Intraspecific variation of long bone cross-sectional properties in Pan troglodytes troglodytes and Gorilla gorilla gorilla.

OBJECTIVES: Morphological intraspecific variation is due to the balance between skeletal plasticity and genetic constraint on the skeleton. Osteogenic responses to external stimuli, such as locomotion, have been well documented interspecifically across the primate order, but less so at the intraspecific level. Here, we examine the differences in cross-sectional variability of the femur, humerus, radius, and tibia in Pan troglodytes troglodytes versus Gorilla gorilla gorilla. We investigate whether there are sex, species, bone, and trait differences in response to variable body size and locomotion. MATERIALS AND METHODS: Adult male and female P. t. troglodytes and G. g. gorilla long bones from the Cleveland Museum of Natural History were scanned with a peripheral quantitative computer tomography system. Scans were taken at the midshaft of each bone according to functional bone length. Coefficients of variation were used to provide a size-independent measure of variation. We applied a Bonferroni correction to account for the multiple pairwise tests. RESULTS: There were limited significant differences between males and females, however, females tended to be more variable than males. Variation in Gorilla, when significant, was greater than in Pan, although significant differences were limited. There were no differences between bone variability in male and female Gorilla, and female Pan. DISCUSSION: Increased female variability may be due to more variable locomotor behavior, particularly during periods of pregnancy, lactation, and caring for an offspring compared to consistent locomotion over the life course by males. Body size may be a contributing factor to variability; more work is needed to understand this relationship.

Oldest evidence of abundant C4 grasses and habitat heterogeneity in eastern Africa.

The assembly of Africa's iconic C4 grassland ecosystems is central to evolutionary interpretations of many mammal lineages, including hominins. C4 grasses are thought to have become ecologically dominant in Africa only after 10 million years ago (Ma). However, paleobotanical records older than 10 Ma are sparse, limiting assessment of the timing and nature of C4 biomass expansion. This study uses a multiproxy design to document vegetation structure from nine Early Miocene mammal site complexes across eastern Africa. Results demonstrate that between ~21 and 16 Ma, C4 grasses were locally abundant, contributing to heterogeneous habitats ranging from forests to wooded grasslands. These data push back the oldest evidence of C4 grass-dominated habitats in Africa-and globally-by more than 10 million years, calling for revised paleoecological interpretations of mammalian evolution.

The evolution of hominoid locomotor versatility: Evidence from Moroto, a 21 Ma site in Uganda.

Living hominoids are distinguished by upright torsos and versatile locomotion. It is hypothesized that these features evolved for feeding on fruit from terminal branches in forests. To investigate the evolutionary context of hominoid adaptive origins, we analyzed multiple paleoenvironmental proxies in conjunction with hominoid fossils from the Moroto II site in Uganda. The data indicate seasonally dry woodlands with the earliest evidence of abundant C4 grasses in Africa based on a confirmed age of 21 million years ago (Ma). We demonstrate that the leaf-eating hominoid Morotopithecus consumed water-stressed vegetation, and postcrania from the site indicate ape-like locomotor adaptations. These findings suggest that the origin of hominoid locomotor versatility is associated with foraging on leaves in heterogeneous, open woodlands rather than forests.

A chimpanzee enamel-diet δ13C enrichment factor and a refined enamel sampling strategy: Implications for dietary reconstructions.

Reconstructing diets from stable carbon isotopic signals in enamel bioapatite requires the application of a δ13C enamel-diet enrichment factor, or the isotopic offset between diet and enamel, which has not been empirically determined for any primate. In this study, an enamel-diet enrichment factor (ε∗enamel-diet) of 11.8 ± 0.3‰ is calculated for chimpanzees (Pan troglodytes) at Ngogo in Kibale National Park, Uganda, based on a comprehensive isotopic assessment of previously analyzed dietary plant data and new isotopic analyses of enamel apatite. Different enamel sampling methods are evaluated to determine the potential influence of weaning on isotopic enamel values and dietary interpretations. The new chimpanzee enrichment factor and a sampling strategy that excludes teeth that formed before weaning completion are applied to all known chimpanzee δ13Cenamel data, either previously published or newly derived in this study, resulting in a dietary range of almost 6‰ across all chimpanzees sampled. This new chimpanzee enamel-diet enrichment factor is then used to reassess dietary reconstructions of 12 fossil hominin species whose isotopic enamel signatures have been determined. Results reveal hominin diets that are isotopically more positive than previously reconstructed, highlighting the widespread contribution of 13C-enriched C4/crassulacean acid metabolism (CAM) resources in fossil hominin diets and emphasizing the broad use of these resources during human evolution. These findings stress the importance of ascertaining and employing an appropriate enrichment factor for dietary reconstructions of specific taxa as well as standardizing the sampling protocol for tooth enamel in isotopic paleodietary reconstructions.

New hominoid fossils from Moroto II, Uganda and their bearing on the taxonomic and adaptive status of Morotopithecus bishopi.

The early Miocene site of Moroto II, Uganda has yielded some of the oldest known hominoid fossils. A new partial mandible (UMP MORII 03'551) is notable for its long tooth row and large, narrow M2 with well-developed cristids - a morphological combination previously unknown for large bodied catarrhines of the Early Miocene and suggesting folivory. The tooth proportions are compatible with belonging to the same taxon as the maxilla UMP 62-11, the holotype of Morotopithecus bishopi; likewise, the long tooth row and vertical planum of UMP MORII 03'551 suggest that it may represent the same taxon as mandible(s) UMP 66-01 and UMP 62-10. Canine size strongly suggests UMP MORII 03'551 is a female. Comparisons of the tooth crown morphology and tooth row proportions, relative enamel thickness, enamel-dentine junction morphology, long-period line periodicity, and dental wear patterns support significant morphological, developmental, and inferred dietary differentiation, and therefore generic-level distinctiveness, among Afropithecus, Morotopithecus and the Proconsul clade. An isolated M1 (UMP MORII 03'559) is morphologically dissimilar, and much smaller than the actual or inferred size of molars in UMP MORII 03'551, UMP 66-01 and UMP 62-10, supporting the presence of two hominoid taxa at Moroto II, M. bishopi and a smaller bodied proconsulid. Given the high level of body mass dimorphism inferred for Morotopithecus and other early Miocene catarrhines, the known postcrania from Moroto II could be attributable to either taxon. However, UMP MORII 03'551 and the femora UMP MORII 94'80 derive from the same stratigraphic interval, while the isolated M1 was deposited later, increasing the likelihood that the mandible and femora are from the same individual. These new fossils expand our understanding of the taxonomic and adaptive diversity of early Miocene catarrhines.

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.

Evidence for rapid faunal change in the early Miocene of East Africa based on revised biostratigraphic and radiometric dating of Bukwa, Uganda.

Field expeditions to Bukwa in the late 1960s and early 1970s established that the site had a small but diverse early Miocene fauna, including the catarrhine primate Limnopithecus legetet. Initial potassium-argon radiometric dating indicated that Bukwa was 22 Ma, making it the oldest of the East African early Miocene fossil localities known at the time. In contrast, the fauna collected from Bukwa was similar to other fossil localities in the region that were several million years younger. This discrepancy was never resolved, and due to the paucity of primate remains at the site, little subsequent research took place. We have collected new fossils at Bukwa, reanalyzed the existing fossil collections, and provided new radiometric dating. 40Ar/39Ar incremental heating ages on lavas bracketing the site indicate that the Bukwa fossils were deposited ∼19 Ma, roughly 3 Ma younger than the original radiometric age. Our radiometric dating results are corroborated by a thorough reanalysis of the faunal assemblage. Bukwa shares taxa with both stratigraphically older localities (Tinderet, Napak) and with stratigraphically younger localities (Kisingiri, Turkana Basin) perfectly corresponding to our revised radiometric age. This revised age for Bukwa is important because it indicates that significant faunal turnover may have occurred in East Africa between 20 and 19 Ma. Bukwa samples immigrant taxa such as large suids, large ruminants, and ochotonids that are absent from stratigraphically older but well-sampled localities in the region, such as Tinderet (∼20 Ma) and Napak (20 Ma). Further age refinements for Bukwa and the entire East African early Miocene sequence will help to constrain the timing of this faunal turnover event, of particular importance in paleoanthropology since this temporal sequence also provides us with what is currently our best window into the early evolution of cercopithecoid and hominoid primates.

Long bone cross-sectional properties reflect changes in locomotor behavior in developing chimpanzees.

OBJECTIVES: Recent studies indicate that the locomotor behavior of wild chimpanzees changes during development. Before transitioning to quadrupedal knuckle-walking in adulthood, young chimpanzees engage in a significant amount of upper limb loading suspensory behavior. We investigated whether these dramatic changes in locomotion influence the strength and shape of chimpanzee long bones. MATERIALS AND METHODS: We examined changes in chimpanzee arboreal locomotion over the course of development using behavioral data collected on wild chimpanzees. We measured the midshaft geometric properties of femora and humeri of wild-caught individuals housed in museum collections using micro computed tomographic scans. RESULTS: Chimpanzees spent less time moving arboreally as they aged. Femoral/humeral strength ratios also increased with age, as predicted by the changing loading environment during development. Additional analyses revealed that femoral shape, but not humeral shape, varied across chimpanzee age classes. Adult femora were more elliptical compared with those of infants. This change in adult femora is consistent with the observation that adult chimpanzees spend most of their time moving terrestrially and consequently experience a less variable loading environment than do infants. DISCUSSION: Taken together, these findings contribute to our understanding of how ontogenetic changes in function affect form. As similar changes may have characterized the behavioral and skeletal ontogeny of extinct hominoids including hominins, these findings furnish a potential means to make inferences about the behavior of fossil taxa based on the structural properties of their bones.

Dentognathic remains of an Afropithecus individual from Kalodirr, Kenya.

We describe here the well-preserved dentognathic remains of an Afropithecus individual from the early Miocene site of Kalodirr in northern Kenya. The specimen includes a nearly complete dentition in which most of the crowns are undamaged and unworn. The new information gleaned from this specimen adds to our knowledge of this genus in several ways. Afropithecus exhibits an atypical pattern of canine dimorphism, and is probably more easily sexed by the pattern of variation in its upper premolars. Both phenomena are likely related to the modification of its antemolar dentition for the purpose of sclerocarp harvesting. The new fossils clarify the role in this adaptation of the premolars, which appear specialized for initiating and propagating cracks in large food items bearing protective coats. Comparison with other Miocene apes suggests that the closest known relative of Afropithecus is Nacholapithecus, to which Equatorius is more distantly related. Morotopithecus shares some primitive traits with Afropithecus, but lacks the derived features shared by the latter and Nacholapithecus.

Lumbar vertebral body bone microstructural scaling in small to medium-sized strepsirhines.

Bone mass, architecture, and tissue mineral density contribute to bone strength. As body mass (BM) increases any one or combination of these properties could change to maintain structural integrity. To better understand the structural origins of vertebral fragility and gain insight into the mechanisms that govern bone adaptation, we conducted an integrative analysis of bone mass and microarchitecture in the last lumbar vertebral body from nine strepsirhine species, ranging in size from 42 g (Microcebus rufus) to 2,440 g (Eulemur macaco). Bone mass and architecture were assessed via µCT for the whole body and spherical volumes of interest (VOI). Allometric equations were estimated and compared with predictions for geometric scaling, assuming axial compression as the dominant loading regime. Bone mass, microarchitectural, and vertebral body geometric variables predominantly scaled isometrically. Among structural variables, the degree of anisotropy (Tb.DA) was the only parameter independent of BM and other trabecular architectural variables. Tb.DA was related to positional behavior. Orthograde primates had higher average Tb.DA (1.60) and more craniocaudally oriented trabeculae while lorisines had the lowest Tb.DA (1.25), as well as variably oriented trabeculae. Finally, lorisines had the highest ratio of trabecular bone volume to cortical shell volume (∼3x) and while there appears to be flexibility in this ratio, the total bone volume (trabecular + cortical) scales isometrically (BM(1.23) , r(2) = 0.93) and appears tightly constrained. The common pattern of isometry in our measurements leaves open the question of how vertebral bodies in strepsirhine species compensate for increased BM.

New Oligocene primate from Saudi Arabia and the divergence of apes and Old World monkeys.

It is widely understood that Hominoidea (apes and humans) and Cercopithecoidea (Old World monkeys) have a common ancestry as Catarrhini deeply rooted in Afro-Arabia. The oldest stem Catarrhini in the fossil record are Propliopithecoidea, known from the late Eocene to early Oligocene epochs (roughly 35-30 Myr ago) of Egypt, Oman and possibly Angola. Genome-based estimates for divergence of hominoids and cercopithecoids range into the early Oligocene; however, the mid-to-late Oligocene interval from 30 to 23 Myr ago has yielded little fossil evidence documenting the morphology of the last common ancestor of hominoids and cercopithecoids, the timing of their divergence, or the relationship of early stem and crown catarrhines. Here we describe the partial cranium of a new medium-sized (about 15-20 kg) fossil catarrhine, Saadanius hijazensis, dated to 29-28 Myr ago. Comparative anatomy and cladistic analysis shows that Saadanius is an advanced stem catarrhine close to the base of the hominoid-cercopithecoid clade. Saadanius is important for assessing competing hypotheses about the ancestral morphotype for crown catarrhines, early catarrhine phylogeny and the age of hominoid-cercopithecoid divergence. Saadanius has a tubular ectotympanic but lacks synapomorphies of either group of crown Catarrhini, and we infer that the hominoid-cercopithecoid split happened later, between 29-28 and 24 Myr ago.

Early Miocene hippopotamids (Cetartiodactyla) constrain the phylogenetic and spatiotemporal settings of hippopotamid origin.

The affinities of the Hippopotamidae are at the core of the phylogeny of Cetartiodactyla (even-toed mammals: cetaceans, ruminants, camels, suoids, and hippos). Molecular phylogenies support Cetacea as sister group of the Hippopotamidae, implying a long ghost lineage between the earliest cetaceans (approximately 53 Ma) and the earliest hippopotamids (approximately 16 Ma). Morphological studies have proposed two different sister taxa for hippopotamids: suoids (notably palaeochoerids) or anthracotheriids. Evaluating these phylogenetic hypotheses requires substantiating the poorly known early history of the Hippopotamidae. Here, we undertake an original morphological phylogenetic analysis including several "suiform" families and previously unexamined early Miocene taxa to test previous conflicting hypotheses. According to our results, Morotochoerus ugandensis and Kulutherium rusingensis, until now regarded as the sole African palaeochoerid and the sole African bunodont anthracotheriid, respectively, are unambiguously included within the Hippopotamidae. They are the earliest known hippopotamids and set the family fossil record back to the early Miocene (approximately 21 Ma). The analysis reveals that hippopotamids displayed an unsuspected taxonomic and body size diversity and remained restricted to Africa during most of their history, until the latest Miocene. Our results also confirm the deep nesting of Hippopotamidae within the paraphyletic Anthracotheriidae; this finding allows us to reconstruct the sequence of dental innovations that links advanced selenodont anthracotheriids to hippopotamids, previously a source of major disagreements on hippopotamid origins. The analysis demonstrates a close relationship between Eocene choeropotamids and anthracotheriids, a relationship that potentially fills the evolutionary gap between earliest hippopotamids and cetaceans implied by molecular analyses.

Specimen size and porosity can introduce error into microCT-based tissue mineral density measurements.

The accurate measurement of tissue mineral density, rho(m), in specimens of unequal size or quantities of bone mineral using polychromatic microCT systems is important, since studies often compare samples with a range of sizes and bone densities. We assessed the influence of object size on microCT measurements of rho(m) using (1) hydroxyapatite rods (HA), (2) precision-manufactured aluminum foams (AL) simulating trabecular bone structure, and (3) bovine cortical bone cubes (BCt). Two beam-hardening correction (BHC) algorithms, determined using a 200 and 1200 mg/cm(3) HA wedge phantom, were used to calculate rho(m) of the HA and BCt. The 200 mg/cm(3) and an aluminum BHC algorithm were used to calculate the linear attenuation coefficients of the AL foams. Equivalent rho(m) measurements of 500, 1000, and 1500 mg HA/cm(3) rods decreased (r(2)>0.96, p<0.05 for all) as HA rod diameter increased in the 200 mg/cm(3) BHC data. Errors averaged 8.2% across these samples and reached as high as 29.5%. Regression analyses suggested no size effects in the 1200 mg/cm(3) BHC data but differences between successive sizes still reached as high as 13%. The linear attenuation coefficients of the AL foams increased up to approximately 6% with increasing volume fractions (r(2)>0.81, p<0.05 for all) but the strength of the size-related error was also BHC dependent. Equivalent rho(m) values were inversely correlated with BCt cube size (r(2)>0.92, p<0.05). Use of the 1200 mg/cm(3) BHC ameliorated the size-related artifact compared to the 200 mg/cm(3) BHC but errors with this BHC were still significant and ranged between 5% and 12%. These results demonstrate that object size, structure, and BHC algorithm can influence microCT measurements of rho(m). Measurements of rho(m) of specimens of unequal size or quantities of bone mineral must be interpreted with caution unless appropriate steps are taken to minimize these potential artifacts.

A fossil hominoid proximal femur from Kikorongo Crater, southwestern Uganda.

The external morphology of a fragmentary right proximal femur from southwestern Uganda is described here. Discovered in the Kikorongo Crater of Queen Elizabeth National Park in 1961, this specimen was informally assigned to Homo sapiens (although never described) and tentatively dated to the late Pleistocene. However, because aspects of the external morphology of the femur align the fossil with the African great apes, we suggest that the Kikorongo femur may be the first postcranial fossil of the genus Pan. Like the African apes, the Kikorongo specimen lacks both an obturator externus groove and an intertrochanteric line. It has a short femoral neck with a circular cross section, and a narrow and deep superior notch. Using resampling statistics and discriminant function analysis, the Kikorongo femur clustered with the genus Pan, as opposed to Gorilla or Homo. However, if the specimen is from Pan, it would be large for this taxon. Furthermore, features that clearly distinguish the external morphology of Plio-Pleistocene hominin proximal femora from African ape femora, such as the shape of the femoral neck in cross section and femoral neck length, have converged in Holocene humans and African apes. Unfortunately, the internal morphology of the femoral neck of the Kikorongo fossil was not discernable. Although we hypothesize that the Kikorongo femur is from the genus Pan, there is such variability in the proximal femora of modern humans that, although it would be an unusual human, it remains possible that this fossil represents H. sapiens.

A new pliopithecoid genus from the early Miocene of Uganda.

A partial face and mandible from the early Miocene site of Napak IX in Uganda are described here as a new genus and species of catarrhine primate, Lomorupithecus harrisoni gen. et sp. nov. The face is among the most complete specimens known for a Miocene small-bodied catarrhine. Several aspects of its anatomy indicate that the new species is a stem catarrhine, and as such, it may provide valuable information pertaining to the primitive catarrhine cranial morphotype. Lomorupithecus is most similar in its facial anatomy to members of the Pliopithecoidea, and these similarities could be interpreted in three ways. They could be symplesiomorphies, which would support the traditional view of the primitive catarrhine cranial morphotype; they could be synapomorphies reflecting a phylogenetic position of Lomorupithecus within Pliopithecoidea; or they could represent convergence. Phylogenetic analysis of Lomorupithecus along with 35 other primates indicates that it is a pliopithecoid. As such, it would be the oldest and only Afro-Arabian member of this otherwise Eurasian clade.

The phylogenetic position of Morotopithecus.

The phylogenetic relationship of the Ugandan Miocene hominoid Morotopithecus bishopi to fossil and living hominoids remains to be determined. In a cladistic approach to this question, we used three published Miocene character sets as the basis of a phylogenetic analysis: J. Hum. Evol. 29 (1995) 101; Function, Phylogeny, and Fossils: Miocene Hominoid Evolution and Adaptations, 1997, 389. Because these datasets often describe the same anatomy using different characters and states, three different datasets were created to reflect these alternatives. In addition, new postcranial characters describable in Morotopithecus were added to each of the above datasets and a fourth dataset was created using only postcranial characters. The most parsimonious tree(s) recovered in all analyses consistently placed Morotopithecus as a sister taxon to the extant great apes, with Hylobates sister to this clade. Morotopithecus was also consistently more derived than Proconsul, Afropithecus, and Kenyapithecus (as defined prior to the description of Equatorius), but less derived than Oreopithecus, Sivapithecus (only craniodentally) and Dryopithecus. These results imply that Morotopithecus is more derived than Hylobates. However, gibbons are believed to have branched off by at least 18 Ma while Morotopithecus is dated at >20.6 Ma. Possible explanations include: (1) the dating of the Morotopithecus material is too old; (2) the Hylobates divergence time has been underestimated; (3) the great ape condition, and not that of Hylobates, is primitive for hominoids; (4) the similarities of Morotopithecus and great apes are homoplasies. Given current evidence, the first possibility is unlikely, but it is not possible to choose definitively between the latter three possibilities. This conclusion is supported by the fact that despite the consistencies of the analyses, the addition of Morotopithecus and the use of different characters had a large effect on the placement of other Miocene taxa. This raises questions as to the robustness of the connections between Miocene taxa and extant hominoids since different results can be achieved by changing either a few characters, or by adding a single taxon. Many of the characters used to estimate phylogeny may need to be reassessed before a reliable assessment of the phylogenetic position of Morotopithecus can be achieved.

Taxonomic affinities of the Eppelsheim femur.

The taxonomic affinities of the Eppelsheim femur, known as Paidopithex, have been unclear for more than a century. Over the years, due to similarities with Pliopithecus, some authors have considered it a large pliopithecid, while others refer to it as Dryopithecus. The issue could not be resolved, because no definitive Dryopithecus femora were available. With the discovery of the Dryopithecus laietanus skeleton from Can Llobateres (CLl 18800), it has become possible to test the attribution of the Eppelsheim femur to Dryopithecus on the basis of direct morphological and metrical comparisons. By means of allometric techniques, we show that the Eppelsheim and D. laietanus femora fit different hindlimb morphologies with regard to relative length and relative head/neck size, with Paidopithex significantly differing from Dryopithecus, but more closely resembling Pliopithecus. Paidopithex also differs from Dryopithecus in other important aspects, such as its lower neck/shaft angle, lack of elevation of the femoral head above the greater trochanter, more posteriorly oriented lesser trochanter, and proximal shaft diameter thicker anteroposteriorly than mediolaterally. In these features, Paidopithex most closely resembles Pliopithecus in spite of differences in body mass (ca. 22 kg vs. ca. 10 kg, respectively). These features suggest that Paidopithex used a primitive locomotor pattern associated with arboreal quadrupedalism, instead of the more derived pattern displayed by Dryopithecus. Currently available evidence confirms that the attribution of Paidopithex to Dryopithecus can be rejected. Paidopithex could be a large and otherwise unknown pliopithecid, but the possibility cannot be ruled out that it represents a third kind of catarrhine.

A comparison of the femoral head and neck trabecular architecture of Galago and Perodicticus using micro-computed tomography (microCT).

Innovations in micro-computed tomography (microCT) in the medical field have resulted in the development of techniques that allow the precise quantification of bone density and fabric related parameters of trabecular bone. For the purpose of this study, the technique was applied to a small sample of Perodicticus potto and Galago senegalensis femora to see if differences in loading environment elicit the predicted effects on trabecular structure. While the overall bone volume was approximately three times larger in the potto, there was no significant difference in the apparent volume density in the two taxa. When regional differences in the proximal femur were examined, the cancellous bone of the femoral head of Perodicticus potto and Galago senegalensis, while not differing in volume density, showed differences in trabecular orientation, with the potto having more randomly oriented trabeculae than the bushbaby. This was as hypothesized, given that the bushbaby submits its femora to more stereotypical loading environments than the potto. In the femoral neck, the cancellous bone was not only more randomly oriented, it was also denser in the potto compared with the bushbaby. This suggests that trabecular morphology may be extremely sensitive to certain differences in the loading environment and that this information, combined with information on cortical bone structure and external geometry, will result in a more complete understanding of how bone shape and composition correspond to loading and locomotor patterns. Ultimately, a synthesis of these different lines of evidence may have considerable applications in paleontological studies that attempt to reconstruct bone use from morphology.

New Sivapithecus postcranial specimens from the Siwaliks of Pakistan.

Several new postcranial elements of Sivapithecus from the Siwaliks of Pakistan are described. These include a distal femur from the U-level of the Dhok Pathan Formation, a navicular from the Chinji Formation, and seven manual and pedal phalanges from the Nagri Formation. The functional morphology of these elements adds new detail to the reconstruction of Sivapithecus positional behavior. Femoral cross-sectional geometry indicates that the shaft was adapted to support mediolaterally directed loading. Femoral condylar asymmetry and a broad but shallow trochlea are distinctly ape-like, revealing capabilities for both rotation and withstanding eccentric loading in the knee. The navicular is characterized by features relating to a broad mid-tarsus and broad distal articulations for the cuneiforms. It also lacks a navicular tubercle as in Pongo. These features suggest that the foot was capable of a powerful grip on large supports, with an inversion/supination capability that would permit foot placement in a variety of positions. The morphology of the new phalanges, including evidence for a relatively large pollex, similarly suggests powerful grasping, consistent with prior evidence from the hallux and tarsus. The functional features of the new specimens permit refinement of previous interpretations of Sivapithecus positional capabilities. They suggest a locomotor repertoire dominated by pronograde activities and also such antipronograde activities as vertical climbing and clambering, but not by antipronograde suspensory activities as practiced by extant apes.