Scavenging has been a key topic in human evolution for decades, mainly focused on the 'hunting vs. scavenging' debate and the role of meat in the diet of early hominins. Scavenging is frequently considered a marginal activity by archaeologists; however, recent theoretical developments and experimental observations in the field of carrion ecology suggest that this is a misconception that should be addressed. Here, we analyze hominin scavenging in the framework of optimal foraging theory and compare the assumptions made in archaeology and paleoanthropology based on current ecological knowledge. We discuss the constraints and opportunities for hominins as facultative scavengers across ecological contexts, both in Africa and beyond. Hominins exhibited several anatomical, physical, and behavioral adaptations that enabled them to detect carcasses from a long distance, reach them relatively quickly, confront other scavengers, if necessary, and process the carcass to obtain food. Carrion should be considered a high-quality resource that is ubiquitous and more predictable than previously assumed. Particularly relevant is the abundant carrion from sources other than predation, especially from megaherbivores. Moreover, hominins likely benefited from carrion partitioning strategies and interspecific facilitation-especially with vultures-which have traditionally been overlooked in archaeological and paleoanthropological research. Thus, we propose that carrion was a valuable resource exploited by all hominin species and populations to varying degrees, especially during periods of food shortage in seasonal environments. We conclude that the 'hunting vs scavenging' dichotomy is no longer supported as hominins are omnivorous, exploiting animal food through hunting or scavenging depending on environmental conditions and their technological and cognitive capacities.

The Sima del Elefante cave is one of the archaeopaleontological sites located in the karstic complex of Sierra de Atapuerca (Burgos, Spain), which is a remarkable locality for documenting an extensive sequence of human evidence. Within the stratigraphic sequence of the Sima del Elefante site, Level TE7 has yielded one of the oldest human remains recovered to date in western Europe. A hominin mid-face has recently been discovered (specimen ATE7-1), attributed to Homo aff. erectus. In this study, the fossil amphibian and reptile remains directly associated with this hominin are analyzed, described, and used to perform quantitative paleoecological reconstructions. The resulting herpetofaunal assemblage comprises a total of eight species: five anurans (Alytes gr. Alytes obstetricans/Alytes almogavarii, Pelodytes punctatus, Bufo gr. Bufo bufo, Epidalea calamita, and Rana temporaria), two lizards (small-sized Lacertidae indet. and Anguis fragilis), and two snakes (Coronella cf. Coronella austriaca and Vipera sp.). The paleoecological reconstruction was performed using the Mutual Ecogeographic Range method in conjunction with the Uncertain Distribution Area-Occupied Distribution Area technique to obtain temperature and precipitation estimates, and habitat weighting was used to infer the surrounding environment. The climate of TE7 was reconstructed as cool and humid, with an overall pattern concordant with a present-day continental Mediterranean climate, exhibiting similar temperatures and higher precipitation than today for the Sierra de Atapuerca location. The associated landscape evidenced a more humid environment, primarily made up of woodlands and open humid habitats with aquatic and periaquatic areas. The results obtained provide novel insights into the ecological tolerances of early hominins and thus contribute to the growing body of knowledge regarding their subsistence capabilities and behaviors.

The Omo-Turkana Basin is one of three major regions for the study of hominin evolution in Africa. It has yielded a rich hominin fossil record of 1231 specimens, around a third of the record for the whole of Africa for the period from the Messinian through the Calabrian. Here, we consider the fossil hominin record of the Omo-Turkana Basin as an object of study in its own right and show the contribution that an analysis of such an exhaustive record can make. The data come from 117 publications allowing the most complete, accurate, and up-to-date synthesis of this record. Our analysis provides a quantitative perspective on the biases affecting this record, such as skeletal element abundance representation, chronostratigraphic distribution, and difficulties in taxonomic assignment. It also provides historical perspective, illustrating the major contribution made by the Omo-Turkana hominin fossil record to our knowledge of human evolution. We provide a synthetic overview of the taxa represented and discuss the chronological distribution of taxonomic groups in the basin including the relative abundance of Paranthropus and Homo (2/3 and 1/3, respectively) during their long period of coexistence. Integrating the data makes it possible to address difficult questions that have been underinvestigated until now. For example, contrary to the prevailing view, the genus Homo is well represented in the Omo-Turkana Basin between 2.7 and 2 Ma. Additionally, we show that the hominin fossil record of the Upper Burgi and KBS Members is atypical, both in terms of skeletal element abundance and taxonomy. Neither paleoenvironments nor taphonomic or collecting biases can fully explain this anomaly.

Chasing prey to exhaustion on foot-persistence hunting-has been hypothesized to play a key role in the evolution of human endurance, energetics, and thermoregulation. However, direct physiological data from such pursuits are lacking, particularly from open arid environments where persistence hunting may have originated. We report the first physiological measurements from an unacclimatized ultrarunner and hunter, accompanied by three acclimatized hunters, during six persistence pursuits of Oryx gazella in the Namib desert. The pursuits spanned 49.5 km on foot in 7.8 hours over 3 days. Two pursuits were successful: A healthy oryx was caught after 2 hours (31% of time running), and an injured oryx after 1 hour of walking only. Over the 3 days, persistence hunting yielded 1882-3727 kcal per man-hour, surpassing net return rates reported for hunter-gatherers and horticulturalists and net energy equivalent to the daily needs of 50 active individuals. Daily energy expenditure (5024 kcal day) and water turnover (11.4 l day) were high but within ranges reported for subsistence populations in hot climates. Water loss remained below the critical dehydration threshold. Despite high air (36-41 °C) and ground (43-60 °C) temperatures, the hunter maintained a safe core temperature (≤39 °C), while the oryx reached hyperthermic levels (44 °C). These findings suggest that persistence hunting in arid environments is energetically viable and thermally manageable, even for unacclimatized individuals. The relatively low proportion of running-and its complete absence when pursuing disadvantaged prey-supports the potential feasibility of persistence hunting even among hominins without advanced endurance running adaptations.

The locomotor behavior and substrate preferences of early colobines are a matter of debate. Some researchers advocate a partly terrestrial origin with the subsequent parallel evolution of arboreality in different clades during the Plio-Pleistocene, while others suggest that arboreality had already evolved earlier in the Miocene. Microcolobus, from the Late Miocene of Nakali (ca. 10 Ma, Kenya), is one of the earliest known and, for the most part, best-documented African Miocene fossil colobines. A large collection of postcranial remains was recovered at Nakali during the fieldwork of the Joint Japan-Kenya Paleontological Expedition, permitting us to test hypotheses regarding the locomotor behavior, substrate preferences, and taxonomy of the Nakali colobines. Here, we present novel interpretations of the functional adaptations of the elbow anatomy of previously published partial skeletons from Nakali (KNM-NA 47916 and KNM-NA 47915) and the description of 12 additional humeral, ulnar, and radial specimens that we attributed to Microcolobus. The elbow specimens from Nakali are morphologically homogenous except one humeral specimen that demonstrates equivocal evidence of functional and taxonomical diversity. Quantitative and qualitative analyses confirm arboreal adaptations of Microcolobus, corroborating hypotheses supporting early adaptations for arboreal locomotor substrate preferences in colobines. The elbow anatomy of Microcolobus can be qualitatively and quantitatively distinguished from that of the Late Miocene colobines Cercopithecoides bruneti and Paracolobus enkorikae. Microcolobus also present several anatomical features seen in the small African colobine Procolobus verus, reflecting a frequently flexed elbow with moderate pronosupination abilities and a stable humeroulnar joint consistent with the use of flexed-elbow climbing and arboreal quadrupedal running.

The appearance of the Protoaurignacian in Europe around 42,000 years ago is widely believed to result from a major dispersal of anatomically modern Homo sapiens out of the Levant, a view primarily supported by perceived similarities between Mediterranean Protoaurignacian and Levantine Ahmarian stone tools. However, no quantitative technological comparison has yet thoroughly tested this connection. Here, we present the first systematic evaluation of lithic technology from Protoaurignacian assemblages in Italy and from the northern Ahmarian and post-Ahmarian layers at the reference sequence of Ksar Akil (Lebanon). Using attribute analysis and multivariate statistics, we assessed technological similarities and differences across different stages of the core reduction sequence. Our results demonstrate very limited affinities and distinct technological trajectories between the two regions. While the northern Ahmarian at Ksar Akil is characterized by bidirectional volumetric core reduction aimed at blade production, the Protoaurignacian exhibits a strong emphasis on bladelet production from unidirectional cores. Although lithic miniaturization trends are observed in both regions, the post-Ahmarian layers at Ksar Akil primarily produced twisted bladelets from burins and carinated cores-a feature uncommon in the Protoaurignacian. These findings challenge the hypothesis of a Levantine origin for the Protoaurignacian and, more broadly, suggest that technological convergence-driven by the growing importance of multicomponent projectile technology and increased mobility-played a central role. Thus, our study underscores the need to reconsider diffusionist explanations and emphasizes the central role of internal cultural innovation among foraging groups settled in different regions of the Old World in shaping the emergence of the Upper Paleolithic.

The onset of the Upper Paleolithic period in Eurasia is marked by the systematic production of pointed blades, commonly associated with the Marine Isotope Stage 3 expansion of modern humans. Consequently, many studies have concentrated on the geographical origins and mechanisms of spread of these blade industries across Eurasia, while comparatively less attention has been devoted to regional cultural dynamics and adaptive strategies. This research focuses on the initial stages of the Upper Paleolithic in the Negev Desert, a key region in the Levant that served as a major crossroads between Africa and Eurasia. The study integrates new cultural and environmental data from the Boker sites, originally excavated by Marks and re-excavated in 2015-2016. New radiocarbon (C) and optically stimulated luminescence chronologies indicate that the Boker sites were occupied over a prolonged period, from approximately 45,000 to 30,000 calibrated years before present (cal BP). The earliest occupations at Boker are attributed to the Initial Upper Paleolithic (IUP), with later phases associated with the Early Upper Paleolithic Ahmarian technocomplex. Lithic assemblages from the Boker sites reveal a consistent use of unidirectional blade technology, suggesting that the desert Ahmarian industry evolved locally from the IUP. However, a notable shift is observed in percussion techniques, transitioning from the use of hard hammerstones in the IUP to soft hammerstones in the Early Upper Paleolithic. This technological change is interpreted as a result of cultural diffusion from the Mediterranean woodland Ahmarian, which predates its southern counterpart. Paleoenvironmental evidence from the Boker sites indicates that conditions were more favorable than those of the present-day Negev, as reflected by the presence of vegetation and water sources. These factors likely contributed to the repeated Upper Paleolithic occupations of this relatively small area. Charred archaeobotanical remains reveal a mix of drought-tolerant halophytic species and relics of Mediterranean thermophilous plants, suggesting a cooler and wetter climate. The discovery of dark, organic-rich layers near the Early Ahmarian occupation at Boker A supports the hypothesis that a salt pan existed in proximity to a freshwater source-an environmental setting that would have been attractive to both humans and animals.

Exploring the paleoecology of extinct species of the genus Theropithecus, which was widespread and diversified in Africa during the Pliocene and the Pleistocene, is crucial to have a full picture of the evolutionary history of this taxon. It also gives insights into the fundamental ecological range of the genus, now restricted to the Ethiopian highlands in a reduced set of habitats. This study aims to investigate the diet of Theropithecus atlanticus (n = 8) from the Plio-Pleistocene site of Ahl al Oughlam (Morocco) through dental microwear texture analysis. We used a recently developed software program, 'trident,' that quantifies the heterogeneity of texture parameters through subsampled wear surfaces to improve dietary discriminations. We considered a comparative dataset of extant primates (n = 104) with different diets, including the leaf-eater Colobus guereza, the grass-eater Theropithecus gelada, and the generalists Papio hamadryas and Chlorocebus aethiops. In addition, we compared with specimens of theropiths (n = 80) from the eastern African Shungura Formation (Lower Omo Valley, Ethiopia) that are nearly contemporaneous to T. atlanticus. Our results indicate that T. atlanticus was mostly feeding on grasses, as do its extant relatives. Some similarities with opportunistic species suggest it might have consumed a substantial amount of challenging foods. This dental microwear texture is in line with open, arid, and harsh environments and the presence and abundance of C3 grasses at Ahl al Oughlam detected by enamel stable isotope analysis on various mammals including theropiths. This contrasts with the environment described on nearly contemporaneous specimens from the Lower Omo Valley where grasses were abundant but composed of C4 ones.

Functional interpretation of fossils documenting our early locomotor evolution relies on biomechanical analyses of modern humans and comparative samples of nonhuman primates. Force plate studies have explored the forces that nonhuman primate limbs exchange with the ground while walking bipedally or quadrupedally, and kinematic studies have detailed limb and joint motion profiles during various forms of locomotion. Much less is known about patterns of hind limb muscle use as revealed through electromyography despite the fact that the interpretation of many features of fossils has been related to aspects of muscle use. To better understand how muscle function may relate to the interpretation of fossil material, the current study focuses on chimpanzees and presents electromyography-based activity profiles for the majority of their hind limb muscles during knuckle-walking, bipedalism, and vertical climbing. Taking advantage of the long history of the Stony Brook Primate Locomotion Lab, this study has compiled electromyographic data from eight chimpanzee subjects to both document hind limb muscle activity patterns and explore variability in those recruitment profiles. The collected data indicate that while there are many commonalities in patterns of muscle use, there is also a fair amount of intersubject variation. Overall, the locomotor adaptive strategy of chimpanzees appears to emphasize versatility and maneuverability rather than energy-efficient locomotion. It is hoped that these data will broaden perspectives on how patterns of muscle use may influence the interpretation of fossils.

Japanese macaques (Macaca fuscata) are capable of bipedal locomotion but exhibit fundamental differences from human walking, including a flexed limb posture and a single-peaked ground reaction force profile. One key factor underlying these differences is their limited hip extension, likely constrained by muscle architecture. This study examines the anatomical restrictions on hip extension in Japanese macaques and spider monkeys (Ateles spp.), which achieve greater hip extension during bipedal walking. We measured passive joint moments before and after the sequential dissection of hip flexor muscles in cadaveric specimens using a custom-built measurement device to quantify passive hip joint resistance across different muscle groups. Our results reveal species-specific differences in hip mobility: Japanese macaques exhibit greater passive resistance to hip extension, primarily due to the adductor muscle group and joint capsule, whereas spider monkeys show lower resistance, with the iliopsoas playing a more substantial role in restricting hip extension. The findings emphasize the role of passive elastic muscle constraints in shaping primate bipedalism and offer comparative insights into the evolution of human locomotion.

Known for over a century, the Late Miocene mammalian faunas of Veles, North Macedonia, have long been recognized for their scientific importance. However, hominid remains had not been previously reported from this fossil-rich area. Here, we report the discovery of an isolated upper molar from the vicinity of Veles-most likely from the Belushka locality-which constitutes the first known record of a Late Miocene hominid from the Republic of North Macedonia, and provide a review of the associated mammalian assemblages. The molar-probably an M-exhibits a relatively large crown, broad and low cusps, inferred thick enamel, low dentine horns, and short, thick roots that are not markedly divergent. These characteristics, along with its inferred middle Turolian (MN12) age and Balkan provenance, suggest that the Veles molar belongs to an indeterminate member of the hominid tribe Graecopithecini. Together with the Graecopithecus freybergi mandible from Pyrgos Vassilissis, Greece, and an isolated graecopithecin upper premolar from Azmaka, Bulgaria, the Veles specimen provides rare additional evidence for the survival of apes well into the Turolian of the Balkan Peninsula. Furthermore, the Veles fossil faunas now document the co-occurrence of apes (Graecopithecini indet.) and monkeys (Mesopithecus), a pattern rarely observed in the Eurasian Miocene fossil record.

Denisovans contributed notably to the genomes of present-day East and Southeast Asians. However, the relationship between the inhabited paleohabitats and the adaptive genetic traits related to infections in modern humans remains underexplored. This study uses geospatial techniques to analyze climatic factors associated with three Denisovan archaeological sites linked to nine specimens. Additionally, past and present climates and biomes, as well as the geographic distributions of eight infectious agents and disease vector groups, were modeled and compared with the modern genetic heritage of Denisovans. Findings reveal that the identified Denisovans inhabited subarctic and monsoon-influenced temperate climates, occupying boreal and seasonal forest biomes in the three studied archaeological sites. Sites such as Denisova Cave and Baishiya Karst Cave exhibited low climatic suitability for Ascaris lumbricoides, visceral leishmaniasis, and Nipah virus. Plasmodium vivax- and Aedes albopictus-like vectors plausibly were also not present. Conversely, Denisova Cave and Baishiya Karst Cave exhibit high climatic suitability for Ixodes persulcatus and Lyme borreliosis when Denisovans inhabited these sites. The paleoenvironment of the Laotian Cobra Cave site-with the exception of Nipah henipavirus-was suitable for all modeled pathogens and vectors. From the studied vectors and diseases, I. persulcatus and Lyme borreliosis are missing from Melanesia, where the region's humans have the highest Denisovan legacy. This suggests that Denisovans from humid continental climates, such as those near Cobra Cave, may have contributed alleles providing adaptive advantages against ascariasis and mosquito-borne diseases in environments where modern human populations with high Denisovan genetic legacy reside.

The timing of Homo erectus and Acheulean emergence in the Early Pleistocene is important to the understanding of the evolution of the genus Homo but is currently insufficiently resolved. Relevant chronologies are often based on a combination of radioisotopic dates and other age indicators such as magnetostratigraphy and biochronology. Here, we report a newly recognized normal magnetozone at ∼1.6 Ma in the Konso Formation, southern Main Ethiopian Rift, Ethiopia. This magnetozone occurs at one of the Konso Fm localities, KGA19, that spans the >1.75- to <1.45-Ma time period otherwise ill-represented in the Formation. We describe the lithostratigraphy and tephrostratigraphy of the KGA19 sedimentary sequence and demonstrate that the Konso Fm Kayle Tuff-1 and the KGA19 Bench Tuff are correlative to the Turkana Basin Orange and Morutot tuffs, respectively. Paleomagnetic analyses of the western sector of KGA19 revealed a normal polarity interval at ∼1.6 Ma, with its age based on Ar/Ar dates and tephrostratigraphic correlations. This suggests that the KGA19 normal magnetozone represents the Gilsá excursion otherwise documented in limited marine cores, Chinese Loess sequences, and few lava flows. A review of these records and sediment accumulation rates of fossil-bearing sequences of eastern Africa suggests that short events such as the Gilsá excursion would be detected if sampling resolution is sufficiently high relative to sedimentation rates. In light of these findings and considerations, evaluation of the Melka Kunture magnetostratigraphy suggests that the age of the Garba IV H. erectus and Acheulean is ∼1.6 Ma, not >1.95 to ∼2 Ma.

Neandertals are known to have occupied Eurasia for over 250,000 years and were well adapted to the environmental conditions of the last ice ages. However, the dietary habits of these ancient humans remain debated, with conflicting evidence suggesting that they may have been primarily carnivorous, omnivorous, or even herbivorous. Traditional isotope analyses have provided some insights into Neandertal diets, but their limitations (preservation and baseline effect) have sparked the need for new approaches. These limitations are well known on the Iberian Peninsula, and while recent results of carbon, oxygen, radiogenic strontium, and zinc isotopes and trace element ratios allow the reconstruction of the Gabasa (Spain) Neandertal diet, some questions still remain unanswered. Our study explores the potential of using calcium and stable strontium isotopes (δCa and δSr, respectively) to supplement previous analyses performed on the Gabasa Neandertal. Based on the low δCa and δSr values observed for the Neandertal specimen, our results suggest a hypercarnivorous diet that included low but non-negligible quantities of bone, although it is not possible to rule out the possible impact of milk consumption. Overall, our work argues that Ca, Sr, and Zn stable isotopes can supplement conventional isotope studies and offer a more comprehensive picture of human diets, including that of Neandertals.

The coevolution of the hands and feet in modern humans has been a subject of significant interest due to their unique morphological features that differentiate humans from other primates and their implications in human evolution. This study aims to investigate the degree of correlated responses to selection between hands and feet and to determine whether one of the autopods has exerted a greater influence on this coevolution, focusing on their homologous elements and morphological traits. We analyzed the 38 long bones of the hands and feet from 96 modern human specimens, employing a comprehensive methodological framework that includes morphological analysis, assessments of modularity, integration, and covariation patterns under random selection. Additionally, Bayesian analyses were conducted to test whether foot morphology drives hand morphology or vice versa. Our findings indicate a high degree of morphological integration between the hands and feet, revealing a trend of increasing correlation from the first to the fifth ray. Consistent with previous studies, our Bayesian model provides robust evidence that the feet drive the morphological coevolution of human autopods, likely in response to functional selection pressures associated with bipedalism. However, our results also highlight that the intertwined evolutionary trajectories of the hands and feet are not a simple unidirectional model, underscoring the complexity of morphological integration and the diverse coevolutionary patterns among different rays, reflecting their specialized functions and evolutionary adaptations.

The Olduvai Gorge Naisiusiu Beds and Ndutu Beds are significant for understanding the cultural and biological evolution of Homo sapiens. However, the timing and span of deposition of these beds is poorly understood. We present a chronology based on luminescence dates for sedimentary drill core samples and one ostrich eggshell (OES) radiocarbon date from the Naisiusiu type section outcrop. The Naisiusiu Beds type section rests on the Olduvai protogorge floor, on a post-Ndutu incisional surface that eroded through older Olduvai Beds (∼80 m) into the top of the Bed I Basalt. The estimated minimum thickness of the Naisiusiu Beds within the gorge is >13.3 m. Three lithostratigraphic units were identified in the 9.2-m-thick type section: The lower unit comprises alternating fluvial/hyperconcentrated flow sediments and yields an OES date of 49,728 ± 1378 cal year BP; the middle unit is composed of predominantly fluvial sediments containing Later Stone Age (LSA) artifacts and faunal remains, dating between 34.2 ± 2.8 ka and 24.0 ± 2.0 ka; and the upper unit comprises volcaniclastic hyperconcentrated flow deposits with interstratified carbonate-cemented surfaces, dating between 24.0 ± 2.0 ka and 19.6 ± 1.6 ka. Slower accretion rates of the middle unit (7.3 cm/ka) are associated with repeated fluvial cutting and filling. Upper Naisiusiu stacks of hyperconcentrated flows and intervening hiatuses have faster accretion rates (54.9 cm/ka). The OES radiocarbon date of 49,728 ± 1378 cal year BP (modeled age = 49,498 ± 2385 BP) from the top part of the lower unit indicates that the type section extends back to >50 ka. The LSA assemblage is associated with sediments dated to 34.2 ± 2.8 ka using luminescence. A date of 62.7 ± 5.6 ka on the youngest Middle Stone Age occurrence in the underlying Ndutu Beds at Type Locality 26 provides a maximum age for the Middle Stone Age/LSA transition at the Olduvai Gorge.

Assigning an age to the nearly complete cranium found in the Petralona Cave in Greece is of outstanding importance because this fossil has a key position in European human evolution. This topic has been debated since its discovery more than 60 years ago, highlighting the difficulties in applying physical dating methods to prehistoric samples. Previous results obtained on various types of samples yielded a large age range between about 170 and 700 ka, precluding any consensus on the age of the human fossil. On the other hand, the original stratigraphic position of the cranium also remains enigmatic, in spite of all the efforts provided by various researchers. Here, we present new U-series dates performed on the calcite that grew directly on the cranium, which is the only sample able to provide crucial information on the age of the fossil. The results yield a finite age suggesting that the Petralona cranium has a minimum age of 286 ± 9 ka. Other speleothems and calcitic coatings were sampled in three main locations in the cave; among them, samples came from the 'Mausoleum' where the cranium was supposedly found cemented to a wall. The data show that the calcite covering the cranium is not contemporaneous with that of the Mausoleum wall, despite what was previously thought. The different possibilities, depending on whether or not the cranium was attached to the wall, are discussed in the paper. From a morphological point of view, the Petralona hominin forms part of a distinct and more primitive group than Homo sapiens and Neanderthals, and the new age estimate provides further support for the coexistence of this population alongside the evolving Neanderthal lineage in the later Middle Pleistocene of Europe.