The Middle/Later Stone Age transition and cultural dynamics of late Pleistocene East Africa

The Middle to Later Stone Age (MSA/LSA) transition is a prominent feature of the African archeological record that began in some places ~30,000–60,000 years ago, historically associated with the origin and/or dispersal of “modern” humans. Unlike the analogous Middle to Upper Paleolithic transition in Eurasia and associated Neanderthal extinction, the African MSA/LSA record remains poorly documented, with its potential role in explaining changes in the behavioral diversity and geographic range of Homo sapiens largely unexplored. I review archeological and biogeographic data from East Africa, show regionally diverse pathways to the MSA/LSA transition, and emphasize the need for analytical approaches that document potential ancestor‐descendent relationships visible in the archeological record, needed to assess independent invention, population interaction, dispersal, and other potential mechanisms for behavioral change. Diversity within East Africa underscores the need for regional, rather than continental‐scale narratives of the later evolutionary history of H. sapiens.     

Neandertals and moderns mixed,and it matters

Twenty‐five years ago, the Middle‐to‐Upper Paleolithic transition in Europe could be represented as a straightforward process subsuming both the emergence of symbolic behavior and the replacement of Neandertals by modern humans. The Aurignacian was a proxy for the latter, during which enhanced cognitive capabilities explained ornaments and art. The few instances of Neandertal symbolism were deemed to long postdate contact and dismissed as “imitation without understanding,” if not geological contamination. Such views were strengthened by the recent finding that, in southern Africa, several features of the European Upper Paleolithic, including bone tools, ornaments, and microliths, emerged much earlier. Coupled with genetic suggestions of a recent African origin for extant humans, fossil discoveries bridging the transition between “archaics” and “moderns” in the realm of anatomy (Omo‐Kibish, Herto) seemingly closed the case. Over the last decade, however, taphonomic critiques of the archeology of the transition have made it clear that, in Europe, fully symbolic sapiens behavior predates both the Aurignacian and moderns. And, in line with evidence from the nuclear genome rejecting strict replacement models based on mtDNA alone, the small number of early modern specimens that passed the test of direct dating present archaic features unknown in the African lineage, suggesting admixture at the time of contact. In the realm of culture, the archeological evidence also supports a Neandertal contribution to Europe's earliest modern human societies, which feature personal ornaments completely unknown before immigration and are characteristic of such Neandertal‐associated archeological entities as the Châtelperronian and the Uluzzian. The chronometric data suggest that, north of the Ebro divide, the entire interaction process may have been resolved within the millennium centered around 42,000 calendar years ago. Such a rapid absorption of the Neandertals is consistent with the size imbalance between the two gene reservoirs and further supports significant levels of admixture.     

Late archaic and modern Homo sapiens from Europe, Africa, and Southwest Asia: Craniometric comparisons and phylogenetic implications

The craniometric affinities among Neandertals. Upper Palcolithic Europeans, early anatomically modern Southwest Asians, and archaic and modern Africans are investigated using univariate and multivariate methods. For the first time, it is possible to analyse the North African finds Dar-es-Soltane 5, Nazlet Khater, and Wadi Kubbaniya. It was not possible to include the Neandertals from Central Europe due to their poor state of preservation. The results point to, first, a basic distinction between Neandertals on the one hand and modern humans from all geographic regions on the other, and, secondly, to great similarities between modern African and European populations. Late archaic sapiens specimens from Africa were more similar to Upper Paleolithic Europeans than were the Neandertals. The results do not support the hypothesis that a regional evolution giving rise to modern humans took place in Europe. The results are, however, consistent with the hypothesis that modern populations originated in Africa and spread to Europe from there.     

The Big Deal about Blades: Laminar Technologies and Human Evolution

Despite the rapid expansion of archaeological knowledge of the Paleolithic over the past several decades, some generalized interpretive frameworks inherited from previous generations of researchers are remarkably tenacious. One of the most persistent of these is the assumed correlation between blade technologies, Upper Paleolithic industries, and anatomically (and behaviorally) modern humans. In this paper, we review some of the evidence for the production of early blade technologies in Eurasia and Africa dating to the late Lower and the Middle Paleolithic. The basic techniques for blade production appeared thousands of years before the Upper Paleolithic, and there is no justification for linking blades per se to any particular aspect of hominid anatomy or to any major change in the behavioral capacities of hominids. It is true that blades came to dominate the archaeological records of western Eurasia and Africa after 40,000 years ago, perhaps as a consequence of increasing reliance on complex composite tools during the Upper Paleolithic. At the same time, evidence from other regions of the world demonstrates that evolutionary trends in Pleistocene Eurasia were historically contingent and not universal. [Middle Paleolithic, Upper Paleolithic, blade technology, human evolution, hominid behavior and capacities]     

Existe-t-il un Paléolithique inférieur au Japon ?

All Lower Paleolithic sites discovered by Mr. Shinïchi FUJIMURA from 1981 to 2000 were falsified by himself. We wonder if Lower Paleolithic sites really exist (corresponding to a period of 30,000 years ago in Japan) in the archipelago of Japan. We examined cultures of sites likely dating to the Early Upper Paleolithic and Lower Paleolithic discovered in the archipelago of Japan. While Japan was continuous with the continent in the Mindel glaciation (400,000 years ago) or in the Riss glaciation (200,000 years ago), Homo erectus or Homo heidelbergensis with the lithic industry of the Acheulean arrived in the archipelago of Japan. Thereafter, they survived on the islands of Japan cut the continent to the Upper Paleolithic while keeping the industry.     

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

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

Genetics and modern human origins

The past decade has brought considerable debate on the subject of modern human origins. The nature of the transition from Homo erectus to archaic Homo sapiens to modern H. sapiens has been examined primarily in terms of the relative contribution of archaic populations to later moderns, both within and among geographic regions. The recent African origin model proposes that modern humans appeared first in Africa between 100,000 and 200,000 years ago, and then spread through the rest of the Old World, replacing preexisting populations.^1–6 This model has been referred to by a variety of names, including “replacement”, “Garden of Eden”, “Noah's Ark”, and “out of Africa”. The recent African origin model contrasts with the multiregional model, which proposes a species-wide transition to modern humans throughout the Old World during the past million years or more.^7–10 Indeed, some proponents of the multiregional model advocate placing Homo erectus and all subsequent species of Homo in the evolutionary species Homo sapiens.¹¹ This contrasts with the view that there were multiple hominid species during the Middle Pleistocene. The debate continues.^12,13 Although the multiregional model is often portrayed as proposing a simultaneous transition to anatomically modern humans in different geographic regions, it explicitly allows for varying degrees of continuity across time and space.¹⁰ This model, in the broad sense, does not rule out the possibility that modern human morphology appeared first in Africa and then spread through the rest of the Old World through gene flow. However, not all advocates of the multiregional model adhere to this specific subset of the general model.⁹ Comparison of the African and multiregional models is complicated by considering other, less extreme, hypotheses. Some versions of the recent African origin model imply a speciation event associated with the initial origin of modern humans. Another version, which suggests the possibility of some admixture between “moderns” leaving Africa and preexisting “archaics” elsewhere in the Old World,^14,15 is similar to some variants of the multiregional model, which also suggest that modern morphology appeared first in Africa, but involved admixture with other Old World populations.¹⁶ The major difference between these views appears to be the extent of admixture, although the exact level is never specified. A further complication is the possibility that multiple dispersals from Africa produced a more complicated pattern of worldwide variation.¹⁷     

The Origin of Homo sapiens in the Light of Different Research Methods

The origins of Homo sapiens is a central issue of modern paleoanthropology. The available fossil material serves as a basis for postulating different hypotheses and models, but as is widely appreciated, anthropologists have yet to reach a consensus about human origins. It seems possible that the main reasons behind such an irreducible divergence of opinions are different methodological approaches rather than the analyses of fossil material. Some scientists would say that it is the fragmentary nature of fossil material which accounts for the debates about the origins of H. sapiens. Had the debate been only a matter of the empirical considerations many disagreements concerning H. sapiens would have probably disappeared long ago. But since the controversies are imbued in methodological reality the closure of the debate is not to be expected soon. There are three research methods: morphological, archaeological and genetic. Each approach has a specific definition of H. sapiens at its disposal, which largely accounts for the different scenarios for the origin of our species. Any debate concerning this problem must therefore begin with a discussion about the research methods. The controversy about our origins thus appears to be of secondary importance. The present paper aims at presenting the methodological controversy in relation to the origins of H. sapiens. The discussion about the genealogy of H. sapiens certainly is in urgent need of a new, more integrated way of approaching the past.     

Les origines de l’art et les théories sur l’évolution humaine : le cas français

In recent years, we have witnessed an international debate about the question of the origins of art. On the one hand, some specialists have suggested that art appeared for the first time at the beginning of the Upper Paleolithic associated to the emergence of Homo sapiens sapiens. From this point of view, Paleolithic art as well as other hallmarks of behavioral modernity were exclusive to anatomically modern humans. On the other hand, some scholars have put into question the traditional paradigm concerning the origins of art and have suggested that artistic objects arose over a long period of time among different species, including Neanderthals. In order to contextualize this debate, we analyze in this article the history of the different interpretations and controversies concerning the question of the origins of art. Taking as reference the French case, we examine the connections between the different theories about art's origins suggested by Pleistocene art specialists during the last century and the dominant paradigms in human paleontology during the same period. Informed by one another, the question of the origins of art and that of human evolution seems to be inextricable linked.     

Recent work on the Paleolithic of Central Asia

Central Asia has played an important role in Paleolithic archeology since the discovery of Teshik-Tash in 1938. Since the 1970s, attention has turned to Lower Paleolithic studies following the important discoveries at Kul'dara and other deeply stratified loess sites in southern Tajikistan. The sites, dated to as early as 800,000 years ago, are not the earliest evidence of hominids in Asia, but they do reflect early adaptations to arid mid-latitude environments. The stone-tool industries from these early sites are composed of pebble and flake implements; they do not include bifaces. Along with the archeological material in the loess is an impressive paleoenvironmental record with good chronological control. In other parts of Central Asia, there are reported elements of the Acheulean technocomplex at sites such as Sel'ungur and Yangadazha, but that claim cannot be substantiated. In the Upper Pleistocene, important research questions include the extent and meaning of Middle Paleolithic variability, the relation of Middle to Upper Paleolithic cultures, and the effect of glacial or interglacial climatic oscillations on Paleolithic settlement and adaptation.     

The absolute dating of Upper Pleistocene subSaharan fossil hominids and their place in human evolution

In the evolution of anatomically modern man and his subspecies most specialists have concentrated on investigating geographical areas other than Africa as the possible area of origin.In this study 20 fossil hominids and associated faunal remains from South and East Africa were dated by microanalysis, radiocarbon, and amino-acid dating in order to see whether modern man appears later, was sympatric, or even predated Neandertal man.These dates indicate that anatomically modern man occurs sympatrically and possibly even predates the Rhodesian group of Neandertals in Africa. Modern man might also be contemporary to and possibly even predate the occurrence of Neandertal in Europe.This would indicate that modern man did not evolve from but possibly gave rise to the Neandertals as off-shoots.Two possibilities for the evolution of modern man are suggested. First, that Homo sapiens capensis evolved about 90,000 to 100,000 years ago from possibly Homo erectus by way of a “basic” Homo sapiens and later gave rise to Homo sapiens rhodesiensis, Homo sapiens afer, and possibly Homo sapiens palestinus around 50,000 years ago with Homo neanderthalensis and Homo sapiens capensis evolving separately from Homo erectus. In this case Homo neanderthalensis would be a different species from Homo sapiens which includes Homo sapiens capensis, Homo sapiens rhodesiensis, Homo sapiens afer, and possibly Homo sapiens palestinus.Secondly, Homo sapiens capensis evolved by way of a “basic” Homo sapiens with Homo sapiens rhodesiensis and Homo sapiens palestinus branching off from Homo sapiens capensis around 50,000 years ago. Before that, around 90,000 to 100,000 years ago Homo sapiens capensis evolved first and was then followed by Homo sapiens neanderthalensis from a “basic” Homo sapiens stock, but diverged. This means, all Neandertals, Homo sapiens capensis, Homo sapiens sapiens and Homo sapiens afer can be considered as subspecies of Homo sapiens.The author favors the first scheme since on relative dating grounds the existence of Neandertal man in Europe before the earliest date of Homo sapiens capensis and a “basic” Homo sapiens seems to be fairly well documented. Irrespective of either one of these possibilities, modern man evolved in Africa and seems to have migrated into Europe and other parts of the world.New absolute dating techniques are mentioned in detail like the new radiocarbon-collagen method and amino acid dating.     

The emergence and intensification of early hunter‐gatherer niche construction

Hunter‐gatherers, especially Pleistocene examples, are not well‐represented in archeological studies of niche construction. However, as the role of humans in shaping environments over long time scales becomes increasingly apparent, it is critical to develop archeological proxies and testable hypotheses about early hunter‐gatherer impacts. Modern foragers engage in niche constructive behaviors aimed at maintaining or increasing the productivity of their environments, and these may have had significant ecological consequences over later human evolution. In some cases, they may also represent behaviors unique to modern Homo sapiens. Archeological and paleoenvironmental data show that African hunter‐gatherers were niche constructors in diverse environments, which have legacies in how ecosystems function today. These can be conceptualized as behaviorally mediated trophic cascades, and tested using archeological and paleoenvironmental proxies. Thus, large‐scale niche construction behavior is possible to identify at deeper time scales, and may be key to understanding the emergence of modern humans.     

The Neanderthal lower arm

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

Changes in the ‘Connectedness’ and Resilience of Paleolithic Societies in Mediterranean Ecosystems

Human predator‐prey relationships changed dramatically in the Mediterranean Basin between 250,000 and 9,000 years ago. Many of these changes can be linked to increases in Paleolithic human population densities. Small game species are particularly diagnostic of increases in human hunting pressure and are a major source of evidence for demographic change after 40–45,000 years ago. Biomass-corrected data on prey choice also indicate increasing use of those species that possess higher reproductive efficiencies. Step-wise, apparently irreversible shifts in human predatory niche are apparent in the Mediterranean Basin, beginning with the earliest Upper Paleolithic in the east and spreading westward. Evidence of demographic pressure and greater use of resilient prey populations is followed by technological innovations to exploit these animals more efficiently. The zooarchaeological findings suggest that Middle and Lower Paleolithic reproductive units probably were not robust at the micropopulation scale, due to the rather narrow set of behavioral responses that characterized social groups at the time, and thus localized extinctions at the micropopulation level were likely to have been common. Upper Paleolithic groups were the quintessential colonizers and, in addition, uniquely good at holding on to habitat gained. Upper Paleolithic archaeological “cultures” had shorter histories of existence than those of earlier periods, but they were even more widespread geographically. The demographic robustness of the Upper Paleolithic systems may stem from wholesale strategies for evening-out or sharing risk and volatility in technology. Micropopulations were larger and often denser on landscapes, more connected via cooperative ties, and thus more robust.     

Stress physiologique et état de santé des plus anciens Hommes anatomiquement modernes du sud-est de l’Europe (données dentaires,couche 6-1, Buran-Kaya III,Crimée)

Buran-Kaya III site, in Crimea, provides an exceptional stratigraphic sequence, extending from Middle Paleolithic to Neolithic. Among the archaeological layers allocated to the Gravettian (sensu lato), more than 170 anthropological remains have been discovered in the layer 6-1. They represent the oldest Upper Paleolithic modern humans from Eastern Europe in a well-documented archaeological context. The aim of this article, based on dental remains and enamel dental hypoplasias in particular, is to evaluate the health of the modern humans, which lived more than 32,000 years BP ago in Crimea. This study puts into the light the low frequency of hypoplasias (occurring in the two youngest individuals). This article complete the rare studies concerning the presence of this physiological stress during the Early Upper Paleolithic and confirm the scarcity of enamel dental hypoplasias and dental pathologies, such as caries, at this period.     

The upper paleolithic of Europe: An overview

The Upper Paleolithic of Europe, 40,000–10,000 years ago, presents one of the richest, most complex records for the anatomy and cultural adaptations of fossil hominids in the world. New chronological information points to roughly simultaneous appearance of certain Upper Paleolithic technological traits in both SE and SW Europe, while growing evidence suggests a significant degree of biological and cultural continuity between the Middle and Upper Paleolithic. On the other hand, there is considerable evidence that evolution continued to operate in both domains throughout the course of the late Upper Pleistocene, apparently in adaptive relationship to the major environmental changes of the Upper Pleniglacial and Tardiglacial. Spectacular developments in the realms of art and ideology may be understood in the special biogeographical, social, and economic conditions of Europe at the height of the Last Ice Age; both ended rather abruptly with the onset of the Holocene as the landscapes of Europe underwent pervasive upheavals.     

Fossil Humankind and Other Anthropoid Primates of China

More than 70 sites have yielded human fossils in China. They are attributed to Homo sapiens erectus and Homo sapiens sapiens. The earliest one is possibly about 1.7 Ma. A series of common morphological features, including shovel-shaped incisors and flatness of the face, characterize them. There is a morphological mosaic between H. s. erectus and H. s. sapiens in China. The existence of common features and the morphological mosaic suggest continuity of human evolution in China. That there are a few features which are more commonly seen in the Neanderthal lineage, occurring in a few Chinese fossil skulls, probably suggests gene flow between China and the West. Based on them, in 1998 I proposed an hypothesis—continuity with hybridization—for human evolution in China. The hypothesis is supported by paleolithic archeology, and it supports the multiregional evolution hypothesis of modern human origins. The anatomically modern humans of East Asia originated most probably in China. Although some nonhuman anthropoid primates of China—Gigantopithecus, Sivapithecus, Ramapithecus and Lufengpithecus—have been suggested as the direct ancestors of human beings, the discovery of more specimens and further studies do not support these suggestions. Therefore, it is most probable that the transition between apes and humans did not occur in China.     

Genetic analysis of lice supports direct contact between modern and archaic humans

Parasites can be used as unique markers to investigate host evolutionary history, independent of host data. Here we show that modern human head lice, Pediculus humanus, are composed of two ancient lineages, whose origin predates modern Homo sapiens by an order of magnitude (ca. 1.18 million years). One of the two louse lineages has a worldwide distribution and appears to have undergone a population bottleneck ca. 100,000 years ago along with its modern H. sapiens host. Phylogenetic and population genetic data suggest that the other lineage, found only in the New World, has remained isolated from the worldwide lineage for the last 1.18 million years. The ancient divergence between these two lice is contemporaneous with splits among early species of Homo, and cospeciation analyses suggest that the two louse lineages codiverged with a now extinct species of Homo and the lineage leading to modern H. sapiens. If these lice indeed codiverged with their hosts ca. 1.18 million years ago, then a recent host switch from an archaic species of Homo to modern H. sapiens is required to explain the occurrence of both lineages on modern H. sapiens. Such a host switch would require direct physical contact between modern and archaic forms of Homo.     

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.     

Throwing in the Middle and Upper Paleolithic: inferences from an analysis of humeral retroversion

When in evolutionary history did long-range projectile weapons become an important component of hunting toolkits? The archeological evidence for the development of projectile weaponry is complex and generally indirect, and has led to different conclusions about the origin and spread of this technology. Lithic evidence from the Middle Stone Age (MSA) has led some researchers to suggest that true long- range projectile weaponry developed in Africa perhaps as early as 80,000 years ago, and was part of the subsistence toolkit carried by modern humans who expanded out of Africa after 50,000 years ago. Alternatively, temporal patterns in the morphology of pointed lithics has led others to posit an independent, convergent origin of projectile weaponry in Africa, the Near East, and Europe during the interval between 50,000-40,000 years ago. By either scenario, projectile weapons would not have been a component of the hunting arsenal of Neandertals, but may have been in use by European early modern humans and thus, projectile technology may have entered into the competitive dynamics that existed between these two groups. The origins of projectile weapons can be addressed, in part, through analyses of the skeletal remains of the prehistoric humans who made and used them. Habitual behavior patterns—including those related to the production and use of technology—can be imprinted on the skeleton through both genetic and epigenetic pathways. Recent studies in the field of sports medicine indicate that individuals who engage in habitual throwing have increased humeral retroversion angles in their throwing arms and a greater degree of bilateral asymmetry in retroversion angles than do non-throwers. This contribution investigates humeral torsion through analysis of the retroversion angle in samples of Eurasian Neandertals, European early modern humans of the middle and late Upper Paleolithic, and comparative samples of recent humans. This analysis was conducted under the assumption that if throwing-based projectile weaponry was used by early modern Europeans but not Neandertals, Upper Paleolithic samples should be similar to recent human groups engaged in habitual throwing in the degree of humeral retroversion in the dominant limb and in bilateral asymmetry in this feature. Neandertals on the other hand, would not be expected to show marked asymmetry in humeral retroversion. Consistent with other studies, Neandertals exhibit increased retroversion angles (decreased humeral torsion or a more posteriorly oriented humeral head) relative to most modern human samples, although this appears more likely related to body form and overall activity levels than to habitual throwing. Although Neandertals with bilaterally preserved humeri sufficient for measurement are rare (consisting of only two males and one female), levels of bilateral asymmetry in humeral retroversion are low, suggesting a lack of regular throwing. While patterning across fossil and comparative samples in levels of humeral retroversion was not clear cut, males of both the middle and late Upper Paleolithic demonstrate a high level of bilateral asymmetry, comparable to or in excess of that seen in samples of throwing athletes. This may indicate habitual use of throwing-based projectile weaponry by middle Upper Paleolithic times. Small sample sizes and relatively great variance in the fossil samples makes these results, however, suggestive rather than conclusive.