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Birch tar is the oldest synthetic substance made by early humans. The earliest such artefacts are associated with Neanderthals. According to traditional interpretations, their study allows understanding Neanderthal tool behaviours, skills and cultural evolution. However, recent work has found that birch tar can also be produced with simple processes, or even result from fortuitous accidents. Even though these findings suggest that birch tar per se is not a proxy for cognition, they do not shed light on the process by which Neanderthals produced it, and, therefore, cannot evaluate the implications of that behaviour. Here, we address the question of how tar was made by Neanderthals. Through a comparative chemical analysis of the two exceptional birch tar pieces from Königsaue (Germany) and a large reference birch tar collection made with Stone Age techniques, we found that Neanderthals did not use the simplest method to make tar. Rather, they distilled tar in an intentionally created underground environment that restricted oxygen flow and remained invisible during the process. This degree of complexity is unlikely to have been invented spontaneously. Our results suggest that Neanderthals invented or developed this process based on previous simpler methods and constitute one of the clearest indicators of cumulative cultural evolution in the European Middle Palaeolithic.

Acknowledgements: The authors thank G. Marciani and O. Jöris for comments on archaeology; C. Jeong, M. Spyrou and K. Prüfer for comments on genetics; M. O’Reilly for graphical support for Fig. 5 and Extended Data Fig. 9; the entire IT and laboratory teams at the Department of Archaeogenetics of MPI-SHH for technical assistance; M. Meyer and S. Nagel for support with single-stranded library preparation; K. Post, P. van Es, J. Glimmerveen, M. Medendorp, M. Sier, S. Dikstra, M. Dikstra, R. van Eerden, D. Duineveld and A. Hoekman for providing access to human specimens from the North Sea (The Netherlands); M. D. Garralda and A. Estalrrich for providing access to human specimens from La Riera (Spain); J. Górski and M. Zając for providing access to human specimens from Maszycka cave; C. Di Patti for providing access to human specimens from San Teodoro 2 (Italy); P. Blaževičius for providing access to the Donkalnis human remains and the new radiocarbon dates; the Italian Ministry of Culture and Soprintendenza Archeologia Belle Arti e Paesaggio for the Provinces of Verona, Rovigo, and Vicenza for granting access to the human remains of Tagliente 2; F. Fontana, who carries out investigations of the Riparo Tagliente site (Italy); the Friuli Venezia Giulia Superintendency for providing access to the human tooth Pradis 1; and the Soprintendenza Archeologia Belle Arti e Paesaggio for the Provinces of Barletta-Andria-Trani and Foggia for providing access to the Paglicci human remains. This project has received funding by the European Research Council under the European Union’s Horizon 2020 research and innovation programme under grant agreements no. 803147-RESOLUTION (to S.T.), no. 771234-PALEoRIDER (to W.H.), no. 864358 (to K.M.), no. 724703 and no. 101019659 (to K.H.). K.H. is also supported by the Deutsche Forschungsgemeinschaft (DFG FOR 2237). E.A. has received funding from the Van de Kamp fonds. PACEA co-authors of this research benefited from the scientific framework of the University of Bordeaux’s IdEx Investments for the Future programme/GPR Human Past. A.G.-O. is supported by a Ramón y Cajal fellowship (RYC-2017-22558). L. Sineo, M.L. and D.C. have received funding from the Italian Ministry of University and Research (MUR) PRIN 2017 grants 20177PJ9XF and 20174BTC4R_002. H. Rougier received support from the College of Social and Behavioral Sciences of CSUN and the CSUN Competition for RSCA Awards. C.L.S. and T. Saupe received support from the European Union through the European Regional Development Fund (project no. 2014-2020.4.01.16-0030) and C.L.S. received support from the Estonian Research Council grant PUT (PRG243). S. Shnaider received support from the Russian Science Foundation (no. 19-78-10053)., Modern humans have populated Europe for more than 45,000 years1,2. Our knowledge of the genetic relatedness and structure of ancient hunter-gatherers is however limited, owing to the scarceness and poor molecular preservation of human remains from that period3. Here we analyse 356 ancient hunter-gatherer genomes, including new genomic data for 116 individuals from 14 countries in western and central Eurasia, spanning between 35,000 and 5,000 years ago. We identify a genetic ancestry profile in individuals associated with Upper Palaeolithic Gravettian assemblages from western Europe that is distinct from contemporaneous groups related to this archaeological culture in central and southern Europe4, but resembles that of preceding individuals associated with the Aurignacian culture. This ancestry profile survived during the Last Glacial Maximum (25,000 to 19,000 years ago) in human populations from southwestern Europe associated with the Solutrean culture, and with the following Magdalenian culture that re-expanded northeastward after the Last Glacial Maximum. Conversely, we reveal a genetic turnover in southern Europe suggesting a local replacement of human groups around the time of the Last Glacial Maximum, accompanied by a north-to-south dispersal of populations associated with the Epigravettian culture. From at least 14,000 years ago, an ancestry related to this culture spread from the south across the rest of Europe, largely replacing the Magdalenian-associated gene pool. After a period of limited admixture that spanned the beginning of the Mesolithic, we find genetic interactions between western and eastern European hunter-gatherers, who were also characterized by marked differences in phenotypically relevant variants.

Objectives Cuncaicha, a rockshelter site in the southern Peruvian Andes, has yielded archaeological evidence for human occupation at high elevation (4,480 masl) during the Terminal Pleistocene (12,500-11,200 cal BP), Early Holocene (9,500-9,000 cal BP), and later periods. One of the excavated human burials (Feature 15-06), corresponding to a middle-aged female dated to ~8,500 cal BP, exhibits skeletal osteoarthritic lesions previously proposed to reflect habitual loading and specialized crafting labor. Three small tools found in association with this burial are hypothesized to be associated with precise manual dexterity. Materials and methods Here, we tested this functional hypothesis through the application of a novel multivariate methodology for the three-dimensional analysis of muscle attachment surfaces (entheses). This original approach has been recently validated on both lifelong-documented anthropological samples as well as experimental studies in nonhuman laboratory samples. Additionally, we analyzed the three-dimensional entheseal shape and resulting moment arms for muscle opponens pollicis. Results Results show that Cuncaicha individual 15-06 shows a distinctive entheseal pattern associated with habitual precision grasping via thumb-index finger coordination, which is shared exclusively with documented long-term precision workers from recent historical collections. The separate geometric morphometric analysis revealed that the individual's opponens pollicis enthesis presents a highly projecting morphology, which was found to strongly correlate with long joint moment arms (a fundamental component of force-producing capacity), closely resembling the form of Paleolithic hunter-gatherers from diverse geo-chronological contexts of Eurasia and North Africa. Discussion Overall, our findings provide the first biocultural evidence to confirm that the lifestyle of some of the earliest Andean inhabitants relied on habitual and forceful precision grasping tasks.

Small flake industries are a commonly identified component of Lower Paleolithic archaeological assemblages in Eurasia. Utilized as blanks for tools, at many sites, their functions are often poorly understood. Here we present a preliminary traceological analysis of lithics from Marathousa 1 (MAR-1; Megalopolis, Greece). MAR-1 dates to ca. 400-500 ka BP and is one of the oldest open-air sites in South-Eastern Europe. It has yielded a lithic assemblage made almost exclusively of small-sized flakes found in direct association with diverse megafauna including elephants, hippo and deer. For this preliminary study, a total of 223 artifacts were sampled for a taphonomical analysis and 13 for a functional analysis. The lithic artifacts from MAR-1 are exceptionally well-preserved and are only slightly affected by chemical alterations. They are therefore ideal for a techno-morpho-functional analysis. Use-wear traces confirm on-site butchery. Our results also confirm that plants were worked at the site, whereas technological traces, rarely observed on lithics from this age, can also be seen on a number of specimens. Whereas both retouched and unretouched tools contribute significantly to the MAR-1 toolkit, shapes are varied, and at this phase of the study do not appear morphologically or technologically standardized. However, backing (natural or retouched) opposite to a sharp edge can be observed on numerous specimens. When compared to the sparse functional data available on small tools from Lower Paleolithic Europe and the Levant, small flake tools include a wide variety of techno-functional types. To fully begin to understand this diversity, lithic toolkits must be considered in relation to the rest of the assemblage and the accompanying contextual data, including information from technological, archaeozoological, and palaeoenvironmental datasets.

The Middle and Late Pleistocene is arguably the most interesting period in human evolution. This broad period witnessed the evolution of our own lineage, as well as that of our sister taxon, the Neanderthals, and related Denisovans. It is exceptionally rich in both fossil and archaeological remains, and uniquely benefits from insights gained through molecular approaches, such as paleogenetics and paleoproteomics, that are currently not widely applicable in earlier contexts. This wealth of information paints a highly complex picture, often described as 'the Muddle in the Middle,' defying the common adage that 'more evidence is needed' to resolve it. Here we review competing phylogenetic scenarios and the historical and theoretical developments that shaped our approaches to the fossil record, as well as some of the many remaining open questions associated with this period. We propose that advancing our understanding of this critical time requires more than the addition of data and will necessitate a major shift in our conceptual and theoretical framework.

A number of different approaches are currently available to digitally restore the symmetry of a specimen deformed by taphonomic processes. These tools include mirroring and retrodefor-mation to approximate the original shape of an object by symmetrisation. Retrodeformation has the potential to return a rather faithful representation of the original shape, but its power is limited by the availability of bilateral landmarks. A recent protocol proposed by Schlager and colleagues (2018) overcomes this issue by using bilateral landmarks and curves as well as semilandmarks. Here we applied this protocol to the Middle Pleistocene human cranium from Steinheim (Germany), the holotype of an abandoned species named Homo steinheimensis. The peculiar morphology of this fossil, associated with the taphonomic deformation of the entire cranium and the lack of a large portion of the right side of the face, has given rise to different hypotheses over its phylogenetic position. The reconstruction presented here sheds new light on the taphonomic origin of some features observed on this crucial specimen and results in a morphology consistent with its attribution to the Neanderthal lineage. © 2021 by the authors. Licensee MDPI, Basel, Switzerland.

The aim of this paper is to evaluate the craniometric affinities of the only Cuncaicha cranial specimen with other early, middle, and late Holocene South American samples. To do so, the sku...

Two fossilized human crania (Apidima 1 and Apidima 2) from Apidima Cave, southern Greece, were discovered in the late 1970s but have remained enigmatic owing to their incomplete nature, taphonomic distortion and lack of archaeological context and chronology. Here we virtually reconstruct both crania, provide detailed comparative descriptions and analyses, and date them using U-series radiometric methods. Apidima 2 dates to more than 170 thousand years ago and has a Neanderthal-like morphological pattern. By contrast, Apidima 1 dates to more than 210 thousand years ago and presents a mixture of modern human and primitive features. These results suggest that two late Middle Pleistocene human groups were present at this site—an early Homo sapiens population, followed by a Neanderthal population. Our findings support multiple dispersals of early modern humans out of Africa, and highlight the complex demographic processes that characterized Pleistocene human evolution and modern human presence in southeast Europe. Detailed comparative analyses of two fossil crania from Apidima Cave, Greece, indicate that two late Middle Pleistocene human groups were present at this site; first an early Homo sapiens population followed by a Neanderthal population.

The face is the most distinctive feature used to identify others. Modern humans have a short, retracted face beneath a large globular brain case that is distinctively different from that of our closest living relatives. The face is a skeletal complex formed by 14 individual bones housing parts of the digestive, respiratory, visual, and olfactory systems. A key to understanding the origin and evolution of the human face is to analyze the faces of extinct taxa in the hominin clade over the last 6 million years. Yet as new fossils are recovered, and the number of hominin species grows, the question of how and when the modern human face originated remains unclear. By examining key features of the facial skeleton, here we evaluate the evolutionary history of the modern human face in the context of its development, morphology, and function, and suggest that its appearance is the result of a combination of biomechanical, physiological and social influences.

The fossil record of early Homo sapiens in the African Pleistocene remains sparse. In contrast to its prominent position regarding the cultural evolution of our species, southern Africa plays a secondary role in narratives regarding human biological origins. Reasons for this are a limited and fragmentary fossil record from the Middle Stone Age (MSA), further complicated by a number of human remains coming from contexts lacking chronostratigraphic information. Similar to the southern African MSA overall, the rich archeological deposits of Sibudu stand in opposition to its scarce record of hominin fossils. Here, we report on three human teeth (SIB-1, 2, 3) from securely stratified MSA deposits at Sibudu dating between > 77 and 64 ka. The teeth include two lower deciduous molars (Ldm2) with heavy occlusal wear and one fragment. We focus on describing the find and archeological context, followed by an initial assessment of the fossils and their contextualization within the African record. The juvenile teeth derive from rich and well-stratified archeological deposits, associated with a Howiesons Poort industry at ~ 64 ka from PGS3 (SIB-3) and pre-Still Bay occupations in strata Casper and Danny at > 77 ka (SIB-1, 2). The latter constitute the oldest human fossils from Sibudu. Metric and morphological analyses of the Ldm2s (SIB-2, 3) find a combination of archaic traits (e.g., mid-trigonid crest) and crown dimensions that overlap with ranges of both Pleistocene and recent Homo sapiens. These results match with a population of Homo sapiens that lies chronologically between the earliest members of the species and recent humans.

Significance The microbiome plays key roles in human health, but little is known about its evolution. We investigate the evolutionary history of the African hominid oral microbiome by analyzing dental biofilms of humans and Neanderthals spanning the past 100,000 years and comparing them with those of chimpanzees, gorillas, and howler monkeys. We identify 10 core bacterial genera that have been maintained within the human lineage and play key biofilm structural roles. However, many remain understudied and unnamed. We find major taxonomic and functional differences between the oral microbiomes of Homo and chimpanzees but a high degree of similarity between Neanderthals and modern humans, including an apparent Homo-specific acquisition of starch digestion capability in oral streptococci, suggesting microbial coadaptation with host diet., The oral microbiome plays key roles in human biology, health, and disease, but little is known about the global diversity, variation, or evolution of this microbial community. To better understand the evolution and changing ecology of the human oral microbiome, we analyzed 124 dental biofilm metagenomes from humans, including Neanderthals and Late Pleistocene to present-day modern humans, chimpanzees, and gorillas, as well as New World howler monkeys for comparison. We find that a core microbiome of primarily biofilm structural taxa has been maintained throughout African hominid evolution, and these microbial groups are also shared with howler monkeys, suggesting that they have been important oral members since before the catarrhine–platyrrhine split ca. 40 Mya. However, community structure and individual microbial phylogenies do not closely reflect host relationships, and the dental biofilms of Homo and chimpanzees are distinguished by major taxonomic and functional differences. Reconstructing oral metagenomes from up to 100 thousand years ago, we show that the microbial profiles of both Neanderthals and modern humans are highly similar, sharing functional adaptations in nutrient metabolism. These include an apparent Homo-specific acquisition of salivary amylase-binding capability by oral streptococci, suggesting microbial coadaptation with host diet. We additionally find evidence of shared genetic diversity in the oral bacteria of Neanderthal and Upper Paleolithic modern humans that is not observed in later modern human populations. Differences in the oral microbiomes of African hominids provide insights into human evolution, the ancestral state of the human microbiome, and a temporal framework for understanding microbial health and disease.

KNM-OG 45500 is a hominin fossil composed of parts of a frontal bone, left temporal bone, and cranial vault pieces. Since its discovery along the Olorgesailie Formation (Kenya) in 2003, it has been associated with the Homo erectus hypodigm. The specimen, derived from a geological context dated to ca. 900 Ka BP, has been described as a very small individual of probable female sex. However, despite its status as an important hominin specimen, it has not been used in a quantitative comparative framework because of its fragmentary condition. Here, we undertake a virtual reconstruction of the better-preserved fragment, the frontal bone. We additionally apply geometric morphometric analyses, using a geographically diverse fossil and modern human sample, in order to investigate the morphological affinities of KNM-OG 45500. Our results show that the frontal shape of KNM-OG 45500 exhibits similarities with Early Pleistocene fossils from Eurasia and Africa that are assigned to H. erectus sensu lato (s.l.). Its size, on the other hand, is notably smaller than most other Homo erectus fossils and modern humans and similar to the specimens from Dmanisi (Georgia) and to Homo naledi. Taken together, our analyses of the frontal bone suggest a taxonomic attribution of KNM-OG 45500 to H. erectus s.l. and extend even further the range of size variability associated with this taxon around 900 Ka BP.

BACKGROUND: Recent advances in sequencing have facilitated large-scale analyses of the metagenomic composition of different samples, including the environmental microbiome of air, water, and soil, as well as the microbiome of living humans and other animals. Analyses of the microbiome of ancient human samples may provide insights into human health and disease, as well as pathogen evolution, but the field is still in its very early stages and considered highly challenging. - RESULTS: The metagenomic and pathogen content of Egyptian mummified individuals from different time periods was investigated via genetic analysis of the microbial composition of various tissues. The analysis of the dental calculus’ microbiome identified Red Complex bacteria, which are correlated with periodontal diseases. From bone and soft tissue, genomes of two ancient pathogens, a 2200-year-old Mycobacterium leprae strain and a 2000-year-old human hepatitis B virus, were successfully reconstructed. - CONCLUSIONS: The results show the reliability of metagenomic studies on Egyptian mummified individuals and the potential to use them as a source for the extraction of ancient pathogen DNA. Background Results - Sample information and dating - General metagenomic assessment - Mycobacterium leprae (individual Abusir1630) - Hepatitis B virus (individual Abusir1543) - Oral microbiome assessment Discussion Conclusions Methods - Sample extraction and radiocarbon dating - Sample extraction and library preparation - Metagenomic screening - Authentication of ancient DNA - Content of endogenous DNA (SourceTracker2) - Data processing of sample Abusir1630b (M. leprae) -- Read processing, mapping, and variant calling -- SNP typing -- Anthropological analysis -- Phylogeny -- Beast analysis -- Temporal signal - Data processing individual Abusir1543 (hepatitis B virus) -- Read processing, mapping, and variant calling -- Phylogeny -- Recombination analysis -- Beast analysis -- Temporal signal

Summary Systematic tool production and use is one of humanity’s defining characteristics, possibly originating as early as >3 million years ago.1, 2, 3 Although heightened manual dexterity is considered to be intrinsically intertwined with tool use and manufacture, and critical for human evolution, its role in the emergence of early culture remains unclear. Most previous research on this question exclusively relied on direct morphological comparisons between early hominin and modern human skeletal elements, assuming that the degree of a species’ dexterity depends on its similarity with the modern human form. Here, we develop a new approach to investigate the efficiency of thumb opposition, a fundamental component of manual dexterity, in several species of fossil hominins. Our work for the first time takes into account soft tissue as well as bone anatomy, integrating virtual modeling of musculus opponens pollicis and its interaction with three-dimensional bone shape form. Results indicate that a fundamental aspect of efficient thumb opposition appeared approximately 2 million years ago, possibly associated with our own genus Homo, and did not characterize Australopithecus, the earliest proposed stone tool maker. This was true also of the late Australopithecus species, Australopithecus sediba, previously found to exhibit human-like thumb proportions. In contrast, later Homo species, including the small-brained Homo naledi, show high levels of thumb opposition dexterity, highlighting the increasing importance of cultural processes and manual dexterity in later human evolution., Highlights • Increased thumb opposition efficiency was present about 2 million years ago • This evolutionary advantage was less pronounced in Australopithecus • This dexterity is shared with all recent hominins, including Homo naledi, Karakostis et al. integrate virtual muscle modeling with three-dimensional bone shape analysis to investigate biomechanical efficiency for thumb opposition in the fossil human record. They report the earliest evidence of increased manual dexterity, a vital component of human-like tool use, in thumb bones dated to about 2 million years ago.

Previous research found sexual dimorphism in the bony labyrinth of a Greek population sample (Osipov et al. 2013). This study intends to investigate the nature of this structure’s sexual dimorphism across populations of diverse geographic origin and to identify the effect of inter-population variation on the accuracy of determining sex using the bony labyrinth. Three population samples of known sex were analyzed originating from Europe (n = 30), Africa (n = 38), and North America (n = 30). The discriminant function developed in Osipov et al. (2013) was applied, and new function equations for sex estimation were developed. In addition, we used principal component analyses for investigating population differences, while bivariate tests were used to compare across population samples, sexes, and anatomical sides. A significant level of sexual dimorphism was found in all population samples, being driven by both size and shape differences. Discriminant functions for sex estimation were developed for all three population samples combined (71.4% accuracy) as well as separately (70–80% accuracy). The German sample was the least sexually dimorphic, whereas the North American sample exhibited the greatest sexual dimorphism. The size and shape of the bony labyrinth also significantly differed across population samples. The bony labyrinth is found to be sexually dimorphic across distinct population groups. Due to significant differences across our population samples, the accuracy of the previously proposed method for sex determination (Osipov et al., 2013) was relatively low. For this purpose, this study presented new functions, whose accuracy was tested in three distinct population samples.

We present an updated tectono-stratigraphic development model of the Megalopolis Basin (MB), which is an intra-montane basin, located in the actively extending domain of the Hellenic Arc, based on re-interpretation of borehole data, field mapping and stratigraphic – sedimentological reconnaissance. The Megalopolis basin develops on the hanging-wall of the W. Mainalo Fault System, which accommodates the deformation associated with the exhumation of the PQ metamorphics in the window of Assea, east of the MB. During the early stages of basin development, NNW-SSE normal faults controlled its eastern margin. These interacted with and gradually dismembered the ENE-WSW ones that were related to pre-Pliocene extension.The establishment of ENE-WSW Quaternary extension in the southern Peloponnese is associated with major, range-bounding NNW-SSE faults, such as the Sparta F. that controls the eastern margin of the Taygetos horst. Fault growth and consequent uplift of the Taygetos horst affected the southern reaches of the Megalopolis Basin, through the development of an intra-basinal high (Leontari horst) that split the southern portion of the MB in two sub-basins, while the activity on the faults on the eastern margin of the MB ceases. The switching-off of the eastern margin was taken over by the faults that control the western margin of the MB, such as the prominent NNW-SSE, east-throwing Ellinitsa Fault and the Lykaion Fault System; the latter is a rather elusive structure, owing to the nature of the affected formations (erodible Pindos clastics) and dense forest cover that obscure fault exposures.The successive stages of fault evolution are reflected in, and largely control, the sedimentation type(s) in the MB. The initial Pliocene lacustrine conditions were gradually replaced by extensive fluviatile sedimentation (Megalopolis Fm), which interfingers with more focused, lacustrine deposits in the basin centre (Marathousa Fm.), when the Pleistocene Megalopolis Lake developed. The subsequent deposition of the (mainly) fluvial Apiditsa and Ellinitsa formations, follows the gradual starvation of sediment feeding from the E-ESE, (which fed the Megalopolis Fm) and marks the onset of fault activity in the southern and western parts of the MB. The establishment of the modern Alfeios drainage, initially deposited floodplain sediments, subsequently to cut into them and form terraces, following episodic(?) drops of its base-level, owing to alternating climatic conditions and/or surges of fault activity. Finally, the breaching of the basement salient in the NNW (Karytaina gorge), led to the establishmnent of the present-day base level, with Alfeios cutting into its more recent deposits.This research was conducted under the auspices of the Ephoreia of Paleoanthropology and Speleology, Greek Ministry of Culture, and was supported by the European Research Council (CROSSROADS).

Tool use is one of humanity’s defining characteristics, possibly originating as early as > 3 million years ago. Although heightened manual dexterity is considered to be intrinsically intertwined with tool use and manufacture and critical for human evolution, its role in the emergence of early culture remains unclear. Most previous research on this question exclusively relied on direct morphological comparisons between early hominin and modern human skeletal elements, assuming that the degree of a species’ dexterity depends on its similarity with the modern human form. Here we develop a new approach to investigate the biomechanical efficiency of thumb opposition, a fundamental component of manual dexterity, in several species of fossil hominins. Our work for the first time takes into account soft tissue as well as bone anatomy, integrating virtual modeling of the muscle opponens pollicis and its interaction with three-dimensional bone shape form. Results indicate that efficient thumb opposition appeared approximately 2 million years ago, likely associated with our own genus Homo, and did not characterize Australopithecus, the earliest proposed stone tool-maker. This was true also of the late Australopithecus species, A. sediba, previously found to exhibit human-like thumb morphology. In contrast, later Homo species, including the small-brained H. naledi, show high levels of thumb opposition dexterity, highlighting the increasing importance of cultural processes and manual dexterity in later human evolution.

The mining activities in the Middle Pleistocene lacustrine basin of Megalopolis (Peloponnesus, Greece) have exposed expanded sections of lacustrine sediments. In particular, the northernmost mine of Kyparissia has yielded numerous vertebrate fossils, recovered during field surveys and small-scale rescue excavations. The stratified specimens indicate the presence of at least two fossiliferous horizons, which are stratigraphically closely situated. The fossils are generally widely dispersed, but they are much more abundant along the western margin of the palaeolake. Four sites showing denser concentration of fossils were located and partially excavated during the 2004–2008 fieldwork. An additional site was identified and excavated in 2012. The fossils were found in organic-rich sediments (mostly lignites), indicating that they were deposited during a warm and humid (i.e., interglacial) time period, in a richly vegetated environment. The recovered fauna is diverse, dominated by hippopotamuses, deer and elephants. The present study presents the chelonian, avian and suiform finds. The following taxa are recognised: Emys orbicularis, Testudo marginata, Sus scrofa, Hippopotamus antiquus, as well as a diverse sample of aquatic birds. This faunal composition indicates a temperate, woodland/forest environment, with continuous presence of a large water body, whilst the presence of the darter (Anhinga sp.) points to relatively warm conditions. Biochronologically it points to an age within the early part of the Middle Pleistocene.

The Villafranchian ursids of Greece are scanty, represented by a few isolated teeth and bones. During our last field campaigns in the Mygdonia Basin (Macedonia, Greece) we discovered an almost complete cranium, as well as some cranial, dental and postcranial remains, which are presented in this article. The new material originates from the locality Tsiotra Vryssi (TSR), dated to the late Villafranchian (1.8–1.2 Ma). The specimens are described and compared with ursids from various European localities; their morphological characters and dimensions place it with Ursus etruscus Cuvier, 1823, while the dental features are most similar to the specimens from Pietrafitta (Italy). The TSR cranium was CT-scanned in order to create a 3D virtual model and study its internal anatomy. Its observed endocranial traits exhibit primitive conditions, confirming the basal position of U. etruscus among the Pleistocene European ursids. Most of these features are shared with U. arctos and clearly differ from U. deningeri and U. spelaeus. Finally, we provide a revision of the known Greek material of U. etruscus and the biostratigraphic distribution of the Miocene–Pleistocene ursids of Greece. The earliest secure appearance of Ursus etruscus in Greece is found in the locality Dafnero (Western Macedonia), dated to the end of the middle Villafranchian at ~2.0 Ma, whereas its probable last occurrence is known from the locality Apollonia 1 of the Mygdonia Basin, dated to the latest Villafranchian at ~1.2 Ma.

The lacustrine deposits exposed at the Lower Palaeolithic site Marathousa 1 (Megalopolis, S. Greece) and intercalated between lignite Seam II and III yielded a collection of small vertebrate remains. The assemblage includes fish and small mammals; the mammal assemblage encompasses a variety of species, dominated by voles (arvicolids) of the genera Arvicola and Microtus. Other rodents are represented by a dipod cf. Alactaga and a murid of the genus Apodemus. In addition, there is a number of insectivore remains that refer to the family Soricidae, and more specifically to the genus Crocidura. The unrooted Arvicola molars show a positive‘Mimomys’ enamel differentiation with a mean SDQ value of 122, indicating a late Middle Pleistocene age (ca. 400 ka.) (MIS 12/11). The Marathousa 1 assemblage is clearly younger than the Choremi assemblages sampled ca. 20 years ago in the same basin. In the Choremi assemblage, Mimomys, the predecessor of the genus Arvicola, is the predominant arvicolid.

In anthropological sciences, muscle attachments are typically utilized for reconstructing the physical activities of past human populations. This approach relies on the concept that entheseal bone morphology is influenced by cumulative biomechanical stress. A fundamental criterion for assessing the stage of entheseal change involves the proportion of elevated bone area. However, it is not yet clear if bone elevation is associated with biomechanical forces exerted during physical activity, while the histology of the entheses of the human hand, the least-bodyweight-bearing anatomical area, is not fully investigated. Multiple previous studies on entheses have concluded that the concentration of calcified fibrocartilage reflects the level of applied forces. On this basis, if hand entheseal surface elevation was associated with the level of biomechanical stress, then a greater amount of calcified fibrocartilage would be expected in entheses' central and more projecting bone areas. More importantly, individual entheses with a greater proportion of elevated bone areas would present a higher total concentration of calcified fibrocartilage. To test these hypotheses, this pilot study conducted a histological quantitative analysis on two thumb entheses of four fully-documented body donors. Across individuals, all central entheseal regions presented greater calcified fibrocartilage, while the entheses showing additional bone elevation in their marginal areas comprised substantially higher total values. The observations of this small-scale pilot study support the concept that interindividual differences in entheseal bone morphology are related to varying levels of biomechanical loading. Furthermore, they encourage future research to compile larger sample sizes, for comparing individuals with distinct lifelong activities. Anat Rec, 302:1093-1103, 2019. © 2018 Wiley Periodicals, Inc.

Objectives The purpose of this study was to put forth a precise landmark-based technique for reconstructing the three-dimensional shape of human entheseal surfaces, to investigate whether the shape of human entheses is related to their size. The effects of age-at-death and bone length on entheseal shapes were also assessed. Materials and methods The sample comprised high-definition three-dimensional models of three right hand entheseal surfaces, which correspond to 45 male adult individuals of known age. For each enthesis, a particular landmark configuration was introduced, whose precision was tested both within and between observers. The effect of three-dimensional size, age-at-death, and bone length on shape was investigated through shape regression. Results The method presented high intra-observer and inter-observer repeatability. All entheses showed significant allometry, with the area of opponens pollicis demonstrating the most substantial relationship. This was particularly due to variation related to its proximal elongated ridge. The effect of age-at-death and bone length on entheses was limited. Discussion The introduced methodology can set a reliable basis for further research on the factors affecting entheseal shape. Using both size and shape, variables can provide further information on entheseal variation and its biomechanical implications. The low entheseal variation by age verifies that specimens under 50 years of age are not substantially affected by age-related changes. The lack of correlation between entheseal shape and bone length or age implies that other factors may regulate entheseal surfaces. Future research should focus on multivariate shape patterns among entheses and their association with occupation.

Farming was first introduced to southeastern Europe in the mid-7th millennium BCE - brought by migrants from Anatolia who settled in the region before spreading throughout Europe. However, the dynamics of the interaction between the first farmers and the indigenous hunter-gatherers remain poorly understood because of the near absence of ancient DNA from the region. We report new genome-wide ancient DNA data from 204 individuals-65 Paleolithic and Mesolithic, 93 Neolithic, and 46 Copper, Bronze and Iron Age-who lived in southeastern Europe and surrounding regions between about 12,000 and 500 BCE. We document that the hunter-gatherer populations of southeastern Europe, the Baltic, and the North Pontic Steppe were distinctive from those of western Europe, with a West-East cline of ancestry. We show that the people who brought farming to Europe were not part of a single population, as early farmers from southern Greece are not descended from the Neolithic population of northwestern Anatolia that was ancestral to all other European farmers. The ancestors of the first farmers of northern and western Europe passed through southeastern Europe with limited admixture with local hunter-gatherers, but we show that some groups that remained in the region mixed extensively with local hunter-gatherers, with relatively sex-balanced admixture compared to the male-biased hunter-gatherer admixture that we show prevailed later in the North and West. After the spread of farming, southeastern Europe continued to be a nexus between East and West, with intermittent steppe ancestry, including in individuals from the Varna I cemetery and associated with the Cucuteni-Trypillian archaeological complex, up to 2,000 years before the Steppe migration that replaced much of northern Europe9s population.