Evidence of diverse animal exploitation during the Middle Paleolithic at Ghar-e Boof (southern Zagros).

Although Middle Paleolithic (MP) hominin diets consisted mainly of ungulates, increasing evidence demonstrates that hominins at least occasionally consumed tortoises, birds, leporids, fish, and carnivores. Until now, the MP zooarchaeological record in the Zagros Mountains has been almost exclusively restricted to ungulates. The narrow range of hominin prey may reflect socioeconomic decisions and/or environmental constraints, but could also result from a research bias favoring the study of large prey, since archaeologists have undertaken no systematic taphonomic analyses of small game or carnivores in the region. Here, we report on the first comprehensive taphonomic analysis of an MP faunal assemblage from Ghar-e Boof (∼ 81-45 kyr), a Late Pleistocene site in the southern Zagros of Iran. Anthropogenic bone surface modifications point to hominins as the main agent of accumulation. Hominins preyed primarily on ungulates, particularly wild goat. However, we also found evidence for MP hominin exploitation of carnivores and tortoises at the site. Although small game represents only a minor portion of the diet, our results suggest that the hunting behavior of MP hominins in the Zagros was more diverse than previously thought, similar to what we find elsewhere in Eurasia.

First direct evidence of lion hunting and the early use of a lion pelt by Neanderthals.

During the Upper Paleolithic, lions become an important theme in Paleolithic art and are more frequent in anthropogenic faunal assemblages. However, the relationship between hominins and lions in earlier periods is poorly known and primarily interpreted as interspecies competition. Here we present new evidence for Neanderthal-cave lion interactions during the Middle Paleolithic. We report new evidence of hunting lesions on the 48,000 old cave lion skeleton found at Siegsdorf (Germany) that attest to the earliest direct instance of a large predator kill in human history. A comparative analysis of a partial puncture to a rib suggests that the fatal stab was delivered with a wooden thrusting spear. We also present the discovery of distal lion phalanges at least 190,000 old from Einhornhöhle (Germany), representing the earliest example of the use of cave lion skin by Neanderthals in Central Europe. Our study provides novel evidence on a new dimension of Neanderthal behavioral complexity.

Early evidence for bear exploitation during MIS 9 from the site of Schöningen 12 (Germany).

A cutmarked bear metatarsal and phalanx from the German open-air sites of Schöningen 12 II-1 and 12 B, respectively, correlated with the interglacial optimum of MIS 9 (ca. 320 ka), provide early evidence for the exploitation of bear skins. Archaeological sites with evidence of bear exploitation from the Lower Paleolithic are rare, with only Boxgrove (United Kingdom) and Bilzingsleben (Germany) yielding cutmarked bear bones indicating skinning. We interpret these finds as evidence for bear hunting and primary access since bear skins are best extracted shortly after the animal's death. The very thin cutmarks found on the Schöningen specimens indicate delicate butchering and show similarities in butchery patterns to bears from other Paleolithic sites. The Eurasian Lower Paleolithic record does not show any evidence for the exploitation of bear meat; only Middle Paleolithic sites, such as Biache-Saint-Vaast (France; ca. 175 ka) and Taubach (Germany; ca. 120 ka), yield evidence for the exploitation of both skin and meat from bear carcasses. Bear skins have high insulating properties and might have played a role in the adaptations of Middle Pleistocene hominins to the cold and harsh winter conditions of Northwestern Europe.

Integrating faunal and botanical remains using multivariate statistics to reconstruct (pre)historic subsistence developments.

Archaeological faunal and botanical remains are often treated and published separately to understand past subsistence practices. This distinction is an arbitrary one based on methodological differences, especially since we know from ethnological sources that animal husbandry and crop cultivation are usually interdependent in agricultural systems. Here, we use correspondence, detrended correspondence, and canonical correspondence analyses to integrate these different lines of evidence. We customise this method by: • Adjusting criteria to select and prepare data for integration. • Including independent parameters such as chronology and mean annual precipitation to study relationships. • Presenting additional visualisations of data to aid interpretation. The customised method we present can be applied to any time period, geographical region or research question, as long as botanical and faunal data are available. By analysing these data in an integrative way, we can improve our knowledge of subsistence and agriculture, which in turn can provide a context to better understand social and political changes in past societies.

Did climate determine Late Pleistocene settlement dynamics in the Ach Valley, SW Germany?

The loss of Neanderthal groups across Western and Central Europe during Oxygen Isotope Stage (OIS) 3 has held the attention of archaeologists for decades. The role that climatic change, genetic interbreeding, and interspecies competition played in the extinction of Neanderthal groups is still debated. Hohle Fels is one of several important Middle and Upper Paleolithic sites from the Ach Valley in southwestern Germany which documents the presence of Neanderthals and modern humans in the region. Chronological and stratigraphic records indicate that these two groups occupied the site with little to no overlap or interaction. This provides the opportunity to examine the behavioural variability of Swabian Neanderthal populations without the complication of cross-cultural influence. In this study we contribute a terrestrial paleoenvironmental record derived from the small mammal material from Hohle Fels Cave to the ever-growing archaeological record of this period. By reconstructing the climate and landscape of the Ach Valley during this time we can identify the effect that the OIS 3 environment had on the presence of Neanderthals in the region. Based on indicator taxa and the habitat weighing method, the small mammal record, which includes rodents, insectivores, and bats, from Hohle Fels shows that the earliest Neanderthal occupation took place on a landscape characterized by substantial woodland and forest, rivers and ponds, as well as moist meadows and grasslands. A gradual increase in cold tundra and arctic environments is clear towards the end of the Middle Paleolithic, extending to the end of the early Aurignacian which may correlate with the onset of the Heinrich 4 event (~42,000 kya). Our taphonomic analysis indicates the material was accumulated primarily by opportunistic predators such as the great grey owl, snowy owl, and European eagle owl, and therefore reflects the diversity of landscapes present around the site in the past. Importantly, at the time Neanderthals abandoned the Ach Valley we find no indication for dramatic climatic deterioration. Rather, we find evidence of a gradual cooling of the Swabian landscape which may have pushed Neanderthal groups out of the Ach Valley prior to the arrival of modern human Aurignacian groups.

Paleolithic subsistence strategies and changes in site use at Klissoura Cave 1 (Peloponnese, Greece).

Klissoura Cave 1 in southern Greece preserves a long archaeological sequence that spans roughly 90,000 years and includes Middle Paleolithic, Uluzzian, Upper Paleolithic, and Mesolithic deposits. The site provides a unique opportunity to examine diachronic change and shifts in the intensity of site use across the Late Pleistocene. There is an overall picture of the intensified use of faunal resources at the site, evidenced by a shift from large to small game, and to small fast-moving taxa in particular. This trend is independent of climatic change and fluctuations in site use, and most likely reflects a broader, regional growth of hominin populations. At the same time, multiple lines of evidence (e.g., input of artifacts and features, sedimentation mechanisms, and intensification of faunal resources) indicate that the intensity of site use changed, with a sharp increase from the Middle Paleolithic to Aurignacian. This allows us to address a fundamental issue in the study of human evolution: differences in population size and site use between Neandertals and modern humans. At Klissoura Cave 1, the increase in occupation intensity might be related to population growth or larger group size, but it might also be due to changes in season of site use, more favorable environmental conditions at the beginning of the Upper Paleolithic, and/or changes in the composition of people occupying the site. These explanations are not necessarily mutually exclusive, and indeed the data support a combination of factors. Ascribing the increase in occupation intensity to larger Upper Paleolithic populations more broadly is difficult, particularly because there is little consensus on this topic elsewhere in Eurasia. The data are complicated and vary greatly between sites and regions. This makes Klissoura Cave 1, as the only currently available case study in southeastern Europe, a critical example in understanding the range of variation in demography and site use across the Middle to Upper Paleolithic transition.

New cranium of the large cercopithecid primate Theropithecus oswaldi leakeyi (Hopwood, 1934) from the paleoanthropological site of Makuyuni, Tanzania.

The Pleistocene hominin site of Makuyuni, near Lake Manyara, Tanzania, is known for fossils attributable to Homo and Acheulean artifacts (Ring et al., 2005; Kaiser et al., 2010; Frost et al., 2012). Here we describe the fossil primate material from the Manyara Beds, which includes the first nearly complete female cranium of Theropithecus oswaldi leakeyi and a proximal tibia from the same taxon. The cranium is dated to between 633 and 780 Ka and the tibia to the Pleistocene. The T. oswaldi lineage is one of the most important among Neogene mammals of Africa: it is both widespread and abundant. The size of the dentition, cranium, and tibia all confirm the previously recognized trend of increasing body size in this lineage and make their taxonomic assignments secure. The morphology of this specimen provides new insights into the evolution of this lineage through time, as well as its geographic variation and sexual dimorphism. The cranium also shows damage consistent with a mammalian carnivore, most likely a felid. The identification of this material as representing T. o. leakeyi agrees with the Middle Pleistocene age estimates for the MK4 locality in particular and the Manyara Beds in general.

Excavations at Schöningen and paradigm shifts in human evolution.

The exceptional preservation at Schöningen together with a mixture of perseverance, hard work, and sheer luck led to the recovery of unique finds in an exceptional context. The 1995 discovery of numerous wooden artifacts, most notably at least 10 carefully made spears together with the skeletons of at least 20 to 25 butchered horses, brought the debate about hunting versus scavenging among late archaic hominins and analogous arguments about the purportedly primitive behavior of Homo heidelbergensis and Neanderthals to an end. Work under H. Thieme's lead from 1992 to 2008 and results from the current team since 2008 demonstrate that late H. heidelbergensis or early Neanderthals used sophisticated artifacts made from floral and faunal materials, in addition to lithic artifacts more typically recovered at Lower Paleolithic sites. The finds from the famous Horse Butchery Site and two dozen other archaeological horizons from the edges of the open-cast mine at Schöningen provide many new insights into the technology and behavioral patterns of hominins about 300 ka BP during MIS 9 on the Northern European Plain. An analysis of the finds from Schöningen and their contexts shows that the inhabitants of the site were skilled hunters at the top of the food chain and exhibited a high level of planning depth. These hominins had command of effective means of communication about the here and now, and the past and the future, that allowed them to repeatedly execute well-coordinated and successful group activities that likely culminated in a division of labor and social and economic patterns radically different from those of all non-human primates. The unique preservation and high quality excavations have led to a major paradigm shift or "Schöningen Effect" that changed our views of human evolution during the late Lower Paleolithic. In this respect, we can view the behaviors documented at Schöningen as a plausible baseline for the behavioral sophistication of archaic hominins of the late Middle Pleistocene and subsequent periods.

Bone taphonomy of the Schöningen "Spear Horizon South" and its implications for site formation and hominin meat provisioning.

This paper presents the faunal remains from the new excavation area at the Lower Paleolithic site of Schöningen. The focus of the study is on the southern extension of the main find horizon (Spear Horizon South), which includes the layer that yielded the famous Schöningen spears (13 II-4). Taxonomic data corroborate previous studies, that hominins primarily hunted Equus mosbachensis, a large Pleistocene horse. Equid body part representation at the site suggests that the animals were hunted and butchered locally. There is no evidence for density-mediated attrition in the assemblage. Weathering damage is uncommon, though there is ample evidence that carnivores had access to the bone. Carnivore bite sizes were measured and compared to experimental data provided by previous authors. Based on relationships between bite size and carnivore behavior and body size, we conclude that the primary modifying agents were large carnivores (i.e., wolves or saber-toothed cats). Previous studies show that carnivores often had secondary access to the remains, after hominins. Cut marks are commonly arranged haphazardly on the bones. This may indicate that multiple hominins participated in the butchery of horse skeletons, or that they were butchered over the course of hours or days. Cut marks on axial elements are more "orderly," which probably reflects the physical logistics of orienting one's body in relation to a large carcass. These data differ from sites formed by Middle and Upper Paleolithic hominins, which might suggest that in later times, a system of organized meat provisioning was already in place. Taken together, the faunal evidence from the Spear Horizon South indicates that late Lower Paleolithic hominins using the site understood the behaviors of different prey species, hunted socially to take down large game, and successfully competed with large carnivores on the landscape for primary access to ungulate remains.

The European saber-toothed cat (Homotherium latidens) found in the "Spear Horizon" at Schöningen (Germany).

The 300,000 year old Lower Paleolithic site Schöningen 13 II-4 became world famous with the discovery of the oldest well-preserved and complete wooden spears. Through ongoing excavations, new archaeological discoveries of scientific importance are still being made from the same archaeological layer where the spears were found. In this context, remains of a rare carnivore species, the European saber-toothed cat (Homotherium latidens), were recovered. Here we present five teeth and one humerus fragment that are unambiguously from two individual saber-toothed cats. The humerus is a unique specimen; it shows evidence of hominin impacts and use as a percussor. The Homotherium remains from Schöningen are the best documented finds of this species in an archaeological setting and they are amongst the youngest specimens of Homotherium in Europe. The presence of this species as a carnivore competitor would certainly have impacted the lives of late Middle Pleistocene hominins. The discovery illustrates the possible day-to-day challenges that the Schöningen hominins would have faced and suggests that the wooden spears were not necessarily only used for hunting, but possibly also as a weapon for self-defense.