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Fossil gibbons are exceedingly rare, with much of the hylobatid fossil record and, consequently, hylobatid evolutionary history remaining unknown. Kapi ramnagarensis was described as a stem hylobatid on the basis of an isolated lower right M 3 from ∼13.0-12.5 Ma deposits surrounding Ramnagar (J&K), India. This interpretation was recently challenged, with alternative hypotheses suggesting that it is instead a stem catarrhine or a strangely derived pliopithecoid that has converged on hylobatid morphology. A series of morphological features were said to distinguish Kapi from fossil and extant hylobatids; notably, however, none of these features were examined or compared using quantitative analyses. Here, we further examine the dental morphology of Kapi, providing quantitative analyses to critically evaluate the hypothesis that Kapi represents a stem catarrhine or pliopithecoid rather than a stem hylobatid. Results demonstrate that none of the claimed differences between Kapi and hylobatids hold up under closer scrutiny, and multivariate discriminant analyses taking size and shape into account strongly support Kapi as a hylobatid with high posterior probabilities. Although only represented by a single lower molar, Kapi remains the most compelling candidate for the earliest known hylobatid in the fossil record and thus likely documents the simultaneous arrival of lesser and great apes to Asia during the Middle Miocene., Competing Interests: Declaration of competing interest The authors declare no competing interests., (Copyright © 2024 Elsevier Ltd. All rights reserved.)

The phylogenetic relationships of the small-bodied catarrhine Pliobates cataloniae (∼11.6 Ma, NE Iberian Peninsula) have been controversial since its original description. However, the recent report of additional dentognathic remains has supported its crouzeliid pliopithecoid status. Based on the available hypodigm, the molar enamel-dentine junction (EDJ) shape of P. cataloniae is compared with that of other pliopithecoids from the same basin as well as both extinct and extant hominoids to further evaluate its pliopithecoid affinities. We also quantitatively compare the EDJ shape among these taxa by means of landmark-based three-dimensional geometric morphometrics using principal component analysis (PCA), canonical variate analysis, and between-group PCA. Permutation tests are performed to test whether Pliobates variation exceeds that of extant hominoid genera. Results indicate that Pliobates is similar in molar EDJ shape to other pliopithecoids, particularly crouzeliids. The variation displayed by Pliobates upper molars is less marked at the EDJ level than at the outer enamel surface, probably owing to differential enamel wear and intraspecific differences in enamel thickness. Multivariate analyses of EDJ shape show that all pliopithecoids (including Pliobates) cluster together in the PCAs, canonical variate analyses, and between-group PCAs and occupy a different portion of the morphospaces from extinct and extant hominoids. Posterior and typicality probabilities strongly support the classification of Pliobates as a pliopithecoid, wheras permutation tests fail to reject the single-genus hypothesis for the P. cataloniae hypodigm. We conclude that P. cataloniae is a crouzeliid pliopithecoid, as recently supported by cladistic analyses of craniodental characters, and that previous cladistic results that supported a stem hominoid status are attributable to postcranial convergences with crown hominoids. Our results further highlight the potential of three-dimensional geometric morphometrics analyses of the EDJ shape for better informing fossil primate alpha-taxonomy by means of quantitatively testing hypotheses about tooth shape variation., Competing Interests: Declaration of competing interest There is no conflict of interest., (Copyright © 2024 Elsevier Ltd. All rights reserved.)

Whether the size of the prefrontal cortex (PFC) in humans is disproportionate when compared to other species is a persistent debate in evolutionary neuroscience. This question has left the study of over/under-expansion in other structures relatively unexplored. We therefore sought to address this gap by adapting anatomical areas from the digital atlases of 18 mammalian species, to create a common interspecies classification. Our approach used data-driven analysis based on phylogenetic generalized least squares to evaluate anatomical expansion covering the whole brain. Our main finding suggests a divergence in primate evolution, orienting the stereotypical mammalian cerebral proportion toward a frontal and parietal lobe expansion in catarrhini (primate parvorder comprising old world monkeys, apes, and humans). Cerebral lobe volumes slopes plotted for catarrhini species were ranked as parietal∼frontal > temporal > occipital, contrasting with the ranking of other mammalian species (occipital > temporal > frontal∼parietal). Frontal and parietal slopes were statistically different in catarrhini when compared to other species through bootstrap analysis. Within the catarrhini's frontal lobe, the prefrontal cortex was the principal driver of frontal expansion. Across all species, expansion of the frontal lobe appeared to be systematically linked to the parietal lobe. Our findings suggest that the human frontal and parietal lobes are not disproportionately enlarged when compared to other catarrhini. Nevertheless, humans remain unique in carrying the most relatively enlarged frontal and parietal lobes in an infraorder exhibiting a disproportionate expansion of these areas.

Pliopithecoids are a diverse group of Miocene catarrhine primates from Eurasia. Their positional behavior is still unknown, and many species are known exclusively from dentognathic remains. Here, we describe a proximal radius (IPS66267) from the late Miocene of Castell de Barberà (Vallès-Penedès Basin, NE Iberian Peninsula) that represents the first postcranial specimen of the pliopithecoid Barberapithecus huerzeleri. A body mass estimate based on the radius is compared with dental estimates, and its morphology is compared with that of extant and fossil anthropoids by qualitative means as well as by landmark-based three-dimensional geometric morphometrics. The estimated body mass of ∼5 kg for IPS66267 closely matches the dental estimates for the (female) holotype, thereby discounting an alternative attribution to the large-bodied hominoid recorded at Castell de Barberà. In multiple features (oval and moderately tilted head with a pronounced lateral lip and a restricted articular area for the capitulum; proximodistally expanded proximal radioulnar joint; and short, robust, and anteroposteriorly compressed neck), the specimen differs from hominoids and resembles instead extant nonateline monkeys and stem catarrhines. The results of the morphometric analysis further indicate that the Barberapithecus proximal radius shows closer similarities with nonsuspensory arboreal cercopithecoids and the dendropithecid Simiolus. From a locomotor viewpoint, the radius of Barberapithecus lacks most of the features functionally related to climbing and/or suspensory behaviors and displays instead a proximal radioulnar joint that would have been particularly stable under pronation. On the other hand, the Barberapithecus radius differs from other stem catarrhines in the less anteroposteriorly compressed and less tilted radial head with a deeper capitular fovea, suggesting a somewhat enhanced mobility at the elbow joint. We conclude that pronograde arboreal quadrupedalism was the main component of the locomotor repertoire of Barberapithecus but that, similar to other crouzeliids, it might have displayed better climbing abilities than pliopithecids., Competing Interests: Declaration of competing interest The authors declare no conflict of interest., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

The evolution of novel vertebral morphologies observed in humans and other extant hominoids may be related to changes in the magnitudes and/or patterns of covariation among traits. To examine this, we tested magnitudes of integration in the vertebral column of cercopithecoids and hominoids, including humans. Three-dimensional surface scans of 14 vertebral elements from 30 Cercopithecus, 32 Chlorocebus, 39 Macaca, 45 Hylobates, 31 Pan, and 86 Homo specimens were used. A resampling method was used to generate distributions of integration coefficient of variation scores for vertebral elements individually using interlandmark distances. Interspecific comparisons of mean integration coefficient of variation were conducted using Mann-Whitney U tests with Bonferroni adjustment. The results showed that hominoids generally had lower mean integration coefficient of variation than cercopithecoids. In addition, humans showed lower mean integration coefficient of variation than other hominoids in their last thoracic and lumbar vertebrae. Cercopithecoids and Hylobates showed relatively lower mean integration coefficient of variation in cervical vertebrae than in thoracolumbar vertebrae. Pan and Homo showed relatively lower mean integration coefficient of variation in the last thoracic and lumbar vertebrae in the thoracolumbar region, except for the L1 of Pan. The results suggest fewer integration-mediated constraints on the evolution of vertebral morphology in hominoids when compared with cercopithecoids. The weaker magnitudes of integration in lumbar vertebrae in humans when compared with chimpanzees likewise suggest fewer constraints on the evolution of novel lumbar vertebrae morphology in humans., Competing Interests: Declaration of competing interest None., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

Recent studies on hominin craniofacial evolution have focused on phenotypic integration or covariation among traits. Covariation is thought to significantly affect evolutionary trajectories, shaping the ways in which hominins and other primates could have evolved. However, the ways in which covariation itself evolves are not well understood. This study aims to investigate the role of phylogeny, development, body size, and positional behavior in shaping the strength of covariation in strepsirrhine and catarrhine primate crania (n = 1009, representing 11 genera). These factors may have been catalysts for change in the magnitude of covariation, and they have changed significantly during primate evolution and particularly hominin evolution. Modern humans in particular have slow developmental trajectories, large bodies, and a unique form of locomotion in the form of orthograde bipedalism. Variance of eigenvalues, mean integration, mean evolvability, and mean conditional evolvability was estimated and their relationship to the various factors described earlier was assessed using phylogenetic and nonphylogenetic analyses. Results indicate that some phylogenetic signal is present, but it is not equivalent across integration statistics or cranial regions. In particular, these results suggest that closely related species are more similar than more distantly related species in evolvability of the cranial base and integration of the face. Two divergent patterns were also identified, in which covariation and evolvability of the cranial base are linked to developmental rate, but those of the face are linked to body size. Neither locomotion nor posture appears related to covariation or evolvability of the primate cranium. These results suggest that overall low covariation observed in the hominin cranium may be a result of separate trends in different cranial regions., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

Pliopithecoids are an extinct group of catarrhine primates from the Miocene of Eurasia. More than 50 years ago, they were linked to hylobatids due to some morphological similarities, but most subsequent studies have supported a stem catarrhine status, due to the retention of multiple plesiomorphic features (e.g., the ectotympanic morphology) relative to crown catarrhines. More recently, some morphological similarities to hominoids have been noted, raising the question of whether they could be stem members of this clade. To re-evaluate these competing hypotheses, we examine the morphology of the semicircular canals of the bony labyrinth of the middle Miocene pliopithecid Epipliopithecus vindobonensis. The semicircular canals are suitable to test between these hypotheses because (1) they have been shown to embed strong phylogenetic signal and reliably discriminate among major clades; (2) several potential hominoid synapomorphies have been identified previously in the semicircular canals; and (3) semicircular canal morphology has not been previously described for any pliopithecoid. We use a deformation-based (landmark-free) three-dimensional geometric morphometric approach to compare Epipliopithecus with a broad primate sample of extant and extinct anthropoids. We quantify similarities in semicircular canal morphology using multivariate analyses, reconstruct ancestral morphotypes by means of a phylomorphospace approach, and identify catarrhine and hominoid synapomorphies based on discrete characters. Epipliopithecus semicircular canal morphology most closely resembles that of platyrrhines and Aegyptopithecus due to the retention of multiple anthropoid symplesiomorphies. However, Epipliopithecus is most parsimoniously interpreted as a stem catarrhine more derived than Aegyptopithecus due to the possession of a crown catarrhine synapomorphy (i.e., the rounded anterior canal), combined with the lack of other catarrhine and any hominoid synapomorphies. Some similarities with hylobatids and atelids are interpreted as homoplasies likely related to positional behavior. The semicircular canal morphology of Epipliopithecus thus supports the common view that pliopithecoids are stem catarrhines., Competing Interests: Declaration of competing interest The authors declare no conflict of interest., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

Objectives: Epigenetic mechanisms influence the development and maintenance of complex phenotypes and may also contribute to the evolution of species-specific phenotypes. With respect to skeletal traits, little is known about the gene regulation underlying these hard tissues or how tissue-specific patterns are associated with bone morphology or vary among species. To begin exploring these topics, this study evaluates one epigenetic mechanism, DNA methylation, in skeletal tissues from five nonhuman primate species which display anatomical and locomotor differences representative of their phylogenetic groups., Materials and Methods: First, we test whether intraspecific variation in skeletal DNA methylation is associated with intraspecific variation in femur morphology. Second, we identify interspecific differences in DNA methylation and assess whether these lineage-specific patterns may have contributed to species-specific morphologies. Specifically, we use the Illumina Infinium MethylationEPIC BeadChip to identify DNA methylation patterns in femur trabecular bone from baboons (n = 28), macaques (n = 10), vervets (n = 10), chimpanzees (n = 4), and marmosets (n = 6)., Results: Significant differentially methylated positions (DMPs) were associated with a subset of morphological variants, but these likely have small biological effects and may be confounded by other variables associated with morphological variation. Conversely, several species-specific DMPs were identified, and these are found in genes enriched for functions associated with complex skeletal traits., Discussion: Overall, these findings reveal that while intraspecific epigenetic variation is not readily associated with skeletal morphology differences, some interspecific epigenetic differences in skeletal tissues exist and may contribute to evolutionarily distinct phenotypes. This work forms a foundation for future explorations of gene regulation and skeletal trait evolution in primates., (© 2020 Wiley Periodicals, Inc.)

A diversity of pliopithecoids is known from Miocene localities in Europe, but until recently, this group was relatively poorly represented in China. However, new discoveries have shown that Chinese pliopithecoids were taxonomically diverse and geographically widespread. The earliest pliopithecoids in China (and Eurasia) are Dionysopithecus and Platodontopithecus from the Early Miocene of Sihong, Jiangsu (∼19-18 Ma). During the Middle Miocene (∼15-12 Ma), several species of pliopithecoids are recorded at localities in Gansu Province (Laogou), Inner Mongolia (Damiao), Xinjiang Uygur Autonomous Region (Tieersihabahe), and Ningxia Hui Autonomous Region (Tongxin). Finally, a late-surviving anapithecine crouzeliid, Laccopithecus robustus, is known from the Late Miocene (∼7 Ma) of Shihuiba in Yunnan, which postdates the extinction of pliopithecoids in Europe (during MN 10). Paleontological investigations at a late Early Miocene locality near Fanchang in Anhui Province have yielded a large sample of isolated teeth (more than one hundred) of a previously unknown species of pliopithecoid. The associated micromammals indicate an age contemporaneous with the Shanwang Formation in Shandong Province (MN 3-4, ∼18-17 Ma). All of the permanent teeth are represented except for I 2 . With its unique suite of dental features, the Fanchang pliopithecoid can be attributed to a new species and genus. Shared derived features of the lower molars confirm that the Fanchang pliopithecoid has its closest affinities with European crouzeliids, but a number of primitive traits indicate that it is a stem member of the clade. The evidence points to China as an important center for the early diversification of pliopithecoids. Contrary to previous zoogeographic scenarios, the occurrence of an early crouzeliid in China implies that the Pliopithecidae and Crouzeliidae may have diverged from a stem pliopithecoid in Asia during the Early Miocene before their arrival in Europe., Competing Interests: Declaration of competing interest The authors declare that there is no conflict of interest., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

Objectives: This study aimed to test the taxonomic utility of the catarrhine supraorbital region using 3D geometric morphometrics, with the aim of establishing its potential use in elucidating the position of more debated hominin groups., Materials and Methods: 230 3D coordinates were used to record the supraorbital morphology of two datasets: one containing 460 non-hominin catarrhine primates from species and subspecies of Gorilla, Pan, Papio, and Macaca; and the other containing 55 Pleistocene hominins from Homo, Australopithecus, and Paranthropus. Principal component analyses in tangent, form, and allometry-free shape space were used to assess differentiation of taxa, with biological distinctiveness of taxa being established using step-wise discriminant analysis with subsampling., Results: Results indicated that the recorded supraorbital morphology could be used to separate non-hominin catarrhine primate genera, species, and subspecies, although accuracy was found to decrease with decreasing Linnaean rank. In addition, analyses in tangent space were found to produce the highest accuracy when classifying primates of known taxonomy. Biological distinctiveness of the middle and later Homo species was comparable to or higher than that of the non-hominin primates, and relatively lower for the earlier groups of Homo., Discussion: This study indicates that the supraorbital region preserves taxonomic information that can be used to delineate between closely related groups, both within hominins and wider catarrhine primates. Therefore, this region may be used to provide insight when assessing the taxonomic affiliation of disputed hominin specimens., (© 2019 Wiley Periodicals, Inc.)

Previous investigations of the primate talo-crural joint (TCJ; specifically on the talus and distal tibia) have demonstrated that substrate preference significantly influences morphology, but this association is not necessarily found in subadults. This has been interpreted as the result of a plastic, behaviorally induced response of bone due to substrate use. In this investigation, we use geometric morphometric and phylogenetic comparative methods to investigate ontogenetic phenotypic plasticity in the catarrhine TCJ. Osteological specimens from four African hominoid and four cercopithecid species, divided into subadult and adult developmental stages based on molar eruption, formed the study group. We tested for phylogenetic signal in the shape of both the talar and tibial articular surfaces, at both developmental stages. We then used phylomorphospaces to examine the evolution of shape differences at each developmental stage for each element, and to determine if substrate usage is associated with shape in this phylogenetic context. A significant phylogenetic signal was found for both articular surfaces in subadults, but not adults. In phylomorphospace, both talar and tibial articular morphologies show an association with substrate preference in adults, but not in subadults. Our results provide confirmation of the significant effect of habitual substrate usage and the consequences of bone remodeling during ontogeny on the shape and presentation of the TCJ. These results also suggest caution when using adult talo-tibial shapes to evaluate phylogenetic relationships as TCJ morphology can be considered as a palimpsest, with substrate usage overwriting phylogenetic information in adult specimens. Anat Rec, 302:1977-1984, 2019. © 2019 American Association for Anatomy., (© 2019 American Association for Anatomy.)

The early Miocene site of Moroto II, Uganda has yielded some of the oldest known hominoid fossils. A new partial mandible (UMP MORII 03'551) is notable for its long tooth row and large, narrow M 2 with well-developed cristids - a morphological combination previously unknown for large bodied catarrhines of the Early Miocene and suggesting folivory. The tooth proportions are compatible with belonging to the same taxon as the maxilla UMP 62-11, the holotype of Morotopithecus bishopi; likewise, the long tooth row and vertical planum of UMP MORII 03'551 suggest that it may represent the same taxon as mandible(s) UMP 66-01 and UMP 62-10. Canine size strongly suggests UMP MORII 03'551 is a female. Comparisons of the tooth crown morphology and tooth row proportions, relative enamel thickness, enamel-dentine junction morphology, long-period line periodicity, and dental wear patterns support significant morphological, developmental, and inferred dietary differentiation, and therefore generic-level distinctiveness, among Afropithecus, Morotopithecus and the Proconsul clade. An isolated M 1 (UMP MORII 03'559) is morphologically dissimilar, and much smaller than the actual or inferred size of molars in UMP MORII 03'551, UMP 66-01 and UMP 62-10, supporting the presence of two hominoid taxa at Moroto II, M. bishopi and a smaller bodied proconsulid. Given the high level of body mass dimorphism inferred for Morotopithecus and other early Miocene catarrhines, the known postcrania from Moroto II could be attributable to either taxon. However, UMP MORII 03'551 and the femora UMP MORII 94'80 derive from the same stratigraphic interval, while the isolated M 1 was deposited later, increasing the likelihood that the mandible and femora are from the same individual. These new fossils expand our understanding of the taxonomic and adaptive diversity of early Miocene catarrhines., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

Kamoyapithecus hamiltoni is a potential early hominoid species described from fragmentary dentognathic specimens from the Oligocene site of Losodok (Turkana Basin, northwestern Kenya). Other catarrhine dental materials have been recovered at Losodok, but were not initially included in the Kamoyapithecus hypodigm. Here we present descriptions of the unpublished canine and incisor specimens from Losodok, and revisit the published specimens in light of recent changes in understanding of hominoid anterior dental evolution. The new fossils include a canine (KNM-LS 18354) that is morphologically compatible with being a female of Kamoyapithecus (KNM-LS 8). Randomization analyses of both Gorilla gorilla and middle Miocene Griphopithecus alpani demonstrate that the size difference between KNM-LS 8 and KNM-LS 18354 is also compatible with their being male and female canines of the same species. Significantly, a canine tip (KNM-LS 18352) attributed to Kamoyapithecus documents the distinctive burin tip morphology now recognized as characterizing Proconsul sensu stricto, which may indicate a close relationship between Kamoyapithecus and Proconsul. We also re-examined the enigmatic KNM-LS 1, a smaller lower canine assumed to derive from Losodok but for which historical provenience data are completely lacking. Elemental data derived from portable X-ray fluorescence spectroscopy show that KNM-LS 1 is almost certainly from Losodok rather than from nearby Miocene sites (i.e., Moruorot, Esha, Kalodirr). KNM-LS 1 displays a nyanzapithecine-like morphology and is shown by randomization analyses to be too small to be associated with the Kamoyapithecus canines. This demonstrates that there is a second hominoid taxon present at Losodok that records one of the earliest occurrences of the Nyanzapithecinae., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

The reduction in dental size and mandibular robusticity is regarded as a major trend in human evolution, traditionally considered the result of the peculiar extra-oral food processing skills of Homo. The use of stone tools and fire would have allowed our ancestors to chew softer food in smaller bite size, thus relaxing the selective pressures to keep a large dentition and a robust lower jaw. This perspective assumes that differences in dental size and mandibular robusticity in hominins represent functional dissimilarities. This study uses a catarrhine comparative approach to test this fundamental assumption of the hypotheses on dental and mandibular reduction in Homo. A sample of extant catarrhines and fossil hominins was used to test for correlations between dental size, mandibular robusticity, and dietary proxies, the latter include diet quality, diet heterogeneity, feeding time, and microwear variables. The effects of phylogeny and body size were considered. Findings support the association between technological developments in Homo and reduction in incisor size and mandibular corpus robusticity, though not for premolar, molar size, and symphyseal robusticity. These results challenge the functional interpretation of postcanine reduction and symphyseal changes in the genus Homo., (© 2019 Wiley Periodicals, Inc.)

Objectives: We tested whether patterns of intraspecific variation in catarrhine vertebral shape are consistent with developmental or functional predictions. Intraspecific variation was compared across column regions, morphological features, and species. Transitional regions and later ossifying morphological features were predicted to exhibit increased variation. The lumbosacral region, biomechanically important morphological features, and species with high locomotor demand and/or dedicated pronogrady were predicted to exhibit decreased variation., Materials and Methods: We used a modified Levene's test to compare intraspecific variation in dimensions of the neural canal, vertebral bodies, and spinous and transverse processes in lower thoracic to proximal sacral vertebrae. The sample included all hominoid genera and one cercopithecoid (Chlorocebus)., Results: We found little difference in variation across regions of the vertebral column. In hominoids, vertebral body dimensions were the least variable, neural canal dimensions the most variable, with spinous and transverse processes generally intermediate. Among species, there was a general though not always significant pattern for Chlorocebus to exhibit the least variation, followed by Homo or Hylobates., Discussion: Patterns of variation across morphological features may reflect the complex interaction of functional constraints, developmental timing, and/or variable biomechanical forces. Pongo's elevated variation in spinous process length suggests a release from functional constraint, consistent with its suspensory locomotion and reduced spinous processes. Interspecific differences in vertebral variation based on locomotor demand or posture are generally consistent with patterns previously reported for vertebral formula and other aspects of morphology. Future research would benefit from an expanded taxonomic sample and more detailed analyses of vertebral modularity and developmental timing., (© 2018 Wiley Periodicals, Inc.)

A new species of the "small-bodied ape" Simiolus is described here that extends the temporal range of the genus to the end of the Middle Miocene. As such, it is one of the few species of fossil primates known from East Africa during a time of significant change in which Old World monkeys and crown hominoids replaced the primitive ape-like primates that had dominated the early Miocene. The dynamics of this important event in our evolutionary history are obscured by the small number of fossil primates known from Africa between 14 and 6 million years ago, as well as persistent ambiguity regarding the phylogenetic status of the ape-like Miocene primates. The new species described here helps to fill this temporal gap, and our analysis of its phylogenetic position suggests that Simiolus and many other Miocene primates were not only ape-like, they were, indeed, stem hominoids. Judging from the available material, the new species may be the smallest known ape., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

Objectives: High-resolution imaging of fossils with X-ray computed microtomography (μCT) has become a very powerful tool in paleontological research. However, fossilized bone, embedding matrix, and dental tissues do not always provide a distinct structural signal with X-rays. We demonstrate the benefits of high-resolution neutron radiation in three different specimens showing problematic contrasts with X-ray μCT., Materials and Methods: We compare neutron with X-ray μCT scans of fossils from two Miocene catarrhines from the Vallès-Penedès Basin: the cranium (IPS58443.1, holotype) of the putative stem hominoid Pliobates cataloniae, to discriminate between bone and matrix; and two lower molars (IPS1724n,o, holotype) of Barberapithecus huerzeleri, to discriminate among dental tissues., Results: X-ray μCT scans of these specimens fail to retrieve any contrast between matrix/bone and enamel/dentine, whereas neutron μCT scans deliver high-contrast images, enabling a proper evaluation of the specimens' internal anatomy., Discussion: Low bone/matrix intensity difference with X-ray μCT scans in IPS58443.1 is due to the extreme similarity in chemical composition between the matrix and the fossilized tissues, and the presence of high-density elements. In IPS1724, it is attributable to the convergence of enamel and dentine compositions during fossilization. On the contrary, neutron radiation returns very different contrasts for different isotopes of the same element and easily penetrates most metals. Neutron-based μCT scans therefore enable a correct definition of the bone/sediment and enamel/dentine interfaces, and hence a better segmentation of the images stack. We conclude that neutron radiation represents a successful alternative for high-resolution µCT of small-sized fossils that are problematic with X-rays., (© 2018 Wiley Periodicals, Inc.)

Despite its long history of scientific study, the causes underlying the extinction of the insular hominoid Oreopithecus bambolii are still a matter of ongoing debate. While some authors consider intense tectonism and invading species the cause of its extinction ca. 6.7 Ma, others propose climatic change as the main contributing factor. We rely on long-term patterns of tooth wear and hypsodonty of the Baccinello and Fiume Santo herbivore-faunas to reconstruct changes in habitat prior to, during and after the extinction. While a mosaic of habitats was represented in Baccinello V1 (as shown by a record of browsers, mixed feeders and species engaged in grazing), more closed forests (higher proportion of browsers, shortage of mixed feeders and lack of grazers) characterised Baccinello V2. Finally, there was a partial loss of canopy cover and development of open-patches and low-abrasive grasses in Baccinello V3 (as denoted by new records of taxa involved in grazing)-although still dominated by a forested habitat (since browse was a component in all diets). Our results provide evidence for two perceptible shifts in climate, one between 8.1 and 7.1 Ma and other ca. 6.7 Ma, though this latter was not drastic enough to lead to intensive forest loss, substantially alter the local vegetation or affect Oreopithecus lifestyle-especially if considering the growing evidence of its versatile diet. Although the disappearance of Oreopithecus is complex, our data reject the hypothesis of environmental change as the main factor in the extinction of Oreopithecus and Maremma fauna. When our results are analysed together with other evidence, faunal interaction and predation by invading species from mainland Europe seems to be the most parsimonious explanation for this extinction event. This contrasts with European hominoid extinctions that were associated with major climatic shifts that led to environmental uniformity and restriction of the preferred habitats of Miocene apes., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

Body mass is an important component of any paleobiological reconstruction. Reliable skeletal dimensions for making estimates are desirable but extant primate reference samples with known body masses are rare. We estimated body mass in a sample of extinct platyrrhines and Fayum anthropoids based on four measurements of the articular surfaces of the humerus and femur. Estimates were based on a large extant reference sample of wild-collected individuals with associated body masses, including previously published and new data from extant platyrrhines, cercopithecoids, and hominoids. In general, scaling of joint dimensions is positively allometric relative to expectations of geometric isometry, but negatively allometric relative to expectations of maintaining equivalent joint surface areas. Body mass prediction equations based on articular breadths are reasonably precise, with %SEEs of 17-25%. The breadth of the distal femoral articulation yields the most reliable estimates of body mass because it scales similarly in all major anthropoid taxa. Other joints scale differently in different taxa; therefore, locomotor style and phylogenetic affinity must be considered when calculating body mass estimates from the proximal femur, proximal humerus, and distal humerus. The body mass prediction equations were applied to 36 Old World and New World fossil anthropoid specimens representing 11 taxa, plus two Haitian specimens of uncertain taxonomic affinity. Among the extinct platyrrhines studied, only Cebupithecia is similar to large, extant platyrrhines in having large humeral (especially distal) joints. Our body mass estimates differ from each other and from published estimates based on teeth in ways that reflect known differences in relative sizes of the joints and teeth. We prefer body mass estimators that are biomechanically linked to weight-bearing, and especially those that are relatively insensitive to differences in locomotor style and phylogenetic history. Whenever possible, extant reference samples should be chosen to match target fossils in joint proportionality., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

Afropithecus turkanensis (17-17.5 Ma; Kalodirr, Buluk, Locherangan, Moruorot, Nabwal Hills; Kenya) and Morotopithecus bishopi (20.6 Ma; Moroto II; Uganda) are both large-bodied catarrhines from the early Miocene of eastern Africa with relatively primitive cranial and postcanine dental morphology. They are primarily differentiated by a temporal separation of ∼3.6 million years and by postcranial samples suggesting that M. bishopi was capable of orthograde postures and below-branch arboreality, while A. turkanensis was most likely a pronograde quadruped. Several researchers dispute the validity of the postcranial and dating evidence and argue that M. bishopi and A. turkanensis may be congeneric or even conspecific. Although A. turkanensis possesses a derived suite of specialized anterior dentognathic characters that are functionally convergent with extant pitheciins and associated with sclerocarp foraging and maxillary canine dietary function, a similar analysis of M. bishopi anterior dentognathic anatomy is presently lacking. The current study addresses this shortcoming via a detailed morphometric analysis of relevant A. turkanensis and M. bishopi specimens preserving the anterior palate, maxillary canines and incisors. Results indicate that the anterior dentognathic morphologies of A. turkanensis and M. bishopi are distinct and represent significantly dissimilar feeding adaptations. Specifically, M. bishopi lacks the elongated and anteriorly narrow premaxilla, lateral incisors that are more posterior and mesially positioned relative to the central incisors, and pronounced yet evenly distributed mesial curvature of the maxillary canine that are shared by A. turkanensis and extant pitheciins. Given that A. turkanensis anterior dentognathic morphology is functionally convergent with extant pitheciins to the exclusion of M. bishopi, it is likely that M. bishopi and A. turkanensis have dissimilar feeding adaptations. Although a systematic analysis is required to verify these species at the generic and species level, the absence of any substantial morphological similarity in their anterior dentognathic anatomy is most consistent with the interpretation that M. bishopi and A. turkanensis represent, at the least, different species., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

Songhor is an early Miocene fossil locality in Kenya known for its diverse primate assemblage that includes catarrhine species belonging to the genera Kalepithecus, Limnopithecus, Dendropithecus, Rangwapithecus, and Proconsul. Expeditions to Songhor since the 1930s have recovered unassociated catarrhine postcranial remains from both the fore- and hindlimbs, including multiple elements from the feet. In this study, we describe KNM-SO 31233, a complete left hallucal metatarsal (Mt1), along with several other fragmentary Mt1 specimens (KNM-SO 1080, 5129, 5141, 22235). These fossils were compared to extant catarrhines and platyrrhines, as well as available fossil Miocene catarrhine Mt1s. Morphometric data were obtained from 3D surface renderings and subjected to a number of analyses to assess their phenetic affinity with the comparative sample, make predictions of body mass, and to infer their functional morphology. The size and shape of the Songhor Mt1s are diverse, exhibiting a large robust morph (KNM-SO 5141) similar in size but not in shape to extant African apes, medium-sized morphs (KNM-SO 1080, 5129 and 22235), and a smaller, slender one (KNM-SO 31233) that has a shape resembling arboreal quadrupedal leaping monkeys and suspensory atelines and hylobatids. KNM-SO 31233 is unlike other known fossil Mt1s, and in general, none of the Songhor Mt1s resembled any single extant anthropoid clade or species. The morpho-functional diversity of Songhor Mt1s is consistent with an extensive morphological and phylogenetic catarrhine diversity in the early part of the Miocene epoch., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

The Late Miocene sequence at Haritalyangar, Himachal Pradesh, India, has produced abundant remains of the hominid Sivapithecus and the sivaladapids Sivaladapis and Indraloris. Also recovered from these sediments is an isolated and worn upper molar that was made the holotype of Krishnapithecus krishnaii and assigned to the Pliopithecoidea. However, the heavy wear and absence of definitive pliopithecoid features on the tooth rendered the assignment to this superfamily unconvincing. Here, we describe two lower molars from Haritalyangar that bear unmistakable pliopithecoid features and that are plausibly assignable to the same species as the type specimen of K. krishnaii. They convincingly demonstrate for the first time the presence of the Pliopithecoidea in South Asia. The new molars also reveal that K. krishnaii was perhaps the largest known pliopithecoid and that it possessed highly derived postcanine dental morphology. Because of its highly derived nature, it is difficult to determine its relationships within Pliopithecoidea, but a sister taxon relationship with either the Dionysopithecidae or Pliopithecinae is equally plausible; it is only distantly related to the Crouzeliinae. It is sufficiently distinct, however, from all other pliopithecoids to warrant placement in a separate family., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

The early stages of phylogenetic inference from morphological data involve a sequence of choices about which analytical methods to employ. At each stage, the selection of one method over another can dramatically impact tree inference. Phylogenetic hypotheses are sensitive to decisions relating to which taxa and characters to select for analysis, whether and how to delimit character states, which taxa to use as outgroups, and how to account for character dependence. Using extant hominoids as a test case, I quantify the degree to which phylogenetic inferences are sensitive to the choice of method used to transform continuously scaled variables into categorical traits. I demonstrate that the character coding strategy significantly impacts hypotheses of character state identity and phylogenetic branching patterns. To avoid biasing evolutionary hypotheses, I recommend that continuously scaled characters be analyzed without prior discretization., (© 2017 S. Karger AG, Basel.)

The pterion, on the lateral aspect of the cranium, is where the zygomatic, frontal, sphenoid, squamosal, and parietal bones approach and contact. The configuration of these bones distinguishes New and Old World anthropoids: most extant platyrrhines exhibit contact between the parietal and zygomatic bones, while all known catarrhines exhibit frontal-alisphenoid contact. However, it is thought that early stem-platyrrhines retained the apparently primitive catarrhine condition. Here we re-evaluate the condition of key fossil taxa using μCT (micro-computed tomography) imaging. The single known specimen of Tremacebus and an adult cranium of Antillothrix exhibit the typical platyrrhine condition of parietal-zygomatic contact. The same is true of one specimen of Homunculus, while a second specimen has the 'catarrhine' condition. When these new data are incorporated into an ancestral state reconstruction, they support the conclusion that pterion frontal-alisphenoid contact characterized the last common ancestor of crown anthropoids and that contact between the parietal and zygomatic is a synapomorphy of Platyrrhini., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

Limnopithecus is a small-bodied catarrhine genus that is widespread throughout early Miocene sites in East Africa. Although two species of this genus have been described - Limnopithecus legetet (type species) and Limnopithecus evansi - they are poorly known anatomically and their systematic positions remain unresolved. Here, we provide detailed descriptions and comparisons for two well-preserved maxillary specimens that we attribute to L. evansi. These specimens come from the type locality of the species, Songhor in western Kenya, and add greatly to our knowledge of its dentognathic morphology. Together, they preserve the entire unilateral upper dentition, with overlapping elements demonstrating conspecificity, and provide new information about I(2) morphology and aspects of the palate, nasal aperture, and maxillary sinuses. Detailed morphological comparisons suggest that specimens referred to Limnopithecus from Songhor, Koru, and Rusinga share a unique I(2) morphology not found in any other early Miocene catarrhine. This argues in favor of congeneric status for L. evansi and L. legetet. Moreover, features such as a broad palate, premolar morphology, and the relative proportions of the premolars of L. evansi distinguish it from Lomorupithecus harrisoni, another early Miocene catarrhine from Napak, Uganda. This finding challenges a recently proposed taxonomic interpretation that Lomorupithecus and L. evansi are conspecific. Our results underscore the distinctiveness of L. evansi and Lo. harrisoni, thereby reaffirming the validity of the taxon Lo. harrisoni and indicating that the Songhor and Napak catarrhine communities were relatively distinct, despite their apparent contemporaneity., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

Morphology has been widely used for inferring the phylogenies of numerous taxonomic groups. Recent molecular studies performed on extant non-human primates, however, have cast doubt on the reliability of cranial and postcranial characters for characterizing evolutionary affinities. Because molecular evidence is often not available for fossil specimens, detecting phylogenetic signals in anatomical features is of great relevance. Here we have analyzed molar (M1 and M2) crown shape by means of geometric morphometrics in a large sample of both extant and fossil Miocene catarrhine primates to detect the phylogenetic signal in molar morphometry. Results support that molar shape carries a strong phylogenetic signal, mostly at the superfamily level but also to some extent at the family level. Dietary factors, however, appear to have less influence, especially for M2. The Miocene Pliopithecoidea, Cercopithecoidea, and Hominoidea superfamilies clearly grouped according to the expected taxonomic affinities with the extant groups, although some discrepancies were found depending on the tooth considered. Our findings suggest that although molar crown shape can be used as a reliable proxy for establishing taxonomic affinities of catarrhine fossil primates with extant groups, a significant amount of interspecific variation exists, indicative of derived adaptations at the genus or species level., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

Pliopithecoids represent a monophyletic group of putative stem catarrhines whose evolutionary history is incompletely known. They have been recorded from Europe and Asia, between the late Early Miocene and the Late Miocene. Asian pliopithecoids are less well documented than their European counterparts, often being represented by a fragmentary fossil record. New discoveries are therefore critical to reconstruct the evolutionary history of the whole group. Here, we describe two isolated molars from Ban San Klang, a late Middle Miocene locality in northern Thailand, which confirms the presence of pliopithecoids in Southeast Asia. The lower molar had originally been described as being that of a dendropithecoid, but it was later recognized as pertaining to a pliopithecoid. The discovery, in the same locality, of an additional upper molar attributed to the same species confirms the pliopithecoid status of this taxon and highlights its distinctiveness with respect to other known Asian pliopithecoids. However, the mosaic of primitive and autapomorphic features characterizing this Thai fossil, as well as its limited anatomical representation, preclude us from assigning it to either of the known pliopithecid subfamilies. Nevertheless, it represents the only pliopithecoid in Southeast Asia and displays a mosaic of unique characters which emphasizes the peculiarity of that province, as suggested previously with respect to its hominoid primate., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

For more than 80 years, Proconsul has held a pivotal position in interpretations of catarrhine evolution and hominoid diversification in East Africa. The majority of what we 'know' about Proconsul, however, derives from abundant younger fossils found at the Kisingiri localities on Rusinga and Mfangano Islands rather than from the smaller samples found at Koru--the locality of the type species, Proconsul africanus--and other Tinderet deposits. One outcome of this is seen in recent attempts to expand the genus "Ugandapithecus" (considered here a junior subjective synonym of Proconsul), wherein much of the Tinderet sample was referred to that genus based primarily on differentiating it from the Kisingiri specimens rather than from the type species, P. africanus. This and other recent taxonomic revisions to Proconsul prompted us to undertake a systematic review of dentognathic specimens attributed to this taxon. Results of our study underscore and extend the substantive distinction of Tinderet and Ugandan Proconsul (i.e., Proconsul sensu stricto) from the Kisingiri fossils, the latter recognized here as a new genus. Specimens of the new genus are readily distinguished from Proconsul sensu stricto by morphology preserved in the P. africanus holotype, but also in I(1)s, lower incisors, upper and lower canines, and especially mandibular characteristics. A number of these differences are more advanced among Kisingiri specimens in the direction of crown hominoids. Proconsul sensu stricto is characterized by a suite of unique features that strongly unite the included species as a clade. There have been decades of contentious debate over the phylogenetic placement of Proconsul (sensu lato), due in part to there being a mixture of primitive and more advanced morphology within the single genus. By recognizing two distinct clades that, in large part, segregate these character states, we believe that better phylogenetic resolution can be achieved., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

Changes in lifestyles and body weight affected mammal life-history evolution but little is known about how they shaped species' sensory systems. Since auditory sensitivity impacts communication tasks and environmental acoustic awareness, it may have represented a deciding factor during mammal evolution, including apes. Here, we statistically measure the influence of phylogeny and allometry on the variation of five cochlear morphological features associated with hearing capacities across 22 living and 5 fossil catarrhine species. We find high phylogenetic signals for absolute and relative cochlear length only. Comparisons between fossil cochleae and reconstructed ape ancestral morphotypes show that Australopithecus absolute and relative cochlear lengths are explicable by phylogeny and concordant with the hypothetized ((Pan,Homo),Gorilla) and (Pan,Homo) most recent common ancestors. Conversely, deviations of the Paranthropus oval window area from these most recent common ancestors are not explicable by phylogeny and body weight alone, but suggest instead rapid evolutionary changes (directional selection) of its hearing organ. Premodern (Homo erectus) and modern human cochleae set apart from living non-human catarrhines and australopiths. They show cochlear relative lengths and oval window areas larger than expected for their body mass, two features corresponding to increased low-frequency sensitivity more recent than 2 million years ago. The uniqueness of the "hypertrophied" cochlea in the genus Homo (as opposed to the australopiths) and the significantly high phylogenetic signal of this organ among apes indicate its usefulness to identify homologies and monophyletic groups in the hominid fossil record.

Suspension plays a major adaptive role in shaping primate postcranial morphology, which therefore enables this positional behavior to be inferred in extinct taxa. The proximal humerus stands as a key region for inferring forelimb suspensory capabilities because its morphology can be effectively linked, from a functional viewpoint, to differences in suspension use between primate taxa. Here we provide an assessment of the suspensory capabilities of two pliopithecoids (Epipliopithecus vindobonensis and Pliopithecus antiquus) by means of a 3D geometric morphometric analysis of proximal humeral shape. The comparative sample includes proximal humeri from eight extant anthropoid genera, as well as other extinct catarrhines (the propliopithecoid Aegyptopithecus zeuxis, the stem hominoid Nyanzapithecus vancouveringorum, and an unascribed small catarrhine, GSP 28062, from the Middle Miocene of Pakistan). Body mass estimates based on allometric regressions of humeral head superoinferior diameter are also provided. Our results support some degree of forelimb suspensory behaviors for Epipliopithecus and GSP 28062. In contrast, and unlike previous qualitative assessments, our analysis shows that P. antiquus has a distinct glenohumeral morphology, much closer to that displayed by generalized arboreal quadrupeds with no evidence of suspensory adaptations (as in Aegyptopithecus and stem hominoids from Africa)., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

Uncovering links between skeletal morphology and locomotor behavior is an essential component of paleobiology because it allows researchers to infer the locomotor repertoire of extinct species based on preserved fossils. In this study, we explored ulnar shape in anthropoid primates using 3D geometric morphometrics to discover novel aspects of shape variation that correspond to observed differences in the relative amount of forelimb suspensory locomotion performed by species. The ultimate goal of this research was to construct an accurate predictive model that can be applied to infer the significance of these behaviors. We studied ulnar shape variation in extant species using principal component analysis. Species mainly clustered into phylogenetic groups along the first two principal components. Upon closer examination, the results showed that the position of species within each major clade corresponded closely with the proportion of forelimb suspensory locomotion that they have been observed to perform in nature. We used principal component regression to construct a predictive model for the proportion of these behaviors that would be expected to occur in the locomotor repertoire of anthropoid primates. We then applied this regression analysis to Pliopithecus vindobonensis, a stem catarrhine from the Miocene of central Europe, and found strong evidence that this species was adapted to perform a proportion of forelimb suspensory locomotion similar to that observed in the extant woolly monkey, Lagothrix lagothricha., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

Inferring the evolutionary history of the hominins is necessarily reliant on comparative analyses of fossilized skeletal anatomy. However, the reliability of different primate skeletal regions for recovering phylogenetic relationships is currently poorly understood. Historically, postcranial variation has largely been conceived of as reflecting locomotory and postural adaptation. The shape of the os coxae is central to such discussions given the divergent morphology displayed by the bipedal hominin pelvis relative to other primate taxa. While previous cladistic studies have suggested that postcranial and cranial datasets do not differ in terms of their propensity for homoplasy, methodological issues such as the numbers of characters and their quantification make it difficult to evaluate these findings. Here, we circumvent these problems by constructing morphological distance matrices based on cranial, mandibular and os coxae three-dimensional shape. Statistical comparisons of these morphological distance matrices against a single genetic distance matrix for 11 catarrhine taxa show that cranial and os coxae shape reflect genetic relationships better than the mandible when humans are included, and that the cranium and os coxae do not differ statistically in terms of their genetic correlations. When humans were excluded from the analyses, all three anatomical regions were equally strongly correlated with genetic distance. Moreover, a second analysis focusing solely on os coxae variation of 16 taxa demonstrated that os coxae shape correctly recovers catarrhine taxonomic relationships at the sub-family level, even when humans are included. Taken together, our results suggest that there is no a priori reason to favor cranial shape data over os coxae morphology when inferring the genetic relationships of extant or extinct primate taxa. Morphological similarities between humans and other primates differ depending on the skeletal element, suggesting that combining skeletal elements into a single analysis may provide more accurate reconstructions of genetic relationships., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

The upper ankle joint forms a single articular plane between organism and the foot and substrate. Singular warp analysis shows that its shape reflects substrate use. This study explores whether the differences in shape are genetic with a developmental trajectory evident during ontogeny or epigenetic and the result of substrate use by the individual. A total of 418 matched distal tibial and proximal talar landmarked surfaces from adult and subadult specimens from 12 diverse catarrhine taxa were studied. Specimens were grouped by molar eruption (M1, M2, and M3) for comparative analysis. Generalized Procrustes analysis, multivariate regression, relative warp analysis, singular warp analysis, and permutation tests were used. Singular warp analysis for the entire cohort was highly significant in the first singular warp, with the adult taxa sorting by substrate use. All 173 subadults clustered with an adult "arboreal" shape profile. Among Hominoidea, adults (M3) sorted by substrate use with Pan paniscus and Hylobatidae assuming an "arboreal" shape separate from Pan troglodytes and the remaining taxa with "terrestrial" shape. Cercopithecoid adults sorted by substrate use as well, with the M3 specimens of Papio hamadryas and Macaca thibetana demonstrating a "terrestrial" shape. Differences in mode of locomotion did not affect the findings in the first singular warp. Results confirmed the convergence of talo-crural shape among superfamilies based on substrate use and divergence in shape within Pan and Macaca, based on substrate use. The shape differences among adults (M3) are consistent with a plastic response to the behavioral stimulus of substrate use., (© 2014 Wiley Periodicals, Inc.)

Two catarrhine mandibles and five isolated teeth have been discovered from Early Miocene localities in Western Kenya. One mandible comes from the well-known locality of Songhor whereas the other is from a newly discovered locality, Lower Kapurtay, located near Songhor. The mandibles both can clearly be assigned to the species Rangwapithecus gordoni based on molar morphology, which is unique among Early Miocene catarrhines. The isolated specimens can be assigned to Rangwapithecus based on their similarities in morphology to the homologues preserved in the two mandibles. These specimens provide important new information about the dentognathic morphology of Rangwapithecus, which is described in detail. The mandible from Songhor (KNM-SO 22228) represents the first definitive female mandible of Rangwapithecus. The Lower Kapurtay mandible (KNM-KT 31234) appears to be male but is much smaller than another recently described male mandible of this species (KNM-SO 17500) and the type maxilla (KNM-SO 700). These specimens enable a reassessment of the attributions of all other mandibles and isolated lower teeth of Rangwapithecus, and we present a complete hypodigm of the mandibular and lower dental material for the species. Finally, we provide some additions to the diagnosis of Rangwapithecus gordoni based on previously unknown morphology., (Copyright © 2013 Wiley Periodicals, Inc.)

Rudabánya is rare among Eurasian Miocene fossil primate localities in preserving both a hominid and pliopithecoid, and as such provides the unique opportunity to reconstruct the nature of sympatry and niche partitioning in these taxa. Rudapithecus and Anapithecus have similar locomotor and positional behavior and overlapping body mass ranges. While prior analyses of molar occlusal anatomy and microwear identify Rudapithecus as a soft-object frugivore, reconstructing the dietary behavior of Anapithecus has been more problematic. This taxon has been interpreted to be more folivorous by some, and more frugivorous by others. Here, we use high-resolution polynomial curve fitting (HR-PCF) to quantify and evaluate the mesiodistal and cervico-incisal curvatures of the incisor crowns of Rudapithecus and Anapithecus to identify diet-specific morphological variation in these taxa. Results are consistent with the interpretation that Anapithecus and Rudapithecus were primarily frugivorous and had diets that included similar resource types. However, Anapithecus may have consumed greater amounts of foliage, similar to extant mixed folivore-frugivores (i.e., Gorilla gorilla gorilla, Symphalangus syndactylus), while Rudapithecus generated elevated compressive loads in the incisor region consistent with a specialized role for the anterior dentition in food processing (i.e., removal of tough protective fruit pericarps). We interpret these findings in light of the paleoecology at Rudabánya and conclude that, if these taxa were indeed sympatric, Anapithecus may have used additional leaf consumption as a seasonal fallback resource to avoid direct competition with Rudapithecus. Conversely, Rudapithecus may have relied on less preferred and harder fruiting resources as a seasonal fallback resource during periods of fruit scarcity., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

A well-preserved calcaneus referrable to Proteopithecus sylviae from the late Eocene Quarry L-41 in the Fayum Depression, Egypt, provides new evidence relevant to this taxon's uncertain phylogenetic position. We assess morphological affinities of the new specimen using three-dimensional geometric morphometric analyses with a comparative sample of primate calcanei representing major extinct and extant radiations (n = 58 genera, 106 specimens). Our analyses reveal that the calcaneal morphology of Proteopithecus is most similar to that of the younger Fayum parapithecid Apidium. Principal components analysis places Apidium and Proteopithecus in an intermediate position between primitive euprimates and crown anthropoids, based primarily on landmark configurations corresponding to moderate distal elongation, a more distal position of the peroneal tubercle, and a relatively "unflexed" calcaneal body. Proteopithecus and Apidium are similar to cercopithecoids and some omomyiforms in having an ectal facet that is more tightly curved, along with a larger degree of proximal calcaneal elongation, whereas other Fayum anthropoids, platyrrhines and adapiforms have a more open facet with less proximal elongation. The similarity to cercopithecoids is most plausibly interpreted as convergence given the less tightly curved ectal facets of stem catarrhines. The primary similarities between Proteopithecus and platyrrhines are mainly in the moderate distal elongation and the more distal position of the peroneal tubercle, both of which are not unique to these groups. Proteopithecus and Apidium exhibit derived anthropoid features, but also a suite of primitive retentions. The calcaneal morphology of Proteopithecus is consistent with our cladistic analysis, which places proteopithecids as a sister group of Parapithecoidea., (Copyright © 2013 Wiley Periodicals, Inc.)

The distal component of the talo-crural joint, the talus, was compared, using geometric morphometrics, in 219 specimens from nine extant taxa to identify differences in shape and the factors influencing them. The specimens were laser scanned, digitally reconstructed, and landmarked. The whole talus, proximal and distal articular facet subgroups were analyzed using Generalized Procrustes analysis, linear regression, principal component analysis, analysis of percent variance, dot-product vector analysis, and pair-wise permutation tests to evaluate shape, and were visualized by TPS deformation of an exemplar surface. Significant percentages of shape variation among taxa were due to body mass, talar size, superfamily, and substrate preference. Shape and presentational morphology associated with these factors were documented, along with the similarities and differences among individual taxa. Nearly all taxa were significantly different in overall, proximal and distal shapes. The most important factors influencing whole talar shape were log centroid size and substrate preference. Substrate preference was also the most important factor defining proximal articular morphology and unrelated to other factor such as mass, while distal articular morphology was influenced by superfamily (head angle and shape). Results demonstrated that substrate preference and superfamily significantly influenced distal presentation, while substrate preference influenced proximal articular shape., (Copyright © 2013 Wiley Periodicals, Inc.)

Primate fossil assemblages often have metacarpals and phalanges from which functional/behavioral interpretations may be inferred. For example, intrinsic hand proportions can indicate hand function and substrate use. But, estimates of intrinsic hand proportions from unassociated hand elements can be imperfect due to digit misattribution. Although isolated metacarpals can be identified to a specific digit, phalanges are difficult to assign to a specific ray. We used a resampling approach to evaluate how estimates of intrinsic hand proportions are affected by such uncertainty. First, the phalangeal index-intermediate phalanx length plus proximal phalanx length divided by metacarpal length-for the third digit was calculated for associated specimens of terrestrial, semiterrestrial, and arboreal taxa. We then used resampling procedures to generate distributions of "composite digits" based on resampled ratios in which phalanges from the second, fourth, and fifth rays, and from different individuals, were chosen randomly. Results confirm that the phalangeal index for associated third digits significantly discriminates groups. We also found that resampled ratios had significantly lower means, indicating that using composite digits is prone to systematic underestimation. Resampled ratios also generated distributions with greater variance around the means that obscured distinctions between groups, although significant differences between the most arboreal and terrestrial taxa are maintained. We conclude that using unassociated phalanges to calculate a phalangeal index is prone to sampling bias. Nevertheless, a resampling approach has the potential to inform estimates of hand proportions for fossil taxa, provided that the comparative sample is constrained to mimic the fossil composition., (Copyright © 2013 Wiley Periodicals, Inc.)

This study tests the hypothesis that decreased canine crown height in catarrhines is linked to (and arguably caused by) decreased jaw gape. Associations are characterized within and between variables such as upper and lower canine height beyond the occlusal plane (canine overlap), maximum jaw gape, and jaw length for 27 adult catarrhine species, including 539 living subjects and 316 museum specimens. The data demonstrate that most adult male catarrhines have relatively larger canine overlap dimensions and gapes than do conspecific females. For example, whereas male baboons open their jaws maximally more than 110% of jaw length, females open about 90%. Humans and hylobatids are the exceptions in that canine overlap is nearly the same in both the sexes and so is relative gape (ca. 65% for humans and 110% for hylobatids). A correlation analysis demonstrates that a large portion of relative gape (maximum gape/projected jaw length) is predicted by relative canine overlap (canine overlap/jaw length). Relative gape is mainly a function of jaw muscle position and/or jaw muscle-fiber length. All things equal, more rostrally positioned jaw muscles and/or shorter muscle fibers decrease gape and increase bite force during the power stroke of mastication, and the net benefit is to increase the mechanical efficiency during chewing. Similarly, more caudally positioned muscles and/or longer muscle fibers increase the amount of gape and decrease bite force. Overall, the data support the hypothesis that canine reduction in early hominins is functionally linked to decreased gape and increased mechanical efficiency of the jaws., (Copyright © 2012 Wiley Periodicals, Inc.)

The specialized grasping feet of primates, and in particular the nature of the hallucal grasping capabilities of living strepsirrhines and tarsiers (i.e., 'prosimians'), have played central roles in the study of primate origins. Prior comparative studies of first metatarsal (Mt1) morphology have documented specialized characters in living prosimians that are indicative of a more abducted hallux, which in turn is often inferred to be related to an increased ability for powerful grasping. These include a well-developed peroneal process and a greater angle of the proximal articular surface relative to the long axis of the diaphysis. Although known Mt1s of fossil prosimians share these characters with living non-anthropoid primates, Mt1 morphology in the earliest crown group anthropoids is not well known. Here we describe two Mt1s from the Fayum Depression of Egypt - one from the latest Eocene (from the ∼34 Ma Quarry L-41), and one from the later early Oligocene (from the ∼29-30 Ma Quarry M) - and compare them with a sample of extant and fossil primate Mt1s. Multivariate analyses of Mt1 shape variables indicate that the Fayum specimens are most similar to those of crown group anthropoids, and likely belong to the stem catarrhines Catopithecus and Aegyptopithecus specifically, based on analyses of size. Also, phylogenetic analyses with 16 newly defined Mt1 characters support the hypotheses that "prosimian"-like Mt1 features evolved along the primate stem lineage, while crown anthropoid Mt1 morphology and function is derived among primates, and likely differed from that of basal stem anthropoids. The derived loss of powerful hallucal grasping as reflected in the Mt1 morphology of crown anthropoids may reflect long-term selection for improved navigation of large-diameter, more horizontal branches at the expense of movement in smaller, more variably inclined branches in the arboreal environment., (Copyright © 2012 Elsevier Ltd. All rights reserved.)

A fundamental adaptation to orthograde posture and locomotion amongst living hominoid primates is a numerically reduced lumbar column, which acts to stiffen the lower back and reduce injuries to the intervertebral discs. A related and functionally complementary strategy of spinal stability is a caudal position of the diaphragmatic vertebra relative to the primitive condition found in nonhominoid primates and most other mammals. The diaphragmatic vertebra marks the transition in vertebral articular facet (zygapophysis) orientation, which either resists (prediaphragmatic) or allows (postdiaphragmatic) trunk movement in the sagittal plane (i.e., flexion and extension). Unlike most mammals, which have dorsomobile spines (long lumbar columns and cranially placed diaphragmatic vertebrae) for running and leaping, hominoids possess dorsostable spines (short lumbar columns and caudally placed diaphragmatic vertebrae) adapted to orthogrady and antipronogrady. In contrast to humans and other extant hominoids, all known early hominin partial vertebral columns demonstrate cranial displacement of the diaphragmatic vertebra. To address this difference, variation in diaphragmatic placement is assessed in a large sample of catarrhine primates. I show that while hominoids are characterized by modal common placement of diaphragmatic and last rib-bearing vertebrae in general, interspecific differences in intraspecific patterns of variation exist. In particular, humans and chimpanzees show nearly identical patterns of diaphragmatic placement. A scenario of hominin evolution is proposed in which early hominins evolved cranial displacement from the ancestral hominid condition of common placement to achieve effective lumbar lordosis during the evolution of bipedal locomotion., (Copyright © 2012 Wiley Periodicals, Inc.)

Here, we report on an isolated pliopithecid M3/ (IPS35028) from locality ACM/C3-B2 (12.0 Ma, MN7) of the late Middle Miocene stratigraphic series of Abocador de Can Mata (ACM, Vallès-Penedès Basin, NE Iberian Peninsula). This tooth is about 0.2 million years older than the remains of Pliopithecus canmatensis (11.8-11.7 ma), recorded from several localities from the ACM series. The unusual occlusal features of IPS35028, together with the lack of homologous material for several pliopithecid species, preclude a precise taxonomic attribution of the C3-B2 specimen, which does not fit the morphology of any known pliopithecid M3/. In particular, although an attribution to P. canmatensis would seem reasonable on the basis of size, identical geographic provenance, and similar age, the morphology of IPS35028 appears too primitive compared to the M1/ and M2/ of the former taxon. Instead, the C3-B2 pliopithecid displays several primitive features shared with the dionysopithecine Dionysopithecus and the pliopithecine Pliopithecus piveteaui. It therefore seems more likely that IPS35028 represents a previously unknown pliopithecid taxon, although a formal taxonomic recognition of its probable distinct status is not advisable, given the scarcity of the currently available material. Alternatively, this taxon might be more closely related to small-bodied African catarrhines (such as dendropithecids). However, the morphology of the ACM specimen is not particularly similar to that of the M3/ of these African taxa. Hence, based on age and geographic provenance, an attribution of IPS35028 to the Pliopithecidae is favored here., (Copyright © 2011 Wiley Periodicals, Inc.)

Ecological factors have been shown to be important for brain size evolution. In this comparative study among catarrhine primates, we examine two different ways in which seasonality may be related to brain size. First, seasonality may impose energetic constraints on the brain because it forces animals to deal with periods of food scarcity (Expensive Brain hypothesis). Second, seasonality may act as a selective pressure to increase brain size, as behavioral flexibility helps to overcome periods of food scarcity (Cognitive Buffer hypothesis). Controlling for phylogeny, we found a strong negative relationship between brain size (relative to body mass) and the degree of experienced seasonality, as estimated by the variation in net energy intake. However, we also found a significant positive relationship between relative brain size and the effect of so-called cognitive buffering, proxied by the difference between environmental seasonality and the seasonality in net energy intake actually experienced by the animals. These results show that both energetic constraints of seasonal habitats as well as cognitive buffering affect brain size evolution, leaving environmental seasonality uncorrelated to brain size. With this study we show the importance of simultaneously considering both costs and benefits in models of brain size evolution., (© 2011 The Author(s). Evolution © 2011 The Society for the Study of Evolution.)

Phalanges are considered to be highly informative in the reconstruction of extinct primate locomotor behavior since these skeletal elements directly interact with the substrate during locomotion. Variation in shaft curvature and relative phalangeal length has been linked to differences in the degree of suspension and overall arboreal locomotor activities. Building on previous work, this study investigated these two skeletal characters in a comparative context to analyze function, while taking evolutionary relationships into account. This study examined the correspondence between proportions of suspension and overall substrate usage observed in 17 extant taxa and included angle of curvature and relative phalangeal length. Predictive models based on these traits are reported. Published proportions of different locomotor behaviors were regressed against each phalangeal measurement and a size proxy. The relationship between each behavior and skeletal trait was investigated using ordinary least-squares, phylogenetic generalized least-squares (pGLS), and two pGLS transformation methods to determine the model of best-fit. Phalangeal curvature and relative length had significant positive relationships with both suspension and overall arboreal locomotion. Cross-validation analyses demonstrated that relative length and curvature provide accurate predictions of relative suspensory behavior and substrate usage in a range of extant species when used together in predictive models. These regression equations provide a refined method to assess the amount of suspensory and overall arboreal locomotion characterizing species in the catarrhine fossil record., (Copyright © 2011 Wiley-Liss, Inc.)

Recent excavations in northwestern Kenya have recovered a vertebrate fauna of late early or early late Oligocene age. Among the mammal remains, a fragmentary lower jaw and an isolated upper molar have been attributed to a small primate, Lokonepithecus manai gen. et sp. nov. Lokonepithecus is a primitive member of the Parapithecidae and possibly most closely related to Apidium from the Fayum. The new primate from Kenya is the youngest parapithecid known and its occurrence in the Oligocene of Kenya suggests that sub-Saharan Africa probably played a major role in the evolutionary history of several groups of mammals., (Copyright © 2011 Elsevier Ltd. All rights reserved.)

Taillessness is a distinctive synapomorphy of the Hominoidea that has implications for interpretation of the locomotor behaviors and phylogenetic affinities of the clade's earliest members. However, difficulties persist in confidently identifying taillessness in the catarrhine fossil record, stemming largely from our limited knowledge of the anatomical features with which the tail is associated. Here, we compare the morphology of the sacrum, the sole bony link between the tail and the rest of the body, among extant tailless hominoids and a broad sample of extant cercopithecoids known to vary in tail length (i.e., 'very short', 'short', and 'long') in order to identify morphological correlates of tail length. We examine three features of the sacrum, including the shape of the sacrum's caudal articular surface (CAS), the sacrocaudal articulation (SCA) angle, and the lateral expansion of the last sacral vertebra's transverse processes. Compared with all other taxa, 'long'-tailed cercopithecoids have significantly more circularly-shaped CASs, more acute SCA angles, and more laterally expanded transverse processes of the last sacral vertebra. Tailless hominoids have significantly more elliptically-shaped CASs and less laterally expanded transverse processes than all tailed cercopithecoids, but in the latter parameter, they only differ significantly from 'long'-tailed cercopithecoids. Cercopithecoids with 'short' and 'very short' tails are intermediate between tailless hominoids and 'long'-tailed cercopithecoids with respect to CAS shape and lateral expansion of the transverse processes. SCA angle distinguishes clearly among all three cercopithecoid tail length groups. The results of this study provide evidence for significant differences in sacral morphology among extant catarrhines known to differ in tail length, and have implications for making inferences about tail length and function in extinct catarrhines., (Copyright © 2011 Elsevier Ltd. All rights reserved.)

At a given body mass, folivorous colobines have smaller postcanine teeth than frugivorous cercopithecines. This distinction is a notable exception to the general tendency for folivorous primates to have relatively larger postcanine tooth rows than closely related frugivores. The reason for this anomalous pattern is unclear, but one potential explanation is that the difference in facial size between these two subfamilies confounds the comparison-i.e., it may be that the large postcanine teeth of cercopithecines are a consequence of their large faces. The goal of this study was to test this hypothesis. Phylogenetic comparative methods were used to examine the relationships among postcanine area, facial size, and body mass in 29 anthropoid primates, including eight colobines and eight cercopithecines. Results indicate that there is a strong and highly significant partial correlation between postcanine area and facial size when body mass is held constant, which supports the hypothesis that facial size has an important influence on postcanine size. Moreover, colobines have larger postcanine teeth relative to facial size than cercopithecines. Surprisingly, when facial size is held constant, the partial correlation between postcanine area and body mass is weak and nonsignificant. These results suggest that facial size may be more appropriate than body mass for size-adjusting postcanine measurements in some contexts. A phylogenetic comparative test of the association between diet and relative postcanine size (scaled using facial size) confirms that folivorous anthropoids are characterized by relatively large postcanine teeth in comparison to closely related nonfolivores., (2011 Wiley-Liss, Inc.)

Two contrasting patterns of lumbar vertebral morphology generally characterize anthropoids. "Long-backed" monkeys are distinguished from "short-backed" apes [Benton: The baboon in medical research, Vol. 2 (1967:201)] with respect to several vertebral features thought to afford greater spinal flexibility in the former and spinal rigidity in the latter. Yet, discussions of spinal mobility are lacking important functional insight that can be gained by analysis of the zygapophyses, the spine's synovial joints responsible for allowing and resisting intervertebral movements. Here, prezygapophyseal articular facet (PAF) shape in the thoracolumbar spine of Papio, Hylobates, Pongo, Gorilla, and Pan is evaluated in the context of the "long-backed" versus "short-backed" model. A three-dimensional geometric morphometric approach is used to examine how PAF shape changes along the thoracolumbar vertebral column of each taxon and how PAF shape varies across taxa at corresponding vertebral levels. The thoracolumbar transition in PAF shape differs between Papio and the hominoids, between Hylobates and the great apes, and to a lesser extent, among great apes. At the level of the first lumbar vertebra, the PAF shape of Papio is distinguished from that of hominoids. At the level of the second lumbar vertebra, there is variation to some extent among all taxa. These findings suggest that morphological and functional distinctions in primate vertebral anatomy may be more complex than suggested by a "long-backed" versus "short-backed" dichotomy., (Copyright 2010 Wiley-Liss, Inc.)

Comparisons of joint surface curvature at the base of the thumb have long been made to discern differences among living and fossil primates in functional capabilities of the hand. However, the complex shape of this joint makes it difficult to quantify differences among taxa. The purpose of this study is to determine whether significant differences in curvature exist among selected catarrhine genera and to compare these genera with hominin fossils in trapeziometacarpal curvature. Two 3D approaches are used to quantify curvatures of the trapezial and metacarpal joint surfaces: (1) stereophotogrammetry with nonuniform rational B-spline (NURBS) calculation of joint curvature to compare modern humans with captive chimpanzees and (2) laser scanning with a quadric-based calculation of curvature to compare modern humans and wild-caught Pan, Gorilla, Pongo, and Papio. Both approaches show that Homo has significantly lower curvature of the joint surfaces than does Pan. The second approach shows that Gorilla has significantly more curvature than modern humans, while Pongo overlaps with humans and African apes. The surfaces in Papio are more cylindrical and flatter than in Homo. Australopithecus afarensis resembles African apes more than modern humans in curvatures, whereas the Homo habilis trapezial metacarpal surface is flatter than in all genera except Papio. Neandertals fall at one end of the modern human range of variation, with smaller dorsovolar curvature. Modern human topography appears to be derived relative to great apes and Australopithecus and contributes to the distinctive human morphology that facilitates forceful precision and power gripping, fundamental to human manipulative activities.

Interpretations of the postcranial anatomy of East African early and middle Miocene large-bodied hominoids (e.g., Proconsul, Afropithecus, Turkanapithecus, Nacholapithecus) have suggested that these diverse primates utilized positional behaviors dominated by arboreal quadrupedalism. Preliminary descriptions of the Equatorius africanus partial skeleton (KNM-TH 28860) and other forelimb specimens, however, have argued that this animal relied more on terrestrial locomotion compared to its contemporaries, possibly similar to extant large papionin monkeys. In this paper, we reevaluate this interpretation by examining intrinsic hand proportions based on the lengths of the third proximal phalanx and fifth metacarpal in Equatorius in reference to a large sample of extant catarrhine primate taxa. We focused on the lengths of these hand bones because the ratio between phalanx and metacarpal lengths has been previously documented to discriminate terrestrial from arboreal mammalian taxa, including primates. The Equatorius hand displays semi-terrestrial hand proportions with a relatively shorter proximal phalanx compared to most arboreal monkeys. Its proximal phalanx, however, is relatively longer than those of habitually terrestrial monkeys (e.g., Theropithecus, Papio). Accordingly, although Equatorius retains some arboreal quadrupedal characteristics, these results corroborate the previous inference that it engaged in more terrestrial locomotion than earlier Miocene apes such as Proconsul. We suggest that the postcranial skeleton of Equatorius evinces the earliest signs of semi-terrestriality in the hominoid fossil record. It is likely that the terrestrial specialization utilized by living hominoids, e.g., knuckle-walking, evolved separately.