Biochronology of South African hominin-bearing sites: A reassessment using cercopithecid primates.

Despite recent advances in chronometric techniques (e.g., Uranium-Lead [U-Pb], cosmogenic nuclides, electron spin resonance spectroscopy [ESR]), considerable uncertainty remains regarding the age of many Plio-Pleistocene hominin sites, including several in South Africa. Consequently, biochronology remains important in assessments of Plio-Pleistocene geochronology and provides direct age estimates of the fossils themselves. Historically, cercopithecid monkeys have been among the most useful taxa for biochronology of early hominins because they are widely present and abundant in the African Plio-Pleistocene record. The last major studies using cercopithecids were published over 30 y ago. Since then, new hominin sites have been discovered, radiometric age estimates have been refined, and many changes have occurred in cercopithecid taxonomy and systematics. Thus, a biochronological reassessment using cercopithecids is long overdue. Here, we provide just such a revision based on our recent study of every major cercopithecid collection from African Plio-Pleistocene sites. In addition to correlations based on shared faunal elements, we present an analysis based on the dentition of the abundant cercopithecid Theropithecus oswaldi, which increases in size in a manner that is strongly correlated with geological age (r2 ∼0.83), thereby providing a highly accurate age-estimation tool not previously utilized. In combination with paleomagnetic and U-Pb data, our results provide revised age estimates and suggest that there are no hominin sites in South Africa significantly older than ∼2.8 Ma. Where conflicting age estimates exist, we suggest that additional data are needed and recall that faunal estimates have ultimately proved reliable in the past (e.g., the age of the KBS Tuff).

Description and taxonomic assessment of fossil Cercopithecidae from the Pliocene Galili Formation (Ethiopia).

The Mount Galili Formation in the Afar region, Ethiopia, samples a critical time in hominin evolution, 4.4 to 3.8 Ma, documenting the last appearance of Ardipithecus and the origin of Australopithecus. This period is also important in the evolution of cercopithecids, especially the origin of Theropithecus in general and Theropithecus oswaldi lineage in particular. Galili has provided a total of 655 cercopithecid specimens that include crania, mandibles, isolated teeth and postcrania. All the fossils were recovered from the Lasdanan (5.3-4.43 Ma), Dhidinley (4.43-3.9 Ma) and Shabeley Laag (∼3.92-3.8 Ma) Members. Here, we described and analyzed 362 fossils employing both qualitative and quantitative methods. Descriptions of the material were supplemented with dental metrics and cranial shape analysis using three-dimensional geometric morphometrics. Results indicate the presence of at least six cercopithecid taxa: Theropithecus oswaldi serengetensis (n = 28), Theropithecus sp. (n = 2), three non-Theropithecus papionin groups (n = 134) and one colobine-size group (n = 58). The T. o. serengetensis represents the earliest form of the lineage, documented from ∼3.9 Ma Galili sediments. The three Galili papionins include a smaller taxon, a medium-sized taxon comparable to Pliopapio alemui and a large papionin overlapping in size with Soromandrillus, Gorgopithecus and Dinopithecus. The majority of Galili colobines have closest affinities to Kuseracolobus aramisi and some overlap with other taxa. Papionins dominate the Galili cercopithecid collection, although colobines are still fairly common (approximately 25% of the sample). Thus, Galili sample is like Kanapoi (4.2-4.1 Ma) and Gona (5.2-3.9 Ma) localities but distinct from Aramis, suggesting paleoecological similarity to the former sites. On the other hand, Theropithecus is less abundant at Galili than geologically younger Hadar (3.4-3.2 Ma) and Woranso-Mille (3.8-3.6 Ma) sites. Whether this difference is due to sampling, time or landscape variation requires further investigation.

The earliest most complete skeleton of Theropithecus.

The middle Pliocene site of Woranso-Mille in the Afar Region of Ethiopia has yielded numerous significant early hominin fossils representing multiple, coexisting taxa. Here we report on another significant discovery, the oldest partial skeleton of the papionin, Theropithecus. The specimen was recovered from the Aralee Issie collection area over multiple field seasons from 2004 through 2019. The specimen was unearthed in situ from the fluvial facies of the Mesgid Dora Tuff dated to 3.66-3.57 Ma. The partial skeleton, ARI-VP-1/26, is that of a subadult male lacking a skull. In the absence of unambiguously associated craniodental remains, the male status of the specimen was established from the dimensions of the long bones in comparison to those of other Theropithecus from Woranso-Mille. ARI-VP-1/26 is noteworthy because it preserves partial hand and foot skeletons, including a complete set of metacarpals from the left side. The theropith status of ARI-VP-1/26 was established based on the detailed anatomy of the postcranial skeleton, especially the proximal and distal humerus, proximal radius, and proximal femur. The morphology of the postcranium of ARI-VP-1/26 is consistent with that of recognized Theropithecus from Woranso-Mille and, specifically, with specimens recognized as Theropithecus oswaldi cf. darti from other sites. The ratio of the lengths of the first metacarpal to metacarpals 2-5 in ARI-VP-1/26 is intermediate between that seen in extant Papio and Theropithecus. In Theropithecus gelada and Theropithecus brumpti, the pairing of pollical and indical metacarpals of near equal length contributes to the species' high opposability index and is associated with a 'manual grazing' feeding habit. Cercopithecids constitute 43% of the identified vertebrates at Aralee Issie, and T. oswaldi cf. darti is the most common mammalian species. The monkeys of Aralee Issie lived in an open shrubland habitat, but the specific reasons for their high prevalence at the site are unclear.

Ontogenetic allometry and scaling in catarrhine crania.

In studies of ontogenetic allometry, ontogenetic scaling has often been invoked to explain cranial morphological differences between smaller and larger forms of closely related taxa. These scaled variants in shape have been hypothesized to be the result of the extension or truncation of common growth allometries. In this scenario, change in size is the determining factor, perhaps under direct selection, and changes in cranial shapes are byproducts, not under direct selection themselves. However, many of these conclusions are based on studies that used bivariate generalizations of shape. Even among multivariate analyses of growth allometries, there are discrepancies as to the prevalence of ontogenetic scaling among primates, how shared the trajectories need to be, and which taxa evince properties of scaled variants. In this investigation, we use a large, comparative ontogenetic sample, geometric morphometric methods, and multivariate statistical tests to examine ontogenetic allometry and evaluate if differences in cranial shape among closely related catarrhines of varying sizes are primarily driven by size divergence, that is, ontogenetic scaling. We then evaluate the hypothesis of size as a line of least evolutionary resistance in catarrhine cranial evolution. We found that patterns of ontogenetic allometry vary among taxa, indicating that ontogenetic scaling sensu stricto does not often account for most morphological differences and that large and small taxa within clades are generally not scaled variants. The presence of a variety of ontogenetic pathways for the evolution of cranial shapes provides indirect evidence for selection acting directly on the cranial shape, rather than on size alone.

Cercopithecid fossils from Kanapoi, West Turkana, Kenya (2007-2015).

Recent fieldwork at Kanapoi has expanded the sample of fossil cercopithecids, facilitating a re-appraisal of their taxonomy. The assemblage now includes at least one species of cercopithecin, two papionins, and two colobines. The guenon Nanopithecus browni is similar in dental size to extant Miopithecus. We tentatively re-affirm the identification of Parapapio cf. ado and confirm the presence of Theropithecus. The colobines include a small form tentatively attributed to Kuseracolobus and a second larger species. The Kanapoi fossils represent the oldest occurrences of guenons in Africa and of the important genus Theropithecus, the most abundant and widespread primate in the Neogene of Africa. In the assemblage, Parapapio cf. ado is the most abundant form, comprising the majority of specimens. All of the other taxa are comparatively rare. Colobines make up a small part of the Kanapoi fossil assemblage compared to most other contemporary sites, including Allia Bay, Kenya, where, like Kanapoi, Australopithecus anamensis has been found. The presence of Theropithecus is consistent with the presence of some relatively open habitat at Kanapoi. While the ecological preferences of the small cercopithecin are unknown, most guenons are associated with relatively wooded habitats, as are most colobines, suggesting the availability of at least some wooded areas.

Fossil Cercopithecidae from the Early Pliocene Sagantole Formation at Gona, Ethiopia.

The Early Pliocene Sagantole Fm. in the Gona Project area, Afar State, Ethiopia, is noted for discoveries of the early hominin Ardipithecus ramidus. A large series of fossil cercopithecid primates dated to between 4.8 and 4.3 Ma has also been collected from these sediments. In this paper, we use qualitative analysis and standard dental and postcranial measures to systematically describe the craniodental remains and tentatively allocate postcrania to taxa where we are able to. We then use these data to compare these specimens to fossil assemblages from contemporary sites, interpret their paleobiology, and discuss implications for the paleoecology of the Gona Sagantole Fm. We recognize three cercopithecid species in the Gona Sagantole Fm. Pliopapio alemui makes up approximately two-thirds of the identifiable specimens; nearly all of the rest are allocated to Kuseracolobus aramisi, and a single molar indicates the presence of a second, somewhat larger but morphologically distinct papionin. Among the Early Pliocene cercopithecids from Gona are also a number of postcranial elements. None of the postcranial remains are directly associated with any of the cranial material. Nonetheless, some of the distal humeri and proximal femora can be tentatively allocated to either Pl. alemui or K. aramisi based on a combination of size, as the latter is approximately 50% larger than the former, and morphology. If these assignments are correct, they suggest K. aramisi was primarily arboreal and similar to most extant colobines, whereas Pl. alemui was more mixed in its substrate use, being more terrestrially adapted than K. aramisi, but less so than extant Papio or Theropithecus. Thus, we interpret the predominance of Pl. alemui over K. aramisi is consistent with a somewhat more open environment at Gona than at Aramis.

Ontogeny and phylogeny of the cercopithecine cranium: A geometric morphometric approach to comparing shape change trajectories.

While the analysis of ontogenetic trajectories is common in geometric morphometrics (GM), the simultaneous comparison of several trajectories can be unwieldy and is, in some cases, unable to make use of one of the main advantages of GM, visualization. Furthermore, due to the paucity of the paleontological record, analyses of trajectories are often limited to extant taxa. We address these issues by presenting a method for visualizing the similarities and differences of cranial ontogenetic trajectories among taxa and a method for reconstructing ancestral ontogenetic trajectories, so that these differences can be investigated in a phylogenetic context. We also tested for the presence of phylogenetic signal in the ontogenetic trajectories themselves. Using an ontogenetic series of 522 crania, representing 17 cercopithecine species from 8 genera, we first calculated ontogenetic trajectories of cranial shape change for each species, and then entered these trajectories into a principal components analysis to produce a developmental shape-change trajectory PCA (δPCA). Then, through an augmentation of the phylomorphospace approach, we projected a molecular phylogeny onto the major axes of trajectory shape variation from the δPCA to produce an 'ontophylomorphospace,' using squared-change parsimony to reconstruct interior nodes. Through these procedures, we were able to determine that the δPCAs illustrate patterns of variation in these developmental trajectories in a visually intuitive manner that allows for easier comparisons among taxa. Through examination of the ontophylomorphospace, we found that African papionins exhibit extensive homoplasy in the evolution of cranial ontogenetic trajectories, and that Asian species of Macaca show highly derived ontogenetic trajectories relative to other cercopithecines. Additionally, we found no support for the presence of a phylogenetic signal in cranial ontogenetic trajectories. The δPCA and the ontophylomorphospace are ways in which to visualize and compare complex, multivariate shape transformations, both among extant taxa and over evolutionary time, respectively.

Evolution of the modern baboon (Papio hamadryas): A reassessment of the African Plio-Pleistocene record.

Baboons (Papio hamadryas) are among the most successful extant primates, with a minimum of six distinctive forms throughout Sub-Saharan Africa. However, their presence in the fossil record is unclear. Three early fossil taxa are generally recognized, all from South Africa: Papio izodi, Papio robinsoni and Papio angusticeps. Because of their derived appearance, P. angusticeps and P. robinsoni have sometimes been considered subspecies of P. hamadryas and have been used as biochronological markers for the Plio-Pleistocene hominin sites where they are found. We reexamined fossil Papio forms from across Africa with an emphasis on their distinguishing features and distribution. We find that P. robinsoni and P. angusticeps are distinct from each other in several cranial features, but overlap extensively in dental size. Contrary to previous assessments, no diagnostic cranio-mandibular material suggests these two forms co-occur, and dental variation at each site is comparable to that within P. h. ursinus, suggesting that only one form is present in each case. P izodi, however, may co-occur with P. robinsoni, or another Papio form, at Sterkfontein Member 4. P izodi appears more primitive than P. robinsoni and P. angusticeps. P. robinsoni is slightly distinct from P. hamadryas subspecies in its combination of features while P. angusticeps might be included within one of the modern P. hamadryas varieties (i.e., P. h. angusticeps). No definitive Papio fossils are currently documented in eastern Africa until the Middle Pleistocene, pointing to southern Africa as the geographic place of origin for the genus. These results have implications for Plio-Pleistocene biochronology and baboon evolution.

Trajectory analysis among African hominoids can provide insights into genetic and epigenetic influences during ontogeny.

OBJECTIVES: Prior examination of the ontogeny of Hominoid talo-crural joint morphology using Singular Warp analysis suggested both a genetic and epigenetic signal during development. This question is examined using trajectory analysis and its implications for the Hominin fossil record explored. MATERIALS AND METHODS: Trajectory analysis was used to examine talo-crural joint shape at different molar eruption stages during development among a cohort of 221 specimens of Homo sapiens, Gorilla gorilla, Pan troglodytes, and Pan paniscus. Trajectory length, angle, and shape were compared among species. Trajectories that showed a consistent pattern of shape change were predicted to show a genetic signal, while change in that pattern an epigenetic signal. RESULTS: The trajectories of all four hominoid tali were consistent during M1-M2 development, but there was a change in Pan paniscus alone at M2-M3. The tibial trajectories were again consistent in M1-M2 development, however, only Pan paniscus persisted in this trajectory at M2-M3 while the other three changed. DISCUSSION: Trajectory analysis demonstrated an epigenetic signal in Pan paniscus in the talus and among the other three hominoids in the tibia. Behavioral (epigenetic) changes may impact hominoid ankle shape in extant species. These changes may reflect differences in locomotor repertoire altering shape, as has been suggested in the Hominin fossil record. Further examination of closely related hominoid species may provide insights into potential behavioral influences in the rapid change and diversity of Hominin talo-crural shape in the Plio-pleistocene.

Dietary change among hominins and cercopithecids in Ethiopia during the early Pliocene.

The incorporation of C4 resources into hominin diet signifies increased dietary breadth within hominins and divergence from the dietary patterns of other great apes. Morphological evidence indicates that hominin diet became increasingly diverse by 4.2 million years ago but may not have included large proportions of C4 foods until 800 thousand years later, given the available isotopic evidence. Here we use carbon isotope data from early to mid Pliocene hominin and cercopithecid fossils from Woranso-Mille (central Afar, Ethiopia) to constrain the timing of this dietary change and its ecological context. We show that both hominins and some papionins expanded their diets to include C4 resources as early as 3.76 Ma. Among hominins, this dietary expansion postdates the major dentognathic morphological changes that distinguish Australopithecus from Ardipithecus, but it occurs amid a continuum of adaptations to diets of tougher, harder foods and to committed terrestrial bipedality. In contrast, carbon isotope data from cercopithecids indicate that C4-dominated diets of the earliest members of the Theropithecus oswaldi lineage preceded the dental specialization for grazing but occurred after they were fully terrestrial. The combined data indicate that the inclusion of C4 foods in hominin diet occurred as part of broader ecological changes in African primate communities.

New cranium of the large cercopithecid primate Theropithecus oswaldi leakeyi (Hopwood, 1934) from the paleoanthropological site of Makuyuni, Tanzania.

The Pleistocene hominin site of Makuyuni, near Lake Manyara, Tanzania, is known for fossils attributable to Homo and Acheulean artifacts (Ring et al., 2005; Kaiser et al., 2010; Frost et al., 2012). Here we describe the fossil primate material from the Manyara Beds, which includes the first nearly complete female cranium of Theropithecus oswaldi leakeyi and a proximal tibia from the same taxon. The cranium is dated to between 633 and 780 Ka and the tibia to the Pleistocene. The T. oswaldi lineage is one of the most important among Neogene mammals of Africa: it is both widespread and abundant. The size of the dentition, cranium, and tibia all confirm the previously recognized trend of increasing body size in this lineage and make their taxonomic assignments secure. The morphology of this specimen provides new insights into the evolution of this lineage through time, as well as its geographic variation and sexual dimorphism. The cranium also shows damage consistent with a mammalian carnivore, most likely a felid. The identification of this material as representing T. o. leakeyi agrees with the Middle Pleistocene age estimates for the MK4 locality in particular and the Manyara Beds in general.

Cranial sexual dimorphism in the Kinda baboon (Papio hamadryas kindae).

OBJECTIVES: The smallest extant member of genus Papio, the Kinda baboon exhibits low sexual dimorphism and a distinctive cranial shape. Ontogenetic scaling accounts for most cranial-shape differences within Papio, but studies have shown that the Kinda follows a separate ontogenetic trajectory. If so, its cranial-dimorphism pattern should differ from other subspecies. To evaluate this hypothesis, morphometric analysis was used to investigate cranial dimorphism in Papio. MATERIALS AND METHODS: Three-dimensional landmarks were digitized on 434 adult crania representing six Papio subspecies. Size- and shape-dimorphism magnitudes were quantified using centroid size and Procrustes distances. Patterns of sex- and size-related variation were explored using MAN(C)OVA, multivariate regression, and form-space PCA. Canine dimorphism was investigated using dental metrics. RESULTS: Kinda size and shape dimorphism are significantly lower than in other Papio subspecies. The relative magnitude of Kinda shape dimorphism is similar to other southern baboons; Kinda canine dimorphism is unremarkable. MAN(C)OVA results support subspecies differences in cranial dimorphism and scaling. Allometric and dimorphism vectors differ significantly in some subspecies, and their vector-angle matrices are strongly correlated. The Kinda's allometric vector angles are divergent. Form-space PC3, summarizing size-independent dimorphism, separates the Kinda from other subspecies. DISCUSSION: The Kinda baboon exhibits significantly lower size and shape dimorphism than other baboons, but its relative dimorphism levels are unexceptional. The Kinda differs from other subspecies in patterns of allometry, size-related shape dimorphism, and residual shape dimorphism. Kinda facial shape is "masculinized" relative to size, especially in females, suggesting female sexual selection contributed to the evolution of Kinda dimorphism.

Continuity of mammalian fauna over the last 200,000 y in the Indian subcontinent.

Mammalian extinction worldwide during the Late Pleistocene has been a major focus for Quaternary biochronology and paleoecology. These extinctions have been variably attributed to the impacts of climate change and human interference. However, until relatively recently, research has been largely restricted to the Americas, Europe, and Australasia. We present the oldest Middle-Late Pleistocene stratified and numerically dated faunal succession for the Indian subcontinent from the Billasurgam cave complex. Our data demonstrate continuity of 20 of 21 identified mammalian taxa from at least 100,000 y ago to the present, and in some cases up to 200,000 y ago. Comparison of this fossil record to contemporary faunal ranges indicates some geographical redistribution of mammalian taxa within India. We suggest that, although local extirpations occurred, the majority of taxa survived or adapted to substantial ecological pressures in fragmented habitats. Comparison of the Indian record with faunal records from Southeast and Southwest Asia demonstrates the importance of interconnected mosaic habitats to long-term faunal persistence across the Asian tropics. The data presented here have implications for mammalian conservation in India today, where increasing ecological circumscription may leave certain taxa increasingly endangered in the most densely populated region of the world.

Reassessment of Olduvai Bed I cercopithecoids: A new biochronological and biogeographical link to the South African fossil record.

Fossil monkeys have long been used as important faunal elements in studies of African Plio-Pleistocene biochronology, particularly in the case of the South African karst cave sites. Cercopithecoid fossils have been known from Tanzania's Olduvai Gorge for nearly a century, with multiple taxa documented including Theropithecus oswaldi and Cercopithecoides kimeui, along with papionins and colobines less clearly attributable to species. A small number of large papionin fossils, including a partial male cranium and partial female skull, have been previously identified as an early form of Papio, but noted as distinct from extant baboons as well as other fossil Papio species. In 2013 we reviewed the Olduvai cercopithecoid material at the National Museum of Tanzania, with a particular focus on the specimens from Beds I-IV. Quantitative and qualitative comparisons of the Olduvai papionins largely confirmed previous observations, with one notable exception. The large papionin taxon from Bed I previously recognized as Papio sp. is more properly recognized as Gorgopithecus major, a taxon previously known only from South Africa. Features shared between the Olduvai specimens and G. major include relatively short and concavo-convex tubular nasals, antero-posteriorly curved upper incisor roots, downwardly curved brow ridges in the midline, and robust zygomatic arches. The recognition of G. major at Olduvai Bed I, a well-known horizon with precise radiometric dates, provides an important biochronological and biogeographical link with South African localities Kromdraai A, Swartkrans Member 1 and possibly Swartkrans Members 2-3 and Cooper's A and D.

Constructing cranial ontogenetic trajectories: A comparison of growth, development, and chronological age proxies using a known-age sample of Macaca mulatta.

Recent morphometric research has generated opposing conclusions regarding the ontogenetic trajectories of catarrhine crania, possibly due to the ontogenetic proxies used to calculate them. Therefore, we used three surrogates: size, molar eruption, and chronological age to generate trajectories in a known-age sample to produce ontogenetic trajectories and determine the similarities and differences between them. Forty-three landmarks from an ontogenetic series of 160 Macaca mulatta crania, with associated ages at death, were used to produce ontogenetic trajectories of cranial shape change. These were computed by sex through multivariate regression of Procrustes aligned coordinates against three surrogates for ontogeny: natural log of centroid size (growth), molar eruption stage (development), and chronological age. These trajectories were compared by calculating the angles between them. Each trajectory was also used to produce simulated adults from juveniles, which were then compared with each other and actual adults. The different trajectories are nearly parallel as each of the surrogates track similar aspects of ontogenetic cranial shape change, but chronological age was the most divergent. Simulated adults produced using the developmental stage trajectories were most similar to actual adults. When simulated adults were produced from opposite sex trajectories, they resembled the sex from which the trajectory was produced, not the sex of the juvenile specimen. We discuss properties of the trajectories produced from each of the surrogates, the possible reasons for previously opposing conclusions, how these properties can inform future investigations, and how our investigation bears on analyses of heterochrony.

Early Pliocene Cercopithecidae from Woranso-Mille (Central Afar, Ethiopia) and the origins of the Theropithecus oswaldi lineage.

A large series of fossil cercopithecids has been recovered from the hominid-bearing Woranso-Mille site, Afar State, northeastern Ethiopia. Here we report the taxonomy of those specimens from the Am-Ado, Aralee Issie, Korsi Dora, Makah Mera, and Mesgid Dora collection areas, which are all roughly contemporaneous and dated to between 3.6 and 3.8 million years ago. This series includes a minimum of two cercopithecine and three colobine species. Theropithecus oswaldi cf. darti is by far the most common species in the assemblage, making up over 90% of identifiable cercopithecid specimens. There is also at least one other species of papionin, which cannot be currently assigned to a genus. The colobines are here allocated to Cercopithecoides cf. meaveae and two other species, one small and one large, that cannot be currently assigned to genus. The T. oswaldi cf. darti series from Woranso-Mille is both the earliest and largest identified to date. It documents the earliest occurrence of the T. oswaldi lineage and strongly suggests that parallel evolution of molar morphology has occurred within the genus between T. oswaldi and Theropithecus brumpti. Given the dominance of monkeys at Woranso-Mille, and the preponderance of Theropithecus among cercopithecids, T. o. cf. darti is likely to be the most common mammal present at the 3.6-3.8 million-years-old localities of the Woranso-Mille study area. Some explanations for this unusual occurrence are explored, and implications for the paleoenvironment at Woranso-Mille are also discussed.

The ontogeny of talo-crural appositional articular morphology among catarrhine taxa: adult shape reflects substrate use.

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.

Facial modeling and measurement based upon homologous topographical features.

Measurement of human faces is fundamental to many applications from recognition to genetic phenotyping. While anthropometric landmarks provide a conventional set of homologous measurement points, digital scans are increasingly used for facial measurement, despite the difficulties in establishing their homology. We introduce an alternative basis for facial measurement, which 1) provides a richer information density than discrete point measurements, 2) derives its homology from shared facial topography (ridges, folds, etc.), and 3) quantifies local morphological variation following the conventions and practices of anatomical description. A parametric model that permits matching a broad range of facial variation by the adjustment of 71 parameters is demonstrated by modeling a sample of 80 adult human faces. The surface of the parametric model can be adjusted to match each photogrammetric surface mesh generally to within 1 mm, demonstrating a novel and efficient means for facial shape encoding. We examine how well this scheme quantifies facial shape and variation with respect to geographic ancestry and sex. We compare this analysis with a more conventional, landmark-based geometric morphometric (GMM) study with 43 landmarks placed on the same set of scans. Our multivariate statistical analysis using the 71 attribute values separates geographic ancestry groups and sexes with a high degree of reliability, and these results are broadly similar to those from GMM, but with some key differences that we discuss. This approach is compared with conventional, non-parametric methods for the quantification of facial shape, including generality, information density, and the separation of size and shape. Potential uses for phenotypic and dysmorphology studies are also discussed.

Expanded geographic distribution and dietary strategies of the earliest Oldowan hominins and Paranthropus.

The oldest Oldowan tool sites, from around 2.6 million years ago, have previously been confined to Ethiopia's Afar Triangle. We describe sites at Nyayanga, Kenya, dated to 3.032 to 2.581 million years ago and expand this distribution by over 1300 kilometers. Furthermore, we found two hippopotamid butchery sites associated with mosaic vegetation and a C4 grazer-dominated fauna. Tool flaking proficiency was comparable with that of younger Oldowan assemblages, but pounding activities were more common. Tool use-wear and bone damage indicate plant and animal tissue processing. Paranthropus sp. teeth, the first from southwestern Kenya, possessed carbon isotopic values indicative of a diet rich in C4 foods. We argue that the earliest Oldowan was more widespread than previously known, used to process diverse foods including megafauna, and associated with Paranthropus from its onset.