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.

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.

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.

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.

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.

A 3D taphonomic model of long bone modification by lions in medium-sized ungulate carcasses.

Here, we present the first three-dimensional taphonomic analysis of a carnivore-modified assemblage at the anatomical scale of the appendicular skeleton. A sample of ten carcasses composed of two taxa (zebra and wildebeest) consumed by wild lions in the Tarangire National Park (Tanzania) has been used to determine element-specific lion damage patterns. This study presents a novel software for the 3D spatial documentation of bone surface modifications at the anatomical level. Combined with spatial statistics, the present analysis has been able to conclude that despite variable degrees of competition during carcass consumption, lions generate bilateral patterning consisting of substantial damage of proximal ends of stylopodials and zeugopodials, moderate damage of the distal ends of femora and marginal damage of distal ends of humeri and zeugopodials. Of special interest is, specifically, the patterning of tooth marks on shafts according to element, since these are crucial to determine not only the type of carnivore involved in any given bone assemblage, but also the interaction with other agents (namely, hominins, in the past). Lions leave few tooth marks on mid-shaft sections, mostly concentrated on certain sections and orientations of stylopodials and, to a lesser extent, of the proximal tibia. Redundant occurrence of tooth marks on certain bone sections renders them as crucial to attest lion agency in carcass initial consumption. Indirectly, this can also be used to determine whether hominins ever acquired carcasses at lion kills.

Early Pleistocene faunivorous hominins were not kleptoparasitic, and this impacted the evolution of human anatomy and socio-ecology.

Humans are unique in their diet, physiology and socio-reproductive behavior compared to other primates. They are also unique in the ubiquitous adaptation to all biomes and habitats. From an evolutionary perspective, these trends seem to have started about two million years ago, coinciding with the emergence of encephalization, the reduction of the dental apparatus, the adoption of a fully terrestrial lifestyle, resulting in the emergence of the modern anatomical bauplan, the focalization of certain activities in the landscape, the use of stone tools, and the exit from Africa. It is in this period that clear taphonomic evidence of a switch in diet with respect to Pliocene hominins occurred, with the adoption of carnivory. Until now, the degree of carnivorism in early humans remained controversial. A persistent hypothesis is that hominins acquired meat irregularly (potentially as fallback food) and opportunistically through klepto-foraging. Here, we test this hypothesis and show, in contrast, that the butchery practices of early Pleistocene hominins (unveiled through systematic study of the patterning and intensity of cut marks on their prey) could not have resulted from having frequent secondary access to carcasses. We provide evidence of hominin primary access to animal resources and emphasize the role that meat played in their diets, their ecology and their anatomical evolution, ultimately resulting in the ecologically unrestricted terrestrial adaptation of our species. This has major implications to the evolution of human physiology and potentially for the evolution of the human brain.

Artificial intelligence provides greater accuracy in the classification of modern and ancient bone surface modifications.

Bone surface modifications are foundational to the correct identification of hominin butchery traces in the archaeological record. Until present, no analytical technique existed that could provide objectivity, high accuracy, and an estimate of probability in the identification of multiple structurally-similar and dissimilar marks. Here, we present a major methodological breakthrough that incorporates these three elements using Artificial Intelligence (AI) through computer vision techniques, based on convolutional neural networks. This method, when applied to controlled experimental marks on bones, yielded the highest rate documented to date of accurate classification (92%) of cut, tooth and trampling marks. After testing this method experimentally, it was applied to published images of some important traces purportedly indicating a very ancient hominin presence in Africa, America and Europe. The preliminary results are supportive of interpretations of ancient butchery in some places, but not in others, and suggest that new analyses of these controversial marks should be done following the protocol described here to confirm or disprove these archaeological interpretations.

Lions as Bone Accumulators? Paleontological and Ecological Implications of a Modern Bone Assemblage from Olduvai Gorge.

Analytic models have been developed to reconstruct early hominin behaviour, especially their subsistence patterns, revealed mainly through taphonomic analyses of archaeofaunal assemblages. Taphonomic research is used to discern which agents (carnivores, humans or both) generate the bone assemblages recovered at archaeological sites. Taphonomic frameworks developed during the last decades show that the only large-sized carnivores in African biomes able to create bone assemblages are leopards and hyenas. A carnivore-made bone assemblage located in the short-grassland ecological unit of the Serengeti (within Olduvai Gorge) was studied. Taphonomic analyses of this assemblage including skeletal part representation, bone density, breakage patterns and anatomical distribution of tooth marks, along with an ecological approach to the prey selection made by large carnivores of the Serengeti, were carried out. The results show that this bone assemblage may be the first lion-accumulated assemblage documented, although other carnivores (namely spotted hyenas) may have also intervened through postdepositional ravaging. This first faunal assemblage potentially created by lions constitutes a new framework for neotaphonomic studies. Since lions may accumulate carcasses under exceptional circumstances, such as those documented at the site reported here, this finding may have important consequences for interpretations of early archaeological and paleontological sites, which provide key information about human evolution.

Differential predation by age and sex classes in blue wildebeest in Serengeti: study of a modern carnivore den in Olduvai Gorge (Tanzania).

Age and sex selection of prey is an aspect of predator ecology which has been extensively studied in both temperate and African ecosystems. This dimension, along with fecundity, survival rates of prey and mortality factors other than predation are important in laying down the population dynamics of prey and have important implications in the management of species. A carnivore den located in the short-grassland ecological unit of the Serengeti was studied. Sex- and age- class (using five age categories) of the wildebeest remains recovered were analyzed through horn morphology, biometrics of the bones and tooth wear patterns. We compared our results with previous studies from lion and hyaena kills through multivariate analyses. Seasonality of the accumulation was analyzed through tooth histology. PCA and CVA results show that age class selection by predators depends on season, habitat-type, and growth rate of the wildebeest population. Female-biased predation was found to contradict classical hypotheses based on territorial male behaviour. The lion and spotted hyaena showed strong selection on age classes, contrary to previous studies. Migratory wildebeest sex ratio is regulated through differential predation by seasons and female deaths in the wet season are a trade-off for population stability. These data are crucial for an effective management of the species and the new method created may be useful for different carnivore species and their prey.

The "bear" essentials: actualistic research on Ursus arctos arctos in the Spanish Pyrenees and its implications for paleontology and archaeology.

Neotaphonomic studies of large carnivores are used to create models in order to explain the formation of terrestrial vertebrate fossil faunas. The research reported here adds to the growing body of knowledge on the taphonomic consequences of large carnivore behavior in temperate habitats and has important implications for paleontology and archaeology. Using photo- and videotrap data, we were able to describe the consumption of 17 ungulate carcasses by wild brown bears (Ursus arctos arctos) ranging the Spanish Pyrenees. Further, we analyzed the taphonomic impact of these feeding bouts on the bones recovered from those carcasses. The general sequence of consumption that we charted starts with separation of a carcass's trunk; viscera are generally eaten first, followed by musculature of the humerus and femur. Long limb bones are not broken open for marrow extraction. Bears did not transport carcasses or carcass parts from points of feeding and did not disperse bones appreciably (if at all) from their anatomical positions. The general pattern of damage that resulted from bear feeding includes fracturing, peeling, crenulation, tooth pitting and scoring of axial and girdle elements and furrowing of the upper long limb bones. As predicted from observational data, the taphonomic consequences of bear feeding resemble those of other non-durophagus carnivores, such as felids, and are distinct from those of durophagus carnivores, such as hyenids. Our results have paleontological and archaeological relevance. Specifically, they may prove useful in building analogical models for interpreting the formation of fossil faunas for which bears are suspected bone accumulators and/or modifiers. More generally, our comparative statistical analyses draw precise quantitative distinctions between bone damage patterns imparted respectively by durophagus (modelled here primarily by spotted hyenas [Crocuta crocuta] and wolves [Canis lupus]) and non-durophagus (modelled here by brown bears and lions [Panthera leo]) carnivorans.

First partial skeleton of a 1.34-million-year-old Paranthropus boisei from Bed II, Olduvai Gorge, Tanzania.

Recent excavations in Level 4 at BK (Bed II, Olduvai Gorge, Tanzania) have yielded nine hominin teeth, a distal humerus fragment, a proximal radius with much of its shaft, a femur shaft, and a tibia shaft fragment (cataloged collectively as OH 80). Those elements identified more specifically than to simply Hominidae gen. et sp. indet are attributed to Paranthropus boisei. Before this study, incontrovertible P. boisei partial skeletons, for which postcranial remains occurred in association with taxonomically diagnostic craniodental remains, were unknown. Thus, OH 80 stands as the first unambiguous, dentally associated Paranthropus partial skeleton from East Africa. The morphology and size of its constituent parts suggest that the fossils derived from an extremely robust individual who, at 1.338±0.024 Ma (1 sigma), represents one of the most recent occurrences of Paranthropus before its extinction in East Africa.

Disentangling Early Stone Age palimpsests: determining the functional independence of hominid- and carnivore-derived portions of archaeofaunas.

Determining the extent to which hominid- and carnivore-derived components of fossil bone palimpsests formed independently of each other can provide valuable information to paleoanthropologists interested in reconstructing the foraging adaptations of hominids. Because stone tool cutmarks, hammerstone percussion marks, and carnivore tooth marks are usually only imparted on bone during nutrient extraction from a carcass, these bone surface modifications are particularly amenable to the types of analyses that might meet this goal. This study compares the percentage of limb bone specimens that preserve evidence of both hominid- and carnivore-imparted bone damage from actualistic control samples and several Plio-Pleistocene archaeofaunas, including new data from Swartkrans Member 3 (South Africa). We argue that this procedure, which elucidates the degree of hominid-carnivore independence in assemblage formation, will allow researchers to extract for focused analyses high integrity components (hominid and carnivore) from presumably low integrity sites. Comparisons suggest that the hominid- and carnivore-derived components from sites in Olduvai Gorge Bed II (Tanzania), the ST Site Complex at Peninj (Tanzania), and Swartkrans Member 3 formed largely independent of each other, while data from the FLK 22 Zinjanthropus (FLK Zinj) site (Olduvai Gorge Bed I) indicate significant interdependence in assemblage formation. This contrast suggests that some Early Stone Age assemblages (e.g., the Olduvai Gorge Bed II sites, the Peninj ST Site Complex, and Swartkrans Member 3) are probably more useful than others (e.g., FLK Zinj) for assessing the maximal carcass-acquiring abilities of early hominids; in such assemblages as those in the former set, sole hominid-contribution is more confidently discerned and isolated for analysis than in assemblages such as FLK Zinj.

Beyond leopards: tooth marks and the contribution of multiple carnivore taxa to the accumulation of the Swartkrans Member 3 fossil assemblage.

The ca. 1.0 myr old fauna from Swartkrans Member 3 (South Africa) preserves abundant indication of carnivore activity in the form of tooth marks (including pits) on many bone surfaces. This direct paleontological evidence is used to test a recent suggestion that leopards, regardless of prey body size, may have been almost solely responsible for the accumulation of the majority of bones in multiple deposits (including Swartkrans Member 3) from various Sterkfontein Valley cave sites. Our results falsify that hypothesis and corroborate an earlier hypothesis that, while the carcasses of smaller animals may have been deposited in Swartkrans by leopards, other kinds of carnivores (and hominids) were mostly responsible for the deposition of large animal remains. These results demonstrate the importance of choosing appropriate classes of actualistic data for constructing taphonomic inferences of assemblage formation. In addition, they stress that an all-encompassing model of assemblage formation for the hominid-bearing deposits of the Sterkfontein Valley is inadequate and that each must be evaluated individually using not just analogical reasoning but also incorporating empirical data generated in the preserved fossil samples.