Rope making in the Aurignacian of Central Europe more than 35,000 years ago.

Evidence for the manufacture and use of fiber technology such as rope and twine is rare in the Paleolithic, despite the widely held view that such artifacts were in regular use during the Pleistocene. On the basis of the discovery of a more than 35,000-year-old perforated baton made from mammoth ivory at Hohle Fels Cave in Ach Valley of southwestern Germany together with experimental studies, we are now able to demonstrate one way people of the early Upper Paleolithic manufactured rope. This work contributes to our understanding of the evolution of technology, cooperative work, and Paleolithic social organization.

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.

The ROCEEH Out of Africa Database (ROAD): A large-scale research database serves as an indispensable tool for human evolutionary studies.

Large scale databases are critical for helping scientists decipher long-term patterns in human evolution. This paper describes the conception and development of such a research database and illustrates how big data can be harnessed to formulate new ideas about the past. The Role of Culture in Early Expansions of Humans (ROCEEH) is a transdisciplinary research center whose aim is to study the origins of culture and the multifaceted aspects of human expansions across Africa and Eurasia over the last three million years. To support its research, the ROCEEH team developed an online tool named the ROCEEH Out of Africa Database (ROAD) and implemented its web-based applications. ROAD integrates geographical data as well as archaeological, paleoanthropological, paleontological and paleobotanical content within a robust chronological framework. In fact, a unique feature of ROAD is its ability to dynamically link scientific data both spatially and temporally, thereby allowing its reuse in ways that were not originally conceived. The data stem from published sources spanning the last 150 years, including those generated by the research team. Descriptions of these data rely on the development of a standardized vocabulary and profit from online explanations of each table and attribute. By synthesizing legacy data, ROAD facilitates the reuse of heritage data in novel ways. Database queries yield structured information in a variety of interoperable formats. By visualizing data on maps, users can explore this vast dataset and develop their own theories. By downloading data, users can conduct further quantitative analyses, for example with Geographic Information Systems, modeling programs and artificial intelligence. In this paper, we demonstrate the innovative nature of ROAD and show how it helps scientists studying human evolution to access datasets from different fields, thereby connecting the social and natural sciences. Because it permits the reuse of "old" data in new ways, ROAD is now an indispensable tool for researchers of human evolution and paleogeography.

Ancient DNA of narrow-headed vole reveal common features of the Late Pleistocene population dynamics in cold-adapted small mammals.

The narrow-headed vole, collared lemming and common vole were the most abundant small mammal species across the Eurasian Late Pleistocene steppe-tundra environment. Previous ancient DNA studies of the collared lemming and common vole have revealed dynamic population histories shaped by climatic fluctuations. To investigate the extent to which species with similar adaptations share common evolutionary histories, we generated a dataset comprised the mitochondrial genomes of 139 ancient and 6 modern narrow-headed voles from several sites across Europe and northwestern Asia covering approximately the last 100 thousand years (kyr). We inferred Bayesian time-aware phylogenies using 11 radiocarbon-dated samples to calibrate the molecular clock. Divergence of the main mtDNA lineages across the three species occurred during marine isotope stages (MIS) 7 and MIS 5, suggesting a common response of species adapted to open habitat during interglacials. We identified several time-structured mtDNA lineages in European narrow-headed vole, suggesting lineage turnover. The timing of some of these turnovers was synchronous across the three species, allowing us to identify the main drivers of the Late Pleistocene dynamics of steppe- and cold-adapted species.

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.

Middle and Upper Paleolithic occupations of Fumane Cave (Italy): a geoarchaeological investigation of the anthropogenic features.

Here we present the results of a microcontextual analysis of purported combustion features recovered from Middle and Upper Paleolithic occupations at the cave site of Fumane, Italy. Our analyses, which integrate micromorphology with organic petrology, show that only a few of the features represent primary, intact hearths; some of them show evidence for various phases of anthropogenic reworking, either through trampling or sweeping and dumping. Several of the features are multi-layered and reflect a complex formation history of various activities related to combustion and site maintenance. Many appear to be the remnants of occupation horizons only partially preserved and peripherally related to combustion. Within several of the intact hearths from the Mousterian, we were able to identify variable fuel sources in different features, implying a degree of flexibility in the fuel-selection strategies of the Neanderthal occupants of Fumane. In this study we design a classification system of the anthropogenic features and also conduct a spatial analysis, through which we can infer diachronic patterns in the frequency and intensity of site occupation and the spatial distribution of activities. We note a decrease in frequency of combustion features throughout the Mousterian which continues into the Uluzzian. The features associated with the Protoaurignacian occupation, in contrast with those from the Mousterian, are multi-layered and well-defined. We argue that these trends, which correspond with other trends in artefact frequency, imply changes in the settlement dynamics of the site during the transition from the last Neanderthal occupation of the cave to the arrival of modern humans.

Using microartifacts to infer Middle Pleistocene lifeways at Schöningen, Germany.

While archeologists usually favor the study of large and diagnostic lithic artifacts, this study illustrates the invaluable contribution of lithic microartifacts for interpreting hominin lifeways. Across a 64 m2 area of the Middle Pleistocene lakeshore site of Schöningen 13 II-3 in Northern Germany, we recovered a total of 57 small and micro flint artifacts, four small debris pieces, three natural fragments and three bone retouchers in close association with the skeleton of an extinct Eurasian straight-tusked elephant (Palaeoloxodon antiquus). This area lacks the type of formal knapped stone tools that would normally constitute the focus of archeological interpretations. By adopting a holistic approach, including morpho-technical analysis, experimental archeology, and use-wear and residue analyses, we demonstrate that these small and microartifacts are resharpening flakes that tell the story of the site. Fifteen resharpening flakes preserve microwear traces of processing wood. Microscopic residues of wood adhered to the former working edges of the tools corroborate this observation. Additionally, hominins used a sharp-edged, natural fragment of flint to process fresh animal tissue, which likely originates from the butchery of the elephant. These results provide unique, 300,000-year-old evidence for the functionally interconnected use of lithic, osseous and wood technologies. Furthermore, we document in-situ transformations of stone tools and the presence of both curational and expedient behaviors, thereby demonstrating the temporal depth of hominin activities at the lakeshore where the elephant died, and in the broader landscape as a whole.

The origin of chert in the Aurignacian of Vogelherd Cave investigated by infrared spectroscopy.

The analyses of raw material provenance offers the possibility of tracing short and long-distance raw material transport. So far, most studies of raw material of flint and chert in Europe have been based on macroscopic analyses. We apply infrared spectroscopy to Aurignacian assemblages from Vogelherd cave and to the Magdalenian site Randecker Maar in southwestern Germany. We compare raw material samples from three chert-bearing areas in Germany with archaeological samples from Vogelherd. Our results show that infrared spectroscopy can distinguish between different raw materials. Our archaeological samples from Vogelherd correspond to the sampled geological cherts in terms of their spectral signature. Our comparison of reference samples and archaeological samples highlights problems in commonly used macroscopic identifications of chert raw materials.

Grey wolf genomic history reveals a dual ancestry of dogs.

The grey wolf (Canis lupus) was the first species to give rise to a domestic population, and they remained widespread throughout the last Ice Age when many other large mammal species went extinct. Little is known, however, about the history and possible extinction of past wolf populations or when and where the wolf progenitors of the present-day dog lineage (Canis familiaris) lived1-8. Here we analysed 72 ancient wolf genomes spanning the last 100,000 years from Europe, Siberia and North America. We found that wolf populations were highly connected throughout the Late Pleistocene, with levels of differentiation an order of magnitude lower than they are today. This population connectivity allowed us to detect natural selection across the time series, including rapid fixation of mutations in the gene IFT88 40,000-30,000 years ago. We show that dogs are overall more closely related to ancient wolves from eastern Eurasia than to those from western Eurasia, suggesting a domestication process in the east. However, we also found that dogs in the Near East and Africa derive up to half of their ancestry from a distinct population related to modern southwest Eurasian wolves, reflecting either an independent domestication process or admixture from local wolves. None of the analysed ancient wolf genomes is a direct match for either of these dog ancestries, meaning that the exact progenitor populations remain to be located.

Microstratigraphic preservation of ancient faunal and hominin DNA in Pleistocene cave sediments.

Ancient DNA recovered from Pleistocene sediments represents a rich resource for the study of past hominin and environmental diversity. However, little is known about how DNA is preserved in sediments and the extent to which it may be translocated between archaeological strata. Here, we investigate DNA preservation in 47 blocks of resin-impregnated archaeological sediment collected over the last four decades for micromorphological analyses at 13 prehistoric sites in Europe, Asia, Africa, and North America and show that such blocks can preserve DNA of hominins and other mammals. Extensive microsampling of sediment blocks from Denisova Cave in the Altai Mountains reveals that the taxonomic composition of mammalian DNA differs drastically at the millimeter-scale and that DNA is concentrated in small particles, especially in fragments of bone and feces (coprolites), suggesting that these are substantial sources of DNA in sediments. Three microsamples taken in close proximity in one of the blocks yielded Neanderthal DNA from at least two male individuals closely related to Denisova 5, a Neanderthal toe bone previously recovered from the same layer. Our work indicates that DNA can remain stably localized in sediments over time and provides a means of linking genetic information to the archaeological and ecological records on a microstratigraphic scale.

A refined proposal for the origin of dogs: the case study of Gnirshöhle, a Magdalenian cave site.

Dogs are known to be the oldest animals domesticated by humans. Although many studies have examined wolf domestication, the geographic and temporal origin of this process is still being debated. To address this issue, our study sheds new light on the early stages of wolf domestication during the Magdalenian period (16-14 ka cal BP) in the Hegau Jura region (Southwestern Germany and Switzerland). By combining morphology, genetics, and isotopes, our multidisciplinary approach helps to evaluate alternate processes driving the early phases of domestication. The isotope analysis uncovered a restricted, low δ15N protein diet for all analyzed Gnirshöhle specimens, while morphological examinations and phylogenetic relationships did not unequivocally assign them to one or the other canid lineage. Intriguingly, the newly generated mitochondrial canid genomes span the entire genetic diversity of modern dogs and wolves. Such high mitochondrial diversity could imply that Magdalenian people tamed and reared animals originating from different wolf lineages. We discuss our results in light of three ecological hypotheses and conclude that both domestication and the existence of a specialized wolf ecomorph are highly probable. However, due to their proximity to humans and a restricted diet, we propose domestication as the most likely scenario explaining the patterns observed herein.

Ecosystem engineering in the Quaternary of the West Coast of South Africa.

Despite advances in our understanding of the geographic and temporal scope of the Paleolithic record, we know remarkably little about the evolutionary and ecological consequences of changes in human behavior. Recent inquiries suggest that human evolution reflects a long history of interconnections between the behavior of humans and their surrounding ecosystems (e.g., niche construction). Developing expectations to identify such phenomena is remarkably difficult because it requires understanding the multi-generational impacts of changes in behavior. These long-term dynamics require insights into the emergent phenomena that alter selective pressures over longer time periods which are not possible to observe, and are also not intuitive based on observations derived from ethnographic time scales. Generative models show promise for probing these potentially unexpected consequences of human-environment interaction. Changes in the uses of landscapes may have long term implications for the environments that hominins occupied. We explore other potential proxies of behavior and examine how modeling may provide expectations for a variety of phenomena.

Bayesian luminescence dating at Ghar-e Boof, Iran, provides a new chronology for Middle and Upper Paleolithic in the southern Zagros.

Ghar-e Boof is a Paleolithic cave site in Iran well known for its rich early Upper Paleolithic Rostamian assemblages. The site is located on the edge of the Dasht-e Rostam plain in the southern Zagros. Recent excavations by the members of the Tübingen-Iranian Stone Age Research Project at Ghar-e Boof also recovered well-stratified Middle Paleolithic assemblages. Here, we provide the first detailed luminescence chronology for the Middle and Upper Paleolithic of the site. More generally, our work is the first luminescence chronology for a Middle and Upper Paleolithic site in the Zagros Mountains region in Iran. The luminescence ages for the Upper Paleolithic of Ghar-e Boof agree with published 14C dates. We applied Bayesian models specifically designed for luminescence dating using the R package 'BayLum' to incorporate the well-established stratigraphic constraints, as well as the published 14C ages with our optically stimulated luminescence (OSL) ages to improve the precision of the chronological framework. The Bayesian chronology shows a significantly improved precision of the OSL ages in particular for the upper part of the sequence where 14C ages were available. The Bayesian OSL ages for the Rostamian horizons, archaeologicalhorizon (AH) III (a-b-c), and AH IV, fall in the range of 37-42 ka (68% credible interval [CI]). Moreover, we determined a series of dates between 45 and 81 ka (68% CI) for the Middle Paleolithic strata from AH IVd to AH VI. Our results point to a demographic shift in the populations responsible for the Middle Paleolithic and the Rostamian within three millennia. This major technological change accompanied by the rise of symbolic artifacts such as personal ornaments, may or may not reflect a replacement of Neanderthals by modern humans. While we are confident that the Rostamian was made by modern humans, available information does not allow us to be sure who made the local Middle Paleolithic.

Early anthropogenic use of hematite on Aurignacian ivory personal ornaments from Hohle Fels and Vogelherd caves, Germany.

The Aurignacian (ca. 43-35 ka) of southwestern Germany is well known for yielding some of the oldest artifacts related to symbolic behaviors, including examples of figurative art, musical instruments, and personal ornaments. Another aspect of these behaviors is the presence of numerous pieces of iron oxide (ocher); however, these are comparatively understudied, likely owing to the lack of painted artifacts from this region and time period. Several Aurignacian-aged carved ivory personal ornaments from the sites of Hohle Fels and Vogelherd contain traces of what appear to be red ocher residues. We analyzed these beads using a combination of macroanalytical and microanalytical methods, including scanning electron microscopy equipped with energy dispersive spectroscopy and Raman spectroscopy. We found that the residue is composed of the iron oxide mineral hematite (Fe2O3). Further analyses on associated archaeological sediments by X-ray diffraction revealed the absence of hematite and other iron oxide mineral phases, suggesting that the hematite residues were intentionally applied to the ivory personal ornaments by human agents. These findings have important implications as they represent evidence for the direct application of ocher on portable symbolic objects by early Homo sapiens in Europe. Furthermore, our results reveal shared behavioral practices from two key Aurignacian sites maintained over several millennia and illuminate aspects of pigment use and symbolic practices during a pivotal time in the cultural evolution of humans.

Breaking through the Aquitaine frame: A re-evaluation on the significance of regional variants during the Aurignacian as seen from a key record in southern Europe

The cultural dynamics that led to the appearance of the Aurignacian have intrigued archaeologists since the start of Paleolithic research. However, cultural reconstructions have often focused on a restricted region of Europe, namely the northern Aquitaine Basin. The Mediterranean Basin, though, is also a region worthy of consideration when testing if the Protoaurignacian was followed by the Early Aurignacian adaptive system. Fumane Cave is a pivotal site for tackling this issue because it contains evidence of repeated human occupations during the time span of the European Aurignacian. Here we investigate the diachronic variability of the lithic assemblages from five cultural units at Fumane Cave using a combination of reduction sequence and attribute analyses. This paper also reassesses the presence and stratigraphic reliability of the organic artifacts recovered at Fumane Cave. Our results show that the features of the Protoaurignacian techno-typology are present throughout the stratigraphic sequence, and by extension, to the onset of Heinrich Event 4. Additionally, the appearance of split-based points in the youngest phase is evidence of extensive networks that allowed this technological innovation to spread across different Aurignacian regions.

Split-based points from the Swabian Jura highlight Aurignacian regional signatures.

The systematic use of antlers and other osseous materials by modern humans marks a set of cultural and technological innovations in the early Upper Paleolithic, as is seen most clearly in the Aurignacian. Split-based points, which are one of the most common osseous tools, are present throughout most regions where the Aurignacian is documented. Using results from recent and ongoing excavations at Geißenklösterle, Hohle Fels and Vogelherd, we nearly tripled the sample of split-based points from 31 to 87 specimens, and thereby enhance our understanding of the technological economy surrounding the production of osseous tools. Aurignacian people of the Swabian Jura typically left spit-based points at sites that appear to be base camps rich with numerous examples of personal ornaments, figurative art, symbolic imagery, and musical instruments. The artifact assemblages from SW Germany highlight a production sequence that resembles that of SW France and Cantabria, except for the absence of tongued pieces. Our study documents the life histories of osseous tools and demonstrates templates for manufacture, use, recycling, and discard of these archetypal artifacts from the Aurignacian. The study also underlines the diversified repertoire of modern humans in cultural and technological realms highlighting their adaptive capabilities.

Regional patterns of diachronic technological change in the Howiesons Poort of southern Africa.

The Howiesons Poort (HP) of southern Africa plays an important role in models on the early behavioral evolution of Homo sapiens. The HP is often portrayed as a coherent MSA industry characterized by early complex material culture. Recent work has emphasized parallel technological change through time across southern Africa potentially driven by ecological adaptations or demographic change. Here we examine patterns of diachronic variation within the HP and evaluate potential causal factors behind these changes. We test previous temporal assessments of the technocomplex at the local and regional level based on high-resolution quantitative data on HP lithic assemblages from Sibudu (KwaZulu-Natal) and comparisons with other southern African sites. At Sibudu, consistent unidirectional change in lithic technology characterizes the HP sequence. The results show a gradual reduction in typical HP markers such as the proportion of blades, backed pieces, and HP cores, as well as declining size of blades and backed artifacts. Quantitative comparisons with seven HP sites in South Africa suggest that lithic technology varies between regions over time instead of following similar changes. Concerning hypotheses of causal drivers, directional changes in lithic technology at Sibudu covary with shifting hunting patterns towards larger-sized bovids and a gradual opening of the vegetation. In contrast, variation in lithic technology shows little association with site use, mobility patterns or demographic expansions. Unlike at Sibudu, diachronic changes at other HP sites such as Diepkloof, Klasies River and Klipdrift appear to be associated with aspects of mobility, technological organization and site use. The regional diachronic patterns in the HP partly follow paleoclimatic zones, which could imply different ecological adaptations and distinct connection networks over time. Divergent and at times decoupled changes in lithic traits across sites precludes monocausal explanations for the entire HP, supporting more complex models for the observed technological trajectories.

Fox dietary ecology as a tracer of human impact on Pleistocene ecosystems.

Nowadays, opportunistic small predators, such as foxes (Vulpes vulpes and Vulpes lagopus), are well known to be very adaptable to human modified ecosystems. However, the timing of the start of this phenomenon in terms of human impact on ecosystems and of the implications for foxes has hardly been studied. We hypothesize that foxes can be used as an indicator of past human impact on ecosystems, as a reflection of population densities and consequently to track back the influence of humans on the Pleistocene environment. To test this hypothesis, we used stable isotope analysis (δ13C, δ15N) of bone collagen extracted from faunal remains from several archaeological sites located in the Swabian Jura (southwest Germany) and covering a time range over three important cultural periods, namely the Middle Palaeolithic (older than 42,000 years ago) attributed to Neanderthals, and the early Upper Palaeolithic periods Aurignacian and Gravettian (42,000 to 30,000 years ago) attributed to modern humans. We then ran Bayesian statistic systems (SIBER, mixSIAR) to reconstruct the trophic niches and diets of Pleistocene foxes. We observed that during the Middle Palaeolithic period, when Neanderthals sparsely populated the Swabian Jura, the niches occupied by foxes suggest a natural trophic behavior. In contrast, during the early Upper Palaeolithic periods, a new trophic fox niche appeared, characterized by a restricted diet on reindeer. This trophic niche could be due to the consumption of human subsidies related to a higher human population density and the resulting higher impact on the Pleistocene environment by modern humans compared to Neanderthals. Furthermore, our study suggests that, a synanthropic commensal behavior of foxes started already in the Aurignacian, around 42,000 years ago.

Author Correction: A 300,000-year-old throwing stick from Schöningen, northern Germany, documents the evolution of human hunting.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

A 300,000-year-old throwing stick from Schöningen, northern Germany, documents the evolution of human hunting.

The poor preservation of Palaeolithic sites rarely allows the recovery of wooden artefacts, which served as key tools in the arsenals of early hunters. Here, we report the discovery of a wooden throwing stick from the Middle Pleistocene open-air site of Schöningen that expands the range of Palaeolithic weaponry and establishes that late Lower Palaeolithic hominins in Northern Europe were highly effective hunters with a wide array of wooden weapons that are rarely preserved in the archaeological record.