The taxonomy of Sahelanthropus tchadensis from a craniometric perspective.

Sahelanthropus tchadensis has raised much debate since its initial discovery in Chad in 2001, given its controversial classification as the earliest representative of the hominin lineage. This debate extends beyond the phylogenetic position of the species, and includes several aspects of its habitual behavior, especially in what regards its locomotion. The combination of ancestral and derived traits observed in the fossils associated with the species has been used to defend different hypotheses related to its relationship to hominins. Here, the cranial morphology of Sahelanthropus tchadensis was assessed through 16 linear craniometric measurements, and compared to great apes and hominins through Principal Component Analysis based on size and shape and shape information alone. The results show that S. tchadensis share stronger morphological affinities with hominins than with apes for both the analysis that include size information and the one that evaluates shape alone. Since TM 266-01-060-1 shows a strong morphological affinity with the remaining hominins represented in the analysis, our results support the initial interpretations that S. tchadensis represents an early specimen of our lineage or a stem basal lineage more closely related to hominins than to Panini.

When is a handaxe a planned-axe? exploring morphological variability in the Acheulean.

The handaxe is an iconic stone tool form used to define and symbolise both the Acheulean and the wider Palaeolithic. There has long been debate around the extent of its morphological variability between sites, and the role that extrinsic factors (especially raw material, blank type, and the extent of resharpening) have played in driving this variability, but there has been a lack of high-resolution examinations of these factors in the same study. In this paper, we present a 2D geometric morphometric analysis of 1097 handaxes from across Africa, the Levant, and western Europe to examine the patterning of this variability and what it can tell us about hominin behaviour. We replicate the findings of previous studies, that handaxe shape varies significantly between sites and entire continental regions, but we find no evidence for raw material, blank type, or resharpening in determining this pattern. What we do find, however, is that markers of reduction trajectory vary substantially between sites, suggesting that handaxes were deployed differently according to hominin need at a given site. We argue this is reflective of a continuum of reduction strategies, from those focused on the maintenance of a sharp cutting edge (i.e. direct use in cutting activities), to those focused on maintaining tip shapes, and perhaps a corresponding production of flakes. Implications for hominin behavioural flexibility are discussed.

A rhinopithecus swarm optimization algorithm for complex optimization problem.

This paper introduces a novel meta-heuristic algorithm named Rhinopithecus Swarm Optimization (RSO) to address optimization problems, particularly those involving high dimensions. The proposed algorithm is inspired by the social behaviors of different groups within the rhinopithecus swarm. RSO categorizes the swarm into mature, adolescent, and infancy individuals. Due to this division of labor, each category of individuals employs unique search methods, including vertical migration, concerted search, and mimicry. To evaluate the effectiveness of RSO, we conducted experiments using the CEC2017 test set and three constrained engineering problems. Each function in the test set was independently executed 36 times. Additionally, we used the Wilcoxon signed-rank test and the Friedman test to analyze the performance of RSO compared to eight well-known optimization algorithms: Dung Beetle Optimizer (DBO), Beluga Whale Optimization (BWO), Salp Swarm Algorithm (SSA), African Vultures Optimization Algorithm (AVOA), Whale Optimization Algorithm (WOA), Atomic Retrospective Learning Bare Bone Particle Swarm Optimization (ARBBPSO), Artificial Gorilla Troops Optimizer (GTO), and Harris Hawks Optimization (HHO). The results indicate that RSO exhibited outstanding performance on the CEC2017 test set for both 30 and 100 dimension. Moreover, RSO ranked first in both dimensions, surpassing the mean rank of the second-ranked algorithms by 7.69% and 42.85%, respectively. Across the three classical engineering design problems, RSO consistently achieves the best results. Overall, it can be concluded that RSO is particularly effective for solving high-dimensional optimization problems.

3.3 million years of stone tool complexity suggests that cumulative culture began during the Middle Pleistocene.

Cumulative culture, the accumulation of modifications, innovations, and improvements over generations through social learning, is a key determinant of the behavioral diversity across Homo sapiens populations and their ability to adapt to varied ecological habitats. Generations of improvements, modifications, and lucky errors allow humans to use technologies and know-how well beyond what a single naive individual could invent independently within their lifetime. The human dependence on cumulative culture may have shaped the evolution of biological and behavioral traits in the hominin lineage, including brain size, body size, life history, sociality, subsistence, and ecological niche expansion. Yet, we do not know when, in the human career, our ancestors began to depend on cumulative culture. Here, we show that hominins likely relied on a derived form of cumulative culture by at least ~600 kya, a result in line with a growing body of existing evidence. We analyzed the complexity of stone tool manufacturing sequences over the last 3.3 My of the archaeological record. We then compare these to the achievable complexity without cumulative culture, which we estimate using nonhuman primate technologies and stone tool manufacturing experiments. We find that archaeological technologies become significantly more complex than expected in the absence of cumulative culture only after ~600 kya.

The endocast morphology of LES1, Homo naledi.

OBJECTIVES: Homo naledi is near the extreme of small brain size within Homo but is easily recognized as Homo in other aspects of endocast morphology. This study adds new evidence of the endocast morphology of H. naledi by describing the Lesedi Hominin 1 (LES1) endocranium from the Lesedi Chamber and compares it to the previously known H. naledi individual Dinaledi Hominin 3 (DH3) as well as other hominin taxa. MATERIALS AND METHODS: We examined interlandmark distances with both univariate and multivariate methods in multiple hominin taxa and both species of Pan. For each distance, we compared groups using adjusted Z-scores (Azs). Our multivariate analyses included both principal component analyses (PCA) and linear discriminant analyses (LDA). RESULTS: DH3 and LES1 each have absolute third frontal convolution measures that enter the upper half of the variation for Homo sapiens, Homo erectus, and Homo neanderthalensis. Examined relative to the cube root of endocranial volume, H. naledi ranks among the highest values in these samples of Homo. Both absolute and relative values for H. naledi specimens are far above Pan, Australopithecus, and Paranthropus, suggesting an expanded Broca's area. CONCLUSIONS: Both qualitative and quantitative analyses show consistency between LES1 and other H. naledi endocasts and confirm the shared morphology of H. naledi with H. sapiens, H. neanderthalensis, and some specimens of H. erectus.

Left ventricular trabeculation in Hominidae: divergence of the human cardiac phenotype.

Although the gross morphology of the heart is conserved across mammals, subtle interspecific variations exist in the cardiac phenotype, which may reflect evolutionary divergence among closely-related species. Here, we compare the left ventricle (LV) across all extant members of the Hominidae taxon, using 2D echocardiography, to gain insight into the evolution of the human heart. We present compelling evidence that the human LV has diverged away from a more trabeculated phenotype present in all other great apes, towards a ventricular wall with proportionally greater compact myocardium, which was corroborated by post-mortem chimpanzee (Pan troglodytes) hearts. Speckle-tracking echocardiographic analyses identified a negative curvilinear relationship between the degree of trabeculation and LV systolic twist, revealing lower rotational mechanics in the trabeculated non-human great ape LV. This divergent evolution of the human heart may have facilitated the augmentation of cardiac output to support the metabolic and thermoregulatory demands of the human ecological niche.

Buronius manfredschmidi-A new small hominid from the early late Miocene of Hammerschmiede (Bavaria, Germany).

The known diversity of European middle and late Miocene hominids has increased significantly during the last decades. Most of these great apes were frugivores in the broadest sense, ranging from soft fruit frugivores most like chimpanzees to hard/tough object feeders like orangutans, varying in size from larger than siamangs (over 17 kg) to larger than most chimpanzees (~60-70 kg). In contrast to the frequent sympatry of hominoids in the early-to-middle Miocene of Africa, in no European Miocene locality more than one hominid taxon has been identified. Here we describe the first case of hominid sympatry in Europe from the 11.62 Ma old Hammerschmiede HAM 5 level, best known from its excellent record of Danuvius guggenmosi. The new fossils are consistent in size with larger pliopithecoids but differ morphologically from any pliopithecoid and from Danuvius. They are also distinguished from early and middle Miocene apes, share affinities with late Miocene apes, and represent a small hitherto unknown late Miocene ape Buronius manfredschmidi. With an estimated body mass of about 10 kg it represents the smallest known hominid taxon. The relative enamel thickness of Buronius is thin and contrasts with Danuvius, whose enamel is twice as thick. The differences between Buronius and Danuvius in tooth and patellar morphology, enamel thickness and body mass are indicative of differing adaptations in each, permitting resource partitioning, in which Buronius was a more folivorous climber.

Phylogenetic differences in the morphology and shape of the central sulcus in great apes and humans: implications for the evolution of motor functions.

The central sulcus divides the primary motor and somatosensory cortices in many anthropoid primate brains. Differences exist in the surface area and depth of the central sulcus along the dorso-ventral plane in great apes and humans compared to other primate species. Within hominid species, there are variations in the depth and aspect of their hand motor area, or knob, within the precentral gyrus. In this study, we used post-image analyses on magnetic resonance images to characterize the central sulcus shape of humans, chimpanzees (Pan troglodytes), gorillas (Gorilla gorilla), and orangutans (Pongo pygmaeus and Pongo abelii). Using these data, we examined the morphological variability of central sulcus in hominids, focusing on the hand region, a significant change in human evolution. We show that the central sulcus shape differs between great ape species, but all show similar variations in the location of their hand knob. However, the prevalence of the knob location along the dorso-ventral plane and lateralization differs between species and the presence of a second ventral motor knob seems to be unique to humans. Humans and orangutans exhibit the most similar and complex central sulcus shapes. However, their similarities may reflect divergent evolutionary processes related to selection for different positional and habitual locomotor functions.

An ape partial postcranial skeleton (KNM-NP 64631) from the Middle Miocene of Napudet, northern Kenya.

An ape partial postcranial skeleton (KNM-NP 64631) was recovered during the 2015-2021 field seasons at Napudet, a Middle Miocene (∼13 Ma) locality in northern Kenya. Bony elements representing the shoulder, elbow, hip, and ankle joints, thoracic and lumbar vertebral column, and hands and feet, offer valuable new information about the body plan and positional behaviors of Middle Miocene apes. Body mass estimates from femoral head dimensions suggest that the KNM-NP 64631 individual was smaller-bodied (c. 13-17 kg) than some Miocene taxa from eastern Africa, including Ekembo nyanzae, and probably Equatorius africanus or Kenyapithecus wickeri, and was more comparable to smaller-bodied male Nacholapithecus kerioi individuals. Similar to many Miocene apes, the KNM-NP 64631 individual had hip and hallucal tarsometatarsal joints reflecting habitual hindlimb loading in a variety of postures, a distal tibia with a large medial malleolus, an inflated humeral capitulum, probably a long lumbar spine, and a long pollical proximal phalanx relative to femoral head dimensions. The KNM-NP 64631 individual departs from most Early Miocene apes in its possession of a more steeply beveled radial head and deeper humeral zona conoidea, reflecting enhanced supinating-pronating abilities at the humeroradial joint. The KNM-NP 64631 individual also differs from Early Miocene Ekembo heseloni in having a larger elbow joint (inferred from radial head size) relative to the mediolateral width of the lumbar vertebral bodies and a more asymmetrical talar trochlea, and in these ways recalls inferred joint proportions for, and talocrural morphology of, N. kerioi. Compared to most Early Miocene apes, the KNM-NP 64631 individual likely relied on more forelimb-dominated arboreal behaviors, perhaps including vertical climbing (e.g., extended elbow, hoisting). Moreover, the Napudet ape partial postcranial skeleton suggests that an arboreally adapted body plan characterized by relatively large (here, based on joint size) forelimbs, but lacking orthograde suspensory adaptations, may not have been 'unusual' among Middle Miocene apes.

How the Sima de los Huesos was won.

Although the first discovery of a human fossil in the Sima de los Huesos took place in 1976, systematic excavations did not begin there until 1984. Since then, this site has been continuously excavated in month-long camps. The site is dated by different radiometric techniques to between 430,000 and 300,000 years ago. Until the 2023 campaign, just over 7000 human fossils have been recovered, constituting the largest collection of fossils prior to Homo sapiens ever discovered. The fossils correspond to a minimum of 29 individuals of both sexes and different ages at death, from preadolescents to a specimen of advanced age. Comparative anatomy and ancient DNA studies both suggest that this is a population closely related to Homo neanderthalensis. The great variety and extraordinary quality of the fossils recovered have allowed us to carry out a series of investigations that have greatly increased our knowledge about the evolution of Homo in the Middle Pleistocene. Among the most important discoveries, it has been possible to establish body size and proportions, the confirmation that the origin of the accumulation of human fossils was of an anthropic nature, that those past humans took care of disabled individuals and who were capable of having an oral language almost as complex and efficient as that of our own species.

Edge chipping patterns in posterior teeth of hominins and apes.

Chip scars in fossil teeth are a lasting evidence that bears on human evolution. Chip dimensions in posterior teeth of hominins, apes and white-lipped peccary (Tayassu pecari) are measured from published occlusal images. The results are plotted as D/Dm vs. h/Dm, where h, D and Dm denote indent distance, chip width and mean tooth crown diameter. The hominin species follow a similar pattern where D/Dm monotonically increases up to h/Dm ≈ 0.3. The behavior for the apes is characterized by two phases. In the first, h/Dm monotonically increases up to h/Dm ≈ 0.26 while in the second (h/Dm ≈ 0.26 to 0.42), D/Dm experiences a drastic change in behavior. The interpretation of chip morphology is assisted by results from controlled spherical indentation tests on extracted human molars. This study shows that in addition to the commonly recognized chipping due to cusp loading, a chip may also initiate from the inner wall of the tooth's central fossa. Accordingly, it is suggested that the chipping in hominins generally initiates from a (worn) cusp while that in apes involves cusp loading up to h/Dm ≈ 0.26 and fossa loading thereafter. The behavior for T. pecari is much similar to that of the apes. The fossa chipping is facilitated by a consumption of hard, large-size diet (e.g., plants, roots, barks and nuts) and presence of broad central fossa, conditions that are met in apes. Finally, a simple expression for the critical chipping force Pch due to fossa loading is developed.

Illuminating the function of the orphan transporter, SLC22A10, in humans and other primates.

SLC22A10 is an orphan transporter with unknown substrates and function. The goal of this study is to elucidate its substrate specificity and functional characteristics. In contrast to orthologs from great apes, human SLC22A10, tagged with green fluorescent protein, is not expressed on the plasma membrane. Cells expressing great ape SLC22A10 orthologs exhibit significant accumulation of estradiol-17ß-glucuronide, unlike those expressing human SLC22A10. Sequence alignments reveal a proline at position 220 in humans, which is a leucine in great apes. Replacing proline with leucine in SLC22A10-P220L restores plasma membrane localization and uptake function. Neanderthal and Denisovan genomes show proline at position 220, akin to modern humans, indicating functional loss during hominin evolution. Human SLC22A10 is a unitary pseudogene due to a fixed missense mutation, P220, while in great apes, its orthologs transport sex steroid conjugates. Characterizing SLC22A10 across species sheds light on its biological role, influencing organism development and steroid homeostasis.

Ethnography and ethnohistory support the efficiency of hunting through endurance running in humans.

Humans have two features rare in mammals: our locomotor muscles are dominated by fatigue-resistant fibres and we effectively dissipate through sweating the metabolic heat generated through prolonged, elevated activity. A promising evolutionary explanation of these features is the endurance pursuit (EP) hypothesis, which argues that both traits evolved to facilitate running down game by persistence. However, this hypothesis has faced two challenges: running is energetically costly and accounts of EPs among late twentieth century foragers are rare. While both observations appear to suggest that EPs would be ineffective, we use foraging theory to demonstrate that EPs can be quite efficient. We likewise analyse an ethnohistoric and ethnographic database of nearly 400 EP cases representing 272 globally distributed locations. We provide estimates for return rates of EPs and argue that these are comparable to other pre-modern hunting methods in specified contexts. EP hunting as a method of food procurement would have probably been available and attractive to Plio/Pleistocene hominins.

The complete sequence and comparative analysis of ape sex chromosomes.

Apes possess two sex chromosomes-the male-specific Y chromosome and the X chromosome, which is present in both males and females. The Y chromosome is crucial for male reproduction, with deletions being linked to infertility1. The X chromosome is vital for reproduction and cognition2. Variation in mating patterns and brain function among apes suggests corresponding differences in their sex chromosomes. However, owing to their repetitive nature and incomplete reference assemblies, ape sex chromosomes have been challenging to study. Here, using the methodology developed for the telomere-to-telomere (T2T) human genome, we produced gapless assemblies of the X and Y chromosomes for five great apes (bonobo (Pan paniscus), chimpanzee (Pan troglodytes), western lowland gorilla (Gorilla gorilla gorilla), Bornean orangutan (Pongo pygmaeus) and Sumatran orangutan (Pongo abelii)) and a lesser ape (the siamang gibbon (Symphalangus syndactylus)), and untangled the intricacies of their evolution. Compared with the X chromosomes, the ape Y chromosomes vary greatly in size and have low alignability and high levels of structural rearrangements-owing to the accumulation of lineage-specific ampliconic regions, palindromes, transposable elements and satellites. Many Y chromosome genes expand in multi-copy families and some evolve under purifying selection. Thus, the Y chromosome exhibits dynamic evolution, whereas the X chromosome is more stable. Mapping short-read sequencing data to these assemblies revealed diversity and selection patterns on sex chromosomes of more than 100 individual great apes. These reference assemblies are expected to inform human evolution and conservation genetics of non-human apes, all of which are endangered species.

Estimating ancestral ranges and biogeographical processes in early hominins.

Historical biogeography provides crucial insights into understanding the evolutionary history of hominins. We applied maximum-likelihood and biogeographical stochastic mapping to infer the ancestral ranges of hominins and estimate the frequency of biogeographical events. These events were inferred using two time-calibrated phylogenetic trees that differ in the position of Australopithecus sediba. Results suggest that regardless of which phylogeny was selected, Northcentral Africa was the preferred ancestral region for the ancestor of the Homo-Pan clade, as well as the ancestor of Sahelanthropus and later hominins. The northern and middle part of eastern Africa was the preferred ancestral region for several clades originating at subsequent deep nodes of the trees (∼5-4 Ma). The choice of tree topology had one important effect on results: whether hominin ancestors appearing after ∼4 Ma were widespread or endemic. These different patterns highlight the biogeographic significance of the phylogenetic relationships of A. sediba. Overall, the results showed that dispersal, local extinction, and sympatry played vital roles in creating the hominin distribution, whereas vicariance and jump dispersal were not as common. The results suggested symmetry in the directionality of dispersals. Distance probably influenced how rapidly taxa colonized a new region, and dispersals often followed the closest path. These findings are potentially impacted by the imperfection of the fossil record, suggesting that the results should be interpreted cautiously.

Virtual reconstruction and geometric morphometric analysis of the Kocabas hominin fossil from Turkey: Implications for taxonomy and evolutionary significance.

The Kocabas specimen comes from a travertine quarry near the homonymous village in the Denizli basin (Turkey). The specimen comprises three main fragments: portions of the right and left parietal and left and right parts of the frontal bone. The fossil was assumed to belong to the Homo erectus s.l. hypodigm by some authors, whereas others see similarities with Middle Pleistocene fossils (Broken Hill 1/Kabwe, Bodo, or Ceprano). Here, we present the first attempt to make a complete reconstruction of the missing medial portion of the frontal bone and a comprehensive geometric morphometric analysis of this bone. We restored the calotte by aligning and mirroring the three preserved fragments. Afterward, we restored the missing portion by applying the thin-plate spline interpolation algorithm of target fossils onto the reconstructed Kocabas specimen. For the geometric morphometric analyses, we collected 80 landmarks on the frontal bone (11 osteometric points, 14 bilateral curve semilandmarks, and 41 surface semilandmarks). The comparative sample includes 21 fossils from different chronological periods and geographical areas and 30 adult modern humans from different populations. Shape analyses highlighted the presence in Kocabas of features usually related to Middle Pleistocene Homo, such as a developed supraorbital torus associated with a relatively short frontal squama and reduced post-toral sulcus. Cluster analysis and linear discriminant analysis classification procedure suggest Kocabas being part of the same taxonomic unit of Eurasian and African Middle Pleistocene Homo. In light of our results, we consider that attributing the Kocabas hominin to H. erectus s.l. may be unwarranted. Results of our analyses are compatible with different evolutionary scenarios, but a more precise chronological framework is needed for a thorough discussion of the evolutionary significance of this specimen. Future work should clarify its geological age, given uncertainties regarding its stratigraphic provenance.

Hominin turnover at Laetoli is associated with vegetation change: Multiproxy evidence from the large herbivore community.

Vegetation change in eastern Africa during the Pliocene would have had an important impact on hominin adaptation and ecology, and it may have been a key driver of hominin macroevolution, including the extinction of Australopithecus and the emergence of Paranthropus and Homo. The Pliocene paleoanthropological site of Laetoli in Tanzania provides an opportunity to investigate the relationship between vegetation change and hominin turnover because it encompasses the time period when grass cover was spreading across eastern Africa and because hominin species turnover occurred locally at Laetoli, with Paranthropus aethiopicus in the Upper Ndolanya Beds (UNB) replacing Australopithecus afarensis in the Upper Laetolil Beds (ULB). However, it remains unresolved how the vegetation of the UNB and the ULB differed from each other. To examine differences between the two stratigraphic units, multiple proxies-hypsodonty, mesowear, and stable carbon isotopes of tooth enamel (δ13Cenamel)-are used to infer the diets of large herbivores and compare the dietary guild structure of the large herbivore communities. All three proxies indicate an increase in the abrasiveness and C4-content in the diets of the large herbivores in the UNB relative to those in the ULB. After inferring the diets of species based on all three proxies, the large herbivore community of the UNB had a greater proportion of grazers and a smaller proportion of mixed feeders than in the ULB but maintained a similar proportion of browsers and frugivores. The ULB community has few modern-day analogs, whereas the UNB community is most closely analogous to those in modern African grasslands. Thus, hominin turnover at Laetoli is associated with an increase in grass cover within a woodland-grassland mosaic and is part of a broader transformation of the herbivore community structure.

Evolutionary and functional analyses of LRP5 in archaic and extant modern humans.

BACKGROUND: The human lineage has undergone a postcranial skeleton gracilization (i.e. lower bone mass and strength relative to body size) compared to other primates and archaic populations such as the Neanderthals. This gracilization has been traditionally explained by differences in the mechanical load that our ancestors exercised. However, there is growing evidence that gracilization could also be genetically influenced. RESULTS: We have analyzed the LRP5 gene, which is known to be associated with high bone mineral density conditions, from an evolutionary and functional point of view. Taking advantage of the published genomes of archaic Homo populations, our results suggest that this gene has a complex evolutionary history both between archaic and living humans and within living human populations. In particular, we identified the presence of different selective pressures in archaics and extant modern humans, as well as evidence of positive selection in the African and South East Asian populations from the 1000 Genomes Project. Furthermore, we observed a very limited evidence of archaic introgression in this gene (only at three haplotypes of East Asian ancestry out of the 1000 Genomes), compatible with a general erasing of the fingerprint of archaic introgression due to functional differences in archaics compared to extant modern humans. In agreement with this hypothesis, we observed private mutations in the archaic genomes that we experimentally validated as putatively increasing bone mineral density. In particular, four of five archaic missense mutations affecting the first ß-propeller of LRP5 displayed enhanced Wnt pathway activation, of which two also displayed reduced negative regulation. CONCLUSIONS: In summary, these data suggest a genetic component contributing to the understanding of skeletal differences between extant modern humans and archaic Homo populations.