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Noncoding DNA is central to our understanding of human gene regulation and complex diseases1,2, and measuring the evolutionary sequence constraint can establish the functional relevance of putative regulatory elements in the human genome3,4,5,6,7,8,9. Identifying the genomic elements that have become constrained specifically in primates has been hampered by the faster evolution of noncoding DNA compared to protein-coding DNA10, the relatively short timescales separating primate species11, and the previously limited availability of whole-genome sequences12. Here we construct a whole-genome alignment of 239 species, representing nearly half of all extant species in the primate order. Using this resource, we identified human regulatory elements that are under selective constraint across primates and other mammals at a 5% false discovery rate. We detected 111,318 DNase I hypersensitivity sites and 267,410 transcription factor binding sites that are constrained specifically in primates but not across other placental mammals and validate their cis-regulatory effects on gene expression. These regulatory elements are enriched for human genetic variants that affect gene expression and complex traits and diseases. Our results highlight the important role of recent evolution in regulatory sequence elements differentiating primates, including humans, from other placental mammals.

Personal ornaments are widely viewed as indicators of social identity and personhood. Ornaments are ubiquitous from the Late Pleistocene to the Holocene, but they are most often found as isolated objects within archaeological assemblages without direct evidence on how they were displayed. This article presents a detailed record of the ornaments found in direct association with an Early Mesolithic buried female infant discovered in 2017 at the site of Arma Veirana (Liguria, Italy). It uses microscopic, 3D, and positional analyses of the ornaments as well as a preliminary perforation experiment to document how they were perforated, used, and what led to their deposit as part of the infant’s grave goods. This study provides important information on the use of beads in the Early Mesolithic, in general, as well as the relationship between beads and young subadults, in particular. The results of the study suggest that the beads were worn by members of the infant’s community for a considerable period before they were sewn onto a sling, possibly used to keep the infant close to the parents while allowing their mobility, as seen in some modern forager groups. The baby was then likely buried in this sling to avoid reusing the beads that had failed to protect her or simply to create a lasting connection between the deceased infant and her community.

The archaeological data on the earliest hominin behavioral subsistence activities in North Africa are derived primarily from the Early Pleistocene site of Ain Boucherit (northeastern Algeria). Ain Boucherit consists of two archaeological layers, Ain Boucherit Upper (AB-Up) and Ain Boucherit Lower (AB-Lw), estimated to ~ 1.9 Ma and ~ 2.4 Ma, respectively. Cutmarked and hammerstone percussed bones associated with Oldowan stone tools were found in both layers, with AB-Lw yielding the oldest in North Africa. The faunal assemblages from both deposits are dominated by small-sized bovids and equids. Evidence of cutmarks and percussion marks in both assemblages shows that hominins exploited animal carcasses, involving skinning, evisceration and defleshing activities. The evidence of meat and marrow acquisition is more abundant at AB-Lw with carnivore activity being scarce. However, the AB-Up assemblage shows more carnivore damage and less hominin-induced tool marks. Ain Boucherit evidence, is similar, in type and chronology, to that provided by the Early Pleistocene sites in East Africa (e.g., the Gona sites), where the oldest evidence of stone tools used in faunal exploitation have been discovered. This paper reports on the ability of early North African Oldowans to compete successfully for accessing animal resources with other predators.

The production of sharp-edged stone flakes is often viewed as a unique adaptation specific to the hominin lineage. The discovery of large stone cores and flakes dated to 3.3 Ma has shown that the earliest evidence of hominin stone flake production differs from the later Oldowan (dated to 2.6 Ma) record. Recent discoveries have shown that capuchin monkeys (Sapajus libidinosus) can unintentionally produce sharp-edged flake assemblages as a by-product of percussive behaviour. Capuchin monkeys are the only wild primate to intentionally strike two stones together (stone-on-stone percussion) to produce quartz powder. In doing so, they unintentionally produce high frequencies of sharp-edged flakes. It has been suggested that these possess the same technological attributes commonly used to describe intentional hominin stone flakes. Here we directly compare the basic morphometric descriptions, technological and 3D morphometric properties of unintentional capuchin flakes with experimentally produced flakes derived from two intentional flaking techniques employed during the Plio-Pleistocene, freehand and passive hammer knapping and a sample of Oldowan freehand flakes from Olduvai Gorge (Tanzania). Our results show that unintentional stone-on-stone flakes overlap in terms of their shape, size and technological attributes with those produced through intentional knapping techniques. We, however, identify attributes that may be critical for differentiating between intentional knapping techniques documented in the early archaeological record and unintentional flakes. Using these data we show that Oldowan flakes more closely resemble intentional freehand techniques.

[INTRODUCTION] Millions of people have received genome and exome sequencing to date, a collective effort that has illuminated for the first time the vast catalog of small genetic differences that distinguish us as individuals within our species. However, the effects of most of these genetic variants remain unknown, limiting their clinical utility and actionability. New approaches that can accurately discern disease-causing from benign mutations and interpret genetic variants on a genome-wide scale would constitute a meaningful initial step towards realizing the potential of personalized genomic medicine., [RATIONALE] As a result of the short evolutionary distance between humans and nonhuman primates, our proteins share near-perfect amino acid sequence identity. Hence, the effects of a protein-altering mutation found in one species are likely to be concordant in the other species. By systematically cataloging common variants of nonhuman primates, we aimed to annotate these variants as being unlikely to cause human disease as they are tolerated by natural selection in a closely related species. Once collected, the resulting resource may be applied to infer the effects of unobserved variants across the genome using machine learning., [RESULTS] Following the strategy outlined above we obtained whole-genome sequencing data for 809 individuals from 233 primate species and cataloged 4.3 million common missense variants. We confirmed that human missense variants seen in at least one nonhuman primate species were annotated as benign in the ClinVar clinical variant database in 99% of cases. By contrast, common variants from mammals and vertebrates outside the primate lineage were substantially less likely to be benign in the ClinVar database (71 to 87% benign), restricting this strategy to nonhuman primates. Overall, we reclassified more than 4 million human missense variants of previously unknown consequence as likely benign, resulting in a greater than 50-fold increase in the number of annotated missense variants compared to existing clinical databases. To infer the pathogenicity of the remaining missense variants in the human genome, we constructed PrimateAI-3D, a semisupervised 3D-convolutional neural network that operates on voxelized protein structures. We trained PrimateAI-3D to separate common primate variants from matched control variants in 3D space as a semisupervised learning task. We evaluated the trained PrimateAI-3D model alongside 15 other published machine learning methods on their ability to distinguish between benign and pathogenic variants in six different clinical benchmarks and demonstrated that PrimateAI-3D outperformed all other classifiers in each of the tasks., [CONCLUSION] Our study addresses one of the key challenges in the variant interpretation field, namely, the lack of sufficient labeled data to effectively train large machine learning models. By generating the most comprehensive primate sequencing dataset to date and pairing this resource with a deep learning architecture that leverages 3D protein structures, we were able to achieve meaningful improvements in variant effect prediction across multiple clinical benchmarks.

Fathers contribute substantially to infant care, yet the mechanisms facilitating paternal bonding and interactions with infants are not as well understood as they are in mothers. Several hormonal changes occur as males transition into parenthood, first in response to a partner's pregnancy, and next in response to interacting with the newborn. These changes may prepare fathers for parenting and help facilitate and maintain paternal care. Experimental studies with monkeys and rodents suggest that paternal care requires elevated estradiol levels, which increase when a male's partner is pregnant and are higher in fathers than non-fathers, but its role in the expression of paternal behaviors throughout infant development is unknown. To assess estradiol's role in paternal care, we analyzed the relationship between paternal estradiol metabolites and 1) offspring age, and 2) paternal care behavior (holding, carrying, huddling, playing, grooming), in wild, red-bellied lemurs (Eulemur rubriventer). We collected 146 fecal samples and 1597 h of behavioral data on 10 adult males who had newborn infants during the study. Estradiol metabolites increased four-fold in expectant males, and in new fathers they fluctuated and gradually decreased with time. Infant age, not paternal behavior, best predicted hormone levels in new fathers. These results suggest that hormonal changes occur in expectant males with facultative paternal care, but they do not support the hypothesis that estradiol is directly associated with the day-to-day expression of paternal care. Future research should explore estradiol's role in facilitating behaviors, including infant-directed attention and responsiveness, or preparing fathers for infant care generally.

The rich diversity of morphology and behavior displayed across primate species provides an informative context in which to study the impact of genomic diversity on fundamental biological processes. Analysis of that diversity provides insight into long-standing questions in evolutionary and conservation biology and is urgent given severe threats these species are facing. Here, we present high-coverage whole-genome data from 233 primate species representing 86% of genera and all 16 families. This dataset was used, together with fossil calibration, to create a nuclear DNA phylogeny and to reassess evolutionary divergence times among primate clades. We found within-species genetic diversity across families and geographic regions to be associated with climate and sociality, but not with extinction risk. Furthermore, mutation rates differ across species, potentially influenced by effective population sizes. Lastly, we identified extensive recurrence of missense mutations previously thought to be human specific. This study will open a wide range of research avenues for future primate genomic research.

Palaeobiological and archeological excavations at the site of Ewass Oldupa, found in the western Plio-Pleistocene rift basin of Oldupai Gorge (also Olduvai Gorge), Tanzania, revealed rich fossiliferous levels and the earliest remains of human activity at Oldupai Gorge, dated to 2 million years ago. This paper provides zooarchaeological taxonomic, taphonomic, and behavioral analyses, applying several methods to explore the setting in which the assemblage was formed. We identified agency behind bone surface modifications, such as cut, tooth and percussion marks, and determined the frequency of carnivore tooth marks as well as their distribution on both discrete specimens and across species. In addition, our work revealed co-occurrence of modifications to include butchering marks and carnivore tooth marks. Ravaging levels were estimated as percentage. The faunal accumulation from Ewass Oldupa contains two cut marked specimens, together with low degrees of percussion and carnivore tooth marks, moderate ravaging, and diagenetic changes suggestive of water flow. Thus, multiple lines of evidence indicate a palimpsest accumulation. Taxonomic diversity is high, with up to 22 taxa representing diverse habitats, ranging from open grassland to wooded bushlands, as well as moist mosaics during Bed I. Overall, this archaeo-faunal assemblage speaks to increased behavioral versatility among Oldowan hominins and interactions with the carnivore guild.

Research on the Algerian localities of Ain Boucherit and Ain Hanech led to the dating of the oldest stone tools in North Africa, but also to the recovery of new material of the rare three-toed horse Hipparion ambiguum (Equidae) from its type locality of Ain Boucherit and the dating of the youngest North African hipparion record at Ain Hanech (1.7 Ma). Here we review the latest occurrences of the hipparions in the Old World and the problems concerning the taxonomical classification of the Pleistocene African hipparions. The decline and extinction of the hipparions (genus Hipparion, various subgenera) in the Old World occurred with a North-South gradient. Around 2.5 Ma, hipparions went extinct in Europe, West Asia and the Indian Subcontinent and the last dated record from North Africa is 1.7 Ma. There is evidence that at this time, there were still three species in Africa south of the Sahara and one in China. The last Chinese species went extinct before 1 Ma and the last African species between 0.6 and 0.4 Ma. Similar reduction of the diversity and extinction from North to South happened at different times in many other groups of vertebrates and was recognized already in the 18th century, when this pattern was explained with global cooling. Present data on temperature and atmospheric CO2 show a general decreasing trend and interfering Milankovich type cyclicities of different periodicities causing punctuated events major environmental change. The progressive demise of the hipparions seems to be related to such events. We discuss problems concerning the taxonomy of the last hipparions. The name Eurygnathohippus cornelianus is widespread, but the species name is a junior synonym. We classify this taxon with robust metapodials and elongated protocones as Hipparion (Eurygnathohippus) steytleri. The last North African hipparions are not a single species, but three: the large and very gracile H. (E.) libycum, the gracile H. (E.) ambiguum of intermediate size, and the small

Wonderwerk Cave, in South Africa, is an exceptional site that has yielded a large collection of small mammal fossils in a stratigraphic sequence reaching back ca. 2 million years. Taphonomic studies undertaken to date, show that Tytonidae (likely Tyto alba) was the dominant predator during the Earlier Stone Age. They produced masses of pellets that formed a dense carpet-like surface that covered the cave floor at intervals throughout the sequence. This paper compares the taphonomic signatures of five different Earlier Stone Age small mammal assemblages from Wonderwerk Cave, including assemblages not studied before, as well as a modern pellet assemblage collected from inside the cave. These samples were examined using taphonomic signatures, bone density and spatial distribution which confirm that the main predator in all periods of cave occupation were members of the Family Tytonidae, most likely Barn owls. The Wonderwerk small mammals have enabled us to clarify site formation processes and confirm that there was no transport or mixing of fossils, neither spatially (re-sedimentation) nor chronologically (reworking). This has confirmed the integrity of the stratigraphic sequence in the cave, reinforcing interpretations of palaeoecology, and elucidating intensity of occupation by hominins versus predators, and the behaviour of the predators vis a vis their prey.

Two well‑preserved, subadult 800 ky scapulae from Gran Dolina belonging to Homo antecessor, provide a unique opportunity to investigate the ontogeny of shoulder morphology in Lower Pleistocene humans. We compared the H. antecessor scapulae with a sample of 98 P. troglodytesand 108 H. sapiens representatives covering seven growth stages, as well as with the DIK‑1‑1 (Dikika; Australopithecus afarensis), KNM‑ WT 15000 (Nariokotome; H. ergaster), and MH2 (Malapa; A. sediba) specimens. We quantified 15 landmarks on each scapula and performed geometric morphometric analyses. H. sapiens scapulae are mediolaterally broader with laterally oriented glenoid fossae relative to Pan and Dikika shoulder blades. Accordingly, H. antecessor scapulae shared more morphological affinities with modern humans, KNM‑WT 15000, and even MH2. Both H. antecessor and modern Homo showed significantly more positive scapular growth trajectories than Pan (slopes: P. troglodytes = 0.0012; H. sapiens = 0.0018; H. antecessor= 0.0020). Similarities in ontogenetic trajectories between the H. antecessor and modern human data suggest that Lower Pleistocene hominin scapular development was already modern human‑like. At the same time, several morphological features distinguish H. antecessor scapulae from modern humans along the entire trajectory. Future studies should include additional Australopithecus specimens for further comparative assessment of scapular growth trends.

Adaptations of the lower back to bipedalism are frequently discussed but infrequently demonstrated in early fossil hominins. Newly discovered lumbar vertebrae contribute to a near-complete lower back of Malapa Hominin 2 (MH2), offering additional insights into posture and locomotion in Australopithecus sediba. We show that MH2 possessed a lower back consistent with lumbar lordosis and other adaptations to bipedalism, including an increase in the width of intervertebral articular facets from the upper to lower lumbar column (‘pyramidal configuration’). These results contrast with some recent work on lordosis in fossil hominins, where MH2 was argued to demonstrate no appreciable lordosis (‘hypolordosis’) similar to Neandertals. Our three-dimensional geometric morphometric (3D GM) analyses show that MH2’s nearly complete middle lumbar vertebra is human-like in overall shape but its vertebral body is somewhat intermediate in shape between modern humans and great apes. Additionally, it bears long, cranially and ventrally oriented costal (transverse) processes, implying powerful trunk musculature. We interpret this combination of features to indicate that A. sediba used its lower back in both bipedal and arboreal positional behaviors, as previously suggested based on multiple lines of evidence from other parts of the skeleton and reconstructed paleobiology of A. sediba.

Located within the Beni Fouda intramontane basin, the Oued Boucherit area (Guelta Zerga, North East Algeria) hosts a unique succession of archaeological and palaeontological deposits spanning from the late Pliocene to the Early Pleistocene. While the recent work by Sahnouni et al., (2018) mostly focused on the two Oldowan archaeological levels AB-Lw and AB-Up, we present here an updated overview of the entire Oued Boucherit sequence, resulting from a long-term multidisciplinary investigation that started in 1992. In particular, we report for the first time a description of the oldest palaeontological levels dated to >3 Ma, and of the uppermost archaeological level hosting Acheulean artefacts. The age-depth model initially proposed by Sahnouni et al. (2018) was further extended to chronologically constrain the whole sequence, which spans from ∼3.93 Ma at the bottom to ∼1.65 Ma at the top. The Oued Boucherit sedimentary sequence records the earliest evolutionary steps of lithic technology, with early Oldowan assemblages at Ain Boucherit dated to about 2.44 Ma (AB-Lw) and 1.92 Ma (AB-Up) Ma, followed by the ∼1.77-1.72 Ma assemblages of Ain Hanech and El Kherba showing more derived features, and the appearance of the Acheulean technology, dated to ∼1.67 Ma. This chonostratigraphic framework indicates that the Oued Boucherit area (<1 km) hosts the oldest Oldowan and Acheulean assemblages identified so far in North Africa. It is also especially striking that their chronology is somewhat close to the earliest Oldowan and Acheulean evidence found in East Africa and South Africa. Moreover, at least six fossiliferous levels ranging from ∼3.78 to 1.77-1.72 Ma have been identified and stratigraphically positioned within the sequence. They represent a unique opportunity to study not only the evolution of large mammal faunal assemblages in North Africa, but may also give some key insights about the evolution of palaeoecological and palaeoenvironmental conditions in the region thr

[Background] An individual’s microbiome changes over the course of its lifetime, especially during infancy, and again in old age. Confounding factors such as diet and healthcare make it difficult to disentangle the interactions between age, health, and microbial changes in humans. Animal models present an excellent opportunity to study age- and sex-linked variation in the microbiome, but captivity is known to influence animal microbial abundance and composition, while studies of free-ranging animals are typically limited to studies of the fecal microbiome using samples collected non-invasively. Here, we analyze a large dataset of oral, rectal, and genital swabs collected from 105 free-ranging rhesus macaques (Macaca mulatta, aged 1 month-26 years), comprising one entire social group, from the island of Cayo Santiago, Puerto Rico. We sequenced 16S V4 rRNA amplicons for all samples., [Results] Infant gut microbial communities had significantly higher relative abundances of Bifidobacterium and Bacteroides and lower abundances of Ruminococcus, Fibrobacter, and Treponema compared to older age groups, consistent with a diet high in milk rather than solid foods. The genital microbiome varied widely between males and females in beta-diversity, taxonomic composition, and predicted functional profiles. Interestingly, only penile, but not vaginal, microbiomes exhibited distinct age-related changes in microbial beta-diversity, taxonomic composition, and predicted functions. Oral microbiome composition was associated with age, and was most distinctive between infants and other age classes., [Conclusions] Across all three body regions, with notable exceptions in the penile microbiome, while infants were distinctly different from other age groups, microbiomes of adults were relatively invariant, even in advanced age. While vaginal microbiomes were exceptionally stable, penile microbiomes were quite variable, especially at the onset of reproductive age. Relative invariance among adults, including elderly individuals, is contrary to findings in humans and mice. We discuss potential explanations for this observation, including that age-related microbiome variation seen in humans may be related to changes in diet and lifestyle.

Lumbar lordosis is a key element of the upright posture, being interpreted as a consequence of bipedal locomotion. There is consensus that the generic modern human pattern of metameric vertebral body wedging is sexually dimorphic in modern humans. However, recently published studies have compared this pattern with other hominins, such as Neanderthals. These tried to establish whether the (male) Neanderthal lumbar vertebrae express a pattern that falls within or outside the range of modern human males. In the present study, data collected by 3D landmarks of the lumbar vertebrae of modern humans from different geographic regions and Neanderthals (Ntotal = 505 individual vertebrae) are used to clarify this problem, observing a similarity of the generic human pattern but with some interspecific differences in the pattern in the upper and lower lumbar vertebrae. Thus, the vertebral bodies L1-L3 of Neanderthals are more ventrally-wedged than in male modern humans, whilst the L4-L5 vertebral bodies of Neanderthal show a more progressive increase of dorsal wedging than in modern human males. The obtained results support modularity of the lumbar spine with different patterns in its upper and lower parts, and add that human geographic variability must be taken into account when carrying out comparisons of Neanderthals and modern humans. Our findings make clear the need to investigate this matter in more detail including complementary methods. Finally, key issues concerning the choice of measurement are discussed and recommendations made about how much inference can be made about complex systems such as the lumbar spine based on single linear measurements.

The phylogenetic relationships between hominins of the Early Pleistocene epoch in Eurasia, such as Homo antecessor, and hominins that appear later in the fossil record during the Middle Pleistocene epoch, such as Homo sapiens, are highly debated. For the oldest remains, the molecular study of these relationships is hindered by the degradation of ancient DNA. However, recent research has demonstrated that the analysis of ancient proteins can address this challenge. Here we present the dental enamel proteomes of H. antecessor from Atapuerca (Spain) and Homo erectus from Dmanisi (Georgia), two key fossil assemblages that have a central role in models of Pleistocene hominin morphology, dispersal and divergence. We provide evidence that H. antecessor is a close sister lineage to subsequent Middle and Late Pleistocene hominins, including modern humans, Neanderthals and Denisovans. This placement implies that the modern-like face of H. antecessor—that is, similar to that of modern humans—may have a considerably deep ancestry in the genus Homo, and that the cranial morphology of Neanderthals represents a derived form. By recovering AMELY-specific peptide sequences, we also conclude that the H. antecessor molar fragment from Atapuerca that we analysed belonged to a male individual. Finally, these H. antecessor and H. erectus fossils preserve evidence of enamel proteome phosphorylation and proteolytic digestion that occurred in vivo during tooth formation. Our results provide important insights into the evolutionary relationships between H. antecessor and other hominin groups, and pave the way for future studies using enamel proteomes to investigate hominin biology across the existence of the genus Homo.

Our knowledge of ancient human population structure in sub-Saharan Africa, particularly prior to the advent of food production, remains limited. Here we report genome-wide DNA data from four children—two of whom were buried approximately 8,000 years ago and two 3,000 years ago—from Shum Laka (Cameroon), one of the earliest known archaeological sites within the probable homeland of the Bantu language group1,2,3,4,5,6,7,8,9,10,11. One individual carried the deeply divergent Y chromosome haplogroup A00, which today is found almost exclusively in the same region12,13. However, the genome-wide ancestry profiles of all four individuals are most similar to those of present-day hunter-gatherers from western Central Africa, which implies that populations in western Cameroon today—as well as speakers of Bantu languages from across the continent—are not descended substantially from the population represented by these four people. We infer an Africa-wide phylogeny that features widespread admixture and three prominent radiations, including one that gave rise to at least four major lineages deep in the history of modern humans.

Changes in potential regulatory elements are thought to be key drivers of phenotypic divergence. However, identifying changes to regulatory elements that underlie human-specific traits has proven very challenging. Here, we use 63 reconstructed and experimentally measured DNA methylation maps of ancient and present-day humans, as well as of six chimpanzees, to detect differentially methylated regions that likely emerged in modern humans after the split from Neanderthals and Denisovans. We show that genes associated with face and vocal tract anatomy went through particularly extensive methylation changes. Specifically, we identify widespread hypermethylation in a network of face- and voice-associated genes (SOX9, ACAN, COL2A1, NFIX and XYLT1). We propose that these repression patterns appeared after the split from Neanderthals and Denisovans, and that they might have played a key role in shaping the modern human face and vocal tract.

We describe new specimens of the Miocene moschid Hispanomeryx, from the early Vallesian sites of Castell de Barberà (CB) and Ecoparc de Can Mata (ECM), Vallès-Penedès Basin, representing the first Iberian record of Hispanomeryx outside the inner Miocene basins. Fossils from ECM constitute Hispanomeryx lacetanus, sp. nov., the first Hispanomeryx to be described with a ‘Palaeomeryx fold (external postprotocristid).’ Fossils from CB constitute the first unambiguous record of Hispanomeryx aragonensis outside its type locality (La Ciesma-1B, Ebro Basin, Spain) and allow us to describe for the first time the upper molars of this species. Our phylogenetic analysis of Hispanomeryx places H. andrewsi from China as the sister group of a clade containing two European lineages. One of the latter places H. daamsi in a clade with the sister species H. duriensis and H. lacetanus, sp. nov., and the other groups together the two forms of H. aragonensis (type series from La Ciesma-1B plus the new material from CB). Emended diagnoses for the genus Hispanomeryx and all previously described species are presented. The new Hispanomeryx material from the Vallès-Penedès Basin constitutes the first unambiguous forest-dwelling Hispanomeryx ever recorded in Iberia, demonstrating that the previous lack of Hispanomeryx fossils in the Vallès-Penedès Basin was due to insufficient sampling rather than to paleoecological absence. It is therefore apparent that Hispanomeryx was as environmentally plastic as its relative Micromeryx, inhabiting similar habitats that varied from dry and open to humid and forested.

Bone marrow and grease constitute an important source of nutrition and have attracted the attention of human groups since prehistoric times. Marrow consumption has been linked to immediate consumption following the procurement and removal of soft tissues. Here, we present the earliest evidence for storage and delayed consumption of bone marrow at Qesem Cave, Israel (~420 to 200 ka). By using experimental series controlling exposure time and environmental parameters, combined with chemical analyses, we evaluated bone marrow preservation. The combination of archaeological and experimental results allowed us to isolate specific marks linked to dry skin removal and determine a low rate of marrow fat degradation of up to 9 weeks of exposure. This is the earliest evidence of such previously unidentified behavior, and it offers insights into the socio-economy of the human groups who lived at Qesem and may mark a threshold to new modes of Palaeolithic human adaptation.

Recent studies suggest that closely related species can accumulate substantial genetic and phenotypic differences despite ongoing gene flow, thus challenging traditional ideas regarding the genetics of speciation. Baboons (genus Papio) are Old World monkeys consisting of six readily distinguishable species. Baboon species hybridize in the wild, and prior data imply a complex history of differentiation and introgression. We produced a reference genome assembly for the olive baboon (Papio anubis) and whole-genome sequence data for all six extant species. We document multiple episodes of admixture and introgression during the radiation of Papio baboons, thus demonstrating their value as a model of complex evolutionary divergence, hybridization, and reticulation. These results help inform our understanding of similar cases, including modern humans, Neanderthals, Denisovans, and other ancient hominins.

East Africa has provided the earliest known evidence for Oldowan stone artifacts and hominin-induced stone tool cutmarks dated to ~2.6 million years (Ma) ago. The ~1.8-million-year-old stone artifacts from Ain Hanech (Algeria) were considered to represent the oldest archaeological materials in North Africa. Here we report older stone artifacts and cutmarked bones excavated from two nearby deposits at Ain Boucherit estimated to ~1.9 Ma ago, and the older to ~2.4 Ma ago. Hence, the Ain Boucherit evidence shows that ancestral hominins inhabited the Mediterranean fringe in northern Africa much earlier than previously thought. The evidence strongly argues for early dispersal of stone tool manufacture and use from East Africa or a possible multiple-origin scenario of stone technology in both East and North Africa.

Our most recent fossil relatives, the Neanderthals, had a large brain and a very heavy body compared to modern humans. This type of body requires high levels of energetic intake. While food (meat and fat consumption) is a source of energy, oxygen via respiration is also necessary for metabolism. We would therefore expect Neanderthals to have large respiratory capacities. Here we estimate the pulmonary capacities of Neanderthals, based on costal measurements and physiological data from a modern human comparative sample. The Kebara 2 male had a lung volume of about 9.04 l; Tabun C1, a female individual, a lung volume of 5.85 l; and a Neanderthal from the El Sidrón site, a lung volume of 9.03 l. These volumes are approximately 20% greater than the corresponding volumes of modern humans of the same body size and sex. These results show that the Neanderthal body was highly sensitive to energy supply.

Feeding behaviors may differ between past and current predators due to differences in the environments inhabited by these species at different times. We provide an example of this behavioral variability in spotted hyena (Crocuta crocuta), for which our analysis of a late Pleistocene micromammal assemblage indicates that hyenas preyed upon small rodents, a feeding habit that is rarely observed today among hyenas. The Bois Roche cave site is situated at the edge of a low bluff overlooking the floodplain of a small stream in Cherves-Richemont (Charente, France). The deposits are dated by electron spin resonance (ESR) to about 69.7 ± 4.1 Ka. Excavations at the site recovered fossil bones and teeth of large and small mammals, together with hyena coprolites. Water screening of the sediments produced large accumulations of rodent remains with low taxonomic diversity. Small mammal bones were recovered from hyena coprolites as well. Descriptions of small mammal bone modification, both from the sediments and coprolites, are reported here. The analysis yielded a distinct taphonomic pattern representative of large carnivores (over 30 kg), which differs from any other modern or fossil predator-accumulated microfaunal assemblage taphonomically analyzed to date. To our knowledge, previous studies of hyena diet have not recorded high concentrations of a single-rodent prey species. We conclude that the low species diversity of this small mammal assemblage most likely relates to a local abundance of the prey species due to an outbreak in the rodent population, rather than from specialist predator behavior and hunting technique.

[Objectives]: According to eco‐geographic rules, humans from high latitude areas present larger and wider trunks than their low‐latitude areas counterparts. This issue has been traditionally addressed on the pelvis but information on the thorax is largely lacking. We test whether ribcages are larger in individuals inhabiting high latitudes than in those from low latitudes and explored the correlation of rib size with latitude. We also test whether a common morphological pattern is exhibited in the thorax of different cold‐adapted populations, contributing to their hypothetical widening of the trunk., [Materials and methods]: We used 3D geometric morphometrics to quantify rib morphology of three hypothetically cold‐adapted populations, viz. Greenland (11 individuals), Alaskan Inuit (8 individuals) and people from Tierra del Fuego (8 individuals), in a comparative framework with European (Spain, Portugal and Austria; 24 individuals) and African populations (South African and sub‐Saharan African; 20 individuals)., [Results]: Populations inhabiting high latitudes present longer ribs than individuals inhabiting areas closer to the equator, but a correlation (p < 0.05) between costal size and latitude is only found in ribs 7–11. Regarding shape, the only cold adapted population that was different from the non‐cold‐adapted populations were the Greenland Inuit, who presented ribs with less curvature and torsion., [Conclusions]: Size results from the lower ribcage are consistent with the hypothesis of larger trunks in cold‐adapted populations. The fact that only Greenland Inuit present a differential morphological pattern, linked to a widening of their ribcage, could be caused by differences in latitude. However, other factors such as genetic drift or specific cultural adaptations cannot be excluded and should be tested in future studies.

The Krapina costal sample was studied by Gorjanović-Kramberger in the early twentieth century. He pointed out unique features in the sample such as the rounder rib cross-section, which was recently confirmed in other Neanderthal specimens. Round rib cross-sections are characteristic of Homo ergaster, suggesting this may be plesiomorphic for Pleistocene Homo, but it is unknown whether Homo antecessor also had this rib shape. Furthermore, the influence of allometry on the cross-sectional shape of ribs is still unknown. The large costal sample from Krapina allows us to address these issues. We quantified cross-section morphology at the midshaft throughout a closed curve of one landmark and nine sliding semilandmarks in the Krapina costal remains (n = 7), as well as in other Neanderthals (n = 50), H. antecessor (n = 3) and modern humans, both fossil (n = 12) and recent (n = 160). We used principal components analysis and mean comparisons to explore interspecific differences, regression analysis to investigate allometry, and partial least squares analysis to examine covariation of cross-section shape and overall rib morphology. Neanderthal cross-sections tended to be larger than those of recent humans except for the Krapina and Tabun remains. Regarding shape, inter-group differences were found only in the diaphragmatic thorax, where Neanderthal and H. antecessor ribs were statistically significantly rounder than those of modern humans. Allometry accounted for covariation of size on shape, but the Neandertal and modern human trajectories had different slopes. While our results based on the Krapina costal sample are similar to previous findings, we also make several new insights: 1) the cross-section morphology observed in Neanderthals was probably present in H. antecessor, albeit less marked; 2) the distinct roundness of Neanderthal cross-sections is not related to size; 3) rounder cross-sections are correlated with ribs presenting less curvature in cranial view and a low

The Kebara 2 individual (K2), an adult Neanderthal male, was found in the early 1980s during excavation at the Kebara cave on Mount Carmel (Israel) and is dated to around 60 ka (Arensburg, 1989; Bar-Yosef and Vandermeersh, 1991; Valladas et al., 1991). Because the remains are fairly complete and well preserved, this individual has been used as a reference Neanderthal male in studies of biological aspects such as body size and shape, locomotion, obstetrics or bioenergetics (Rak and Arensburg, 1987; Ruff, 1991, 2002; Tague, 1992; Churchill, 2006; Froehle and Churchill, 2009; Kramer and Sylvester, 2009; Weaver and Hublin, 2009; Holliday, 2012).

[Objectives]: First ribs bear information about thorax morphology and are usually well preserved, compared to other ribs, in bone/fossil samples. Several studies have addressed ontogeny of the first rib by studying changes in bone microanatomy and rib morphology separately, but no studies have combined both approaches to study how internal and external changes covary during ontogeny. The aim of this project is to fill this gap in our knowledge., [Materials and methods]: We applied 3D geometric morphometrics of sliding semilandmarks to 14 first ribs of Homo sapiens to quantify rib curvature and mid‐shaft cross‐section outline. Ontogenetic variation was addressed throughout a principal component analysis (PCA). Additionally, we made histological sections at the mid‐shaft of the same ribs and studied tissue matrix composition and compartmentalization. Finally, we performed partial least squares (PLS) and regression analyses to study covariation between rib morphology and compartmentalization variables., [Results]: PCA shows that first ribs increase their curvature over the course of ontogeny and the rib midshaft becomes less rounded during ontogeny. In addition, the sternal end becomes more medially oriented during ontogeny and the relative head‐tubercle distance becomes longer. Compartmentalization shows a decrease in the area occupied by mineralized tissues and an increase in the area occupied by non‐mineralized tissues over the course of ontogeny, which covaries with mid‐shaft cross‐section shape., [Conclusions]: Our results show detailed variation in rib morphology along with histological changes in bone tissue compartmentalization and, for the first time, the correlation between the two. This could be related to muscle attachments on the 1st rib and also to changes in breathing mode, from diaphragmatic in perinatals to pulmonary in adults, which could also have implications for understanding thorax evolution.

Bed II is a critical part of early Pleistocene Olduvai Gorge, Tanzania. Its deposits include transitions from humid to more arid conditions (with associated faunal changes), from Homo habilis to erectus, and from Oldowan to Acheulean technology. Bed II (~ 1.8–1.2 Ma) is stratigraphically and environmentally complex, with facies changes, faulting, and unconformities, making site-to-site correlation over the ~ 20 km of exposure difficult. Bed II tuffs are thinner, less evenly preserved, and more reworked than those of Bed I. Five marker tuffs (Tuffs IIA–IID, Bird Print Tuff (BPT)), plus local tephra, were collected from multiple sites and characterized using stratigraphic position, mineral assemblage, and electron probe microanalysis of phenocryst (feldspar, hornblende, augite, titanomagnetite) and glass (where available) composition. Lowermost Bed II tuffs are dominantly nephelinitic, Middle Bed II tuffs (BPT, Tuff IIC) have basaltic components, and upper Bed II Tuff IID is trachytic. The BPT and Tuff IID are identified widely using phenocryst compositions (high-Ca plagioclase and high-Ti hornblende, respectively), though IID was originally (Hay, 1976) misidentified as Tuff IIC at Loc 91 (SHK Annexe) in the Side Gorge. This work helps establish a high-resolution basin-wide paleolandscape context for the Oldowan–Acheulean transition and helps link hominin, faunal and archaeological records.

The disappearance of the earliest human culture, the Oldowan, and its substitution by a new technology, the Acheulean, is one of the main topics in modern Paleoanthropology. Recent research has established that the Acheulean emerged originally in East Africa around 1.7–1.6 million years ago, and from that area expanded across the rest of Africa, Europe and parts of Asia. Despite the great relevance of the Oldowan-Acheulean transition, little is known about the biological and cultural evolutionary mechanisms underlying this process. Traditionally, it has been assumed that this major cultural change was ignited by the emergence of a new human species, Homo ergaster/erectus, and that there was a steady technological evolution during the Oldowan that eventually led to the emergence of the Acheulean handaxes. However, these assumptions are not grounded in the current available evidence, but rooted in culturalhistory paradigms that should now be superseded. Olduvai Gorge (Tanzania) is the site where the traditional view of the OldowanAcheulean transition was established. The aim of the recently launched Olduvai Geochronology and Archaeology Project is to tackle this question by conducting a comprehensive research program at Olduvai, based on the retrieval of fresh data derived from new laboratory and fieldwork research. The multidisciplinary character of this ongoing study is providing an integrative perspective to the analysis of the paleoecology, archaeology, geology and geochronology of the transition to the Acheulean at Olduvai. Using an innovative theoretical perspective that combines interests in cultural change, ecological adaptations, and biological evolution, and state-of-the-art methods in archaeology, geology and taphonomy, this project aims to make Olduvai one of the world’s best references for the understanding of the evolutionary processes that led to the emergence of the Acheulean, the longest lasting culture in the history of humankind.