The role of developmental rate, body size, and positional behavior in the evolution of covariation and evolvability in the cranium of strepsirrhines and catarrhines.

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.

Palaeoecological differences underlie rare co-occurrence of Miocene European primates.

BACKGROUND: The two main primate groups recorded throughout the European Miocene, hominoids and pliopithecoids, seldom co-occur. Due to both their rarity and insufficiently understood palaeoecology, it is currently unclear whether the infrequent co-occurrence of these groups is due to sampling bias or reflects different ecological preferences. Here we rely on the densely sampled primate-bearing sequence of Abocador de Can Mata (ACM) in Spain to test whether turnovers in primate assemblages are correlated with palaeoenvironmental changes. We reconstruct dietary evolution through time (ca. 12.6-11.4 Ma), and hence climate and habitat, using tooth-wear patterns and carbon and oxygen isotope compositions of enamel of the ubiquitous musk-deer Micromeryx. RESULTS: Our results reveal that primate species composition is strongly correlated with distinct environmental phases. Large-bodied hominoids (dryopithecines) are recorded in humid, densely-forested environments on the lowermost portion of the ACM sequence. In contrast, pliopithecoids inhabited less humid, patchy ecosystems, being replaced by dryopithecines and the small-bodied Pliobates toward the top of the series in gallery forests embedded in mosaic environments. CONCLUSIONS: These results support the view that pliopithecoid primates preferred less humid habitats than hominoids, and reveal that differences in behavioural ecology were the main factor underpinning their rare co-occurrence during the European Miocene. Our findings further support that ACM hominoids, like Miocene apes as a whole, inhabited more seasonal environments than extant apes. Finally, this study highlights the importance of high-resolution, local investigations to complement larger-scale analyses and illustrates that continuous and densely sampled fossiliferous sequences are essential for deciphering the complex interplay between biotic and abiotic factors that shaped past diversity.

Habitat use by a primate community in a lowland dipterocarp forest in Danum Valley, Borneo.

Knowledge of niche partitioning with respect to habitat is indispensable to understand the mechanism of coexistence of multiple species. Among primates, however, data are still deficient because repeated survey for a sufficiently long time, covering seasonal changes over a large area, is the only way to clarify habitat segregation within a seasonally fluctuating environment. Southeast Asia is particularly interesting because of the supra-annual, highly unpredictable seasonality in fruiting known as mast fruiting. We conducted repeated route census, habitat monitoring, and group tracking for 25 months in two study sites (ca. 10 km apart) in the largely primary lowland dipterocarp forest of the Danum Valley Conservation Area, eastern Sabah, northern Borneo, Malaysia. The five species of diurnal primates (Bornean orangutan Pongo pygmaeus, Müeller's gibbon Hylobates muelleri, red leaf monkey Presbytis rubicunda, long-tailed macaque Macaca fascicularis, and southern pig-tailed macaque M. nemestrina) did not show horizontal spatial segregation. Red leaf monkeys showed preferences for places with short tree height, but their distribution was not confined to such places. In response to the fruiting peak observed once during the study period, orangutans increased their numbers simultaneously in the two study sites. The average tree height used by the five species was different, but their range overlapped substantially. Compared with other primate communities, the lack of horizontal spatial segregation and the suggested long-distance movement of orangutans seem to be unique characteristics in Borneo, although the use of different forest strata is a widespread phenomenon among primate communities throughout the world.

Functional and developmental influences on intraspecific variation in catarrhine vertebrae.

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.

Understanding climate's influence on the extinction of Oreopithecus (late Miocene, Tusco-Sardinian paleobioprovince, Italy).

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.

Articular scaling and body mass estimation in platyrrhines and catarrhines: Modern variation and application to fossil anthropoids.

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.

Alternative methods for calculating percentage prediction error and their implications for predicting body mass in fossil taxa.

Since body mass covaries with many ecological aspects of an animal, body mass prediction of fossil taxa is a frequent goal of paleontologists. Body mass prediction often relies on a body mass prediction equation (BMPE): a bivariate relationship between a predictor variable (e.g., molar occlusal area, femoral head breadth) and body mass as observed in extant taxa. A variety of metrics have been used to assess the reliability of BMPEs, including percentage prediction error (%PE), which involves predicting body masses of a test sample comprising individuals with associated masses. A mean %PE can be calculated in two ways: 1) as the mean %PE of multiple individual predictions (%MPE), or 2) as the %PE of mean body mass generated from the mean predictor value of multiple individuals (here termed %PEM). Differences between these two approaches have never been formally examined and no formal protocols have been recommended. Using a large sample of cercopithecoid primates (406 individuals from 50 species/subspecies) with associated body masses, body mass is predicted with six previously published interspecific BMPEs. Both %MPE and %PEM are calculated and compared. For all BMPEs, the distributions of differences between %MPE and %PEM exhibit positive skew and have medians significantly greater than zero, indicating that the examined prediction equations are more accurate at predicting mean body mass when they are applied to mean predictor values. The decreased predictive accuracy of %MPE relative to %PEM likely stems from changing the unit of analysis from mean values (in the reference sample) to individual values (in the test sample) when calculating %MPE. Empirical results are supported with a simulated dataset. Implications for body mass prediction in fossil species are discussed.

Buccal dental microwear texture and catarrhine diets.

OBJECTIVES: Two-dimensional dental microwear analyses on occlusal and nonocclusal enamel surfaces have been widely applied to reconstruct the feeding behaviors of extant primates and to infer ecological adaptations in fossil hominins. To date, analyses of dental microwear texture, using three-dimensional, Scale-Sensitive Fractal Analysis approaches has only been applied to occlusal surfaces. Here, for the first time, we apply this 3D proxy to buccal enamel surfaces of catarrhine primates of known feeding ecologies to assess the utility of nonocclusal microwear texture variables as indicators of dietary habits. MATERIALS AND METHODS: Buccal microwear texture attributes were collected from high-resolution second molar casts in a sample of seven extant African catarrhine taxa with differing dietary behaviors. A white-light confocal microscope with a 100× objective lens was used to record six microwear texture variables that assess complexity, anisotropy, heterogeneity, and textural fill volume. RESULTS: The physical properties and variation in hardness of ingested foods is reflected by significant differences in the microwear variables on buccal enamel surfaces between species, which is in agreement with early reports using 2D microwear signatures of the same samples. Species that consume hard brittle items showed high buccal enamel complexity and low anisotropy values, while folivorous species that consume tough foods revealed high anisotropy and low complexity enamel patterns. DISCUSSION: Buccal microwear texture analysis on enamel surfaces clearly reflects diet-related variation in nonhuman primates. Our findings indicate that microwear texture attributes on nonworking enamel surfaces provide an alternative procedure for reconstructing dietary behavior when wear facets on occlusal surfaces are lacking.

Disproportionate Cochlear Length in Genus Homo Shows a High Phylogenetic Signal during Apes' Hearing Evolution.

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.

Co-occurrence of pliopithecoid and hominoid primates in the fossil record: An ecometric analysis.

Both pliopithecoid and hominoid primates were widely distributed throughout Eurasia during the Miocene but are known to have coexisted at only a few localities. It has been speculated that their different habitat preferences permitted only minimal overlap under special environmental conditions. Here we study the context for pliopithecoid and hominoid co-occurrence by assessing taxonomically-based palaeoecological diversity of associated fossil mammals and by direct ecometric analysis based on hypsodonty of mammalian herbivores. Our results show that pliopithecoids persistently inhabited more humid environments compared to the other primate groups studied, suggesting an inability to adapt to changing environmental conditions. The opportunity for hominoids and pliopithecoids to co-occur appears to have been restricted by niche conservatism in the latter group. Our study also indicates that direct ecometric analysis gives a better separation of the ecological preferences of these primate clades than do analyses of taxonomically-based community structure.

Morphological affinities of the proximal humerus of Epipliopithecus vindobonensis and Pliopithecus antiquus: suspensory inferences based on a 3D geometric morphometrics approach.

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).

Inferring the use of forelimb suspensory locomotion by extinct primate species via shape exploration of the ulna.

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.

Adaptive response to sociality and ecology drives the diversification of facial colour patterns in catarrhines.

The faces of Old World monkeys and apes (Catarrhini) exhibit every possible hue in the spectrum of mammal colours. Animal colouration experiences selection for communication, physiology and ecology; however, the relative importance of these factors in producing facial diversity in catarrhines is not known. Here we adopt a comparative approach to test whether facial traits have evolved in tandem with social, geographic and ecological pressures across four catarrhine radiations. Our analyses reveal the underlying correlates of two major axes in the evolution of facial diversity. Facial colour patterns are linked to social factors, such that gregarious and highly sympatric species have evolved more colours in their faces. Facial pigmentation tends to be dominated by ecological factors, and species living in tropical, densely forested and humid habitats in Africa have evolved darker faces. Thus, both sociality and ecology have played a role in producing the highest diversity of faces within mammals.

New evidence for diet and niche partitioning in Rudapithecus and Anapithecus from Rudabánya, Hungary.

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.

Stable isotope ecology of Miocene bovids from Northern Greece and the ape/monkey turnover in the Balkans.

Eurasia was home to a great radiation of hominoid primates during the Miocene. All were extinct by the end of the Miocene in Western Eurasia. Here, we investigate the hypothesis of climate and vegetation changes at a local scale when the cercopithecoid Mesopithecus replaced the hominoid Ouranopithecus along the Axios River, Greece. Because they are herbivorous and were much more abundant than primates, bovids are preferred to primates to study climate change in the Balkans as a cause of hominoid extinction. By measuring carbon stable isotope ratios of bovid enamel, we conclude that Ouranopithecus and Mesopithecus both evolved in pure C3 environments. However, the large range of δ(13)C values of apatite carbonate from bovids combined with their molar microwear and mesowear patterns preclude the presence of dense forested landscapes in northern Greece. Instead, these bovids evolved in rather open landscapes with abundant grasses in the herbaceous layer. Coldest monthly estimated temperatures were below 10°C and warmest monthly temperatures rose close to or above 20°C for the two time intervals. Oxygen isotope compositions of phosphate from bulk samples did not show significant differences between sites but did show between-species variation within each site. Different factors influence oxygen isotope composition in this context, including water provenience, feeding ecology, body mass, and rate of amelogenesis. We discuss this latter factor in regard to the high intra-tooth variations in δ(18)Op reflecting important amplitudes of seasonal variations in temperature. These estimations fit with paleobotanical data and differ slightly from estimations based on climate models. This study found no significant change in climate before and after the extinction of Ouranopithecus along the Axios River. However, strong seasonal variations with relatively cold winters were indicated, conditions quite usual for extant monkeys but unusual for great apes distributed today in inter-tropical regions.

A resampling approach and implications for estimating the phalangeal index from unassociated hand bones in fossil primates.

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.

The human post-fertile lifespan in comparative evolutionary context.

There persist two widely held but mutually inconsistent views on the evolution of post-fertile lifespan of human females. The first, prevalent within anthropology, sees post-fertile lifespan (PFLS) in the light of adaptive processes, focusing on the social and economic habits of humans that selected for a lengthy PFLS. This view rests on the assumption that human PFLS is distinct from that of other species, and focuses on quantifying the selective causes and consequences of that difference. The second view, prevalent within gerontology and comparative biology, emphasizes that PFLS is a phylogenetically widespread trait or that human PFLS is predictable based on life-history allometries. In this view, human PFLS is part of a broad cross-species pattern and its genesis cannot, therefore, rely on human-specific traits. Those who advocate the second view have questioned the "special pleading" for human specific explanations of PFLS, and have argued that human PFLS is quantitatively greater but not qualitatively different than PFLS in many other animals. Papers asking whether human PFLS is explained by the importance of mothers more than grandmothers, whether paternal or maternal grandparents have more of an effect on child survival, or who is providing the excess calories are associated with the first view that assumes the need to explain the existence of human PFLS on the basis of a uniquely human socioecology. Anthropologists largely see human PFLS as derived, while comparative gerontologists point to evidence that it is one instance of a ubiquitous cross-species pattern. The two groups generally occupy non-overlapping research circles, in terms of conferences and journals, and therefore interact little enough to largely avoid the need to reconcile their views, allowing the persistence of misconceptions in each field. Our goal is to identify and address the most important of these misconceptions and thereby make clear that both of these seemingly incongruent views contain valid points. We argue that two distinct but related traits have been lumped together under the same concept of "post-reproductive lifespan," one (post-fertile viability) that is tremendously widespread and another (a post-fertile life stage) that is derived to hominins, and that the differences and connections between these two traits are necessary for understanding human life-history evolution.

The shape and presentation of the Catarrhine talus: a geometric morphometric analysis.

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.

Functional links between canine height and jaw gape in catarrhines with special reference to early hominins.

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.