OBJECTIVES:The current research delves into the use of 3D geometric morphometric for assessing shifts in maturity within both the proximal and distal humeral metaphyses. It mainly focuses on establishing correlations between these shifts and the shape changes observed in the corresponding epiphyses established through radiographic imaging. MATERIAL AND METHODS:The total sample comprises 120 right-side proximal humeral metaphyses and 91 right-side distal humeral metaphyses. The entire sample was categorized into four maturity groups for the humeral distal metaphysis and five maturity groups for the humeral proximal metaphysis based on the appearance and subsequent changes in shape and size. All humeri were scanned using 3D surface scanning devices. Two different 3D template configurations were created to capture the shape changes during the development of the proximal and distal metaphyses. We conducted an analysis of shape (Procrustes shape coordinates) and an analysis on the form space in order to assess both maturity changes of humeral metaphyses and their interrelationship with size. RESULTS:The shape changes explained by PC1 are crucial in establishing the maturational sequence in both the distal and proximal metaphyses. These shape changes reflect the appearance and subsequent changes in shape and size of their corresponding epiphyses. DISCUSSION AND CONCLUSIONS:There is a considerable overlap among different maturity groups related to the gradual nature of the maturational process. However, shape changes in proximal and distal humeral metaphyses are suitable to assess maturity in skeletal specimens.

We present here a new Upper Miocene vertebrate locality, Kayaca, from the Beyağaç Basin in southwestern Türkiye that was discovered after surface survey and was subsequently systematically excavated. The locality yielded a diverse and rich vertebrate fauna notably including several complete cranial specimens. In this article we provide the first results on the taxonomy of the large mammal assemblage, and we discuss the biochronological implications. The Kayaca large mammal assemblage is dominated by hipparionine horses (Hipparion dietrichi morphotype, Cremohipparion sp.) and bovids (Gazella cf. ancyrensis, Sporadotragus parvidens, Tragoportax sp., Palaeoryx pallasi and the rare Gangraia), but includes also rhinocerotids (Miodiceros neumayri, Diceros sp.), suids (Hippopotamodon major), giraffids (Palaeotragus rouenii, Samotherium major), hystricids (Hystrix sp.), orycteropodids (Amphiorycteropus gaudryi), proboscideans (Deinotherium sp., Elephantimorpha indet.) and carnivorans. The latter are identified by some well-preserved specimens belonging to ictitheres (Protictitherium sp.), hyaenids (Adcrocuta eximia), felids (Pontosmilus sp. and a medium-sized felid) but also to the rare mustelid Parataxidea maraghana. The preliminary comparison with already known and well-studied faunal assemblages of Türkiye and the wider region, particularly the neighboring localities of Samos in Greece, suggest an age of ca. 7.4, close to the transition of early/middle Turolian (MN 11/12). Located at the western corner of Asia, Kayaca comprises a key location linking the vertebrate faunas of Asia and Europe and offers valuable insights into the Late Miocene terrestrial ecosystems of the eastern peri-Mediterranean region.

OBJECTIVE:This study follows up on our recent morphological analysis of the juvenile maxilla from Mugharet el'Aliya, Morocco. Although this specimen shows a reportedly archaic morphology, likely due to its large size, 3D shape analyses indicated affinities with early Homo sapiens. Here, we conducted an in-depth comparative investigation of the associated dentition to further clarify this individual's phylogenetic and taxonomic affinities. MATERIALS AND METHODS:Our analyses were based on three kinds of data: (a) external crown dimensions and non-metric features, analyzed via summary statistics; (b) CT scan data enabling the study of internal structures (enamel-dentine junction) via geometric morphometrics; and (c) high-resolution replicas of the external surface of the upper canine enabling the study of perikymata numbers via probability functions. The comparative samples included Middle Pleistocene (Chibanian) Europeans and Africans, Neanderthals, and early and later H. sapiens. RESULTS:Mugharet el'Aliya showed the greatest similarities in external and internal tooth morphology with early and later H. sapiens. Perikymata counts cluster the upper canine with H. sapiens. However, its canine and fourth premolar are megadont at a level generally atypical for H. sapiens. DISCUSSION:Our analyses of the dentition of the Mugharet el'Aliya individual support our previous findings on the morphological analysis of the maxilla, placing this fossil closest to H. sapiens. Our study further strengthens the evidence connecting fossils from the North African Aterian to those from Western Asia, especially Qafzeh. We also provide the first comparative analysis of a permanent upper canine from the Aterian fossil record.

Marathousa 1 (~430 ka BP), located in the Megalopolis Basin, Greece, represents the earliest documented butchery site in the Southern Balkans, providing clear evidence of a direct association between artefacts and remains of Palaeoloxodon antiquus. The lithic assemblage features a distinctive small tools industry, primarily produced from local radiolarite, comprising both simple flakes and retouched tools. Through technological analysis, raw material characterisation, experimental knapping, and statistical analyses, this study explores how Middle Pleistocene hominins organised their technological behaviour as reflected in the lithic assemblage, and how these behaviours were shaped by the resource-rich setting of the Megalopolis Basin, characterised by abundant raw materials, water sources, and faunal availability. Results demonstrate the interplay between freehand and bipolar knapping, reflecting a flexible technological strategy to exploit the available radiolarite. Freehand percussion was mainly used in flake production, while the bipolar technique facilitated initial core reduction and late-stage exhaustion. The consistent microlithisation at the site is also evident in the exploitation of other locally available raw materials, such as limestone, flint, and quartz, supporting previous studies demonstrating small flakes' effectiveness in diverse tasks. Technological patterns at Marathousa 1 broadly correspond to those observed at other Middle Pleistocene small tool sites; however, its distinctive intersection of raw material availability, technological choices and functional demands provides new insights into regional patterns of lithic variability across Eurasia during this period.

The Balkan Peninsula is considered to have acted as a glacial refugium as well as a biogeographic crossroads during the Pleistocene, playing an important role in the survival of biota and population dynamics through time. Furthermore, rather than being a homogeneous habitat, it is thought to have hosted a number of diverse, isolated “refugia within refugia,” providing suitable conditions for the persistence of different plant and animal species, as well as, potentially, Pleistocene hominins. We present the first palynological and charcoal record, although discontinuous, from the middle Pleistocene archaeological site Marathousa 1 (MAR-1; Megalopolis Basin) to reconstruct the local environment along with the first multispecies record of coprophilous fungal spores from Greece to track herbivore activity during the Marine Isotope Stage (MIS) 12. Our data show that during the early and late MIS 12, when the forest cover substantially decreased, mesophilous trees and aquatic vegetation persisted, reflecting wetter and milder conditions at MAR-1. Herbivore presence is documented by ∼473 ka, while its intensification coincides with increased vegetation biomass and local fire activity during the late MIS 12. Our findings suggest that MAR-1 likely served as a glacial refugium for middle Pleistocene hominins, providing essential resources for their survival during MIS 12.

Neanderthals’ lives were historically portrayed as highly stressful, shaped by constant pressures to survive in harsh ecological conditions, thus potentially contributing to their extinction. Recent work has challenged this interpretation, leaving the issue of stress among Paleolithic populations highly contested and warranting in-depth examination. Here, we analyze the frequency of dental enamel hypoplasia, a growth disruption indicator of early life stress, in the largest sample of Neanderthal and Upper Paleolithic dentitions investigated to date for these features. To track potential species-specific patterns in the ontogenetic distribution of childhood stress, we present the first comprehensive Bayesian modelling of the likelihood of occurrence of individual and matched enamel growth disruptions throughout ontogeny. Our findings support similar overall stress levels in both groups but reveal species-specific patterns in its ontogenetic distribution. While Neanderthal children faced increasing likelihoods of growth disruptions starting with the weaning process and culminating in intensity post-weaning, growth disruptions in Upper Paleolithic children were found to be limited around the period of weaning and substantially dropping after its expected completion. These results might, at least in part, reflect differences in childcare or other behavioral strategies between the two taxa, including those that were advantageous for modern humans’ long-term survival.

The Eastern Mediterranean lies directly on the principal migration route for human groups dispersing across Africa, Europe, and Asia. It also encompasses the Balkans, where fauna and flora, as well as hominin populations, are thought to have persisted through glacial periods. Based on this geographic position, the region's palaeoanthropological record would be expected to be among the longest and most continuous, but also the most highly diverse, reflecting complex processes of late survivals, new arrivals, and, potentially, contact between different lineages. While the record from the Near East meets these expectations, little evidence exists to support them in the northern part of the Easter Mediterranean. Here I present new evidence from our recent work in Greece, both in the laboratory and in the field. These new data highlight the regions’ importance as both dispersal corridor and glacial refugium for human populations in the Pleistocene. Results strongly suggest that the paucity of currently available data is due to lack of research, and highlight the potential of the record to shed further light on the complex processes underlying human evolution and dispersals.

The Balkans are considered a major glacial refugium where flora and fauna survived glacial periods and repopulated the rest of Europe during interglacials. While it is also thought to have harboured Pleistocene human populations, evidence linking human activity, paleoenvironmental indicators and a secure temporal placement to glacial periods is scant. Here, we present the first intra-tooth multi-isotope analysis for the European straight-tusked elephant Palaeoloxodon antiquus, on an adult male individual excavated in association with lithic artefacts at the MIS 12 site Marathousa 1 (Megalopolis basin, Greece). The studied find also exhibits anthropogenic modifications, providing direct evidence of hominin presence. We employed strontium, carbon and oxygen isotope analysis on enamel bioapatite to investigate its foraging and mobility behaviour, using a sequential sampling strategy along the tooth growth axis of the third upper molar, to assess ecological changes during the last decade of life. We found a geographically restricted range, in a C3-dominated open woodland environment, and relatively stable conditions over the examined timeframe. Our results show that, despite the severity of the MIS 12 glacial, the Megalopolis basin sustained a mesic habitat, sufficient plant cover and limited seasonal fluctuations in resource availability, pointing to its role as a glacial refugium for both fauna and hominins.

Southern Europe is hypothesized to have acted as a glacial refugium for hominin populations during the Pleistocene. Of particular importance is South-East Europe, which most likely played a dual role, both as refugium and dispersal corridor, especially during the Middle Pleistocene glaciations, when drastic climatic conditions led to major sea level drops in the Aegean. However, little is known about the palaeoenvironmental conditions at the time of hominin presence in this region, making these hypotheses difficult to test. Here we analyze biomarker data and leaf wax stable isotopic compositions of the MIS 12 Lower Palaeolithic site Marathousa 1 (Megalopolis Basin, Greece) to assess the climatic conditions accompanying the time of hominin presence in the area. Our data indicate a major cooling affecting the north Mediterranean/Aegean domain during this time interval, with lowest temperatures recorded between 440-432 ka. The glacial peak is associated with changes in vegetation (i.e., from more forested to more open landscape), reduction of humidity and water availability (i.e., moisture depletion, increased evaporation). Hominins are present at the Marathousa 1 location at the end of this interval (434-432 ka), confirming that the Megalopolis Basin served as a refugium for hunter-gatherer groups during periods of harsh climatic conditions. Additionally, the progressive cooling is associated with an important sedimentary hiatus between 465-440 ka reflected in all circum-Mediterranean records (both marine and continental), indicating a regional impact of the MIS 12 glaciation over surface processes.

Despite extensive archaeological research, our knowledge of the human population history of Upper Paleolithic Europe remains limited, primarily due to the scarce availability and poor molecular preservation of fossil remains. As teeth dominate the fossil record and preserve genetic signatures in their morphology, we compiled a large dataset of 450 dentitions dating between ~47 and 7 thousand years ago (ka), outnumbering existing skeletal and paleogenetic datasets. We tested a range of competing demographic scenarios using a coalescent-based machine learning Approximate Bayesian Computation (ABC) framework that we modified for use with phenotypic data. Mostly in agreement with but also challenging some of the hitherto available evidence, we identified a population turnover in western Europe at ~28 ka, isolates in western and eastern refugia between ~28 and 14.7 ka, and bottlenecks during the Last Glacial Maximum. Methodologically, this study marks the pioneering application of ABC to skeletal phenotypes, paving the way for exciting future research avenues.

The evolution of the human hand is a topic of great interest in paleoanthropology. As the hand can be involved in a vast array of activities, knowledge regarding how it was used by early hominins can yield crucial information on the factors driving biocultural evolution. Previous research on early hominin hands focused on the overall bone shape. However, while such approaches can inform on mechanical abilities and the evolved efficiency of manipulation, they cannot be used as a definite proxy for individual habitual activity. Accordingly, it is crucial to examine bone structures more responsive to lifetime biomechanical loading, such as muscle attachment sites or internal bone architecture. In this study, we investigate the manual entheseal patterns of Australopithecus afarensis, Australopithecus africanus, and Australopithecus sediba through the application of the validated entheses-based reconstruction of activity method. Using a comparative sample of later Homo and three great ape genera, we analyze the muscle attachment site proportions on the thumb, fifth ray, and third intermediate phalanx to gain insight into the habitual hand use of Australopithecus. We use a novel statistical procedure to account for the effects of interspecies variation in overall size and ray proportions. Our results highlight the importance of certain muscles of the first and fifth digits for humanlike hand use. In humans, these muscles are required for variable in-hand manipulation and are activated during stone-tool production. The entheses of A. sediba suggest muscle activation patterns consistent with a similar suite of habitual manual activities as in later Homo. In contrast, A. africanus and A. afarensis display a mosaic entheseal pattern that combines indications of both humanlike and apelike manipulation. Overall, these findings provide new evidence that some australopith species were already habitually engaging in humanlike manipulation, even if their manual dexterity was likely not as high as in later Homo.

An intensive, target -oriented surface survey conducted in the Megalopolis basin during 2012-2013 led to the discovery of several Palaeolithic sites and findspots with lithics and faunal remains, including Marathousa-1, a Lower Palaeolithic open-air elephant -butchering site, dated to ca. 400-500 ka BP. This study presents the results from the techno-typological analysis of 413 lithic artefacts collected as surface or stratified finds during the survey research. The aim of the work is to evaluate the diachronic occupation of the area in relation to the lithic technologies used, the typology of the artefacts, and the raw materials exploited. It was possible to provide a chrono-cultural attribution of 167 artefacts, ranging from the Lower Palaeolithic to the Holocene, with a significant component of the collection attributed to the Middle Palaeolithic. Several diagnostic retouched tools allow us to infer that the open-air sites were occupied comparatively more intensively during the Middle Palaeolithic period. We also provide a detailed account of the lithic assemblage from Kavia cave, a previously unstudied site identified during the survey. The high frequency of artefacts pertaining to the Upper Palaeolithic in Kavia is in line with previously identified settlement and mobility patterns of the Peloponnese, where the occupation of caves becomes more intensive from the Upper Palaeolithic onward, as attested at the sites of Klissoura, Kephalari, and Franchthi. The Upper Palaeolithic component from Kavia Cave adds new data to a meagre sample of known sites from this period. The results from the typological and technological analysis of the lithic assemblages collected during the survey in Megalopolis support the conclusions of the basin's long-term and ongoing research, indicating a relatively continuous hominin presence during the Pleistocene.

Recent investigations in the upper Lower–Middle Pleistocene deposits of the Megalopolis Basin (Greece) led to the discovery of several sites/findspots with abundant faunal material. Here, we provide an updated overview including new results on the micro- and macro-mammal fauna. Important new discoveries comprise partial hippopotamus skeletons from Marathousa 1 and the new Lower Pleistocene site Choremi 6, as well as a second partial elephant skeleton from Marathousa 1, including a complete tusk and the rarely found stylohyoideum. Based on the first results from the newly collected micromammals, we discuss age constraints of the sites, and we provide biostratigraphic/biochronologic remarks on key mammal taxa for the Middle Pleistocene of Greece and southeastern Europe. The presence of mammals highly dependent on freshwater for their survival, together with temperate-adapted ones in several stratigraphic layers of the basin, including those correlated with glacial stages, when conditions were colder and/or drier, indicate the capacity of the basin to retain perennial freshwater bodies under milder climatic conditions, even during the harsher glacial periods of the European Middle Pleistocene, and further support its refugial status. Yet, the smaller dimensions of the Megalopolis hippopotamuses may represent a response to the changing environmental conditions of the epoch, not optimal for hippopotamuses. Overall, the Megalopolis Basin comprises a unique fossil record for southeastern Europe and provides valuable insights into the Middle Pleistocene terrestrial ecosystems of Europe, and hominin adaptations in particular.

The early Iron Age (800 to 450 BCE) in France, Germany and Switzerland, known as the 'West-Hallstattkreis', stands out as featuring the earliest evidence for supra-regional organization north of the Alps. Often referred to as 'early Celtic', suggesting tentative connections to later cultural phenomena, its societal and population structure remain enigmatic. Here we present genomic and isotope data from 31 individuals from this context in southern Germany, dating between 616 and 200 BCE. We identify multiple biologically related groups spanning three elite burials as far as 100 km apart, supported by trans-regional individual mobility inferred from isotope data. These include a close biological relationship between two of the richest burial mounds of the Hallstatt culture. Bayesian modelling points to an avuncular relationship between the two individuals, which may suggest a practice of matrilineal dynastic succession in early Celtic elites. We show that their ancestry is shared on a broad geographic scale from Iberia throughout Central-Eastern Europe, undergoing a decline after the late Iron Age (450 BCE to similar to 50 CE).

The Kocabaş specimen comes from a travertine quarry near the homonymous village in the Denizli basin (Turkey). The specimen comprises three main fragments: portions of the right and left parietal and left and right parts of the frontal bone. The fossil was assumed to belong to the Homo erectus s.l. hypodigm by some authors, whereas others see similarities with Middle Pleistocene fossils (Broken Hill 1/Kabwe, Bodo, or Ceprano). Here, we present the first attempt to make a complete reconstruction of the missing medial portion of the frontal bone and a comprehensive geometric morphometric analysis of this bone. We restored the calotte by aligning and mirroring the three preserved fragments. Afterward, we restored the missing portion by applying the thin-plate spline interpolation algorithm of target fossils onto the reconstructed Kocabaş specimen. For the geometric morphometric analyses, we collected 80 landmarks on the frontal bone (11 osteometric points, 14 bilateral curve semilandmarks, and 41 surface semilandmarks). The comparative sample includes 21 fossils from different chronological periods and geographical areas and 30 adult modern humans from different populations. Shape analyses highlighted the presence in Kocabaş of features usually related to Middle Pleistocene Homo, such as a developed supraorbital torus associated with a relatively short frontal squama and reduced post-toral sulcus. Cluster analysis and linear discriminant analysis classification procedure suggest Kocabaş being part of the same taxonomic unit of Eurasian and African Middle Pleistocene Homo. In light of our results, we consider that attributing the Kocabaş hominin to H. erectus s.l. may be unwarranted. Results of our analyses are compatible with different evolutionary scenarios, but a more precise chronological framework is needed for a thorough discussion of the evolutionary significance of this specimen. Future work should clarify its geological age, given uncertainties regarding its stratigraphic provenance.

In this study we present the results of the archaeozoological and taphonomic analysis of the macromammal assemblage from the Uluzzian levels of three Italian sites: Uluzzo C Rock Shelter, Roccia San Sebastiano cave, and Riparo del Broion. Our archaeozoological data outlined several common elements and differences between sites sharing this culture. The most abundant taxon registered at Roccia San Sebastiano cave is Cervus elaphus; for Riparo del Broion we registered mostly Ursus sp., but we found anthropogenic modifications on different bones of Cervus elaphus. At Uluzzo C, the most abundant taxon is Vulpes vulpes and the most hunted is Cervus elaphus. Anthropogenic modifications are observed mostly on limb bones (long bones, phalanges, and sesamoids) attesting the practice of marrow extraction. We outlined hunting strategies against carnivore species in particular for northern Italy (Riparo del Broion). In southern Italy, the fauna exploitation is focused mostly on cervids and equids, as attested for several coeval Italian sites (Castelcivita, Grotta del Cavallo, Grotta della Cala, and Grotta La Fabbrica). The data obtained provide new relevant information about animal resource exploitation and allow us to improve and complete our understanding of the Uluzzian hunting and subsistence strategies in the Mediterranean area.

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.

The use of reindeer has been a crucial element in the subsistence strategies of past Arctic and Subarctic populations. However, the spatiotemporal occurrence of systematic herding practices has been difficult to identify in the bioarchaeological record. To address this research gap, this study proposes a new virtual anthropological approach for reconstructing habitual physical activity in reindeer, relying on the protocols of the “Validated Entheses based Reconstruction of Activity” (VERA) method. Following blind analytical procedures, we focused on eight muscle attachment sites (“entheses”) in 36 reindeer free ranging in the wild, 21 specimens in captivity (zoo), and eight racing reindeer (habitual runners). Importantly, our analyses accounted for the effects of variation by subspecies, sex, age, and estimated body size. Our results showed clear differences across activity groups, leading to the development of discriminant function equations with cross-validated accuracies ranging from approximately 88 to 100%. The reliability of our functions was additionally confirmed using a blind test involving six zoo individuals not included in the initial dataset. Our findings support the use of the proposed approach for identifying domestication-related activities in zooarchaeological contexts, introducing a valuable tool for locating suspected domestication hotspots and elucidating the nature of past human-reindeer interactions.

Objectives This study uses a virtual framework to examine the left maxillary fragment of the juvenile fossil from Mugharet el'Aliya, Morocco, found in association with an Aterian lithic industry. Previously, this fossil had been ascribed to modern humans or the Neanderthal lineage based on its "archaic"/"Neanderthal-like" features and apparent large size. Here, we conducted a novel 3D shape comparative analysis of the maxillary fragment to clarify its taxonomic affinities with regard to its size and ontogeny. Materials and Methods Eighty Computed Tomography and surface scans representing ontogenetic samples of Homo sapiens and Homo neanderthalensis were used to capture species-specific differences. The toolkit of geometric morphometrics in combination with surface registration and an elastic iterative closest point algorithm were used to create a dataset of meshes with an identical number of corresponding vertices for the maxillae. Multivariate statistics were applied to Procrustes superimposed coordinates derived from the vertices of this dataset. Results Our analysis showed affinities of the Mugharet el'Aliya individual with our H. sapiens sample, especially with a subadult individual from Qafzeh. No size-independent affinities with Neanderthals of comparable dental age could be identified. Discussion Our results add to the evidence connecting fossils from western Asia, especially Qafzeh and Skhul, and the North African Aterian. Furthermore, Mugharet el'Aliya adds to our knowledge of the ontogenetic development of adult morphology that is frequently used to characterize hominin groups, for example, Neanderthals and modern humans.