Exploring the lack of articular ends at the Middle Pleistocene site of Qesem Cave, Israel.

Biased skeletal part representation is a key element for making inferences about transport decisions, carcass procurement, and use patterns in anthropogenic accumulations. In the absence of destructive taphonomic processes, it is often assumed that the abundance of different anatomical portions represents selective transport and discard patterns of human groups. Because body parts may be transported for specific products such as meat, marrow or grease, a pattern that usually attracts attention in many archaeological sites is the low proportions of appendicular epiphyses. Here we present the case of faunal assemblages from the lower stratigraphic sequence of Qesem Cave, Israel, dated to ca. 430 to 300 ka. All bone accumulations are characterized by a biased skeletal profile including mainly long-limb bones and a virtual absence of epiphyses. The assemblages also show density-mediated attrition not linked to fossil-diagenetic processes, a targeted specific destruction to the most greasy articular ends and an almost total absence of carnivore intervention. Our goal here is to explore the processes that entail the destruction of appendicular epiphyses at Qesem Cave, as well as propose viable hypotheses to explain their underrepresentation on-site. Our results shed light on the domestic activities linked to the processing of bones at the site and support the importance of animal grease in the caloric intake of Middle Pleistocene humans.

The evolution of the human trophic level during the Pleistocene.

The human trophic level (HTL) during the Pleistocene and its degree of variability serve, explicitly or tacitly, as the basis of many explanations for human evolution, behavior, and culture. Previous attempts to reconstruct the HTL have relied heavily on an analogy with recent hunter-gatherer groups' diets. In addition to technological differences, recent findings of substantial ecological differences between the Pleistocene and the Anthropocene cast doubt regarding that analogy's validity. Surprisingly little systematic evolution-guided evidence served to reconstruct HTL. Here, we reconstruct the HTL during the Pleistocene by reviewing evidence for the impact of the HTL on the biological, ecological, and behavioral systems derived from various existing studies. We adapt a paleobiological and paleoecological approach, including evidence from human physiology and genetics, archaeology, paleontology, and zoology, and identified 25 sources of evidence in total. The evidence shows that the trophic level of the Homo lineage that most probably led to modern humans evolved from a low base to a high, carnivorous position during the Pleistocene, beginning with Homo habilis and peaking in Homo erectus. A reversal of that trend appears in the Upper Paleolithic, strengthening in the Mesolithic/Epipaleolithic and Neolithic, and culminating with the advent of agriculture. We conclude that it is possible to reach a credible reconstruction of the HTL without relying on a simple analogy with recent hunter-gatherers' diets. The memory of an adaptation to a trophic level that is embedded in modern humans' biology in the form of genetics, metabolism, and morphology is a fruitful line of investigation of past HTLs, whose potential we have only started to explore.

Feathers and food: Human-bird interactions at Middle Pleistocene Qesem Cave, Israel.

The presence of fast-moving small game in the Paleolithic archaeological faunal record has long been considered a key variable to assess fundamental aspects of human behavior and subsistence. Birds occupy a prominent place in this debate not only due to their small size and to the difficulties in capturing them (essentially due to their ability to fly and their elusiveness), but also due to their possible role in the symbolic array in regard to non-nutritional elements (feathers, talons, etc.) and as reflectors of complex human-world relationships. In this study, we attempt to contribute to this topic by presenting taphonomical data of bird specimens from Qesem Cave (Israel), dated between 420 and 200 ka. Human-induced damage, including cut marks, peeling and human gnawing, has been identified on wing bones of Cygnus sp., Columba sp., Corvus ruficollis and Sturnus sp. Our evidence suggests that avian exploitation was not limited to food only-either to complement the human diet or as occasional food item-but also presumably for the use of feathers. While the consumption of birds as a dietary source seems to be evident as early as the Early Pleistocene, the non-alimentary use of inedible elements, such as feathers and talons, appears to be a practice from the Middle Paleolithic onwards. We argue that the combined nutritional and symbolic use of birds is one characteristic of the new mode of adaptation practiced already by the late Lower Paleolithic Acheulo-Yabrudian hominins in the Levant starting 400 ka. The Qesem findings point to the possible emergence of new cognitive and behavioral skills, which are followed in later periods in the Old World. Finally, we discuss the possible ontological and cosmological significance of human-bird interactions to illuminate our hypothesis regarding the emergence of a new perception of human relationships with the world as an integral part of the new Acheulo-Yabrudian mode of adaptation.

Recycling for a purpose in the late Lower Paleolithic Levant: Use-wear and residue analyses of small sharp flint items indicate a planned and integrated subsistence behavior at Qesem Cave (Israel).

The purposeful production of small flakes is integral to the lithic variability of many Middle Pleistocene sites. Inhabitants of the Acheulo-Yabrudian site of Qesem Cave, Israel, systematically recycled 'old' discarded blanks and tools, using them as cores for the production of small sharp tools with distinct technological features. These recycling end-products were produced in significant quantities throughout the human occupation of Qesem Cave, and their outstanding state of preservation made possible a functional analysis with residue detection using Fourier transform infrared spectroscopy (FTIR) and energy-dispersive X-ray spectroscopy (SEM-EDX). An experimental program accompanying the study tested the efficiency of each tool category, and a reference collection of the organic remains was assembled. Our integrated results show that small flakes were used mainly to process soft to medium animal material through precise cutting activities that required accurate longitudinal motions. Several items show clear and exclusive contact with bone while others were used for designated steps in hide treatment processes. Plant and tuber processing are also evidenced although to a lesser extent. We show that the end products of recycling 'old' flakes reflect preconceived technological and functional characteristics produced in a targeted manner to obtain specific tools designated for anticipated practical tasks. We demonstrate the complementary role of the products of recycling within the Qesem Cave tool-kits alongside larger tools in assisting early humans in the different stages of processing animal materials. Moreover, use-wear and residue evidence indicates that Qesem hominins differentiated their activities across space in the cave. We argue that the meticulous realization of specific tasks and the deliberate, repetitive, and skilled production of tools of different sizes and shapes is one characteristic of the new mode of adaptation practiced by Acheulo-Yabrudian hominins in the Levant in order to better manipulate the available resources following the disappearance of mega-herbivores.

Neanderthal and Woolly Mammoth Molecular Resemblance: Genetic Similarities May Underlie Cold Adaptation Suite.

With the ongoing growth of gene-based research in recent decades, examining changes that have taken place in structures over the course of evolution has become increasingly accessible. One intriguing subject at the forefront of evolutionary research is how environmental pressures affect species evolution through epigenetic adaptation. This article presents the available molecular components of adaptation to cold environments in two extinct mammals: the woolly mammoth and the Neanderthal. These two species coexisted in similar geographic and environmental European settings during the Middle and Upper Pleistocene, and both were direct descendants of African ancestors, although both fully evolved and adapted in Europe during the Middle Pleistocene. The authors assessed the degree of resemblance between mammoth and Neanderthal genetic components by reviewing three case studies of relevant gene variants and alleles associated with cold-climate adaptation found in both genomes. Their observations present the likelihood of a molecular resemblance between the suites of cold adaptation traits in the two species.

Neandertals' large lower thorax may represent adaptation to high protein diet.

Humans are limited in their capacity to convert protein into energy. We present a hypothesis that a "bell" shaped thorax and a wide pelvis evolved in Neandertals, at least in part, as an adaptation to a high protein diet. A high protein diet created a need to house an enlarged liver and urinary system in a wider lower trunk. To test the hypothesis, we applied a model developed to identify points of nutritional stress. A ratio of obligatory dietary fat to total animal fat and protein sourced calories is calculated based on various known and estimated parameters. Stress is identified when the obligatory dietary fat ratio is higher than fat content ratios in available prey. The model predicts that during glacial winters, when carbohydrates weren't available, 74%-85% of Neandertals' caloric intake would have had to come from animal fat. Large animals contain around 50% fat calories, and their fat content is diminished during winter, so a significant stressful dietary fat deficit was identified by the model. This deficit could potentially be ameliorated by an increased capability to convert protein into energy. Given that high protein consumption is associated with larger liver and kidneys in animal models, it appears likely that the enlarged inferior section of the Neandertals thorax and possibly, in part, also his wide pelvis, represented an adaptation to provide encasement for those enlarged organs. Behavioral and evolutionary implications of the hypothesis are also discussed. Am J Phys Anthropol 160:367-378, 2016. © 2016 Wiley Periodicals, Inc.

Cooperative hunting and meat sharing 400-200 kya at Qesem Cave, Israel.

Zooarchaeological research at Qesem Cave, Israel demonstrates that large-game hunting was a regular practice by the late Lower Paleolithic period. The 400- to 200,000-year-old fallow deer assemblages from this cave provide early examples of prime-age-focused ungulate hunting, a human predator-prey relationship that has persisted into recent times. The meat diet at Qesem centered on large game and was supplemented with tortoises. These hominins hunted cooperatively, and consumption of the highest quality parts of large prey was delayed until the food could be moved to the cave and processed with the aid of blade cutting tools and fire. Delayed consumption of high-quality body parts implies that the meat was shared with other members of the group. The types of cut marks on upper limb bones indicate simple flesh removal activities only. The Qesem cut marks are both more abundant and more randomly oriented than those observed in Middle and Upper Paleolithic cases in the Levant, suggesting that more (skilled and unskilled) individuals were directly involved in cutting meat from the bones at Qesem Cave. Among recent humans, butchering of large animals normally involves a chain of focused tasks performed by one or just a few persons, and butchering guides many of the formalities of meat distribution and sharing that follow. The results from Qesem Cave raise new hypotheses about possible differences in the mechanics of meat sharing between the late Lower Paleolithic and Middle Paleolithic.

Function, life histories, and biographies of Lower Paleolithic patinated flint tools from Late Acheulian Revadim, Israel.

Flint tools exhibiting modified patinated surfaces ("double patina", or post-patination flaked items) provide a glimpse into Paleolithic lithic recycling, stone economy, and human choices. Different life cycles of such items are visually evident by the presence of fresh new modified surfaces alongside old patinated ones (according to color and texture differences). New modifications testify to a gap in time between the previous life cycle of the patinated flaked item and its new one. The aim of the current study is to reconstruct the functional properties and life cycles of a sample of modified patinated flaked tools from Late Acheulian Revadim, Israel by applying use-wear and residue analyses. The results of the functional study allow a better understanding of the practical reasoning behind the collection and recycling of old flint tools, while additional inputs from theoretical and methodological advancements assist in reconstructing their probable role in the worldviews of the site's inhabitants.

The influence of smoke density on hearth location and activity areas at Lower Paleolithic Lazaret Cave, France.

We analyze the influence of hearth location and smoke dispersal on potential activity areas at Lower Paleolithic Lazaret Cave, France, focusing on archaeostratigraphic unit UA25, where a single hearth was unearthed, and GIS and activity area analysis were performed by the excavators. We simulated smoke dispersal from 16 hypothetical hearth locations and analyzed their effect on potential working spaces. Four activity zones were defined, according to the average smoke exposure recommendations from the World Health Organization (WHO) and Environmental Protection Agency (EPA). We found that the size of the low smoke density area and its distance from the hearth are the main parameters for choosing hearth location. The simulation results show an optimal hearth location zone of about 5 × 5m2, and it is precisely in this zone that the Lower Paleolithic humans of Lazaret Cave placed their hearth. We demonstrate that the optimal hearth location zone correlates not only with the archaeological hearth in UA25 but also with the locations of hearths in other layers. In addition, our smoke density analysis confirmed the detailed GIS and activity area reconstruction conducted by the excavators, strongly reinforcing their interpretation regarding the spatial organization of human behavior at Lazaret Cave.

Late Acheulian Jaljulia - Early human occupations in the paleo-landscape of the central coastal plain of Israel.

The Lower Paleolithic Late Acheulian in the Levant marks a fascinating chapter in human cultural and biological evolution. Nevertheless, many aspects of the Late Acheulian are still undeciphered, hindered by the complex nature of each site on the one hand, a scarcity of wide, multidisciplinary studies on the other, and by difficulties in obtaining absolute chronology for this timeframe. Therefore, subjects such as human subsistence strategies and modes of adaptation, regional diversity, and the possible existence and nature of interactions between hominin groups are largely understudied. The discovery and study of Jaljulia, a large-scale Late Acheulian site at the central Coastal Plain, Israel, add valuable insights to the research of this chapter in human history. Considered to represent recurrent occupations at a favored, water and flint-rich setting, the site has provided extensive lithic assemblages obtained from several localities. Absolute chronology places the human activity on-site at roughly 500-300 ky (and possibly even later), which is suggested to be divided into several main occupation phases. Geomorphological and sedimentological analyses show a change in environmental conditions, from aeolian sand deposition and overlying Hamra soil during the Middle Pleistocene to high energy fluvial regime which transported large gravels in a north-south paleo-channel. Wetland environments, correlating to the human activity on site, developed later due to higher sea levels and a coastline shifts to the eastward, which caused a blockage of the Yarkon stream corridor to the sea by marine sand. In this paper we present results of the study of the site, including geomorphological formation and post-depositional processes, absolute chronology, lithic and faunal analyses. The site's extensive lithic assemblages are currently under study and future investigations are expected to shed more light on the technological nature of Late Acheulian Jaljulia.

Systematic blade production at late Lower Paleolithic (400-200 kyr) Qesem Cave, Israel.

Qesem Cave is assigned to the Acheulo-Yabrudian cultural complex of the late Lower Paleolithic period. The 7.5 m deep stratigraphic sequence is dated to 400-200 ka (thousands of years ago). It is mostly attributed to the Amudian blade-dominated industry, one of the earliest blade production technologies in the world. In this paper, we present the results of a detailed study of five Amudian assemblages from Qesem Cave and suggest two trajectories for the production of blades at the site. We argue that the reduction sequences of blades at Qesem Cave represent an innovative and straightforward technology aimed at the systemic and serial production of predetermined blanks. We suggest that this predetermined blank technology shows planning and intensity that is not significantly different from Middle Paleolithic Mousterian technological systems. Furthermore, this well-organized serial manufacture of cutting implements mainly for butchering might indicates that a significant change in human behavior had taken place by the late Lower Paleolithic period.

Hearth-side socioeconomics, hunting and paleoecology during the late Lower Paleolithic at Qesem Cave, Israel.

The late Lower Paleolithic archaeofaunas of Qesem Cave in the southern Levant span 400-200 ka and associate with Acheulo-Yabrudian (mainly Amudian) industries. The large mammals are exclusively Eurasian in origin and formed under relatively cool, moist conditions. The zooarchaeological findings testify to large game hunting, hearth-centered carcass processing and meat sharing during the late Lower Paleolithic, not unlike the patterns known from Middle and Upper Paleolithic caves in the region. Well-defined hearth features are rarely preserved in Qesem Cave, but the heterogeneous distributions of burned bones indicate areas of frequent hearth rebuilding throughout the occupation sequence. The hominins delayed consumption of high quality body parts until they could be moved to the cave, where hearths were hubs of processing activities and social interaction. Paradoxically, the cut marks on the Qesem bones are both more abundant and more randomly oriented than those observed in Middle and Upper Paleolithic cases in the Levant. These results suggest that several individuals were directly involved in cutting meat from the bones and that the social mechanics of meat sharing during the late Lower Paleolithic at Qesem Cave differed from those typical of both the Middle and Upper Paleolithic in the region.

Middle Pleistocene dental remains from Qesem Cave (Israel).

This study presents a description and comparative analysis of Middle Pleistocene permanent and deciduous teeth from the site of Qesem Cave (Israel). All of the human fossils are assigned to the Acheulo-Yabrudian Cultural Complex (AYCC) of the late Lower Paleolithic. The Middle Pleistocene age of the Qesem teeth (400-200 ka) places them chronologically earlier than the bulk of fossil hominin specimens previously known from southwest Asia. Three permanent mandibular teeth (C(1) -P(4) ) were found in close proximity in the lower part of the stratigraphic sequence. The small metric dimensions of the crowns indicate a considerable degree of dental reduction although the roots are long and robust. In contrast, three isolated permanent maxillary teeth (I(2) , C(1) , and M(3) ) and two isolated deciduous teeth that were found within the upper part of the sequence are much larger and show some plesiomorphous traits similar to those of the Skhul/Qafzeh specimens. Although none of the Qesem teeth shows a suite of Neanderthal characters, a few traits may suggest some affinities with members of the Neanderthal evolutionary lineage. However, the balance of the evidence suggests a closer similarity with the Skhul/Qafzeh dental material, although many of these resemblances likely represent plesiomorphous features.

An integrated study discloses chopping tools use from Late Acheulean Revadim (Israel).

Chopping tools/choppers provide one of the earliest and most persistent examples of stone tools produced and used by early humans. These artifacts appeared for the first time ~2.5 million years ago in Africa and are characteristic of the Oldowan and Acheulean cultural complexes throughout the Old World. Chopping tools were manufactured and used by early humans for more than two million years regardless of differences in geography, climate, resource availability, or major transformations in human cultural and biological evolution. Despite their widespread distribution through time and space in Africa and Eurasia, little attention has been paid to the function of these items, while scholars still debate whether they are tools or cores. In this paper, we wish to draw attention to these prominent and ubiquitous early lithic artifacts through the investigation of 53 chopping tools retrieved from a specific context at Late Acheulean Revadim (Israel). We combined typo-technological and functional studies with a residue analysis aimed at shedding light on their functional role within the tool-kits of the inhabitants of the site. Here we show that most of the chopping tools were used to chop hard and medium materials, such as bone, most probably for marrow extraction. A few of the tools were also used for cutting and scraping activities, while some also served as cores for further flake detachment. The chopping tools exhibit extraordinarily well-preserved bone residues suggesting they were used mainly for bone-breaking and marrow acquisition. We discuss the data and explore the tool versus core debate also in light of a sample of 50 flake cores made on pebbles/cobbles retrieved from the same archeological layer. The results add further pieces to the puzzle of activities carried out at Revadim and add to our knowledge of the production and use of these enigmatic tools and their role in human evolutionary history.

Shaped stone balls were used for bone marrow extraction at Lower Paleolithic Qesem Cave, Israel.

The presence of shaped stone balls at early Paleolithic sites has attracted scholarly attention since the pioneering work of the Leakeys in Olduvai, Tanzania. Despite the persistent presence of these items in the archaeological record over a period of two million years, their function is still debated. We present new results from Middle Pleistocene Qesem Cave on the use of these implements as percussion tools. Use-wear and abundant bone and fat residues found on ten shaped stone balls indicate crushing of fresh bones by thrusting percussion and provide direct evidence for the use of these items to access bone marrow of animal prey at this site. Two experiments conducted to investigate and verify functional aspects proved Qesem Cave shaped stone balls are efficient for bone processing and provide a comfortable grip and useful active areas for repeated use. Notably, the patina observed on the analyzed items precedes their use at the cave, indicating that they were collected by Qesem inhabitants, most probably from older Lower Paleolithic Acheulian sites. Thus, our results refer only to the final phases of the life of the items, and we cannot attest to their original function. As bone marrow played a central role in human nutrition in the Lower Paleolithic, and our experimental results show that the morphology and characteristics of shaped stone ball replicas are well-suited for the extraction of bone marrow, we suggest that these features might have been the reason for their collection and use at Qesem Cave. These results shed light on the function of shaped stone balls and are consistent with the significance of animal fat in the caloric intake of Middle Pleistocene humans as shown by the archeozoological evidence at Qesem Cave and possibly beyond.

Animal residues found on tiny Lower Paleolithic tools reveal their use in butchery.

Stone tools provide a unique window into the mode of adaptation and cognitive abilities of Lower Paleolithic early humans. The persistently produced large cutting tools (bifaces/handaxes) have long been an appealing focus of research in the reconstruction of Lower Paleolithic survival strategies, at the expenses of the small flake tools considered by-products of the stone production process rather than desired end products. Here, we use use-wear, residues and technological analyses to show direct and very early evidence of the deliberate production and use of small flakes for targeted stages of the prey butchery process at the late Lower Paleolithic Acheulian site of Revadim, Israel. We highlight the significant role of small flakes in Lower Paleolithic adaptation alongside the canonical large handaxes. Our results demonstrate the technological and cognitive flexibility of early human groups in the Levant and beyond at the threshold of the departure from Lower Paleolithic lifeways.

Early evidence of stone tool use in bone working activities at Qesem Cave, Israel.

For a long while, the controversy surrounding several bone tools coming from pre-Upper Palaeolithic contexts favoured the view of Homo sapiens as the only species of the genus Homo capable of modifying animal bones into specialised tools. However, evidence such as South African Early Stone Age modified bones, European Lower Palaeolithic flaked bone tools, along with Middle and Late Pleistocene bone retouchers, led to a re-evaluation of the conception of Homo sapiens as the exclusive manufacturer of specialised bone tools. The evidence presented herein include use wear and bone residues identified on two flint scrapers as well as a sawing mark on a fallow deer tibia, not associated with butchering activities. Dated to more than 300 kya, the evidence here presented is among the earliest related to tool-assisted bone working intended for non-dietary purposes, and contributes to the debate over the recognition of bone working as a much older behaviour than previously thought. The results of this study come from the application of a combined methodological approach, comprising use wear analysis, residue analysis, and taphonomy. This approach allowed for the retrieval of both direct and indirect evidence of tool-assisted bone working, at the Lower Palaeolithic site of Qesem Cave (Israel).