The thumb of Miocene apes: new insights from Castell de Barberà (Catalonia, Spain)

Primate hands display a major selective compromise between locomotion and manipulation. The thumb may or may not participate in locomotion, but it plays a central role in most manipulative activities. Understanding whether or not the last common ancestor of humans and Pan displayed extant-ape-like hand proportions (i.e., relatively long fingers and a short thumb) can be clarified by the analysis of Miocene ape hand remains. Here we describe new pollical remains-a complete proximal phalanx and a partial distal phalanx-from the middle/late Miocene site of Castell de Barberà (ca., 11.2-10.5 Ma, Vallès-Penedès Basin), and provide morphometric and qualitative comparisons with other available Miocene specimens as well as extant catarrhines (including humans). Our results show that all available Miocene taxa (Proconsul, Nacholapithecus, Afropithecus, Sivapithecus, Hispanopithecus, Oreopithecus, and the hominoid from Castell de Barberà) share a similar phalangeal thumb morphology: the phalanges are relatively long, and the proximal phalanges have a high degree of curvature, marked insertions for the flexor muscles, a palmarly bent trochlea and a low basal height. All these features suggest that these Miocene apes used their thumb with an emphasis on flexion, most of them to powerfully assist the fingers during above-branch, grasping arboreal locomotion. Moreover, in terms of relative proximal phalangeal length, the thumb of Miocene taxa is intermediate between the long-thumbed humans and the short-thumbed extant apes. Together with previous evidence, this suggests that a moderate-length hand with relatively long thumb-involved in locomotion-is the original hand morphotype for the Hominidae.     

The postcranium of Miocene hominoids: Were dryopithecines merely “dental apes”?

This paper reviews the non-dental morphological configuration of Miocene hominoids with special reference to the hypothesis of linear relationships between certain fossil species and living analogues. Metrical analysis of the wrist shows thatDryopithecus africanus andPliopithecus vindobonensis are unequivocally affiliated with the morphological pattern of quadrupedal monkeys. Similar analyses of the fossil hominoid elbow shows that they are more cercopithecoid-like than hominoid-like. Multivariate analysis of theP. vindobonensis shoulder in the matrix of extant Anthropoidea indicate that this putative hylobatine fossil shows no indication of even the initial development of hominoid features. The total morphological pattern of theD. africanus forelimb as assessed by principal coordinates analysis of allometrically adjusted shape variables has little resemblance toPan. Likewise, the feet and proximal femora of the Miocene fossils are unlike any living hominoid species. Even theD. africanus skull is similar to extant cercopithecoids in several features. Although ancestors cannot be expected to resemble descendants in every way, the striking dissimilarity between Miocene and extant hominoids seems to eliminate the consideration of a direct ancestor-descendant relationship between specific Miocene and modern forms.     

Cranio-dental evidence of a hominin-like hyper-masticatory apparatus in Oreopithecus bambolii. Was the swamp ape a human ancestor?

The phylogeny of Oreopithecus bambolii has been controversial since Johannes Hurzeler first argued that the Late Miocene (Tortonian) primate was a fossil hominin. While most paleontologist currently exclude Oreopithecus from human ancestry, recent postcranial evidence of hominin-like bipedalism and power precision grips in Oreopithecus has rekindled interest in the fossil Italian hominoid.In this study, a comparative review of hominoid cranio-dental morphology indicates that Oreopithecus possessed a suite of hominin-like characteristics that were apparently functionally related to powerful folivorous mastication. Since the oreopithecine dentition exhibited exceptional adaptations for folivory relative to most other extant and extinct hominoids, the accessory development of a hominin-like hyper-masticatory cranio-mandibular apparatus to further enhance plant comminution and digestibility is not surprising. However, the combination of hominin-like locomotive, manual, and masticatory functional attributes appears to substantiate Hurzeler's designation of O. bambolii as a Late Miocene hominin. Additionally, an extensive compilation of hominoid cranio-dental and postcranial characteristics strongly supports a close phylogenetic relationship between Oreopithecus and the earliest African hominins Sahelanthropus and Australopithecus.The wetland paleoecology of the Tortonian island of Tuscany–Sardinia suggests that Oreopithecus was a specialized semiaquatic folivore who apparently waded bipedally into freshwater swamps to feed on aquatic plants. However, the extensive wear on the oreopithecine canines and incisors along with their manual precision grips may indicate that freshwater invertebrates were also included in their diets. Such specialized aquatic feeding behavior by these ancient Italian swamp apes seems to support Alister Hardy's hypothesis that human bipedalism and power precision grips were inherited from Late Miocene hominin ancestors who originally utilized such functional attributes for aquatic foraging in shallow water environments. Additionally, the remarkably hominin-like cranio-dental morphology of O. bambolii suggests that modern omnivorous humans probably inherited a significant number of their cranio-dental characteristics from these highly specialized aquatic plant eating hominins.     

Early Origin for Human-Like Precision Grasping: A Comparative Study of Pollical Distal Phalanges in Fossil Hominins

Background

The morphology of human pollical distal phalanges (PDP) closely reflects the adaptation of human hands for refined precision grip with pad-to-pad contact. The presence of these precision grip-related traits in the PDP of fossil hominins has been related to human-like hand proportions (i.e. short hands with a long thumb) enabling the thumb and finger pads to contact. Although this has been traditionally linked to the appearance of stone tool-making, the alternative hypothesis of an earlier origin—related to the freeing of the hands thanks to the advent of terrestrial bipedalism—is also possible given the human-like intrinsic hand proportion found in australopiths.

Methodology/Principal Findings

We perform morphofunctional and morphometric (bivariate and multivariate) analyses of most available hominin pollical distal phalanges, including Orrorin, Australopithecus, Paranthropous and fossil Homo, in order to investigate their morphological affinities. Our results indicate that the thumb morphology of the early biped Orrorin is more human-like than that of australopiths, in spite of its ancient chronology (ca. 6 Ma). Moreover, Orrorin already displays typical human-like features related to precision grasping.

Conclusions

These results reinforce previous hypotheses relating the origin of refined manipulation of natural objects–not stone tool-making–with the relaxation of locomotor selection pressures on the forelimbs. This suggests that human hand length proportions are largely plesiomorphic, in the sense that they more closely resemble the relatively short-handed Miocene apes than the elongated hand pattern of extant hominoids. With the advent of terrestrial bipedalism, these hand proportions may have been co-opted by early hominins for enhanced manipulative capabilities that, in turn, would have been later co-opted for stone tool-making in the genus Homo, more encephalized than the previous australopiths. This hypothesis remains may be further tested by the finding of more complete hands of unequivocally biped early hominins.     

First hominoid from the Miocene of Ethiopia and the evolution of the catarrhine elbow

The first known fossil ape from the early-middle Miocene of Fejej, Ethiopia, is described here. The specimen, FJ-18SB-68, is a partial ulna from a locality dated by ⁴⁰Ar/³⁹Ar and paleomagnetic methods to a minimum age of 16.18 MYA. Compared to a variety of extant and fossil ulnae, FJ-18SB-68 is most similar to Turkanapithecus, Proconsul, and Pliopithecus, and appears to have been an arboreal quadruped with substantial forearm rotational mobility. Among the extant ulnae, canonical variates analysis successfully discriminates platyrrhines from catarrhines and within the latter, cercopithecoids from hominoids. Basal catarrhines (e.g., Aegyptopithecus) are platyrrhine-like in their morphology. Two basic trends appear to evolve from this generalized template: one with less mobile and more habitually pronated forearms, as seen in living and fossil cercopithecoids (including Victoriapithecus and Paracolobus), and another with greater forearm rotational mobility in fossil and modern hominoids. Primitive Miocene apes, including Proconsul, Turkanapithecus, and FJ-18SB-68, share with extant hominoids a more laterally positioned and laterally facing radial notch and an incipient trochlear keel. This morphology, along with a large insertion area for m. brachialis, suggests a departure from the more habitually pronated hand posture of monkeys and may indicate greater climbing abilities in these arboreally quadrupedal apes. Later Miocene apes, such as Oreopithecus and Dryopithecus share additional morphological features with hominoids, indicating considerable suspensory and climbing capabilities. Am J Phys Anthropol 105:257–277, 1998. © 1998 Wiley-Liss, Inc.     

Hominoid phalanges from the middle Miocene site of Paşalar, Turkey

Eleven proximal and ten intermediate partial or complete hominoid phalanges have been recovered from the middle Miocene site of Pa?alar in Turkey. Based on species representation at Pa?alar, it is likely that most or all of the phalanges belong to Griphopithecus alpani rather than Kenyapithecus kizili, but both species may be represented. All of the complete or nearly complete phalanges appear to be manual, so comparisons to extant and other fossil primate species were limited to manual phalanges. Comparisons were made to extant hominoid and cercopithecoid primate genera expressing a variety of positional repertoires and varying degrees of arboreality and terrestriality. The comparisons consisted of a series of bivariate indices derived from previous publications on Miocene catarrhine phalangeal morphology. The proximal phalanges have dorsally expanded proximal articular surfaces, which is characteristic of cercopithecoids and most other Miocene hominoids, and indicates that the predominant positional behaviors involved pronograde quadrupedalism. Among the extant primates, many of the proximal and intermediate phalangeal indices clearly distinguish more habitually terrestrial taxa from those that are predominantly arboreal, and especially from taxa that commonly engage in suspensory activities. For nearly every index, the values of the Pa?alar phalanges occupy an intermediate position-most similar to values for Pan and, to a lesser extent, Macaca-indicating a generalized morphology and probably the use of both arboreal and terrestrial substrates. At least some terrestrial activity is also compatible with reconstructions of the Pa?alar habitat. Most proximal and intermediate phalanges of other middle and late Miocene hominoids have similar index values to those of the Pa?alar specimens, revealing broadly similar manual phalangeal morphology among many Miocene hominoids.     

Pierolapithecus and the functional morphology of Miocene ape hand phalanges: paleobiological and evolutionary implications

The partial skeleton of Pierolapithecus, which provides the oldest unequivocal evidence of orthogrady, together with the recently described phalanges from Pa?alar most likely attributable to Griphopithecus, provide a unique opportunity for understanding the changes in hand anatomy during the pronogrady/orthogrady transition in hominoid evolution. In this paper, we describe the Pierolapithecus hand phalanges and compare their morphology and proportions with those of other Miocene apes in order to make paleobiological inferences about locomotor evolution. In particular, we investigate the orthograde/pronograde evolutionary transition in order to test whether the acquisition of vertical climbing and suspension were decoupled during evolution. Our results indicate that the manual phalanges of Miocene apes are much more similar to one another than to living apes. In particular, Miocene apes retain primitive features related to powerful-grasping palmigrady on the basal portion, the shaft, and the trochlea of the proximal phalanges. These features suggest that above-branch quadrupedalism, inherited from stem hominoids, constituted a significant component of the locomotor repertories of different hominoid lineages at least until the late Miocene. Nonetheless, despite their striking morphological similarities, several Miocene apes do significantly differ in phalangeal curvature and/or elongation. Hispanopithecus most clearly departs by displaying markedly-curved and elongated phalanges, similar to those in the most suspensory of the extant apes (hylobatids and orangutans). This feature agrees with several others that indicate orang-like suspensory capabilities. The remaining Miocene apes, on the contrary, display low to moderate phalangeal curvature, and short to moderately-elongated phalanges, which are indicative of the lack of suspensory adaptations. As such, the transition from a pronograde towards an orthograde body plan, as far as this particular anatomical region is concerned, is reflected only in somewhat more elongated phalanges, which may be functionally related to enhanced vertical-climbing capabilities. Our results therefore agree with the view that hominoid locomotor evolution largely took place in a mosaic fashion: just as taillessness antedated the acquisition of an orthograde body plan, the emergence of the latter—being apparently related only to vertical climbing—also preceded the acquisition of suspensory adaptations, as well as the loss of primitively-retained, palmigrady-related features.     

Miocene hominoid palatofacial morphology

The palatofacial morphology of Proconsul africanus, P. nyanzae, P. major and Sivapithecus meteai is compared to extant catarrhines. The early Miocene hominoids (Proconsul) are unlike modern great apes, but retain a primitive catarrhine pattern more similar to some extant cercopthecoids. By middle Miocene times the typical hominoid palatofacial morphology can be recognized in at least one species (S. meteai) and this corresponds to the evolution of the postcranium in which the hominoid pattern is also only recognizable by the middle Miocene.     

Tooth Wear Difference Between the Yuanmou Hominoid and <Emphasis Type="Italic">Lufengpithecus</Emphasis>

The Late Miocene hominoids recovered from Lufeng (Lufengpithecus) and Yuanmou of Yunnan Province, China, are among the most numerous hominoid fossils in Eurasia. They have yielded critical evidence for the evolutionary history, biogeography and paleobiology of Miocene hominoids. We examined and compared the wear pattern and differences of 804 molars of the Yuanmou hominoid and Lufengpithecus. Our results indicate that both the upper and lower molars of the Yuanmou hominoids were more heavily worn than those of Lufengpithecus. The wear patterns of the individual molars between the Yuanmou hominoid and Lufengpithecus also are different. The heaviest wear of lower molars of the Yuanmou hominoid occur in M₂, followed by M₁ and M₃. In Lufengpithecus, M₁ and M₃ were more heavily worn than M₂. There are differences in wear between the upper and lower molars for the two hominoids. Among the various factors related to tooth wear, we suggest that the main reason for the tooth wear differences between the Yuanmou hominoid and Lufengpithecus may be that they had different diets. More soft dietary items like leaves and berries were probably consumed by Lufengpithecus, and the Yuanmou hominoid may mainly have feed on harder or frugivorous diets. This result complements findings from previous studies of tooth size proportion, and the development of lower molar shearing crests in the 2 samples. Enamel thickness, living environment, behavior patterns, and population structure also might account for dental wear differences between the Yuanmou hominoid and Lufengpithecus.     

Parallel evolution in the hominoid trunk and forelimb

The evolutionary history of the living hominoids has remained elusive despite years of exploration and the discovery of numerous Miocene fossil ape species. Part of the difficulty can be attributed to the changing nature of our views about the course of hominoid evolution. In the 1950s and 1960s, individual Miocene taxa were commonly viewed as the direct ancestors of specific living ape species, suggesting an early divergence of the modern lineages.^1–5 However, in most cases, the Miocene forms were essentially “dental apes,” resembling extant species in dental and a few cranial features, but possessing more primitive postcranial features that suggested arboreal quadrupedalism rather than suspensory habits. With the introduction of molecular methods of phylogenetic reconstruction and the increasing use of cladistic analysis, it has become apparent that the radiation leading to the modern hominoids was somewhat more recent than had been believed, and that most of the Miocene hominoid species had little to do with the evolutionary history of the living apes. © 1998 Wiley-Liss, Inc.     

Late Miocene hominoid from Niger

African Late Miocene hominoids are rare, having been reported from six localities in Kenya, Ethiopia and Chad ranging in age from 10.5 to 5.5 Ma. We here report the occurrence of a hominoid in Niger associated with a vertebrate fauna which indicates an age of between 11–5 Ma. The Niger fossil locality is 940 km north of the nearest known extant hominoids, 1000 km west of the nearest recorded fossil hominoid from Chad.     

Reevaluation of the lumbosacral region of Oreopithecus bambolii

Functional interpretations of the postcranium of the late Miocene ape Oreopithecus bambolii are controversial. The claim that Oreopithecus practiced habitual terrestrial bipedalism is partly based on restored postcranial remains originally recovered from Baccinello, Tuscany ( Köhler and Moyà-Solà, 1997). The lower lumbar vertebrae of BA#72 were cited as evidence that Oreopithecus exhibits features indicative of a lordotic lumbar spine, including dorsal wedging of the vertebral bodies and a caudally progressive increase in postzygapophyseal interfacet distance. Here, we demonstrate why the dorsal wedging index value obtained by Köhler and Moyà-Solà (1997) for the BA#72 last lumbar vertebra is questionable due to distortion in that region, present a more reliable way to measure postzygapophyseal interfacet distance, and include an additional metric (laminar width) with which to examine changes in the transverse dimensions of the neural arches. We also quantify the external morphology of the BA#72 proximal sacrum, which, despite well-documented links between sacral morphology and bipedal locomotion, and excellent preservation of the sacral prezygapophyses, first sacral vertebral body, and right ala, was not evaluated by Köhler and Moyà-Solà (1997). Measures of postzygapophyseal interfacet distance and laminar width on the penultimate and last lumbar vertebrae of BA#72 reveal a pattern encompassed within the range of living nonhuman hominoids and unlike that of modern humans, suggesting that Oreopithecus did not possess a lordotic lumbar spine. Results further show that the BA#72 sacrum exhibits relatively small prezygapophyseal articular facet surface areas and mediolaterally narrow alae compared with modern humans, indicating that the morphology of the Oreopithecus sacrum is incompatible with the functional demands of habitual bipedal stance and locomotion. The Oreopithecus lumbosacral region does not exhibit adaptations for habitual bipedal locomotion.     

Morphometric analysis of hominoid lower molars from Yuanmou of Yunnan Province, China

Shape analyses of cross-sectional mandibular molar morphology, using Euclidean Distance Matrix Analysis, were performed on 79 late Miocene hominoid lower molars from Yuanmou of Yunnan Province, China. These molars were compared to samples of chimpanzee, gorilla, orangutan,Lufengpithecus lufengensis, Sivapithecus, Australopithecus afarensis, and human mandibular molars. Our results indicate that the cross-sectional shape of Yuanmou hominoid lower molars is more similar to the great apes that to humans. There are few differences between the Yuanmou,L. lufengensis, andSivapithecus molars in cross-sectional morphology, demonstrating strong affinities between these three late Miocene hominoids. All three of the fossil samples show strong similarities to orangutans. From this, we conclude that these late Miocene hominoids are more closely related to orangutants than to either the African great apes or humans.     

An oreopithecid proximal humerus from the middle miocene of Maboko Island,Kenya

A proximal humerus, recently recovered from the middle Miocene of Maboko Island, Kenya, provides the earliest evidence of postcranial structure and adaptation of Oreopithecidae. Provisionally attributed toNyanzapithecus pickfordi (Harrison, 1986), the specimen manifests a globose head, subequally large tuberosities, and a board, shallow bicipital groove. Although readily distinguished from the fundamentally cercopithecoid proximal humeral morphology ofVictoriapithecus (Senut, 1986), the Maboko Island oreopithecid, shows none of the derived features that are characteristic of the proximal humeri of extant hominoids. It is inferred from proximal humeral anatomy that the Maboko Island oreopithecid was an active arboreal scansor with moderate mobility at the shoulder but lacking adaptations for circumduction of the arm. In combination with craniodental evidence, proximal humeral morphology indicates that Oreopithecidae was a clade of hominoids which originated before the last common ancestor of extant apes and went extinct, without issue, in the later Miocene.     

Postcranial functional morphology of Morotopithecus bishopi, with implications for the evolution of modern ape locomotion

The large-bodied hominoid from Moroto, Uganda has until recently been known only from proconsulid like craniodental remains and some vertebrae with modern ape like features. The discovery of two partial femora and the glenoid portion of a scapula demonstrates that the functional anatomy of Morotopithecus differed markedly from other early and middle Miocene hominoids. Previous studies have consistently associated the vertebral remains with a short, stiff back and with orthograde postures. Although the proximal femur more closely resembles the femora of monkeys than of apes and suggests a moderate degree of hip abduction, the distal femur resembles those of extant large bodied apes and suggests a varied loading regime and an arboreal repertoire that may have included substantial vertical climbing. The femoral shaft displays uniformly thick cortical bone, beyond the range of thickness seen in extant primates, and signifies higher axial loading than is typical of most extant primates. The glenoid fossa is broad and uniformly curved as in extant suspensory primates. Overall, Morotopithecus is reconstructed as an arboreal species that probably relied on forelimb-dominated, deliberate and vertical climbing, suspension and quadrupedalism. Morotopithecus thus marks the first appearance of certain aspects of the modern hominoid body plan by at least 20 Ma. If the suspensory and orthograde adaptations linking Morotopithecus to extant apes are synapomorphies, Morotopithecus may be the only well-documented African Miocene hominoid with a close relationship to living apes and humans.     

A partial hominoid humerus from the middle miocene of Castell de Barberà (Vallès-Penedès Basin, Catalonia, Spain)

The hominoid right partial humerus IPS4334, from the middle Miocene (MN 8) of Castell de Barberà (Vallès-Penedès Basin, Catalonia, Spain), is described. It preserves the mid-distal portion of the shaft until the proximal margins of the radial and coronoid fossae, as well as the proximal portion of the olecranon fossa; the capitulum, the trochlea and the two epicondyles are missing. Although morphological comparisons are restricted, available evidence indicates that IPS4334 is more derived towards the modern hominoid condition than the Klein Hadersdorf specimen attributed to Griphopithecus (ca. 13-14 Ma), thus being most similar (except for its larger size and greater robusticity) to the presumably juvenile specimen of Dryopithecus fontani from Saint Gaudens in France (ca. 11-12 Ma). On the basis of shaft measurements and allometric regressions derived for extant hominoids, a body mass estimate around 50 kg is derived for IPS4334. Morphological similarities with the Saint Gaudens specimen, together with the large body mass estimate, suggest a tentative attribution of IPS4334 to cf. D. fontani, which is the largest hominoid taxon so far recorded from the Vallès-Penedès Basin. The larger size and higher robusticity of IPS4334 as compared to the Saint Gaudens specimen might be explained by the juvenile status of the latter and/or sexual dimorphism. When both specimens are considered together with a partial femur from Abocador de Can Mata, D. fontani emerges as a less suspensory ape than the late Miocene Hispanopithecus, the locomotor repertoire of the former emphasizing climbing, but still displaying a significant quadrupedal component.     

The relationships among jaw‐muscle fiber architecture,jaw morphology,and feeding behavior in extant apes and modern humans

The jaw‐closing muscles are responsible for generating many of the forces and movements associated with feeding. Muscle physiologic cross‐sectional area (PCSA) and fiber length are two architectural parameters that heavily influence muscle function. While there have been numerous comparative studies of hominoid and hominin craniodental and mandibular morphology, little is known about hominoid jaw‐muscle fiber architecture. We present novel data on masseter and temporalis internal muscle architecture for small‐ and large‐bodied hominoids. Hominoid scaling patterns are evaluated and compared with representative New‐ (Cebus) and Old‐World (Macaca) monkeys. Variation in hominoid jaw‐muscle fiber architecture is related to both absolute size and allometry. PCSAs scale close to isometry relative to jaw length in anthropoids, but likely with positive allometry in hominoids. Thus, large‐bodied apes may be capable of generating both absolutely and relatively greater muscle forces compared with smaller‐bodied apes and monkeys. Compared with extant apes, modern humans exhibit a reduction in masseter PCSA relative to condyle‐M₁ length but retain relatively long fibers, suggesting humans may have sacrificed relative masseter muscle force during chewing without appreciably altering muscle excursion/contraction velocity. Lastly, craniometric estimates of PCSAs underestimate hominoid masseter and temporalis PCSAs by more than 50% in gorillas, and overestimate masseter PCSA by as much as 30% in humans. These findings underscore the difficulty of accurately estimating jaw‐muscle fiber architecture from craniometric measures and suggest models of fossil hominin and hominoid bite forces will be improved by incorporating architectural data in estimating jaw‐muscle forces. Am J Phys Anthropol 151:120–134, 2013. © 2013 Wiley Periodicals, Inc.     

Anatomy of the hominoid wrist joint: Its evolutionary and functional implications

Observations on the behavior of living hominoids show generic differences in the use and posture of the wrist joint. Both orang-utans and hylobatids usually use the wrist in suspensory behaviors. However, orang-utans emphasize markedly adducted and flexed wrist postures, while hylobatids emphasize violent forearm and wrist rotation. African apes, especially the gorilla, use the wrist more frequently than other hominoids for terrestrial quadrupedal weight-bearing. Humans use the wrist less frequently for supportive purposes than do other hominoids. These behavioral differences correspond to structural specializations in the proximal carpal joint of each of the hominoid genera. Although each of the hominoid genera has apparently modified its proximal carpal joint best to serve its characteristic behaviors, all hominoids share a unique proximal carpal joint that permits approximately 160ℴ of forearm rotation. The hylobatid proximal carpal joint is specialized in exhibiting a marked development of those structures limiting forearm rotation, but it is in most respects the least derived— that is, closest to the nonhominoid anthropoids. Chimpanzees show a proximal carpal joint that is more generalized than those of the other great apes but more derived than that of hylobatids. The human and gorilla proximal wrist joints, on the other hand, show marked modifications for weight-bearing in terrestrial behaviors. Orang-utans have the most derived proximal carpal joint, which in many respects parallels that of the slow-climbing nonhominoid primates. The comparative anatomy and structural specializations of the wrist joint support (a) an early divergence of hylobatids from the common hominoid stock, (b) a common ancestry for gorillas and humans separate from the other hominoids, and (c) a long independent evolutionary period for orang-utans since their divergence from the common hominoid stock, or one that was marked by strong selection pressures for wrist specializations. Unfortunately, the generalized condition of the chimpanzee’s wrist joint and the very derived condition of the orang-utan wrist provide uncertain evidence as to which of the two was first to diverge from the common hominoid stock. Identification of hominoid wrist specializations as reflecting real phylogenetic relationships or parallelisms depends on how well the phytogeny inferred from wrist morphology accords with those arrived at from the study of other 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.