Persistent predators: Zooarchaeological evidence for specialized horse hunting at Schöningen 13II-4.

The Schöningen 13II-4 site is a marvel of Paleolithic archaeology. With the extraordinary preservation of complete wooden spears and butchered large mammal bones dating from the Middle Pleistocene, Schöningen maintains a prominent position in the halls of human origins worldwide. Here, we present the first analysis of the complete large mammal faunal assemblage from Schöningen 13II-4, drawing on multiple lines of zooarchaeological and taphonomic evidence to expose the full spectrum of hominin activities at the site-before, during, and after the hunt. Horse (Equus mosbachensis) remains dominate the assemblage and suggest a recurrent ambush hunting strategy along the margins of the Schöningen paleo-lake. In this regard, Schöningen 13II-4 provides the first undisputed evidence for hunting of a single prey species that can be studied from an in situ, open-air context. The Schöningen hominins likely relied on cooperative hunting strategy to target horse family groups, to the near exclusion of bachelor herds. Horse kills occurred during all seasons, implying a year-round presence of hominins on the Schöningen landscape. All portions of prey skeletons are represented in the assemblage, many complete and in semiarticulation, with little transport of skeletal parts away from the site. Butchery marks are abundant, and adult carcasses were processed more thoroughly than were juveniles. Numerous complete, unmodified bones indicated that lean meat and marrow were not always so highly prized, especially in events involving multiple kills when fat and animal hides may have received greater attention. The behaviors displayed at Schöningen continue to challenge our perceptions and models of past hominin lifeways, further cementing Schöningen's standing as the archetype for understanding hunting adaptations during the European Middle Pleistocene.

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.

The timing and spatiotemporal patterning of Neanderthal disappearance.

The timing of Neanderthal disappearance and the extent to which they overlapped with the earliest incoming anatomically modern humans (AMHs) in Eurasia are key questions in palaeoanthropology. Determining the spatiotemporal relationship between the two populations is crucial if we are to understand the processes, timing and reasons leading to the disappearance of Neanderthals and the likelihood of cultural and genetic exchange. Serious technical challenges, however, have hindered reliable dating of the period, as the radiocarbon method reaches its limit at ∼50,000 years ago. Here we apply improved accelerator mass spectrometry (14)C techniques to construct robust chronologies from 40 key Mousterian and Neanderthal archaeological sites, ranging from Russia to Spain. Bayesian age modelling was used to generate probability distribution functions to determine the latest appearance date. We show that the Mousterian ended by 41,030-39,260 calibrated years bp (at 95.4% probability) across Europe. We also demonstrate that succeeding 'transitional' archaeological industries, one of which has been linked with Neanderthals (Châtelperronian), end at a similar time. Our data indicate that the disappearance of Neanderthals occurred at different times in different regions. Comparing the data with results obtained from the earliest dated AMH sites in Europe, associated with the Uluzzian technocomplex, allows us to quantify the temporal overlap between the two human groups. The results reveal a significant overlap of 2,600-5,400 years (at 95.4% probability). This has important implications for models seeking to explain the cultural, technological and biological elements involved in the replacement of Neanderthals by AMHs. A mosaic of populations in Europe during the Middle to Upper Palaeolithic transition suggests that there was ample time for the transmission of cultural and symbolic behaviours, as well as possible genetic exchanges, between the two groups.

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.