Shanidar, and Vindija. Our results demonstrate that size variation in the Klasies assemblage was greater than in recent humans, consistent with arguments that the Klasies people were more dimorphic than living humans. Variation in the Skhūl, Dolní Věstonice, and Sima de los Huesos mandibular samples is also higher than in the recent human samples, indicating that the Klasies sample was not unusual among middle and late Pleistocene hominins. In contrast, the Neandertal samples (Krapina, Shanidar, and Vindija) do not evince relatively high mandibular and molar variation, which may indicate that the level of dimorphism in Neandertals was similar to that observed in extant humans. These results suggest that the reduced levels of dimorphism in Neandertals and living humans may have developed independently, though larger fossil samples are needed to test this hypothesis. Am J Phys Anthropol, 2009. © 2009 Wiley‐Liss, Inc.     

The late Neandertal supraorbital fossils from Vindija Cave,Croatia: a biased sample?

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 supraorbital samples are due to sampling bias. We focus upon the supraorbital region because past studies have posited this region as particularly indicative of the Vindija sample's transitional nature. Furthermore, the supraorbital region varies significantly with both age and sex.We analyzed four chords and two derived indices of supraorbital torus form as defined by Smith & Ranyard (1980, Am. J. phys. Anthrop.93, pp. 589-610). For each variable, we analyzed relative sample bias of the Krapina and Vindija samples using three sampling methods. In order to test the hypothesis that the Vindija sample contains an over-representation of females and/or young while the Krapina sample is normal or also female/young biased, we determined the probability of drawing a sample of the same size as and with a mean equal to or less than Vindija's from a Krapina-based population. In order to test the hypothesis that the Vindija sample is female/young biased while the Krapina sample is male/old biased, we determined the probability of drawing a sample of the same size as and with a mean equal or less than Vindija's from a generated population whose mean is halfway between Krapina's and Vindija's. Finally, in order to test the hypothesis that the Vindija sample is normal while the Krapina sample contains an over-representation of males and/or old, we determined the probability of drawing a sample of the same size as and with a mean equal to or greater than Krapina's from a Vindija-based population. Unless we assume that the Vindija sample is female/young and the Krapina sample is male/old biased, our results falsify the hypothesis that the metric differences between the Krapina and Vindija samples are due to sample bias.     

Right handedness of Homo heidelbergensis from Sima de los Huesos (Atapuerca, Spain) 500,000 years ago

Handedness is a product of brain specialization, which in turn seems to be responsible for the higher cognitive capabilities of humans, such as language and technology. Handedness in living humans is well established and shows the highest degree of manual specialization. Studies on hand laterality in nonhuman primates, particularly in chimpanzees, remain a matter of controversy as results tend to vary depending on factors such as the tasks performed and the environment in which the individuals live. Studies in several disciplines have attempted to determine where in the course of human evolution handedness established itself, with evidence collected from sources such as paleoneurological analyses, stone tool flaking, zooarchaeological studies and dental wear analyses, the last one of which have proven the most reliable source of information. Here we report an experimental and paleoanthropological study on hand laterality of a sample of 28 hominids from Sima de los Huesos (Atapuerca, Spain), dated at about 500,000 years ago, and compare our results with dental microwear analysis in other fossil samples such as that from Krapina (Croatia), as well as modern traditional societies. Our results indicate that European Middle Pleistocene Homo heidelbergensis was already as right-handed as modern populations.     

Reexamination of the immature hominid maxilla from Tangier,Morocco

Reexamination of the immature Upper Pleistocene hominid maxilla from Mugharet el-'Aliya (Tangier), Morocco is undertaken in light of new evidence on the growth and development of Upper Pleistocene hominids. Metric and qualitative comparisons were made with 17 immature Upper Pleistocene maxillae, and with a recent Homo sapiens sapiens sample. No unambiguous criteria for aligning the maxilla with Neandertals were found, although one character, the degree of maxillary flexion on the zygoma, strongly suggests that this child could be a representative of H. s. sapiens. The probable lack of a canine fossa in Mugharet el-'Aliya 1, the primary criterion used previously to align it with Neandertals, cannot be accurately extrapolated to its adult form from this juvenile. The present evidence suggests that it is inappropriate to refer to this fossil as “Neandertal-like” or as a North African “neandertaloid.” Thus, the Tangier maxilla should not be cited as evidence for the presence of Neandertal facial features in North Africa during the Upper Pleistocene. © 1993 Wiley-Liss, Inc.     

Fibrous Dysplasia in a 120,000+ Year Old Neandertal from Krapina,Croatia

We describe the first definitive case of a fibrous dysplastic neoplasm in a Neandertal rib (120.71) from the site of Krapina in present-day Croatia. The tumor predates other evidence for these kinds of tumor by well over 100,000 years. Tumors of any sort are a rare occurrence in recent archaeological periods or in living primates, but especially in the human fossil record. Several studies have surveyed bone diseases in past human populations and living primates and fibrous dysplasias occur in a low incidence. Within the class of bone tumors of the rib, fibrous dysplasia is present in living humans at a higher frequency than other bone tumors. The bony features leading to our diagnosis are described in detail. In living humans effects of the neoplasm present a broad spectrum of symptoms, from asymptomatic to debilitating. Given the incomplete nature of this rib and the lack of associated skeletal elements, we resist commenting on the health effects the tumor had on the individual. Yet, the occurrence of this neoplasm shows that at least one Neandertal suffered a common bone tumor found in modern humans.     

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.     

Re-analysis of the hominid radii from Cave of Hearths and Klasies River Mouth, South Africa

Two of the few postcranial fragments from the late Early Stone Age and/or the Middle Stone Age of southern Africa are the proximal radii from the Cave of Hearths and Klasies River Mouth. The Cave of Hearths fossil is metrically indistiguishable from both archaic (e.g., Neandertals) and recent humans, and presents a mosaic of primitive and modern features. The primitive include a relatively slender neck and thick cortical bone (the latter of which distinguishes recent humans from archaic, Early Modern, and Upper Paleolithic hominids); the modern includes an anteromedially (rather than medially) facing radial tuberosity. Its extreme collo–diaphyseal angle is unusual, although it can be matched by modern homologues. The neck–shaft angle of some Neandertal and Early Modern radii also appears to match that of the Cave of Hearths specimen. The Klasies River Mouth radius also has thick cortical bone of the neck. It is morphologically indistinguishable from Early Modern and Neandertal homologues. These, and other fossils, suggest a mosaic pattern of evolution in the postcranial skeleton of the late Early Stone Age and/or Middle Stone Age inhabitants of sub-Saharan Africa.     

An Australasian test of the recent African origin theory using the WLH-50 calvarium

This analysis investigates the ancestry of a single modern human specimen from Australia, WLH-50 (Thorne et al., in preparation; Webb, 1989). Evaluating its ancestry is important to our understanding of modern human origins in Australasia because the prevailing models of human origins make different predictions for the ancestry of this specimen, and others like it. Some authors believe in the validity of a complete replacement theory and propose that modern humans in Australasia descended solely from earlier modern human populations found in Late Pleistocene Africa and the Levant. These ancestral modern populations are believed to have completely replaced other archaic human populations, including the Ngandong hominids of Indonesia. According to this recent African origin theory, the archaic humans from Indonesia are classified as Homo erectus, a different evolutionary species that could not have contributed to the ancestry of modern Australasians. Therefore this theory of complete replacement makes clear predictions concerning the ancestry of the specimen WLH-50. We tested these predictions using two methods: a discriminant analysis of metric data for three samples that are potential ancestors of WLH-50 (Ngandong, Late Pleistocene Africans, Levant hominids from Skhul and Qafzeh) and a pairwise difference analysis of nonmetric data for individuals within these samples. The results of these procedures provide an unambiguous refutation of a model of complete replacement within this region, and indicate that the Ngandong hominids or a population like them may have contributed significantly to the ancestry of WLH-50. We therefore contend that Ngandong hominids should be classified within the evolutionary species, Homo sapiens. The Multiregional model of human evolution has the expectation that Australasian ancestry is in all three of the potentially ancestral groups and best explains modern Australasian origins.     

Bone breakage in the Krapina hominid collection

The fragmentary condition of the Krapina Neandertal remains has been offered as one line of evidence for the hypothesis that these hominids were the victims of cannibals seeking marrow and brains. Two other hypotheses regarding the causes of the framentation have been raised: a substantial portion of the breakage in the Krapina collection is attributable to excavation damage; and the rest of the breakage is attributable to sedimentary pressure and to natural rock falls that occurred during the site's prehistory. The purpose of this paper is report on tests of these three hypotheses concerning the cause of breakage in the Krapina material. Microscopic inspection of all Krapina hominid specimens showed that 23% of the material was inadvertantly broken during excavation or during quarrying that took place at the end of the last century. The morphology of the prehistoric breakage is inconsistent with the cannibalism hypothesis and supports the hypothesis that prehistoric breakage was caused by sedimentary pressure and/or roof falls.     

Early Pleistocene human humeri from the Gran Dolina-TD6 site (Sierra de Atapuerca, Spain)

In this report, we present a morphometric comparative study of two Early Pleistocene humeri recovered from the TD6 level of the Gran Dolina cave site in Sierra de Atapuerca, northern Spain. ATD6-121 belongs to a child between 4 and 6 years old, whereas ATD6-148 corresponds to an adult. ATD6-148 exhibits the typical pattern of the genus Homo, but it also shows a large olecranon fossa and very thin medial and lateral pillars (also present in ATD6-121), sharing these features with European Middle Pleistocene hominins, Neandertals, and the Bodo Middle Pleistocene humerus. The morphology of the distal epiphysis, together with a few dental traits, suggests a phylogenetic relationship between the TD6 hominins and the Neandertal lineage. Given the older geochronological age of these hominins (ca. 900 ka), which is far from the age estimated by palaeogenetic studies for the population divergence of modern humans and Neandertals (ca. 400 ka), we suggest that this suite of derived "Neandertal" features appeared early in the evolution of the genus Homo. Thus, these features are not "Neandertal" apomorphies but traits which appeared in an ancestral and polymorphic population during the Early Pleistocene.