Limb and hip morphology of two African colobine monkeys and its relationship to the mechanics of leaping and bounding locomotion.

OBJECTIVES: Although a bounding gait is practiced by a diversity of animals, the morphological characteristics, kinematics, and energetics associated with this locomotor form remain poorly understood. This study focuses on the locomotor anatomy of two species of African colobine monkeys (Piliocolobus badius, a leaper, and Colobus polykomos, a leaper-bounder) in an effort to assess if bounding should be considered a unique primate locomotor category or is better viewed as a behavior on a leaping continuum. MATERIALS AND METHODS: A total of 53 femora, 28 humeri, and 45 ossa coxae from the two species provide comparative morphological data. Free-body models of bounding and leaping are presented to characterize loading conditions. Species differences in morphometric traits are evaluated via parametric and nonparametric tests (i.e., analysis of variance, resampling). RESULTS: C. polykomos exhibits traits that align more closely with putative leaping specializations when compared to P. badius (e.g., large femoral head, long femur, low femoral neck angle), while also possessing certain traits that are not (e.g., long femoral neck and reduced relative femoral robusticity). Consequently, C. polykomos likely experiences absolutely greater joint forces at the hip and higher bending at the femoral neck both when it leaps and bounds, given equivalent accelerations in bounding and leaping. DISCUSSION: Bounding is best described as a form of low-acceleration leaping. If bounding has lower acceleration requirements relative to leaping, C. polykomos achieves locomotor competence with less energy, relatively smaller bending moments, and reduced joint forces.

The anatomy of the hindlimb of Theropithecus brumpti (Cercopithecidae, Papionini): Morphofunctional implications.

Theropithecus brumpti is a primate known from numerous craniodental specimens in the Plio-Pleistocene Shungura Formation (Lower Omo Valley, Ethiopia), but the anatomy of its hindlimb is documented only by a few associated and mostly incomplete postcranial specimens. The adaptations of T. brumpti are still debated, with its substrate preferences and its use of squatting postures recently discussed based on anatomical differences when compared with its extant representative, Theropithecus gelada. Here, we describe an associated femur and tibia (L 869-1 and L 869-2) of a presumed T. brumpti male and a partial foot (L 865-1r and L 865-1t) of a male T. brumpti, dated to ca. 2.6 Ma and ca. 2.32 Ma respectively. Based on univariate and bivariate morphometric analyses, we provide new data on the morphological correlates of substrate preferences and postural behaviors of this fossil species. Our results are in agreement with previous analyses and present T. brumpti as a predominantly terrestrial primate. We demonstrate the presence of osteological correlates associated with the use of squatting behaviors in T. brumpti but also point to significant anatomical differences between this paleontological species and T. gelada. These differences blur the functional value of characters previously identified as diagnostic of T. gelada and its postural behavior. We further document the postcranial distinctiveness of the Theropithecus clade in relation to the Papio clade. This study thus provides new insights into the postcranial anatomy and paleoecology of an abundant fossil primate from the Plio-Pleistocene of eastern Africa.

A new species of fossil guenon (Cercopithecini, Cercopithecidae) from the Early Pleistocene Lower Ngaloba Beds, Laetoli, Tanzania.

The living guenons (Cercopithecini, Cercopithecidae) are speciose and widely distributed across sub-Saharan Africa but are poorly represented in the fossil record. In addition, the craniodental and skeletal similarity of the guenons has hampered the identification of fragmentary material, likely obscuring the taxonomic diversity represented in the fossil record. Here, we describe a new fossil guenon specimen (LAET 75-3703) from the Lower Ngaloba Beds, Laetoli in Tanzania, dated to ∼1.7-1.2 Ma and preserving the lower face and mandible. Comparison to 278 extant guenon specimens, representing all six extant genera, identified several informative traits for distinguishing between the morphologically similar Chlorocebus and Cercopithecus, and these support the attribution of LAET 75-3703 to Chlorocebus. A discriminant function analysis of seven craniodental indices on a subsample of Chlorocebus and Cercopithecus was robust with an overall correct classification rate of 80.4%, and it classified LAET 75-3703 as a member of Chlorocebus with a posterior probability of 92.7%. LAET 75-3703 shares with Chlorocebus the presence of small 'thumbprint' depressions on the maxilla; a tall, narrow, and diamond-shaped nasal aperture; a relatively longer and shallower face; relatively buccolingually broader molars; and a shallow mandible that decreases in depth posteriorly. In addition, LAET 75-3703 is distinguished from all extant guenons, including other species of Chlorocebus, in having a very small P3 relative to M1 area. As such, LAET 75-3703 is assigned to a new species, Chlorocebus ngedere sp. nov. This specimen represents the first cercopithecin from Laetoli, as well as the oldest fossil cercopithecin confidently attributed to a modern genus.

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.

Patterns and magnitudes of craniofacial covariation in extant cercopithecids.

The cranium contains almost all of the vertebrate sensory organs and plays an essential role in vertebrate evolution. Research on the primate cranium has revealed that it is both highly integrated and modular, but studies have historically focused on covariance between the neurocranium and facial skeleton rather than on bones specific to special senses such as vision. The goal of this work is to investigate patterns and magnitudes of craniofacial covariation in extant cercopithecids with particular attention to the orbits. This study takes a quantitative approach using data collected from 38 homologous cranial landmarks across 11 genera of cercopithecid monkeys (Cercopithecidae, N = 291). These data demonstrate that both patterns and magnitudes of craniofacial covariation differ across Cercopithecidae at subfamily, tribe, and genus levels, with the strongest integration in the papionins (and specifically Papio) and significantly weaker covariation in the colobines, particularly Presbytis. Orbital height does not covary with other measurements of the cranium to the same degree as other cranial traits in Cercopithecidae and is highly constrained across the family. This study has important implications for our understanding of the evolution and development of morphological diversity in the cercopithecid cranium and evolution of the primate eye. This study also highlights the potential error of broad assumptions about generalizing patterns and magnitudes of modularity and integration across primates. Additionally, these findings reiterate the importance of trait selection for interpreting fossil taxonomy, as craniofacial covariation may impact phenotypes commonly used to differentiate fossil primate species.

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.

Ecomorphological specialization leads to loss of evolvability in primate limbs.

Primate limb morphology is often described as either generalized, that is, suited to a range of locomotor and positional behaviors, or specialized for unique locomotor behaviors such as brachiation or bipedalism. The evolution of highly specialized limb morphology may result in loss of evolvability, that is, in a decreased capacity of the locomotor skeleton to evolve in response to selection towards alternative ecomorphological niches. Using evolutionary simulations, I show that the highly specialized limb anatomy of hominoids is associated with a significant loss of evolvability, defined as the number of generations to reach alternative adaptive peaks, and in parallel an increased risk of extinction, particularly in simulated evolution toward generalized quadrupedal limb proportions. Loss of evolvability in apes and humans correlates with three factors: (1) decreased correlation among limb bone lengths (i.e., integration), which slows the rate of change along lines of least evolutionary resistance; (2) limb specialization, which places apes and humans in relatively remote areas of morphospace; and (3) increased skeletal size as a proxy for body size. Thus, locomotor over-specialization can lead to evolutionary dead-ends that significantly increase the probability of hominoid populations going extinct before evolving new adaptive morphologies.

Long bone structural proportions and locomotor behavior in Cercopithecidae.

Limb bone articular and diaphyseal proportions have been shown to relate to locomotor behavior in broad comparisons across catarrhines, but comparisons among phylogenetically and functionally more closely related species may be particularly useful in investigating form-function relationships that can be applied to fossil taxa. Here we compare inter- and intra-limb proportions of diaphyseal strength and articular surface area and breadth of the femur and humerus with frequencies of leaping and vertical climbing behavior in 13 cercopithecid species. Leaping frequency is highly positively correlated with femoral/humeral diaphyseal strength, moderately positively correlated with femoral/humeral articular breadth, and less highly correlated with femoral/humeral articular surface area. These results are consistent with predicted higher bending loads as well as joint reaction forces on the femora of leapers. Surface areas may show a weaker association because they also directly impact joint excursion and are thus more influenced by other aspects of locomotion, including climbing. Climbing frequency is positively correlated with humeral head articular surface area/diaphyseal strength, but weakly negatively correlated with femoral head articular surface area/diaphyseal strength. These combined trends lead to a strong negative association between climbing and femoral/humeral head surface area. Femoral/humeral diaphyseal strength and distal articular breadth are not correlated with climbing frequency. The climbing results are consistent with greater shoulder mobility in more frequent vertical climbers. The lack of such a relationship in the femur among these taxa contrasts with earlier findings for catarrhines more generally, including hominoids, and may be a result of different climbing kinematics in cercopithecoids involving less hip abduction than in hominoids. Different use of the forelimb during climbing in cercopithecoids and hominoids may also explain the lack of association between femoral/humeral diaphyseal strength and climbing in the present study, in contrast to comparisons across catarrhines more generally.

A window into the early evolutionary history of Cercopithecidae: Late Miocene evidence from Chad, Central Africa.

Central Africa is known as a major center of diversification for extant Old World Monkeys (OWM) and yet has a poorly documented fossil record of monkeys. Here we report a new colobine monkey (Cercopithecoides bruneti sp. nov.) from the Central African hominin-bearing fossiliferous area of Toros-Menalla, Chad at ca. 7 Ma. In addition to filling a gap in the spatial and temporal record of early OWM evolutionary history, we assess the ecomorphological diversity of early OWM by providing evidence on the onset of a folivorous diet and a partial reacquisition of terrestrial locomotor habits among Miocene colobines. We also support the phylogenetic affinities of the genus Cercopithecoides among the stem group of the extant African colobine monkeys.

Parietal lobe variation in cercopithecid endocasts.

In extant primates, the posterior parietal cortex is involved in visuospatial integration, attention, and eye-hand coordination, which are crucial functions for foraging and feeding behaviors. Paleoneurology studies brain evolution through the analysis of endocasts, that is molds of the inner surface of the braincase. These may preserve imprints of cortical structures, such as sulci, which might be of interest for locating the boundaries of major cortical regions. Old World monkeys (Cercopithecidae) represent an interesting zoological group for evolutionary studies, because of their diverse ecologies and locomotor behaviors. In this study, we quantify parietal lobe variation within the cercopithecid family, in a sample of 30 endocasts including 11 genera and 17 species, by combining landmark-based and landmark-free geometric morphometric analyses. More specifically, we quantitatively assess variation of the parietal proportions based on landmarks placed on reliable anatomical references and of parietal lobe surface morphology through deformation-based methods. The main feature associated with the cercopithecid endocranial variation regards the inverse proportions of parietal and occipital lobes, with colobines, Theropithecus, and Papio displaying relatively larger parietal lobes and smaller occipital lobes compared with cercopithecins. The parietal surface is anteroposteriorly longer and mediolaterally flatter in colobines, while longitudinally shorter but laterally bulging in baboons. Large parietal lobes in colobines and baboons are likely to be independent evolutionary traits, and not necessarily associated with analogous functions or morphogenetic mechanisms.

Primitive Old World monkey from the earliest Miocene of Kenya and the evolution of cercopithecoid bilophodonty.

Old World monkeys (Cercopithecoidea) are a highly successful primate radiation, with more than 130 living species and the broadest geographic range of any extant group except humans. Although cercopithecoids are highly variable in habitat use, social behavior, and diet, a signature dental feature unites all of its extant members: bilophodonty (bi: two, loph: crest, dont: tooth), or the presence of two cross-lophs on the molars. This feature offers an adaptable Bauplan that, with small changes to its individual components, permits its members to process vastly different kinds of food. Old World monkeys diverged from apes perhaps 30 million years ago (Ma) according to molecular estimates, and the molar lophs are sometimes incompletely developed in fossil species, suggesting a mosaic origin for this key adaptation. However, critical aspects of the group's earliest evolution remain unknown because the cercopithecoid fossil record before ∼18 Ma consists of only two isolated teeth, one from Uganda and one from Tanzania. Here we describe a primitive Old World monkey from Nakwai, Kenya, dated at ∼22 Ma, that offers direct evidence for the initial key steps in the evolution of the cercopithecoid dentition. The simple dentition and absence of bilophodonty in the Nakwai monkey indicate that the initial radiation of Old World monkeys was first characterized by a reorganization of basic molar morphology, and a reliance on cusps rather than lophs suggests frugivorous diets and perhaps hard object feeding. Bilophodonty evolved later, likely in response to the inclusion of leaves in the diet.

Wrist morphology reveals substantial locomotor diversity among early catarrhines: an analysis of capitates from the early Miocene of Tinderet (Kenya).

Considerable taxonomic diversity has been recognised among early Miocene catarrhines (apes, Old World monkeys, and their extinct relatives). However, locomotor diversity within this group has eluded characterization, bolstering a narrative that nearly all early catarrhines shared a primitive locomotor repertoire resembling that of the well-described arboreal quadruped Ekembo heseloni. Here we describe and analyse seven catarrhine capitates from the Tinderet Miocene sequence of Kenya, dated to ~20 Ma. 3D morphometrics derived from these specimens and a sample of extant and fossil capitates are subjected to a series of multivariate comparisons, with results suggesting a variety of locomotor repertoires were present in this early Miocene setting. One of the fossil specimens is uniquely derived among early and middle Miocene capitates, representing the earliest known instance of great ape-like wrist morphology and supporting the presence of a behaviourally advanced ape at Songhor. We suggest Rangwapithecus as this catarrhine's identity, and posit expression of derived, ape-like features as a criterion for distinguishing this taxon from Proconsul africanus. We also introduce a procedure for quantitative estimation of locomotor diversity and find the Tinderet sample to equal or exceed large extant catarrhine groups in this metric, demonstrating greater functional diversity among early catarrhines than previously recognised.

Expression Changes of Structural Protein Genes May Be Related to Adaptive Skin Characteristics Specific to Humans.

Human skin is morphologically and physiologically different from the skin of other primates. However, the genetic causes underlying human-specific skin characteristics remain unclear. Here, we quantitatively demonstrate that the epidermis and dermis of human skin are significantly thicker than those of three Old World monkey species. In addition, we indicate that the topography of the epidermal basement membrane zone shows a rete ridge in humans but is flat in the Old World monkey species examined. Subsequently, we comprehensively compared gene expression levels between human and nonhuman great ape skin using next-generation cDNA sequencing (RNA-Seq). We identified four structural protein genes associated with the epidermal basement membrane zone or elastic fibers in the dermis (COL18A1, LAMB2, CD151, and BGN) that were expressed significantly greater in humans than in nonhuman great apes, suggesting that these differences may be related to the rete ridge and rich elastic fibers present in human skin. The rete ridge may enhance the strength of adhesion between the epidermis and dermis in skin. This ridge, along with a thick epidermis and rich elastic fibers might contribute to the physical strength of human skin with a low amount of hair. To estimate transcriptional regulatory regions for COL18A1, LAMB2, CD151, and BGN, we examined conserved noncoding regions with histone modifications that can activate transcription in skin cells. Human-specific substitutions in these regions, especially those located in binding sites of transcription factors which function in skin, may alter the gene expression patterns and give rise to the human-specific adaptive skin characteristics.

The Influence of Leaping Frequency on Secondary Bone in Cercopithecid Primates.

Bone remodeling is at least partially mediated by the mechanical environment created by an animal's behavior. Here, we test the hypothesis that bone remodeling is primarily induced by high magnitude loads, likely encountered during leaping/bounding behaviors. Osteon population density (OPD), osteon cross-sectional area (On.Ar), and relative osteonal area (%HAV) were measured from femoral and humeral midshaft thin sections of four cercopithecids (N = 5 per species) from Taï Forest, Côte d'Ivoire: Colobus polykomos, Piliocolobus badius, Cercopithecus diana, and Cercocebus atys. All species are generalized quadrupeds but vary in leaping frequency and overall activity budget. Differences between taxa with high (C. polykomos and P. badius) and low leaping frequency (C. diana and C. atys) were assessed via a phylogenetically informed generalized linear mixed model using Markov Chain Monte Carlo methods. Femoral OPD and %HAV are greater in the high frequency leapers than in low frequency leapers, suggesting that frequent high magnitude loads engender remodeling, however, there is no similar pattern in the humerus, which presumably also experiences high magnitude loads during leaping. Additionally, OPD and %HAV are greater in the humerus than the femur, despite load magnitude being presumably higher in the femur. These results provide conflicting support for hypotheses about load magnitude and load frequency as they relate to bone remodeling activity. Future work is proposed to parse out the respective effects of load magnitude and frequency on bone remodeling. Anat Rec, 302:1116-1126, 2019. © 2018 Wiley Periodicals, Inc.

Paternity data and relative testes size as measures of level of sperm competition in the Cercopithecoidea.

Historically, the empirical study of the role of sperm competition in the evolution of sexual traits has been problematic through an enforced reliance on indirect proxy measures. Recently, however, a procedure was developed that uses paternity data to measure sperm competition level directly in terms of males/conception (i.e., the number of males that have sperm present in a female's ampulla at conception). When tested on apes and humans (Hominoidea) this measure proved not only to correlate significantly with the traditionally used measure of relative testes size but also to offer a number of advantages. Here we provide a second test of the procedure, this time using paternity data for the Old World monkeys (Cercopithecoidea). We calculate sperm competition levels (males/conception) for 17 species of wild and free-ranging cercopithecoids and then analyze the data against measures of relative testes size. Calculated sperm competition levels correlate strongly with relative testes size both with and without phylogenetic control at both the species and generic levels. The signal-to-noise ratios inherent in both the past measure of relative testes size and the new measure of sperm competition level from paternity data are discussed. We conclude that although both measures are appropriate for the future study of the role of sperm competition in the evolution of sexual traits, when paternity data are available they provide the more direct and meaningful analytical tool. Not least, they potentially allow a first empirical analysis of the role of sperm competition in the evolution of relative testes size that could then be compared with the wealth of theoretical analyses that already exist.

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.

Forelimb morphology and substrate use in extant Cercopithecidae and the fossil primate community of the Hadar sequence, Ethiopia.

It is suggested that joint architecture of the extant cercopithecid forelimb differentiates terrestrial from arboreal quadrupedal species. Linear dimensions of forelimb joint morphology have also been used to assign fossil species to locomotor categories. However, many primates use a mix of terrestrial and arboreal behaviors, which can be problematic when developing models of behavior reconstruction using morphological variation. The current study uses multivariate analyses to identify morphology related to substrate use in primates, including determination of semiterrestriality. Measurements collected from distal humeri and proximal ulnae of 49 extant cercopithecid primate species were selected based on studies indicating that they could individually predict substrate use. Analyses including one-way analysis of variance, principal components, and discriminant functions were conducted to assess their ability to differentiate between arboreal and terrestrial substrate use. The functions created in these analyses are then applied to data from fossil specimens from the Hadar sequence, Ethiopia, sampling both the Hadar and overlying Busidima Formations, to retrodict possible substrate behavior of fossil monkeys at Hadar through time. As this study is designed to identify function and behavior rather than phylogeny, the taxonomic assignment of the fossil specimens is sometimes uncertain, but substrate behavior can still be inferred. Results suggest that substrate use, including semiterrestrial behavior, in extant and extinct primates can be inferred successfully from multivariate analyses based on joint morphology of the monkey elbow. This study reveals that the ecological distribution of primarily terrestrial fossil primate species of the Hadar sequence is comparable to modern-day communities in habitats similar to those reconstructed for the Hadar members.

The evolution of cranial base and face in Cercopithecoidea and Hominoidea: Modularity and morphological integration.

The evolutionary relationship between the base and face of the cranium is a major topic of interest in primatology. Such areas of the skull possibly respond to different selective pressures. Yet, they are often said to be tightly integrated. In this paper, we analyzed shape variability in the cranial base and the facial complex in Cercopithecoidea and Hominoidea. We used a landmark-based approach to single out the effects of size (evolutionary allometry), morphological integration, modularity, and phylogeny (under Brownian motion) on skull shape variability. Our results demonstrate that the cranial base and the facial complex exhibit different responses to different factors, which produces a little degree of morphological integration between them. Facial shape variation appears primarily influenced by body size and sexual dimorphism, whereas the cranial base is mostly influenced by functional factors. The different adaptations affecting the two modules suggest they are best studied as separate and independent units, and that-at least when dealing with Catarrhines-caution must be posed with the notion of strong cranial integration that is commonly invoked for the evolution of their skull shape.

An anatomical and mechanical analysis of the douc monkey (genus Pygathrix), and its role in understanding the evolution of brachiation.

OBJECTIVES: Pygathrix is an understudied Asian colobine unusual among the Old World monkeys for its use of arm-swinging. Little data exists on the anatomy and mechanics of brachiation in this genus. Here, we consider this colobine to gain insight into the parallel evolution of suspensory behavior in primates. MATERIALS AND METHODS: This study compares axial and appendicular morphological variables of Pygathrix with other Asian colobines. Additionally, to assess the functional consequences of Pygathrix limb anatomy, kinematic and kinetic data during arm-swinging are included to compare the douc monkey to other suspensory primates (Ateles and Hylobates). RESULTS: Compared to more pronograde species, Pygathrix and Nasalis share morphology consistent with suspensory locomotion such as its narrower scapulae and elongated clavicles. More distally, Pygathrix displays a gracile humerus, radius, and ulna, and shorter olecranon process. During suspensory locomotion, Pygathrix, Ateles, and Hylobates all display mechanical convergence in limb loading and movements of the shoulder and elbow, but Pygathrix uses pronated wrist postures that include substantial radial deviation during arm-swinging. DISCUSSION: The adoption of arm-swinging represents a major shift within at least three anthropoid clades and little data exist about its transition. Across species, few mechanical differences are observed during arm-swinging. Apparently, there are limited functional solutions to the challenges associated with moving bimanually below branches, especially in more proximal forelimb regions. Morphological data support this idea that the Pygathrix distal forelimb differs from apes more than its proximal end. These results can inform other studies of ape evolution, the pronograde to orthograde transition, and the convergent ways in which suspensory locomotion evolved in primates.

A new record of the capped langur (Trachypithecus pileatus) in China.

The distribution of the capped langur (Trachypithecus pileatus) in China has become controversial since Shortridge's langur (Trachypithecus shortridgei) was upgraded to a full species. The capped langur is considered to be distributed in northeast India, Bangladesh, Bhutan, and northwest Myanmar only (Brandon-Jones et al., 2004; Choudhury, 2008, 2014; Das et al., 2008; Groves, 2001). In our field survey, however, we obtained photos of the capped langur, demonstrating its existence in China.