Ontogenetic changes in limb bone structural proportions in mountain gorillas (Gorilla beringei beringei)

Behavioral studies indicate that adult mountain gorillas (Gorilla beringei) are the most terrestrial of all nonhuman hominoids, but that infant mountain gorillas are much more arboreal. Here we examine ontogenetic changes in diaphyseal strength and length of the femur, tibia, humerus, radius, and ulna in 30 Virunga mountain gorillas, including 18 immature specimens and 12 adults. Comparisons are also made with 14 adult western lowland gorillas (Gorilla gorilla gorilla), which are known to be more arboreal than adult mountain gorillas. Infant mountain gorillas have significantly stronger forelimbs relative to hind limbs than older juveniles and adults, but are nonsignificantly different from western lowland gorilla adults. The change in inter-limb strength proportions is abrupt at about two years of age, corresponding to the documented transition to committed terrestrial quadrupedalism in mountain gorillas. The one exception is the ulna, which shows a gradual increase in strength relative to the radius and other long bones during development, possibly corresponding to the gradual adoption of stereotypical fully pronated knuckle-walking in older juvenile gorillas. Inter-limb bone length proportions show a contrasting developmental pattern, with hind limb/forelimb length declining rapidly from birth to five months of age, and then showing no consistent change through adulthood. The very early change in length proportions, prior to significant independent locomotion, may be related to the need for relatively long forelimbs for climbing in a large-bodied hominoid. Virunga mountain gorilla older juveniles and adults have equal or longer forelimb relative to hind limb bones than western lowland adults. These findings indicate that both ontogenetically and among closely related species of Gorilla, long bone strength proportions better reflect actual locomotor behavior than bone length proportions.     

Long bone articular and diaphyseal structure in Old World monkeys and apes. II: Estimation of body mass

Body mass estimation equations are generated from long bone cross-sectional diaphyseal and articular surface dimensions in 176 individuals and 12 species of hominoids and cercopithecoids. A series of comparisons is carried out to determine the best body mass predictors for each of several taxonomic/locomotor groupings. Articular breadths are better predictors than articular surface areas, while cross-sectional shaft strengths are better predictors than shaft external breadths. Percent standard errors of estimate (%SEEs) and percent prediction errors for most of the better predictors range between 10-20%. Confidence intervals of equations using sex/species means are fairly representative of those calculated using individual data, except for sex/species means equations with very low %SEEs (under about 10%), where confidence intervals (CIs) based on individuals are likely to be larger. Given individual variability, or biological "error," this may represent a lower limit of precision in estimating individual body masses. In general, it is much more preferable to determine at least broad locomotor affinities, and thus appropriate modern reference groups, before applying body mass estimation equations. However, some structural dimensions are less sensitive to locomotor distinctions than others; for example, proximal tibial articular M-L breadth is apparently "locomotor blind" regarding body mass estimation within the present study sample. In other cases where locomotor affiliation is uncertain, mean estimates from different reference groups can be used, while for some dimensions no estimation should be attempted. The techniques are illustrated by estimating the body masses of four fossil anthropoid specimens of Proconsul nyanzae, Proconsul heseloni, Morotopithecus bishopi, and Theropithecus oswaldi.     

Patterns of joint size dimorphism in the elbow and knee of catarrhine primates

Differences in body size between conspecific sexes may incur differences in the relative size and/or shape of load-bearing joints, potentially confounding our understanding of variation in the fossil record. More specifically, larger males may experience relatively greater limb joint stress levels than females, unless an increase in weight-related forces is compensated for by positive allometry of articular surface areas. This study examines variation in limb joint size dimorphism (JSD) among extant catarrhines to: 1) determine whether taxa exhibit JSD beyond that expected to simply maintain geometric similarity between sexes, and 2) test whether taxa differ in JSD (relative to body size dimorphism) with respect to differences in limb use and/or phylogeny. "Joint size" was quantified for the distal humerus and distal femur of 25 taxa. Analysis of variance was used to test for differences between sexes (in joint size ratios) and among taxa (in patterns of dimorphism). Multiple regression was used to examine differences in JSD among taxa after accounting for variation in body size dimorphism (BSD) and body size. Although degrees of humeral and femoral JSD tend to be the same within species, interspecific variation exists in the extent to which both joints are dimorphic relative to BSD. While most cercopithecoids exhibit relatively high degrees of JSD (i.e., positive allometry), nonhuman hominoids exhibit degrees of JSD closer to isometry. These results may reflect a fundamental distinction between cercopithecoids and hominoids in joint design. Overall, the results make more sense (from a mechanical standpoint) when adjustments to BSD are made to account for the larger effective female body mass associated with bearing offspring. In contrast to other hominoids, modern humans exhibit relatively high JSD in both the knee and elbow (despite lack of forelimb use in weight support). Estimates of BSD based on fossil limb bones will vary according to the extant analogue chosen for comparison.     

Evolution of locomotion in Anthropoidea: the semicircular canal evidence

Our understanding of locomotor evolution in anthropoid primates has been limited to those taxa for which good postcranial fossil material and appropriate modern analogues are available. We report the results of an analysis of semicircular canal size variation in 16 fossil anthropoid species dating from the Late Eocene to the Late Miocene, and use these data to reconstruct evolutionary changes in locomotor adaptations in anthropoid primates over the last 35 Ma. Phylogenetically informed regression analyses of semicircular canal size reveal three important aspects of anthropoid locomotor evolution: (i) the earliest anthropoid primates engaged in relatively slow locomotor behaviours, suggesting that this was the basal anthropoid pattern; (ii) platyrrhines from the Miocene of South America were relatively agile compared with earlier anthropoids; and (iii) while the last common ancestor of cercopithecoids and hominoids likely was relatively slow like earlier stem catarrhines, the results suggest that the basal crown catarrhine may have been a relatively agile animal. The latter scenario would indicate that hominoids of the later Miocene secondarily derived their relatively slow locomotor repertoires.     

Body mass in lowland gorillas: a quantitative analysis

Body proportions and tissue composition (e.g., relative contributions of muscle, skin, bone, and adipose to total body mass) were determined through dissection of four adult captive lowland gorillas. The relative contribution of bone varies little among the four animals (10.2-13.4%) despite considerable range in body weights (99.5-211 kg). In tissue composition, three animals have on average 37.3% muscle relative to body mass. Maximum estimates of body fat range between 19.4-44%. Differences in age, sex, and life history events partially explain the observed variation in body proportions and tissue composition among the four animals. Although gorillas are considered extremely sexually dimorphic in body weight and canine size, differences in tissue are not as dramatic as body mass differences suggest. This study found sex differences mostly in the upper body; males have relatively heavier forelimbs, including heavier deltoid, trunk-binding, and deep back muscles compared to the younger female. The old, obese female had one half the muscle tissue of the other three animals (16% vs. 37.3%), and twice the body fat (44%); forelimbs and upper body musculature were relatively well-developed to compensate for the restricted hip-joint movement due to arthritis. Data on the variation in tissue composition and body proportions in gorillas provide a basis for comparison with other hominoids, including humans. For example, compared to highly dimorphic orangutans, gorillas have more muscle, less adipose tissue, lighter forelimbs and heavier hindlimbs. Such analyses complement studies of the skeleton and contribute to our understanding of human evolution and adaptation.     

Forelimb segment length proportions in extant hominoids and Australopithecus afarensis

Forelimb proportions have been used to infer locomotor adaptation in Australopithecus afarensis. However, little is known about proportions among individual forelimb segments in extant or fossil hominoids. The partial A. afarensis skeleton A.L. 438-1 and the more complete skeleton A.L. 288-1 provide the opportunity to assess relative length of the arm, forearm, wrist, and palm. We compare scaling relationships between pairs of forelimb bones of extant hominoids and A. afarensis, and length of individual forelimb elements to a body size surrogate. Hylobatids, and to a lesser extent orangutans, have the longest forelimb bones relative to size, although the carpus varies little among taxa, perhaps due to functional constraints of the wrist. Pan species are unique in having long metacarpals relative to ulnar length, demonstrating that they probably differ from the common chimp-human ancestor, and also that developmental mechanisms can be altered to results in differential growth of individual forelimb segments. A. afarensis has no forelimb bones that are significantly longer than those of humans for its size. It falls within the range of variation seen in modern humans for all comparisons relative to size, but appears to differ from the typical human brachial index due to a slightly shorter humerus and/or slightly longer ulna. It has short metacarpals like humans only among hominoids. Thus, while Pan may have elongated its metacarpus relative to ulnar length, A. afarensis may have reduced the length of its metacarpals and possibly its humerus relative to body size from the primitive condition.     

Forelimb indicators of prey‐size preference in the Felidae

The forelimbs, along with the crania, are an essential part of the prey‐killing apparatus in cats. Linear morphometrics of the forelimbs were used to determine the morphological differences between felids that specialize on large prey, small prey, or mixed prey. We also compared the scaling of felid forelimbs to those of canids to test whether prey capture strategies affect forelimb scaling. Results suggest that large prey specialists have relatively robust forelimbs when compared with smaller prey specialists. This includes relatively more robust humeri and radii, relatively larger distal ends of the humerus, and relatively larger articular areas of the humerus and radius. Large prey specialists also had relatively longer olecranon processes of the ulna and wider proximal paws. These characters are all important for subduing large prey while the cat positions itself for the killing bite. Small prey specialists have relatively longer distal limb elements for swift prey capture, and mixed prey specialists had intermediate values with relatively more robust metacarpals. Arboreal felids also had more robust limbs. They had relatively longer proximal phalanges for better grip while climbing, and a relatively short brachial index (radius to humerus ratio). Additionally, we found that felids and canids differ in forelimb scaling, which emphasizes the dual use of forelimbs for locomotion and prey capture in felids. This morphometric technique worked well to separate prey‐size preference in felids, but did not work as well to separate locomotor groups, as scansorial and terrestrial felids were not clearly distinguished. J. Morphol. 2009. © 2009 Wiley‐Liss, Inc.     

Morphological correlates of tail length in the catarrhine sacrum

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

Ecological divergence and medial cuneiform morphology in gorillas

Gorillas are more closely related to each other than to any other extant primate and are all terrestrial knuckle-walkers, but taxa differ along a gradient of dietary strategies and the frequency of arboreality in their behavioral repertoire. In this study, we test the hypothesis that medial cuneiform morphology falls on a morphocline in gorillas that tracks function related to hallucial abduction ability and relative frequency of arboreality. This morphocline predicts that western gorillas, being the most arboreal, should display a medial cuneiform anatomy that reflects the greatest hallucial abduction ability, followed by grauer gorillas, and then by mountain gorillas. Using a three-dimensional methodology to measure angles between articular surfaces, relative articular and nonarticular areas, and the curvatures of the hallucial articular surface, the functional predictions are partially confirmed in separating western gorillas from both eastern gorillas. Western gorillas are characterized by a more medially oriented, proportionately larger, and more mediolaterally curved hallucial facet than are eastern gorillas. These characteristics follow the predictions for a more prehensile hallux in western gorillas relative to a more stable, plantigrade hallux in eastern gorillas. The characteristics that distinguish eastern gorilla taxa from one another appear unrelated to hallucial abduction ability or frequency of arboreality. In total, this reexamination of medial cuneiform morphology suggests differentiation between eastern and western gorillas due to a longstanding ecological divergence and more recent and possibly non-adaptive differences between eastern taxa.     

Identification and phyletic distribution of the plasma esterases of some anthropoid primates

Using electrophoresis, specific staining, and selective inhibition, we examined and identified esterase components in the plasma of 13 anthropoid species: 5 hominoids, 7 cercopithecoids, and 1 platyrrhine. Most species, unlike humans, exhibited one or more arylesterase and/or carboxylesterase components as well as cholinesterase and albumin-associated arylesterase. A distinctive arylesterase probably indicates pregnancy in cercopithecoids. While some characters (such as the net charge of the cholinesterase molecule) were extremely uniform across all taxa examined, other quantitative and qualitative traits showed intra- and interspecific variation. Comparison of a cladogram with the distribution of traits shows that the least phyletically stable traits are, in general, those that also exhibit intraspecific polymorphism. One genus-specific, and therefore relatively conservative, trait (weak dimer-dimer bonding in the cholinesterase molecule) evidently appeared independently among papionin cercopithecoids and hominoids. The most parsimonious interpretation of this distribution reinforces the notion of a Pan-Gorilla-Homoclade sororally related to Pongo.In general, plasma esterases are an underexploited source of genetic markers in primates.     

Articular to diaphyseal proportions of human and great ape metatarsals

This study proposes a new way to use metatarsals to identify locomotor behavior of fossil hominins. Metatarsal head articular dimensions and diaphyseal strength in a sample of chimpanzees, gorillas, orangutans, and humans (n = 76) are used to explore the relationships of these parameters with different locomotor modes. Results show that ratios between metatarsal head articular proportions and diaphyseal strength of the hallucal and fifth metatarsal discriminate among extant great apes and humans based on their different locomotor modes. In particular, the hallucal and fifth metatarsal characteristics of humans are functionally related to the different ranges of motion and load patterns during stance phase in the forefoot of humans in bipedal locomotion. This method may be applicable to isolated fossil hominin metatarsals to provide new information relevant to debates regarding the evolution of human bipedal locomotion. The second to fourth metatarsals are not useful in distinguishing among hominoids. Further studies should concentrate on measuring other important qualitative and quantitative differences in the shape of the metatarsal head of hominoids that are not reflected in simple geometric reconstructions of the articulation, and gathering more forefoot kinematic data on great apes to better understand differences in range of motion and loading patterns of the metatarsals. Am J Phys Anthropol 143:198–207, 2010. © 2010 Wiley‐Liss, Inc.     

Patterns of sexual dimorphism in the hominoid distal humerus

Basic biomechanical principles predict that body size differences and differences in the positional behavior of primates should impact on the design of the locomotor skeleton. Allometric distortions in joint shape might be expected between sexes if the degree of body size dimorphism is substantial and/or if sex-specific differences exist in behavior. Nevertheless, there are few documented cases of sexual dimorphism in the limb joints of hominoids, despite substantial body size dimorphism and some reports of intersexual differences in positional behavior. This study re-examines sexual dimorphism in the hominoid distal humerus using coordinate data, and distinguishes explicitly between degree of dimorphism (i.e., the magnitude of intersexual differences) and pattern of dimorphism (i.e. , the nature of these differences). Using a variety of multivariate morphometric methods (e.g., canonical variates analysis of Mosimann shape variables; Euclidean Distance Matrix Analysis of both form and pattern difference matrices), we address the following issues: (1) do males and females of different species and subspecies (or ethnic groups for humans) maintain similar joint shapes? (2) are multiple patterns of dimorphism evident in this region of hominoids? (3) are differences and similarities in degree and pattern predicted by phylogenetic propinquity and positional behavior? For the most part, our results support earlier findings that sexual dimorphism in the shape of the anthropoid elbow is slight. Of the eight taxa considered here, only the western lowland gorillas exhibited significant differences in the shape of the distal humerus. Gorilla gorilla gorilla also displays a significantly different pattern of dimorphism from the orang-utan. Pattern differences between Andaman Islanders and both mountain gorillas and the orang-utan also approach statistical significance (P<0.06 and P<0.08, respectively). Overall, and despite marked differences in the degree of dimorphism, the knuckle-walking African apes are more similar in patterns of dimorphism to each other than to other taxa (e.g., gorillas are more similar to orang-utans in degree, but more similar to chimpanzees and bonobos in pattern). We could find no definitive "human pattern" in our results and suspect that this is because human upper limbs face less stringent mechanical constraints since they are relieved of locomotor stresses (but we cannot rule out the possibility of undocumented differences among our human groups in sex-specific, work-related activities). We anticipate finding additional pattern differences among anthropoids in articular dimorphism as we add other taxa to our sample (including fossil hominids), and examine other joint systems.     

The humerus of Aegyptopithecus zeuxis: a primitive anthropoid

Two complete humeri of Aegyptopithecus zeuxis have been recovered from Oligocene deposits in the Fayum Province of Egypt. These new specimens support previous interpretations of the locomotor adaptations of this species and indicate that A. zeuxis was a robust, slowly moving arboreal quadruped. While the previously described distal articular region of the humerus is virtually identical with the same region in many extant ceboids and the Miocene hominoid Pliopithecus vindobonensis, the more proximal parts of the humerus show many primitive "prosimianlike" features not found the limbs of extant anthropoids. The primitive features include the absence of a distinct deltoid plane, a broad shallow bicipital groove, a large brachialis flange, and an entepicondylar foramen. In most features, the humerus of Aegyptopithecus zeuxis is more primitive than the hypothetical last common ancestor of extant cercopithecoids and hominoids based on neontological comparisons. This supports other lines of evidence indicating that the hominoids from the Egyptian Oligocene are morphologically ancestral to both Old World monkeys and apes.     

Partial skeleton of Proconsul nyanzae from Mfangano Island,Kenya

A partial skeleton attributed to Proconsul nyanzae (KNM-MW 13142) is described. The fossils were found at a site on Mfangano Island, Kenya, which dates to 17.9 ± .1 million years ago. KNM-MW 13142 consists of six partial vertebrae (T12-S1), a nearly complete hipbone, most of the right femur and left femoral shaft, a fragmentary tibia and fibula, and a nearly complete talus and calcaneus. This skeleton provides the first pelvic fossil known for any East African Miocene hominoid. The new Proconsul specimen is compared to a large sample of extant anthropoids to determine its functional and phylogenetic affinities. In most aspects of its anatomy, KNM-MW 13142 closely resembles nonhominoid anthropoids. This individual had a long, flexible spine, narrow torso, and habitually pronograde posture, features characteristic of most extant monkeys. Evidence of spinal musculature suggests a generalized condition intermediate between that of cercopithecoids and hylobatids. The hindlimb of KNM-MW 13142 exhibits relatively mobile hip and ankle joints, with structural properties of the femur like those of hominoids. This mix of features implies a pattern of posture and locomotion that is unlike that of any extant primate. Many aspects of the Proconsul nyanzae locomotor skeleton may represent the primitive catarrhine condition. © 1993 Wiley-Liss, Inc.     

Relative strength of the tibia and fibula and locomotor behavior in hominoids

The fibula has rarely been considered in comparative morphological studies, probably due to its relatively minor role in carrying mechanical loads. However, some differences in morphology (and inferred function) of the fibula between humans and apes, and within apes, have been noted and related to differences in positional behavior. Therefore, the study of tibiofibular relations may be useful in characterizing such differences. This study examines cross-sectional geometric (CSG) properties (cortical area and polar section modulus, Z(p)) of the tibia and fibula at mid-diaphysis across a sample (n=87) of humans, chimpanzees, gorillas, orangutans, and gibbons. The fibula is compared against the tibia in the different taxa. The results indicate that the robusticity of the fibula relative to that of the tibia can be explained in terms of differences in positional behavior. In particular, hominoids that are more arboreal (i.e., gibbons, orangutans, and chimpanzees) possess a relatively more robust fibula than do hominoids that are more terrestrial (i.e., gorillas and humans). The difference appears to be a consequence of the more mobile fibula and more adducted position of the hindlimb necessary in an arboreal environment. Apart from providing the first CSG data on the fibula, these results may be helpful in reconstructing the locomotor behavior of fossil hominoids.     

The effects of locomotion on the structural characteristics of avian limb bones

Despite the wide range of locomotor adaptations in birds, little detailed attention has been given to the relationships between the quantitative structural characteristics of avian limb bones and bird behaviour. Possible differences in forelimb relative to hindlimb strength across species have been especially neglected. We generated cross‐sectional, geometric data from peripheral quantitative computed tomography scans of the humerus and femur of 127 avian skeletons, representing 15 species of extant birds in 13 families. The sample includes terrestrial runners, arboreal perchers, hindlimb‐propelled divers, forelimb‐propelled divers and dynamic soarers. The hindlimb‐propelled diving class includes a recently flightless island form. Our results demonstrate that locomotor dynamics can be differentiated in most cases based on cross‐sectional properties, and that structural proportions are often more informative than bone length proportions for determining behaviour and locomotion. Recently flightless forms, for example, are more easily distinguished using structural ratios than using length ratios. A proper phylogenetic context is important for correctly interpreting structural characteristics, especially for recently flightless forms. Some of the most extreme adaptations to mechanical loading are seen in aquatic forms. Penguins have forelimbs adapted to very high loads. Aquatic species differ from non‐aquatic species on the basis of relative cortical thickness. The combination of bone structural strength and relative cortical area of the humerus successfully differentiates all of our locomotor groups. The methods used in this study are highly applicable to fossil taxa, for which morphology is known but behaviour is not. The use of bone structural characteristics is particularly useful in palaeontology not only because it generates strong signals for many locomotor guilds, but also because analysing such traits does not require knowledge of body mass, which can be difficult to estimate reliably for fossil taxa. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 153 , 601–624.     

Patterns of correlation and covariation of anthropoid distal forelimb segments correspond to Hoxd expression territories

Anthropoids in general and hominoids in particular exhibit differential adaptations in forearm and digital skeletal proportions to a diverse array of locomotor modes. Hox genes act as selector genes with spatially regulated expression patterns during development. Their expression in the forelimb appears to define modules that specify differential skeletal growth. Here we explore forelimb skeletal proportions in a large sample of anthropoids from a background provided by Hoxd expression patterns in late-stage murine embryonic forelimbs. Interspecific correlation and principal components analyses of primate forelimb data indicate that morphological variation in anthropoids reflects well-defined developmental modules downstream of Hoxd expression. The phalanges of digit one appear to represent a single growth module, whereas the metacarpals and manual phalanges of the posterior digits correspond to a second, independent, expression territory that extends proximally into the distal zeugopod. In particular, hominoids show very high correlations among the posterior digits and the independence of digit one. In addition, the distal radius is generally highly correlated with the posterior digits and not digit one. Relying on established functional differences among Hox paralogs, we present a model that parsimoniously explains hominoid forearm and digital proportions as a consequence of downstream effects of Hox. We, therefore, suggest that Hox-defined developmental modules have served as evolutionary modules during manual evolution in anthropoids.     

Relative growth,ontogeny, and sexual dimorphism in gorilla(Gorilla gorilla gorilla and G. g. beringei): Evolutionary and ecological considerations

Gorillas are the largest and among the most sexually dimorphic of all extant primates. While gorillas have been incorporated in broad-level comparisons among large-bodied hominoids or in studies of the African apes, comparisons between gorilla subspecies have been rare. During the past decade, however, behavioral, morphological, and molecular data from a number of studies have indicated that the western lowland (Gorilla gorilla gorilla) and eastern mountain (Gorilla gorilla beringei) subspecies differ to a greater extent than has been previously believed. In this study I compare patterns of relative growth of the postcranial skeleton to evaluate whether differences between subspecies result from the differential extension of common patterns of relative growth. In addition, patterns of ontogeny and sexual dimorphism are also examined. Linear skeletal dimensions and skeletal weight were obtained for ontogenetic series of male and female G.g. gorilla (n = 315) and G.g. beringei (n = 38). Bivariate and multivariate methods of analysis were used to test for differences in patterns of relative growth, ontogeny, and sexual dimorphism between sexes of each subspecies and in same-sex comparisons between subspecies. Results indicate males and females of both subspecies are ontogenetically scaled for postcranial proportions and that females undergo an earlier skeletal growth spurt compared to males. However, results also indicate that the onset of the female growth spurt occurs at different dental stages in lowland and mountain gorillas and that mountain gorillas may be characterized by higher rates of growth. Finally, data demonstrate lowland and mountain gorilla females do not differ significantly in adult body size, but mountain gorilla males are significantly larger than lowland gorilla males, suggesting mountain gorillas are characterized by a higher degree of sexual dimorphism in body size. Thus, although lowland and mountain gorillas do not appear to have evolved novel adaptations of the postcranium which correlate with differences in locomotor behavior, the present investigation establishes subspecies differences in ontogeny and sexual dimorphism which may be linked with ecological variation. Specifically, these findings are evaluated in the context of risk aversion models which predict higher growth rates and increased levels of sexual dimorphism in extreme folivores. Am. J. Primatol. 43:1–31, 1997. © 1997 Wiley-Liss, Inc.     

Morphology of the hallucial phalanges in extant anthropoids and fossil hominoids

Pedal phalanges of living anthropoids and several Miocene fossil hominoid taxa were studied to reveal functional adaptations of living anthropoid feet and to infer positional behavior of fossil hominoids. Among the examined living anthropoids, Pan has a very developed (long and robust) hallux. Proconsul and Nacholapithecus, a large hominoid from Nachola, northern Kenya, display a moderately long hallux like Alouatta and Cebus, suggesting the well-developed capability of a hallux-assisted power grip. Allometric analyses revealed that the Miocene hominoids examined (mainly from East Africa) as a whole displayed a different scaling pattern about the width of the proximal articular surface of the hallucial terminal phalanx from that of living anthropoids. Larger-sized hominoids display a wider articular surface than comparable-sized living anthropoids while smaller-sized fossil hominoids do the reverse. Such a difference was less marked for the height of the articular surface. These results may suggest that positional adaptations of Miocene hominoids are not merely resultants of a common body size function that is observed in living anthropods. The wide articular surface of fossil hominoid hallucial terminal phalanges suggests an adaptation for vertical climbing and clinging, in which the hallux is kept perpendicularly to the long axis of the vertical support.     

Articular morphology of the proximal ulna in extant and fossil hominoids and hominins

Extant hominoids share similar elbow joint morphology, which is believed to be an adaptation for elbow stability through a wide range of pronation-supination and flexion-extension postures. Mild variations in elbow joint morphology reported among extant hominoids are often qualitative, where orangutans are described as having keeled joints, and humans and gorillas as having flatter joints. Although these differences in keeling are often linked to variation in upper limb use or loading, they have not been specifically quantified. Many of the muscles important in arboreal locomotion in hominoids (i.e., wrist and finger flexors and extensors) take their origins from the humeral epicondyles. Contractions of these muscles generate transverse forces across the elbow, which are resisted mainly by the keel of the humeroulnar joint. Therefore, species with well-developed forearm musculature, like arboreal hominoids, should have more elbow joint keeling than nonarboreal species. This paper explores the three- and two-dimensional morphology of the trochlear notch of the elbow of extant hominoids and fossil hominins and hominoids for which the locomotor habitus is still debated. As expected, the elbow articulation of habitually arboreal extant apes is more keeled than that of humans. In addition, extant knuckle-walkers are characterized by joints that are distally expanded in order to provide greater articular surface area perpendicular to the large loads incurred during terrestrial locomotion with an extended forearm. Oreopithecus is characterized by a pronounced keel of the trochlear notch and resembles Pongo and Pan. OH 36 has a morphology that is unlike that of extant species or other fossil hominins. All other hominin fossils included in this study have trochlear notches intermediate in form between Homo and Gorilla or Pan, suggesting a muscularity that is less than in African apes but greater than in humans.