Widespread evidence for elephant exploitation by Last Interglacial Neanderthals on the North European plain.

Neanderthals hunted and butchered straight-tusked elephants, the largest terrestrial mammals of the Pleistocene, in a lake landscape on the North European plain, 125,000 years ago, as recently shown by a study of the Last Interglacial elephant assemblage from Neumark-Nord (Germany). With evidence for a remarkable focus on adult males and on their extended utilization, the data from this location are thus far without parallel in the archaeological record. Given their relevance for our knowledge of the Neanderthal niche, we investigated whether the Neumark-Nord subsistence practices were more than a local phenomenon, possibly determined by local characteristics. Analyzing elephant remains from two other Last Interglacial archaeological sites on the North European plain, Gröbern and Taubach, we identified in both assemblages similar butchering patterns as at Neumark-Nord, demonstrating that extended elephant exploitation was a widespread Neanderthal practice during the (early part of the) Last Interglacial. The substantial efforts needed to process these animals, weighing up to 13 metric tons, and the large amounts of food generated suggest that Neanderthals either had ways of storing vast amounts of meat and fat and/or temporarily aggregated in larger groups than commonly acknowledged. The data do not allow us to rule out one of the two explanations, and furthermore both factors, short-term larger group sizes as well as some form of food preservation, may have played a role. What the data do show is that exploitation of large straight-tusked elephants was a widespread and recurring phenomenon amongst Last Interglacial Neanderthals on the North European plain.

Beaver exploitation, 400,000 years ago, testifies to prey choice diversity of Middle Pleistocene hominins.

Data regarding the subsistence base of early hominins are heavily biased in favor of the animal component of their diets, in particular the remains of large mammals, which are generally much better preserved at archaeological sites than the bones of smaller animals, let alone the remains of plant food. Exploitation of smaller game is very rarely documented before the latest phases of the Pleistocene, which is often taken to imply narrow diets of archaic Homo and interpreted as a striking economic difference between Late Pleistocene and earlier hominins. We present new data that contradict this view of Middle Pleistocene Lower Palaeolithic hominins: cut mark evidence demonstrating systematic exploitation of beavers, identified in the large faunal assemblage from the c. 400,000 years old hominin site Bilzingsleben, in central Germany. In combination with a prime-age dominated mortality profile, this cut mark record shows that the rich beaver assemblage resulted from repetitive human hunting activities, with a focus on young adult individuals. The Bilzingsleben beaver exploitation evidence demonstrates a greater diversity of prey choice by Middle Pleistocene hominins than commonly acknowledged, and a much deeper history of broad-spectrum subsistence than commonly assumed, already visible in prey choices 400,000 years ago.

Hunting and processing of straight-tusked elephants 125.000 years ago: Implications for Neanderthal behavior.

Straight-tusked elephants (Palaeoloxodon antiquus) were the largest terrestrial mammals of the Pleistocene, present in Eurasian landscapes between 800,000 and 100,000 years ago. The occasional co-occurrence of their skeletal remains with stone tools has generated rich speculation about the nature of interactions between these elephants and Pleistocene humans: Did hominins scavenge on elephants that died a natural death or maybe even hunt some individuals? Our archaeozoological study of the largest P. antiquus assemblage known, excavated from 125,000-year-old lake deposits in Germany, shows that hunting of elephants weighing up to 13 metric tons was part of the cultural repertoire of Last Interglacial Neanderthals there, over >2000 years, many dozens of generations. The intensity and nutritional yields of these well-documented butchering activities, combined with previously reported data from this Neumark-Nord site complex, suggest that Neanderthals were less mobile and operated within social units substantially larger than commonly envisaged.

Landscape modification by Last Interglacial Neanderthals.

Little is known about the antiquity, nature, and scale of Pleistocene hunter-gatherer impact on their ecosystems, despite the importance for studies of conservation and human evolution. Such impact is likely to be limited, mainly because of low population densities, and challenging to detect and interpret in terms of cause-effect dynamics. We present high-resolution paleoenvironmental and archaeological data from the Last Interglacial locality of Neumark-Nord (Germany). Among the factors that shaped vegetation structure and succession in this lake landscape, we identify a distinct ecological footprint of hominin activities, including fire use. We compare these data with evidence from archaeological and baseline sites from the same region. At Neumark-Nord, notably open vegetation coincides with a virtually continuous c. 2000-year-long hominin presence, and the comparative data strongly suggest that hominins were a contributing factor. With an age of c. 125,000 years, Neumark-Nord provides an early example of a hominin role in vegetation transformation.

Insights into the palaeobiology of an early Homo infant: multidisciplinary investigation of the GAR IVE hemi-mandible, Melka Kunture, Ethiopia.

Childhood is an ontogenetic stage unique to the modern human life history pattern. It enables the still dependent infants to achieve an extended rapid brain growth, slow somatic maturation, while benefitting from provisioning, transitional feeding, and protection from other group members. This tipping point in the evolution of human ontogeny likely emerged from early Homo. The GAR IVE hemi-mandible (1.8 Ma, Melka Kunture, Ethiopia) represents one of the rarely preserved early Homo infants (~ 3 years at death), recovered in a richly documented Oldowan archaeological context. Yet, based on the sole external inspection of its teeth, GAR IVE was diagnosed with a rare genetic disease-amelogenesis imperfecta (AI)-altering enamel. Since it may have impacted the child's survival, this diagnosis deserves deeper examination. Here, we reassess and refute this diagnosis and all associated interpretations, using an unprecedented multidisciplinary approach combining an in-depth analysis of GAR IVE (synchrotron imaging) and associated fauna. Some of the traits previously considered as diagnostic of AI can be better explained by normal growth or taphonomy, which calls for caution when diagnosing pathologies on fossils. We compare GAR IVE's dental development to other fossil hominins, and discuss the implications for the emergence of childhood in early Homo.

A detailed analysis of the spatial distribution of Schöningen 13II-4 'Spear Horizon' faunal remains.

The Middle Pleistocene Schöningen 13II-4 'Spear Horizon' (Germany) is a key site for the study of human evolution, most notably for the discovery of Paleolithic wooden weaponry and evidence for developed hunting strategies. On the other hand, the 'Spear Horizon' offers an excellent opportunity to approach hominin spatial behavior, thanks to the richness of the archeological assemblage, its exceptional preservation, and the vast expanse of the excavated surface. Analyzing how space was used is essential for understanding hominin behavior at this unique open-air site and, from a wider perspective, for approaching how humans adapted to interglacial environments. In this article, we present an exhaustive spatial study of the complete Schöningen 13II-4 'Spear Horizon' faunal assemblage and its archeological context, combining zooarcheology and spatial analysis through the extensive application of geographic information systems. Our results indicate the existence of different activity areas related to changes in the position of the shoreline due to fluctuations of water table levels of the Schöningen paleolake. These activity areas were likely used on a seasonal basis, whereas the spatial patterning observed in the distribution of faunal remains suggests a diversity of behavioral strategies in terms of intensity and/or duration of occupations. This study refines previous interpretations of the site and reconstructs human behavioral adaptations and the occupational changing lakeland environment during the Middle Pleistocene in Europe.

Stable isotopes reveal patterns of diet and mobility in the last Neandertals and first modern humans in Europe.

Correlating cultural, technological and ecological aspects of both Upper Pleistocene modern humans (UPMHs) and Neandertals provides a useful approach for achieving robust predictions about what makes us human. Here we present ecological information for a period of special relevance in human evolution, the time of replacement of Neandertals by modern humans during the Late Pleistocene in Europe. Using the stable isotopic approach, we shed light on aspects of diet and mobility of the late Neandertals and UPMHs from the cave sites of the Troisième caverne of Goyet and Spy in Belgium. We demonstrate that their diet was essentially similar, relying on the same terrestrial herbivores, whereas mobility strategies indicate considerable differences between Neandertal groups, as well as in comparison to UPMHs. Our results indicate that UPMHs exploited their environment to a greater extent than Neandertals and support the hypothesis that UPMHs had a substantial impact not only on the population dynamics of large mammals but also on the whole structure of the ecosystem since their initial arrival in Europe.

Subsistence strategies throughout the African Middle Pleistocene: Faunal evidence for behavioral change and continuity across the Earlier to Middle Stone Age transition.

The African Middle Pleistocene (781-126 ka) is a key period for human evolution, witnessing both the origin of the modern human lineage and the lithic turnover from Earlier Stone Age (ESA) Acheulean bifacial tools to Middle Stone Age (MSA) prepared core and point technologies. This ESA/MSA transition is interpreted as representing changing landscape use with greater foraging distances and more active hunting strategies. So far, these behavioral inferences are mainly based on the extensive stone tool record, with only a minor role for site-based and regional faunal studies. To provide additional insights into these behavioral changes, this paper details a pan-African metastudy of 63 Middle Pleistocene faunal assemblages from 40 sites. A hierarchical classification system identified 26 well-contextualized assemblages with quantitative paleontological and/or zooarcheological data available for detailed comparative analyses and generalized linear mixed modeling. Modeling of ungulate body size classes structured around three dimensions (context, antiquity and technology) illustrates no one-to-one correlation between changes in lithic technology (Acheulean vs. MSA) and changes in prey representation. All assessed faunal assemblages are dominated by medium-sized bovids, and variations between smaller and larger body size classes are linked to site context (cave vs. open-air), with an increase in cave sites during the Middle Pleistocene. Current data do not signal a broadening of the hominin dietary niche during the Middle Pleistocene; no meaningful variation was visible in the exploitation of smaller-sized bovids or dangerous game, with coastal resources exploited when available. Proportions of anthropogenic bone surface modifications, and hence carcass processing intensity, do increase over time although more zooarcheological data is crucial before making behavioral inferences. Overall, this paper illustrates the potential of broad scale comparative faunal analyses to provide additional insights into processes of human behavioral evolution and the mechanisms underlying patterns of technological, chronological and contextual change.

Evidence for close-range hunting by last interglacial Neanderthals.

Animal resources have been part of hominin diets since around 2.5 million years ago, with sharp-edged stone tools facilitating access to carcasses. How exactly hominins acquired animal prey and how hunting strategies varied through time and space is far from clear. The oldest possible hunting weapons known from the archaeological record are 300,000 to 400,000-year-old sharpened wooden staves. These may have been used as throwing and/or close-range thrusting spears, but actual data on how such objects were used are lacking, as unambiguous lesions caused by such weapon-like objects are unknown for most of human prehistory. Here, we report perforations observed on two fallow deer skeletons from Neumark-Nord, Germany, retrieved during excavations of 120,000-year-old lake shore deposits with abundant traces of Neanderthal presence. Detailed studies of the perforations, including micro-computed tomography imaging and ballistic experiments, demonstrate that they resulted from the close-range use of thrusting spears. Such confrontational ways of hunting require close cooperation between participants, and over time may have shaped important aspects of hominin biology and behaviour.

Archaeology and ichnology at Gombore II-2, Melka Kunture, Ethiopia: everyday life of a mixed-age hominin group 700,000 years ago.

We report the occurrence at 0.7 million years (Ma) of an ichnological assemblage at Gombore II-2, which is one of several archaeological sites at Melka Kunture in the upper Awash Valley of Ethiopia, 2000 m asl. Adults and children potentially as young as 12 months old left tracks in a silty substrate on the shore of a body of water where ungulates, as well as other mammals and birds, congregated. Furthermore, the same layers contain a rich archaeological and palaeontological record, confirming that knapping was taking place in situ and that stone tools were used for butchering hippo carcasses at the site. The site gives direct information on hominin landscape use at 0.7 Ma and may provide fresh perspective on the childhood of our ancestors.

Middle Pleistocene protein sequences from the rhinoceros genus Stephanorhinus and the phylogeny of extant and extinct Middle/Late Pleistocene Rhinocerotidae.

BACKGROUND: Ancient protein sequences are increasingly used to elucidate the phylogenetic relationships between extinct and extant mammalian taxa. Here, we apply these recent developments to Middle Pleistocene bone specimens of the rhinoceros genus Stephanorhinus. No biomolecular sequence data is currently available for this genus, leaving phylogenetic hypotheses on its evolutionary relationships to extant and extinct rhinoceroses untested. Furthermore, recent phylogenies based on Rhinocerotidae (partial or complete) mitochondrial DNA sequences differ in the placement of the Sumatran rhinoceros (Dicerorhinus sumatrensis). Therefore, studies utilising ancient protein sequences from Middle Pleistocene contexts have the potential to provide further insights into the phylogenetic relationships between extant and extinct species, including Stephanorhinus and Dicerorhinus. METHODS: ZooMS screening (zooarchaeology by mass spectrometry) was performed on several Late and Middle Pleistocene specimens from the genus Stephanorhinus, subsequently followed by liquid chromatography-tandem mass spectrometry (LC-MS/MS) to obtain ancient protein sequences from a Middle Pleistocene Stephanorhinus specimen. We performed parallel analysis on a Late Pleistocene woolly rhinoceros specimen and extant species of rhinoceroses, resulting in the availability of protein sequence data for five extant species and two extinct genera. Phylogenetic analysis additionally included all extant Perissodactyla genera (Equus, Tapirus), and was conducted using Bayesian (MrBayes) and maximum-likelihood (RAxML) methods. RESULTS: Various ancient proteins were identified in both the Middle and Late Pleistocene rhinoceros samples. Protein degradation and proteome complexity are consistent with an endogenous origin of the identified proteins. Phylogenetic analysis of informative proteins resolved the Perissodactyla phylogeny in agreement with previous studies in regards to the placement of the families Equidae, Tapiridae, and Rhinocerotidae. Stephanorhinus is shown to be most closely related to the genera Coelodonta and Dicerorhinus. The protein sequence data further places the Sumatran rhino in a clade together with the genus Rhinoceros, opposed to forming a clade with the black and white rhinoceros species. DISCUSSION: The first biomolecular dataset available for Stephanorhinus places this genus together with the extinct genus Coelodonta and the extant genus Dicerorhinus. This is in agreement with morphological studies, although we are unable to resolve the order of divergence between these genera based on the protein sequences available. Our data supports the placement of the genus Dicerorhinus in a clade together with extant Rhinoceros species. Finally, the availability of protein sequence data for both extinct European rhinoceros genera allows future investigations into their geographic distribution and extinction chronologies.

Direct terrestrial-marine correlation demonstrates surprisingly late onset of the last interglacial in central Europe.

An interdisciplinary study of a small sedimentary basin at Neumark Nord 2 (NN2), Germany, has yielded a high-resolution record of the palaeomagnetic Blake Event, which we are able to place at the early part of the last interglacial pollen sequence documented from the same section. We use this data to calculate the duration of this stratigraphically important event at 3400 ± 350 yr. More importantly, the Neumark Nord 2 data enables precise terrestrial-marine correlation for the Eemian stage in central Europe. This shows a remarkably large time lag of ca. 5000 yr between the MIS 5e 'peak' in the marine record and the start of the last interglacial in this region.

Systematic butchering of fallow deer (Dama) at the early middle Pleistocene Acheulian site of Gesher Benot Ya'aqov (Israel).

Three assemblages of fallow deer (Dama sp.) bones excavated from the early middle Pleistocene (oxygen isotope stage 18) layers of the Acheulian site of Gesher Benot Ya'aqov, Israel, furnish evidence of systematic and repeated exploitation of complete carcasses by hominins. The excellent state of preservation of the bones and the presence of only minimal signs of carnivore involvement permit an investigation of the role of hominins as the primary agents responsible for the damage to these bones. Hominin expertise in dealing with fallow deer carcasses is manifested by cut marks, percussion marks, and hack marks on the bones. The archaeozoological analysis of the anatomical position and frequency of these marks suggests that carcass processing followed systematic practices that reflect an in-depth knowledge of fallow deer anatomy and a consistent behavioral strategy. These assemblages represent one of the earliest examples of methodological butchering practices in Eurasia. The evidence of carcass processing observed at Gesher Benot Ya'aqov resembles that seen in late Pleistocene sites in Israel, which were inhabited by modern humans. We interpret the Gesher Benot Ya'aqov data as indicating that the Acheulian hunters at the site (1) were proficient communicators and learners and (2) possessed anatomical knowledge, considerable manual skill, impressive technological abilities, and foresight.