Elevated rates of horizontal gene transfer in the industrialized human microbiome.

Industrialization has impacted the human gut ecosystem, resulting in altered microbiome composition and diversity. Whether bacterial genomes may also adapt to the industrialization of their host populations remains largely unexplored. Here, we investigate the extent to which the rates and targets of horizontal gene transfer (HGT) vary across thousands of bacterial strains from 15 human populations spanning a range of industrialization. We show that HGTs have accumulated in the microbiome over recent host generations and that HGT occurs at high frequency within individuals. Comparison across human populations reveals that industrialized lifestyles are associated with higher HGT rates and that the functions of HGTs are related to the level of host industrialization. Our results suggest that gut bacteria continuously acquire new functionality based on host lifestyle and that high rates of HGT may be a recent development in human history linked to industrialization.

Biomechanical and taxonomic diversity in the Early Pleistocene in East Africa: Structural analysis of a recently discovered femur shaft from Olduvai Gorge (bed I).

Recent Plio-Pleistocene hominin findings have revealed the complexity of human evolutionary history and the difficulties involved in its interpretation. Moreover, the study of hominin long bone remains is particularly problematic, since it commonly depends on the analysis of fragmentary skeletal elements that in many cases are merely represented by small diaphyseal portions and appear in an isolated fashion in the fossil record. Nevertheless, the study of the postcranial skeleton is particularly important to ascertain locomotor patterns. Here we report on the discovery of a robust hominin femoral fragment (OH 84) at the site of Amin Mturi Korongo dated to 1.84 Ma (Olduvai Bed I). External anatomy and internal bone structure of OH 84 were analyzed and compared with previously published data for modern humans and chimpanzees, as well as for Australopithecus, Paranthropus and Homo specimens ranging from the Late Pliocene to Late Pleistocene. Biomechanical analyses based on transverse cross-sections and the comparison of OH 84 with another robust Olduvai specimen (OH 80) suggest that OH 84 might be tentatively allocated to Paranthropus boisei. More importantly, the identification of a unique combination of traits in OH 84 could indicate both terrestrial bipedalism and an arboreal component in the locomotor repertoire of this individual. If interpreted correctly, OH 84 could thus add to the already mounting evidence of substantial locomotor diversity among Early Pleistocene hominins. Likewise, our results also highlight the difficulties in accurately interpreting the link between form and function in the human fossil record based on fragmentary remains, and ultimately in distinguishing between coeval hominin groups due to the heterogeneous pattern of inter- and intraspecific morphological variability detected among fossil femora.

Microbial biomarkers reveal a hydrothermally active landscape at Olduvai Gorge at the dawn of the Acheulean, 1.7 Ma.

Landscape-scale reconstructions of ancient environments within the cradle of humanity may reveal insights into the relationship between early hominins and the changing resources around them. Many studies of Olduvai Gorge during Pliocene-Pleistocene times have revealed the presence of precession-driven wet-dry cycles atop a general aridification trend, though may underestimate the impact of local-scale conditions on early hominins, who likely experienced a varied and more dynamic landscape. Fossil lipid biomarkers from ancient plants and microbes encode information about their surroundings via their molecular structures and composition, and thus can shed light on past environments. Here, we employ fossil lipid biomarkers to study the paleolandscape at Olduvai Gorge at the emergence of the Acheulean technology, 1.7 Ma, through the Lower Augitic Sandstones layer. In the context of the expansion of savanna grasslands, our results represent a resource-rich mosaic ecosystem populated by groundwater-fed rivers, aquatic plants, angiosperm shrublands, and edible plants. Evidence of a geothermally active landscape is reported via an unusual biomarker distribution consistent with the presence of hydrothermal features seen today at Yellowstone National Park. The study of hydrothermalism in ancient settings and its impact on hominin evolution has not been addressed before, although the association of thermal springs in the proximity of archaeological sites documented here can also be found at other localities. The hydrothermal features and resources present at Olduvai Gorge may have allowed early hominins to thermally process edible plants and meat, supporting the possibility of a prefire stage of human evolution.

Deep classification of cut-marks on bones from Arroyo del Vizcaíno (Uruguay).

The earliest widely accepted presence of humans in America dates to approximately 17.5 cal kyr BP, at the end of the Last Glacial Maximum (LGM). Among other evidence, this presence is attested by stone tools and associated cut-marks and other bone surface modifications (BSM), interpreted as the result of the consumption of animals by humans. Claims of an older human presence in the continent have been made based on the proposed anthropogenic modification of faunal remains; however, these have been controversial due to the highly subjective nature of the interpretations. Here, we employ advanced deep learning algorithms to objectively increase the accuracy of BSM identification on bones. With several models that exhibit BSM classification accuracies greater than 94%, we use ensemble learning techniques to robustly classify a selected sample of BSM from the approximately 30 kyr BP site of Arroyo del Vizcaíno, Uruguay. Our results confidently show the presence of cut-marks imparted by stone tools on bones at the site. This result supports an earlier presence of humans in the American continent, expanding additional genetic and archaeological evidence of a human LGM and pre-LGM presence in the continent.

Hominin skeletal part abundances and claims of deliberate disposal of corpses in the Middle Pleistocene.

Humans are set apart from other organisms by the realization of their own mortality. Thus, determining the prehistoric emergence of this capacity is of significant interest to understanding the uniqueness of the human animal. Tracing that capacity chronologically is possible through archaeological investigations that focus on physical markers that reflect "mortality salience." Among these markers is the deliberate and culturally mediated disposal of corpses. Some Neandertal bone assemblages are among the earliest reasonable claims for the deliberate disposal of hominins, but even these are vigorously debated. More dramatic assertions center on the Middle Pleistocene sites of Sima de los Huesos (SH, Spain) and the Dinaledi Chamber (DC, South Africa), where the remains of multiple hominin individuals were found in deep caves, and under reported taphonomic circumstances that seem to discount the possibility that nonhominin actors and processes contributed to their formation. These claims, with significant implications for charting the evolution of the "human condition," deserve scrutiny. We test these assertions through machine-learning analyses of hominin skeletal part representation in the SH and DC assemblages. Our results indicate that nonanthropogenic agents and abiotic processes cannot yet be ruled out as significant contributors to the ultimate condition of both collections. This finding does not falsify hypotheses of deliberate disposal for the SH and DC corpses, but does indicate that the data also support partially or completely nonanthropogenic formational histories.

The meta-group social network of early humans: A temporal-spatial assessment of group size at FLK Zinj (Olduvai Gorge, Tanzania).

Humans are the only primates that maintain regular inter-group relationships and meta-group social networks that enable the inter-group flow of individuals. This is the basis of the band/tribe concept in the anthropology of modern foragers. The present work is a theoretical approach to the development of analytical tools to understand group size and the temporal scale of site occupation in the archaeological record. We selected FLK Zinj as one of the oldest examples of a taphonomically-supported anthropogenic site in which both variables (group size and time) could be modelled using a combination of modern forager regression estimates from their camp sizes and estimates derived from the combined use of modern African foragers' meat consumption rates per day per capita during the dry season and minimum estimates of flesh yields provided by the carcass parts preserved at FLK Zinj. This approach provides the basis for a testable hypothesis which should be further tested in other Oldowan sites. An estimate of 18-28 individuals occupying FLK Zinj was made, which is similar to the estimated 16 individuals of one of the 1.5 Ma Ileret Homo erectus footprint trails. It also shows a similar proportional distribution to Dunbar's equations (group size to neocortex ratio) as documented in modern foragers, which suggest that most of the social network of H. erectus was in the meta-group level as is the case of modern foragers. Irrespective of the range of variation discussed for both variables (group size and length of time represented), it is argued that neither small estimates of time nor small group sizes can account for the formation of FLK Zinj.

Dietary options and behavior suggested by plant biomarker evidence in an early human habitat.

The availability of plants and freshwater shapes the diets and social behavior of chimpanzees, our closest living relative. However, limited evidence about the spatial relationships shared between ancestral human (hominin) remains, edible resources, refuge, and freshwater leaves the influence of local resources on our species' evolution open to debate. Exceptionally well-preserved organic geochemical fossils--biomarkers--preserved in a soil horizon resolve different plant communities at meter scales across a contiguous 25,000 m(2) archaeological land surface at Olduvai Gorge from about 2 Ma. Biomarkers reveal hominins had access to aquatic plants and protective woods in a patchwork landscape, which included a spring-fed wetland near a woodland that both were surrounded by open grassland. Numerous cut-marked animal bones are located within the wooded area, and within meters of wetland vegetation delineated by biomarkers for ferns and sedges. Taken together, plant biomarkers, clustered bone debris, and hominin remains define a clear spatial pattern that places animal butchery amid the refuge of an isolated forest patch and near freshwater with diverse edible resources.

Assessment of statistical agreement of three techniques for the study of cut marks: 3D digital microscope, laser scanning confocal microscopy and micro-photogrammetry.

In the last few years, the study of cut marks on bone surfaces has become fundamental for the interpretation of prehistoric butchery practices. Due to the difficulties in the correct identification of cut marks, many criteria for their description and classification have been suggested. Different techniques, such as three-dimensional digital microscope (3D DM), laser scanning confocal microscopy (LSCM) and micro-photogrammetry (M-PG) have been recently applied to the study of cut marks. Although the 3D DM and LSCM microscopic techniques are the most commonly used for the 3D identification of cut marks, M-PG has also proved to be very efficient and a low-cost method. M-PG is a noninvasive technique that allows the study of the cortical surface without any previous preparation of the samples, and that generates high-resolution models. Despite the current application of microscopic and micro-photogrammetric techniques to taphonomy, their reliability has never been tested. In this paper, we compare 3D DM, LSCM and M-PG in order to assess their resolution and results. In this study, we analyse 26 experimental cut marks generated with a metal knife. The quantitative and qualitative information registered is analysed by means of standard multivariate statistics and geometric morphometrics to assess the similarities and differences obtained with the different methodologies.

Computer vision enables taxon-specific identification of African carnivore tooth marks on bone.

Taphonomic works aim at discovering how paleontological and archaeofaunal assemblages were formed. They also aim at determining how hominin fossils were preserved or destroyed. Hominins and other mammal carnivores have been co-evolving, at least during the past two million years, and their potential interactions determined the evolution of human behavior. In order to understand all this, taxon-specific carnivore agency must be effectively identified in the fossil record. Until now, taphonomists have been able to determine, to some degree, hominin and carnivore inputs in site formation, and their interactions in the modification of part of those assemblages. However, the inability to determine agency more specifically has hampered the development of taphonomic research, whose methods are virtually identical to those used several decades ago (lagged by a high degree of subjectivity). A call for more objective and agent-specific methods would be a major contribution to the advancement of taphonomic research. Here, we present one of these advances. The use of computer vision (CV) on a large data set of images of tooth marks has enabled the objective discrimination of taxon-specific carnivore agency up to 88% of the testing sample. We highlight the significance of this method in an interdisciplinary interplay between traditional taphonomic-paleontological analysis and artificial intelligence-based computer science. The new questions that can be addressed with this will certainly bring important changes to several ideas on important aspects of the human evolutionary process.

Configurational approach to identifying the earliest hominin butchers.

The announcement of two approximately 3.4-million-y-old purportedly butchered fossil bones from the Dikika paleoanthropological research area (Lower Awash Valley, Ethiopia) could profoundly alter our understanding of human evolution. Butchering damage on the Dikika bones would imply that tool-assisted meat-eating began approximately 800,000 y before previously thought, based on butchered bones from 2.6- to 2.5-million-y-old sites at the Ethiopian Gona and Bouri localities. Further, the only hominin currently known from Dikika at approximately 3.4 Ma is Australopithecus afarensis, a temporally and geographically widespread species unassociated previously with any archaeological evidence of butchering. Our taphonomic configurational approach to assess the claims of A. afarensis butchery at Dikika suggests the claims of unexpectedly early butchering at the site are not warranted. The Dikika research group focused its analysis on the morphology of the marks in question but failed to demonstrate, through recovery of similarly marked in situ fossils, the exact provenience of the published fossils, and failed to note occurrences of random striae on the cortices of the published fossils (incurred through incidental movement of the defleshed specimens across and/or within their abrasive encasing sediments). The occurrence of such random striae (sometimes called collectively "trampling" damage) on the two fossils provide the configurational context for rejection of the claimed butchery marks. The earliest best evidence for hominin butchery thus remains at 2.6 to 2.5 Ma, presumably associated with more derived species than A. afarensis.

African bovid tribe classification using transfer learning and computer vision.

Objective analytical identification methods are still a minority in the praxis of paleobiological sciences. Subjective interpretation of fossils and their modifications remains a nonreplicable expert endeavor. Identification of African bovids is a crucial element in the reconstruction of paleo-landscapes, ungulate paleoecology, and, eventually, hominin adaptation and ecosystemic reconstruction. Recent analytical efforts drawing on Fourier functional analysis and discrimination methods applied to occlusal surfaces of teeth have provided a highly accurate framework to correctly classify African bovid tribes and taxa. Artificial intelligence tools, like computer vision, have also shown their potential to be objectively more accurate in the identification of taphonomic agency than human experts. For this reason, here we implement some of the most successful computer vision methods, using transfer learning and ensemble analysis, to classify bidimensional images of African bovid teeth and show that 92% of the large testing set of images of African bovid tribes analyzed could be correctly classified. This brings an objective tool to paleoecological interpretation, where bovid identification and paleoecological interpretation can be more confidently carried out.

Taphonomic analysis of the early Pleistocene (2.4 Ma) faunal assemblage from A.L. 894 (Hadar, Ethiopia).

The A.L. 894 site (Hadar, Ethiopia) is, together with OGS 7 (Gona, Ethiopia), one of the oldest archaeological sites documenting a spatial association of stone tools and bones retrieved from an in situ excavation. In contrast with OGS 7, the better preservation of the bone assemblage at A.L. 894 allows the identification of taphonomic processes of bone breakage, thanks to abundant green bone fractures. The presence of tooth marks and the lack of hominin-produced bone modifications together argue against hominins as the responsible agents for bone accumulation and modification. This taphonomic study of A.L. 894 shows lack of evidence for functional associations between stone tools and bones, a pattern documented in several other early Pleistocene sites. Such a pattern underscores the complex phenomena involved in site formation processes, especially in the earliest archaeological assemblages.

New ages for Middle and Later Stone Age deposits at Mumba rockshelter, Tanzania: optically stimulated luminescence dating of quartz and feldspar grains.

The archaeological deposits at Mumba rockshelter, northern Tanzania, have been excavated for more than 70 years, starting with Margit and Ludwig Köhl-Larsen in the 1930s. The assemblages of Middle Stone Age (MSA) and Later Stone Age (LSA) artefacts collected from this site constitute the type sequences for these cultural phases in East Africa. Despite its archaeological importance, however, the chronology of the site is poorly constrained, despite the application since the 1980s of several dating methods (radiocarbon, uranium-series and amino acid racemisation) to a variety of materials recovered from the deposits. Here, we review these previous chronologies for Mumba and report new ages obtained from optically stimulated luminescence (OSL) and infrared stimulated luminescence (IRSL) measurements on single grains of quartz and multi-grain aliquots of potassium (K) feldspar from the MSA and LSA deposits. Measurements of single grains of quartz allowed the rejection of unrepresentative grains and the application of appropriate statistical models to obtain the most reliable age estimates, while measurements of K-feldspars allowed the chronology to be extended to older deposits. The seven quartz ages and four K-feldspar ages provide improved temporal constraints on the archaeological sequence at Mumba. The deposits associated with the latest Kisele Industry (Bed VI-A) and the earliest Mumba Industry (Bed V) are dated to 63.4 ± 5.7 and 56.9 ± 4.8 ka (thousands of years ago), respectively, thus constraining the time of transition between these two archaeological phases to ~60 ka. An age of 49.1 ± 4.3 ka has been obtained for the latest deposits associated with the Mumba Industry, which show no evidence for post-depositional mixing and contain ostrich eggshell (OES) beads and abundant microlithics. The Nasera Industry deposits (Bed III) contain large quantities of OES beads and date to 36.8 ± 3.4 ka. We compare the luminescence ages with the previous chronologies for Mumba, and briefly discuss how the revised chronology fits in the context of existing archaeological records and palaeoclimatic reconstructions for East Africa.

Sabertooth carcass consumption behavior and the dynamics of Pleistocene large carnivoran guilds.

Apex predators play an important role in the top-down regulation of ecological communities. Their hunting and feeding behaviors influence, respectively, prey demography and the availability of resources to other consumers. Among the most iconic-and enigmatic-terrestrial predators of the late Cenozoic are the Machairodontinae, a diverse group of big cats whose hypertrophied upper canines have earned them the moniker "sabertooths." Many aspects of these animals' paleobiology, especially their prey preferences and carcass consumption behavior, remain unsettled. While skeletal anatomy, dental morphology and wear, and isotopic profiles provide important insights, the most direct way to resolve these issues is through the fossil remains of sabertooth prey. Here, we report on a taphonomic analysis of an early Pleistocene faunal assemblage from Haile 21A (Florida, USA) that preserves feeding damage from the lion-sized sabertooth Xenosmilus hodsonae. Patterns of tooth-marking and bone damage indicate that Xenosmilus fully defleshed the carcasses of their prey and even engaged in some minor bone consumption. This has important implications for Pleistocene carnivoran guild dynamics, including the carcass foraging behavior of the first stone-tool-using hominins.

A case of hominin scavenging 1.84 million years ago from Olduvai Gorge (Tanzania).

Meat eating is one of the hallmarks of human evolution. It has been linked to the beginning of stone tool use, to physiological changes leading to crucial anatomical transformations defining our genus, and to new socioreproductive and cognitive behaviors. Uncontroversial evidence of meat eating goes back to 2.6 million years ago; however, little is known about the frequency and timing with which early hominins acquired animal resources. Here, we show that the combination of hunting and scavenging documented in some modern human foragers may have a long evolutionary trajectory. Using a new set of artificial intelligence methods for objective identification, we present direct evidence of an episode of hominins scavenging from large felids-probably lions-discovered at Olduvai Gorge (DS site, Bed I). This casts a new perspective on the diversity of hominin carcass acquisition behaviors and survival strategies, and places some early Pleistocene hominins in ecological proximity to African large carnivore guilds.

New site at Olduvai Gorge (AGS, Bed I, 1.84 Mya) widens the range of locations where hominins engaged in butchery.

Outstanding questions about human evolution include systematic connections between critical landscape resources-such as water and food-and how these shaped the competitive and biodiverse environment(s) that our ancestors inhabited. Here, we report fossil n-alkyl lipid biomarkers and their associated δ13C values across a newly discovered Olduvai Gorge site (AGS) dated to 1.84 million years ago, enabling a multiproxy analysis of the distributions of critical local landscape resources across an explicit locus of hominin activity. Our results reveal that AGS was a seasonally waterlogged, largely unvegetated lakeside site situated near an ephemeral freshwater river surrounded by arid-adapted C4 grasses. The sparse vegetation at AGS contrasts with reconstructed (micro)habitats at the other anthropogenic sites at Olduvai Gorge, suggesting that central-provisioning places depended more heavily on water access than vegetation viz. woody plants as is often observed for modern hunter-gatherers. As hominins at AGS performed similar butchering activities as at other Bed I sites, our results suggest they did not need the shelter of trees and thus occupied a competitive position within the predatory guild.

Neo-taphonomic analysis of the Misiam leopard lair from Olduvai Gorge (Tanzania): understanding leopard-hyena interactions in open settings.

Misiam is a modern wildebeest-dominated accumulation situated in a steep ravine covered with dense vegetation at Olduvai Gorge (Tanzania). It is interpreted here as a leopard lair to which carcasses have been transported for several years. Felid-specific bone damage patterns, felid-typical skeletal part profiles, taxonomic specialization and the physical presence of leopards observed by the authors show that leopards at Misiam can be specialized medium-sized carcass accumulators. Hyenas also intervened at intervals in the modification of the retrieved faunal assemblage. This makes Misiam a carnivore palimpsest. Here, we additionally show that leopards only transport and accumulate carcasses on occasions, that they can seem highly specialized despite being dietary generalists, and that such a behaviour may be prompted by seasonal competition or during the breeding season or both. Misiam is the first open-air leopard lair with a dense bone accumulation reported. There, leopards engaged in intensive accumulation of carcasses during the wet season, when the southern Serengeti short-grass plains undergo the effect of the famous wildebeest migration and this migratory species reaches the gorge. The ecological importance of this behaviour and its relevance as a proxy for reconstructing prehistoric carnivore behaviours are discussed.

Computer vision supports primary access to meat by early Homo 1.84 million years ago.

Human carnivory is atypical among primates. Unlike chimpanzees and bonobos, who are known to hunt smaller monkeys and eat them immediately, human foragers often cooperate to kill large animals and transport them to a safe location to be shared. While it is known that meat became an important part of the hominin diet around 2.6-2 Mya, whether intense cooperation and food sharing developed in conjunction with the regular intake of meat remains unresolved. A widespread assumption is that early hominins acquired animal protein through klepto-parasitism at felid kills. This should be testable by detecting felid-specific bone modifications and tooth marks on carcasses consumed by hominins. Here, deep learning (DL) computer vision was used to identify agency through the analysis of tooth pits and scores on bones recovered from the Early Pleistocene site of DS (Bed I, Olduvai Gorge). We present the first objective evidence of primary access to meat by hominins 1.8 Mya by showing that the most common securely detectable bone-modifying fissipeds at the site were hyenas. The absence of felid modifications in most of the carcasses analyzed indicates that hominins were the primary consumers of most animals accumulated at the site, with hyenas intervening at the post-depositional stage. This underscores the role of hominins as a prominent part of the early Pleistocene African carnivore guild. It also stresses the major (and potentially regular) role that meat played in the diet that configured the emergence of early Homo.

Early Pliocene hominids from Gona, Ethiopia.

Comparative biomolecular studies suggest that the last common ancestor of humans and chimpanzees, our closest living relatives, lived during the Late Miocene-Early Pliocene. Fossil evidence of Late Miocene-Early Pliocene hominid evolution is rare and limited to a few sites in Ethiopia, Kenya and Chad. Here we report new Early Pliocene hominid discoveries and their palaeoenvironmental context from the fossiliferous deposits of As Duma, Gona Western Margin (GWM), Afar, Ethiopia. The hominid dental anatomy (occlusal enamel thickness, absolute and relative size of the first and second lower molar crowns, and premolar crown and radicular anatomy) indicates attribution to Ardipithecus ramidus. The combined radioisotopic and palaeomagnetic data suggest an age of between 4.51 and 4.32 million years for the hominid finds at As Duma. Diverse sources of data (sedimentology, faunal composition, ecomorphological variables and stable carbon isotopic evidence from the palaeosols and fossil tooth enamel) indicate that the Early Pliocene As Duma sediments sample a moderate rainfall woodland and woodland/grassland.

Tracing the spatial imprint of Oldowan technological behaviors: A view from DS (Bed I, Olduvai Gorge, Tanzania).

DS (David's site) is one of the new archaeological sites documented in the same paleolandscape in which FLK 22 was deposited at about 1.85 Ma in Olduvai Gorge. Fieldwork in DS has unearthed the largest vertically-discrete archaeological horizon in the African Pleistocene, where a multi-cluster anthropogenic accumulation of fossil bones and stone tools has been identified. In this work we present the results of the techno-economic study of the lithic assemblage recovered from DS. We also explore the spatial magnitude of the technological behaviors documented at this spot using powerful spatial statistical tools to unravel correlations between the spatial distributional patterns of lithic categories. At DS, lavas and quartzite were involved in different technological processes. Volcanic materials, probably transported to this spot from a close source, were introduced in large numbers, including unmodified materials, and used in percussion activities and in a wide variety of reduction strategies. A number of volcanic products were subject to outward fluxes to other parts of the paleolandscape. In contrast, quartzite rocks were introduced in smaller numbers and might have been subject to a significantly more intense exploitation. The intra-site spatial analysis has shown that specialized areas cannot be identified, unmodified materials are not randomly distributed, percussion and knapping categories do not spatially overlap, while bipolar specimens show some sort of spatial correlation with percussion activities.