Aridity, availability of drinking water and freshwater foods, and hominin and archeological sites during the Late Pliocene-Early Pleistocene in the western region of the Turkana Basin (Kenya): A review.

Although the Turkana Basin is one of the driest regions of the East African Rift, its Plio-Pleistocene sediments are rich in freshwater vertebrates and invertebrates, providing evidence that freshwater resources were available to hominins in this region during the Plio-Pleistocene (4.2-0.7 Ma). Here we provide an overview of the hydroconnectivity of the Turkana Basin. We then review the period during which freshwater river and lake systems expanded into the western region of the Turkana Basin, where hominin and archeological sites have been discovered in sediments dating back to the Late Pliocene-Pleistocene. Freshwater conditions are reconstructed from river and lake sediments and the flora and micro- and macofauna they contain. Data synthesis suggests that drinking water and freshwater foods prevailed in the western region of the Turkana Basin at 4.20-3.98 Ma, 3.70-3.10 Ma, 2.53-2.22 Ma, then between 2.10 and 1.30 Ma and intermittently from 1.27 to 0.75 Ma. Milestones in hominin evolution occurred in this context, such as the first occurrence of Australopithecus anamensis (4.20-4.10 Ma) and Kenyanthropus platyops (3.50 Ma and 3.30-3.20 Ma), the presence of Paranthropus aethiopicus (2.53-2.45 Ma), early Homo (2.33 Ma), Paranthropus boisei (2.25 Ma and 1.77-1.72 Ma) and Homo ergaster/Homo erectus (1.75 Ma, 1.47-1.42 Ma and 1.10-0.90 Ma). Developments in hominin behavior also occurred during this timeframe, including the first known stone tools (3.30 Ma), the oldest Oldowan sites (2.34 Ma and 2.25 Ma) in the Turkana Basin, the earliest known evidence for the emergence of bifacial shaping in eastern Africa (1.80 Ma), and the first known Acheulean site (1.76 Ma). Our synthesis suggests that, diachronic variation in hydroconnectivity played a role on the amount of drinking water and freshwater foods available in the western region of the Turkana Basin, despite regional aridity.

Genome sequences of 36,000- to 37,000-year-old modern humans at Buran-Kaya III in Crimea.

Populations genetically related to present-day Europeans first appeared in Europe at some point after 38,000-40,000 years ago, following a cold period of severe climatic disruption. These new migrants would eventually replace the pre-existing modern human ancestries in Europe, but initial interactions between these groups are unclear due to the lack of genomic evidence from the earliest periods of the migration. Here we describe the genomes of two 36,000-37,000-year-old individuals from Buran-Kaya III in Crimea as belonging to this newer migration. Both genomes share the highest similarity to Gravettian-associated individuals found several thousand years later in southwestern Europe. These genomes also revealed that the population turnover in Europe after 40,000 years ago was accompanied by admixture with pre-existing modern human populations. European ancestry before 40,000 years ago persisted not only at Buran-Kaya III but is also found in later Gravettian-associated populations of western Europe and Mesolithic Caucasus populations.

Protocol combining tree-based Maximum Parsimony and web-like Phylogenetic Networks analyses to investigate reticulate human evolution.

Our protocol combines Maximum Parsimony and Phylogenetic Networks approaches to understand the phylogenetic relationships and evolutionary processes of hominin species that might have shared inheritance from multiple ancestors. By addressing the questions of pattern and process in human phylogeny, the protocol can be used to clarify the taxonomic definition(s) of diverse hominin groups and ascertain whether or not the mode of evolution of genus Homo is reticulate. Using high quality and informative phenotypic data sets is necessary to yield meaningful results. For complete details on the use and execution of this protocol, please refer to Caparros and Prat (2021).

Early Pleistocene hominin subsistence behaviors in South Africa: Evidence from the hominin-bearing deposit of Cooper's D (Bloubank Valley, South Africa).

Evidence of the consumption of meat through hunting or scavenging by Early Pleistocene hominins is scarce, particularly in South Africa. Moreover, the interpretations of taphonomic evidence are subject to an important discussion commonly called the 'hunting-vs-scavenging debate.' Until today, only the Swartkrans Members 1-3 site has yielded a butchered bone assemblage large enough to permit reconstruction of carcass acquisition strategies by Early Pleistocene hominins in South Africa. This leaves an information gap between 1.4 and 1.0 Ma. Here, we provide the first evidence of meat consumption by hominins during this gap, based on the zooarchaeological study of the large mammal bone assemblage recovered from the Cooper's D site, South Africa. Based on skeletal part representation, our results show density-mediated attrition of bovid bones due to predepositional and postdepositional destruction. We argue that this attrition is the result of both abiotic (i.e., decalcification) and biotic (i.e., carnivore ravaging) processes. Bovid mortality profiles point out the involvement of ambush predators such as large felids. Bone surface modifications also indicate that the assemblage has been accumulated mostly by carnivores but with some hominin involvement as well. We observe all the stages of animal carcass processing (skinning, disarticulation, defleshing, marrow extraction) as well as the exploitation of a diversity of prey size classes at both Swartkrans Members 1-3 and Cooper's D. Thus, our study shows the importance of the Cooper's D bone assemblage for understanding Early Pleistocene hominin subsistence behaviors. Moreover, this article highlights the need for including long bone flake specimens in the analysis of large bone assemblages from South African caves to better understand the Early Pleistocene hominin bone damage record.

A Phylogenetic Networks perspective on reticulate human evolution.

We present a methodological phylogenetic reconstruction approach combining Maximum Parsimony and Phylogenetic Networks methods for the study of human evolution applied to phenotypic craniodental characters of 22 hominin species. The approach consists in selecting and validating a tree-like most parsimonious scenario out of several parsimony runs based on various numerical constraints. An intermediate step from tree to network methods is implemented by running an analysis with a reduced apomorphous character dataset that generates multiple parsimonious trees. These most parsimonious trees are then used as input for a Phylogenetic Networks analysis that results in consensus and reticulate networks. We show here that the phylogenetic tree-like definition of the genus Homo is a relative concept linked to craniodental characters that come in support of hypothetical Last Common Ancestors of the most parsimonious scenario and infer that the Homo reticulate network concords with recent findings in paleogenomic research regarding its mode of evolution.

Balancing costs and benefits in primates: ecological and palaeoanthropological views.

Maintaining the balance between costs and benefits is challenging for species living in complex and dynamic socio-ecological environments, such as primates, but also crucial for shaping life history, reproductive and feeding strategies. Indeed, individuals must decide to invest time and energy to obtain food, services and partners, with little direct feedback on the success of their investments. Whereas decision-making relies heavily upon cognition in humans, the extent to which it also involves cognition in other species, based on their environmental constraints, has remained a challenging question. Building mental representations relating behaviours and their long-term outcome could be critical for other primates, but there are actually very little data relating cognition to real socio-ecological challenges in extant and extinct primates. Here, we review available data illustrating how specific cognitive processes enable(d) modern primates and extinct hominins to manage multiple resources (e.g. food, partners) and to organize their behaviour in space and time, both at the individual and at the group level. We particularly focus on how they overcome fluctuating and competing demands, and select courses of action corresponding to the best possible packages of potential costs and benefits in reproductive and foraging contexts. This article is part of the theme issue 'Existence and prevalence of economic behaviours among non-human primates'.

Refining the ecological brain: Strong relation between the ventromedial prefrontal cortex and feeding ecology in five primate species.

To survive in complex and seasonal environments, primates are thought to rely upon cognitive capacities such as decision-making and episodic memory, which enable them to plan their daily foraging path. According to the Ecological Brain hypothesis, feeding ecology has driven the expansion of the brain to support the corresponding development of cognitive skills. Recent works in cognitive neurosciences indicate that cognitive operations such as decision-making or subjective evaluation (which are contextual and dependent upon episodic memory), relied critically upon a small part of the frontal lobe, often referred to as the ventromedial prefrontal cortex (VMPFC). Several authors suggested that this area might be important for foraging, but this has never been tested. In the present study, we quantified the relation between the size of the VMPFC (along with other cerebral measures: the whole brain, the gyrus rectus and the somatosensory cortex) and key socio-ecological variables in five primate species (Macaca mulatta, Macaca fuscata, Gorilla gorilla, Pan troglodytes and Homo sapiens). We hypothesized that the size of the VMPFC would be greater in primates with a large dietary spectrum and complex foraging strategies. We also hypothesized that the impact of feeding ecology would be stronger on this specific region than on other regions (somatosensory cortex) or on more global cerebral measures (e.g., whole brain). In line with these hypotheses, we found that all cerebral measures were more strongly related to feeding ecology than group size, a proxy for social complexity. As expected, the VMPFC volume is more precisely related to feeding ecology than the whole brain, and appears to be critically related to dietary quality. Thus, combining a comparative approach with predictions coming both from behavioral ecology and cognitive neurosciences, our study provides evidence that feeding ecology played a key role in the development of specific cognitive skills, which rely upon the expansion of a specific cortical area.

Isotopic analyses suggest mammoth and plant in the diet of the oldest anatomically modern humans from far southeast Europe.

Relatively high 15N abundances in bone collagen of early anatomically modern humans in Europe have often been interpreted as a specific consumption of freshwater resources, even if mammoth is an alternative high 15N prey. At Buran-Kaya III, access to associated fauna in a secured archaeological context and application of recently developed isotopic analyses of individuals amino acids offer the opportunity to further examine this hypothesis. The site of Buran-Kaya III is located in south Crimea and has provided a rich archaeological sequence including two Upper Palaeolithic layers, from which human fossils were retrieved and directly dated as from 37.8 to 33.1 ka cal BP. Results from bulk collagen of three human remains suggests the consumption of a high 15N prey besides the contribution of saiga, red deer, horse and hare, whose butchered remains were present at the site. In contrast to bulk collagen, phenylalanine and glutamic acid 15N abundances reflect not only animal but also plant protein contributions to omnivorous diet, and allow disentangling aquatic from terrestrial resource consumption. The inferred human trophic position values point to terrestrial-based diet, meaning a significant contribution of mammoth meat, in addition to a clear intake of plant protein.

3.3-million-year-old stone tools from Lomekwi 3, West Turkana, Kenya.

Human evolutionary scholars have long supposed that the earliest stone tools were made by the genus Homo and that this technological development was directly linked to climate change and the spread of savannah grasslands. New fieldwork in West Turkana, Kenya, has identified evidence of much earlier hominin technological behaviour. We report the discovery of Lomekwi 3, a 3.3-million-year-old archaeological site where in situ stone artefacts occur in spatiotemporal association with Pliocene hominin fossils in a wooded palaeoenvironment. The Lomekwi 3 knappers, with a developing understanding of stone's fracture properties, combined core reduction with battering activities. Given the implications of the Lomekwi 3 assemblage for models aiming to converge environmental change, hominin evolution and technological origins, we propose for it the name 'Lomekwian', which predates the Oldowan by 700,000 years and marks a new beginning to the known archaeological record.

The oldest anatomically modern humans from far southeast Europe: direct dating, culture and behavior.

BACKGROUND: Anatomically Modern Humans (AMHs) are known to have spread across Europe during the period coinciding with the Middle to Upper Paleolithic transition. Whereas their dispersal into Western Europe is relatively well established, evidence of an early settlement of Eastern Europe by modern humans are comparatively scarce. METHODOLOGY/PRINCIPAL FINDING: Based on a multidisciplinary approach for the study of human and faunal remains, we describe here the oldest AMH remains from the extreme southeast Europe, in conjunction with their associated cultural and paleoecological background. We applied taxonomy, paleoecology, and taphonomy combined with geomorphology, stratigraphy, archeology and radiocarbon dating. More than 160 human bone remains have been discovered. They originate from a well documented Upper Paleolithic archeological layer (Gravettian cultural tradition) from the site of Buran-Kaya III located in Crimea (Ukraine). The combination of non-metric dental traits and the morphology of the occipital bones allow us to attribute the human remains to Anatomically Modern Humans. A set of human and faunal remains from this layer has been radiocarbon dated by Accelerator Mass Spectrometry. The direct-dating results of human bone establish a secure presence of AMHs at 31,900+240/-220 BP in this region. They are the oldest direct evidence of the presence of AMHs in a well documented archeological context. Based on taphonomical observations (cut marks and distribution of skeletal elements), they represent the oldest Upper Paleolithic modern humans from Eastern Europe, showing post-mortem treatment of the dead as well. CONCLUSION/SIGNIFICANCE: These findings are essential for the debate on the spread of modern humans in Europe during the Upper Paleolithic, as well as their cultural behaviors.

First occurrence of early Homo in the Nachukui Formation (West Turkana, Kenya) at 2.3-2.4 Myr.

Cognitive abilities and techno-economic behaviours of hominids in the time period between 2.6-2.3 Myr have become increasingly well-documented. This time period corresponds to the oldest evidence for stone tools at Gona (Kada Gona, West Gona, EG 10-12, OGS 6-7), Hadar (AL 666), lower Omo valley (Ftji1, 2 & 5, Omo 57, Omo 123) in Ethiopia, and West Turkana (Lokalalei sites -LA1 & LA2C-) in Kenya. In 2002 a new palaeoanthropological site (LA1alpha), 100 meters south of the LA1 archaeological site, produced a first right lower molar of a juvenile hominid (KNM-WT 42718). The relative small size of the crown, its marked MD elongation and BL reduction, the relative position of the cusps, the lack of a C6 and the mild expression of a protostylid, reinforced by metrical analyses, demonstrate the distinctiveness of this tooth compared with Australopithecus afarensis, A. anamensis, A. africanus and Paranthropus boisei, and its similarity to early Homo. The LA1alpha site lies 2.2 m above the Ekalalei Tuff which is slightly younger than Tuff F dated to 2.34+/-0.04 Myr. This juvenile specimen represents the oldest occurrence of the genus Homo in West Turkana.