Extant Colobines Research Articles

Hundreds of Colobus (Cercopithecidae: Primates) fossils from the later Pleistocene of Ethiopia's Middle Awash study area

AbstractObjectivesThe aim of this study is to introduce and systematically assess a new assemblage of colobine fossils (n = 360) recovered from Late Pleistocene (ca. 100,000 years ago) sediments in the Middle Awash research area, Afar Rift, Ethiopia.Materials and MethodsWe describe the colobine fossils relative to extant colobine taxa and Middle Pleistocene fossil samples from “Andalee” and Asbole, Ethiopia using linear dental, cranial, and postcranial metrics, as well as qualitative features. We also document the ontogenetic and sex profiles of this sample.ResultsBased on morphological affinities to extant and fossil Colobus, we refer these 360 individuals to Colobus cf. guereza. The majority of the individuals (76%) fall into young‐ and middle‐adult age categories. Females are slightly better represented than males, although the majority of individuals (84%) could not be assigned to sex.DiscussionDetails of the last half million years of Colobus morphological evolution have remained obscured by a sparse fossil record. The newly recovered, prepared, and curated fossils described herein add significantly to the record of this group. Several morphologies seen in these new fossils fall between the Asbole Colobus sample and extant Colobus guereza, suggesting an ancestral‐descendant phyletic relationship among the three time‐successive samples. The populational variation documented among these samples indicates that the distinctive pattern of sexual dimorphism observed in C. guereza had evolved by 100 ka.

Feeding ecology of the last European colobine monkey, Dolichopithecus ruscinensis

Currently, very little is known about the ecology of extinct Eurasian cercopithecids. Dietary information is crucial in understanding the ecological adaptations and diversity of extinct cercopithecids and the evolution of this family. For example, the colobine genus Dolichopithecus is represented by multiple large-bodied species that inhabited Eurasia during the Pliocene-Early Pleistocene. The available evidence, though limited, suggests semiterrestrial locomotion, which contrasts with most extant African and Asian colobines that exhibit morphological and physiological adaptations for arboreality and folivory. These differences raise questions regarding the dietary specialization of early colobine taxa and how/if that influenced their dispersion out of Africa and into Eurasia. To further our understanding of the ecology of Plio-Pleistocene cercopithecids, we characterized the dental capabilities and potential dietary adaptations of Dolichopithecus ruscinensis through dental topographic and enamel thickness analyses on an M1 from the locality of Serrat d'en Vacquer, Perpignan (France). We also assessed the feeding behavior of D. ruscinensis through dental microwear texture analysis on a broad sample of fossil molars from fossil sites in France, Greece, Bulgaria, and Romania. Dental topographic and enamel thickness analyses suggest that D. ruscinensis could efficiently process a wide range of foods. Results of the dental microwear texture analysis suggest that its diet ranged from folivory to the consumption of more mechanically challenging foods. Collectively, this suggests a more opportunistic feeding behavior for Dolichopithecus than characteristic of most extant colobines.

Masticatory and ingestive effort in Procolobus verus, a small-bodied African colobine.

The olive colobus (Procolobus verus) is the smallest extant colobine. Based on the axiom that folivory is associated with larger body mass, the olive colobus is expected to be less folivorous than its sister taxon Piliocolobus badius, but previous studies show that the opposite is true. Here we test the hypothesis that masticatory and ingestive effort in the olive colobus is greater due to allometric factors related to bite force scaling and throughput of ingested foods. We analyzed oral processing data collected on olive colobus in the Taï Forest, Ivory Coast, between May 2016 and May 2018. We compare these with previously published data on P. badius and Colobus polykomos from Taï. In terms of overall feeding effort, olive colobus invest more effort (i.e., chewing cycles) than the larger colobines. When contrasts are restricted to commonly consumed foods, this greater energetic investment is not consistently observed. Ingestion of young leaves is associated with a reduced number of masticatory cycles in all three colobine species. A slightly elevated average effort in the olive colobus during young leaf feeding suggests this food source is more challenging in smaller monkeys, but mature leaf processing effort is generally the same among Taï colobines. Thus, for olive colobus, leaf ductility may be more problematic than leaf toughness in terms of masticatory effort. While there may be an allometric cost to being a small colobine, food selectivity is an important mitigating factor.

From leaves to seeds? The dietary shift in late Miocene colobine monkeys of southeastern Europe

Extant colobine monkeys are specialized leaf eaters. But during the late Miocene, western Eurasia was home to colobines that were less efficient at chewing leaves than they were at breaking seed shells. To understand the link between folivory and granivory in this lineage, the dietary niche of Mesopithecus delsoni and Mesopithecus pentelicus was investigated in southeastern Europe, where a major environmental change occurred during the late Miocene. We combined dental topographic estimates of chewing efficiency with dental microwear texture analysis of enamel wear facets. Mesopithecus delsoni was more efficient at chewing leaves than M. pentelicus, the dental topography of which matches an opportunistic seed eater. Concurrently, microwear complexity increases in M. pentelicus, especially in the northernmost localities corresponding to present-day Bulgaria. This is interpreted as a dietary shift toward hard foods such as seeds or tubers, which is consistent with the savanna and open mixed forest biomes that covered Bulgaria during the Tortonian. The fact that M. delsoni was better adapted to folivory and consumed a lower amount of hard foods than M. pentelicus suggests that colobines either adapted to folivory before their dispersal to Europe or evolved adaptations to leaf consumption in multiple occurrences.

An assessment of the postcranial skeleton of the Paracolobus mutiwa (Primates: Colobinae) specimen KNM-WT 16827 from Lomekwi, West Turkana, Kenya

The postcranium of a large-bodied colobine monkey attributed to Paracolobus mutiwa from the site of Lomekwi, West Turkana, Kenya, is described. The partial skeleton (KNM-WT 16827) was recovered from locality LO 1, dated to 2.58–2.53 Ma, and preserves postcranial elements including fragments of scapula, humerus, proximal ulna, proximal radius, os coxae, proximal femur, astragalus, and calcaneus. KNM-WT 16827 was identified as P. mutiwa based on cranial similarities to the holotype female maxilla (KNM-ER 3843) and the holotype of Paracolobus chemeroni (KNM-BC 3), but is currently the only specimen of this taxon with associated cranial and postcranial elements. The skeleton is morphologically distinct from other large cercopithecid specimens from the Turkana Basin, including several assigned to Cercopithecoides williamsi, Cercopithecoides kimeui, Rhinocolobus turkanaensis, and Theropithecus oswaldi and differs from KNM-BC 3 in the larger cranium and shorter and more robust long bones. KNM-WT 16827 has forelimb and hindlimb features exhibiting a mixture of traits more associated with terrestrial locomotor behavior, including robust humeral deltoid tuberosity, retroflexed humeral medial epicondyle, deep ulnar trochlear notch, relatively short lower iliac height, prominent femoral greater trochanter, asymmetrical astragalar trochlea, and weak digit flexor grooves on the calcaneus. KNM-WT 16827 is also proportionally distinct from KNM-BC 3 and other Turkana Basin specimens attributed to large-bodied taxa such as C. williamsi, C. kimeui, R. turkanaensis, and T. oswaldi in having relatively shorter limbs and smaller tarsals. The traits shared with P. chemeroni and other extinct taxa are either typical for colobines, or likely due to P. mutiwa and P. chemeroni sharing adaptations for terrestrial locomotion relative to extant colobinans. Although a full cranial assessment is needed, based on its postcranial morphology KNM-WT 16827 is distinct from KNM-BC 3, C. williamsi, R. turkanaensis, Theropithecus, and extant colobines, warranting further analyses to better assess the taxonomic assignment of the specimen.

Fossil Cercopithecidae from the Early Pliocene Sagantole Formation at Gona, Ethiopia

The Early Pliocene Sagantole Fm. in the Gona Project area, Afar State, Ethiopia, is noted for discoveries of the early hominin Ardipithecus ramidus. A large series of fossil cercopithecid primates dated to between 4.8 and 4.3 Ma has also been collected from these sediments. In this paper, we use qualitative analysis and standard dental and postcranial measures to systematically describe the craniodental remains and tentatively allocate postcrania to taxa where we are able to. We then use these data to compare these specimens to fossil assemblages from contemporary sites, interpret their paleobiology, and discuss implications for the paleoecology of the Gona Sagantole Fm. We recognize three cercopithecid species in the Gona Sagantole Fm. Pliopapio alemui makes up approximately two-thirds of the identifiable specimens; nearly all of the rest are allocated to Kuseracolobus aramisi, and a single molar indicates the presence of a second, somewhat larger but morphologically distinct papionin. Among the Early Pliocene cercopithecids from Gona are also a number of postcranial elements. None of the postcranial remains are directly associated with any of the cranial material. Nonetheless, some of the distal humeri and proximal femora can be tentatively allocated to either Pl. alemui or K. aramisi based on a combination of size, as the latter is approximately 50% larger than the former, and morphology. If these assignments are correct, they suggest K. aramisi was primarily arboreal and similar to most extant colobines, whereas Pl. alemui was more mixed in its substrate use, being more terrestrially adapted than K. aramisi, but less so than extant Papio or Theropithecus. Thus, we interpret the predominance of Pl. alemui over K. aramisi is consistent with a somewhat more open environment at Gona than at Aramis.

Evolution of the bitter taste receptor TAS2R38 in colobines.

Bitter taste perception enables the detection of potentially toxic molecules and thus evokes avoidance behavior in vertebrates. It is mediated by bitter taste receptors, TAS2Rs. One of the best-studied TAS2R is TAS2R38. Phenylthiocarbamide (PTC) perception and TAS2R38 receptors vary across primate species, and this variation may be related to variation in dietary preferences. In particular, we previously found that the low sensitivity of TAS2R38s in Asian colobines likely evolved as an adaptation to their leaf-eating behavior. However, it remains unclear whether this low PTC sensitivity is a general characteristic of the subfamily Colobinae, a primate group that feeds predominantly on leaves. We performed genetic analyses, functional assays with mutant proteins, and behavioral analyses to evaluate the general characteristics of TAS2R38 in colobines. We found that PTC sensitivity is lower in TAS2R38s of African colobines than in TAS2R38s of omnivorous macaques. Furthermore, two amino acids shared between Asian and African colobines were responsible for low sensitivity to PTC, suggesting that the last common ancestor of extant colobines had this phenotype. We also detected amino acid differences between TAS2R38s in Asian and African colobines, indicating that they evolved independently after the separation of these groups.

Mesopithecus pentelicus from the Turolian locality of Kryopigi (Kassandra, Chalkidiki, Greece)

New material of the Mio-Pliocene colobine Mesopithecus from the Turolian locality of Kryopigi (Greece) is described here. It includes a complete skull with the atlas attached and other dental and postcranial elements representing at least five individuals (four males and one female). The material is compared with Mesopithecus delsoni, Mesopithecus pentelicus, Mesopithecus monspessulanus and intermediate forms from more than a dozen Turolian localities of the Greco-Iranian province. These comparisons support the attribution of the Kryopigi material to M. pentelicus. The chronostratigraphic distribution of Mesopithecus species and intermediate forms suggests that the Kryopigi fauna could be dated as younger than the Perivolaki locality with M. delsoni/pentelicus (7.1–7.3 Ma, MN12) and older than the Dytiko localities with M. aff. pentelicus, M. cf. pentelicus and M. cf. monspessulanus (?middle MN13). The dimensions of the atlas are within the distribution of extant colobines. The skull shows bite-marks, probably caused by the hyaena Adcrocuta eximia.

A new fossil cercopithecid tibia from Laetoli and its implications for positional behavior and paleoecology

Detailed analyses and comparisons of postcranial specimens of Plio-Pleistocene cercopithecids provide an opportunity to examine the recent evolutionary history and locomotor diversity in Old World monkeys. Studies examining the positional behavior and substrate preferences of fossil cercopithecids are also important for reconstructing the paleoenvironments of Plio-Pleistocene hominin sites. Here we describe a new fossil cercopithecid tibia (EP 1100/12) from the Australopithecus afarensis-bearing Upper Laetolil Beds (∼3.7 Ma) of Laetoli in northern Tanzania. The fossil tibia is attributed to cf. Rhinocolobus sp., which is the most common colobine at Laetoli. In addition to qualitative comparisons, the tibial shape of EP 1100/12 was compared to that of 190 extant cercopithecids using three-dimensional landmarks. Discriminant function analyses of the shape data were used to assess taxonomic affinity and shape variation relating to positional behavior. EP 1100/12 clustered with extant colobines, particularly the large-bodied genera Nasalis and Rhinopithecus. Comparisons reveal that EP 1100/12 belongs to a large-bodied monkey that engaged in arboreal pronograde quadrupedalism. These findings add further support to previous inferences that woodland and forest environments dominated the paleoenvironment of the Upper Laetolil Beds, which supported the diverse community of cercopithecids at Laetoli. The inferred paleoecology and the presence of large-bodied arboreally-adapted monkeys at Laetoli show that A. afarensis had access to a range of diverse habitats, including woodlands and forests. This supports the possibility that A. afarensis, with its potential range of positional capabilities, was able to utilize arboreal settings for food acquisition and refuge from predators.

Was Mesopithecus a seed eating colobine? Assessment of cracking, grinding and shearing ability using dental topography

Extant colobine monkeys have been historically described as specialized folivores. However, reports on both their behavior and dental metrics tend to ascribe a more varied diet to them. In particular, several species, such as Pygathrix nemaeus and Rhinopithecus roxellana, are dedicated seasonal seed eaters. They use the lophs on their postcanine teeth to crack open the hard endocarp that protects some seeds. This raises the question of whether the bilophodont occlusal pattern of colobine monkeys first evolved as an adaptation to folivory or sclerocarpic foraging. Here, we assess the sclerocarpic foraging ability of the oldest European fossil colobine monkey, Mesopithecus. We use computed microtomograpy to investigate the three-dimensional (3D) dental topography and enamel thickness of upper second molars ascribed to the late Miocene species Mesopithecus pentelicus from Pikermi, Greece. We compare M. pentelicus to a sample of extant Old World monkeys encompassing a wide range of diets. Furthermore, we combine classic dietary categories such as folivory with alternative categories that score the ability to crack, grind and shear mechanically challenging food. The 3D dental topography of M. pentelicus predicts an ability to crack and grind hard foods such as seeds. This is consistent with previous results obtained from dental microwear analysis. However, its relatively thin enamel groups M. pentelicus with other folivorous cercopithecids. We interpret this as a morphological trade-off between the necessity to avoid tooth failure resulting from hard food consumption and the need to process a high amount of leafy material. Our study demonstrates that categories evaluating the cracking, grinding or shearing ability, traditional dietary categories, and dental topography combine well to make a powerful tool for the investigation of diet in extant and extinct primates.

Substrate use and postural behavior in free‐ranging snub‐nosed monkeys (Rhinopithecus bieti) in Yunnan

Extant colobines represent a predominantly arboreal primate radiation, but there are outliers such as some species of snub-nosed monkeys (Rhinopithecus spp.) that utilize both arboreal and terrestrial substrates. We quantified the degree of terrestriality and the distribution of individuals across canopy levels, the extrinsic and intrinsic determinants and constraints of strata association as well as use of postures in relation to behavior and food sources in a wild population of Yunnan snub-nosed monkeys (Rhinopithecus bieti) in the Samage Forest, China. The subjects spent 80.5% of the observation time in the trees and 19.5% on the ground. Ground use occurred most frequently in the context of travelling (61%) and less often in the context of feeding and resting (30% combined). Adult males frequented the ground significantly more often than other age-sex classes. Age-sex differences were also apparent in substrate preferences, which likely reflect body weight constraints. Adult males were significantly more associated with solid substrates and less with terminal branches than adult females and juveniles. Juveniles were seen more often on unstable ground (i.e. on thin and oblique branches) and in the upper canopy. Substrate use also varied with behavior: feeding was much more common in the upper stratum, outer canopy, on highly inclined and thin branches, as compared to resting. Severe weather also influenced canopy use in that tree crowns were used less often during periods of snowfall. The most frequently exhibited resting and feeding postures was sitting. Standing and suspensory postures were most often used in the context of feeding on lichens.

A functional multivariate analysis of Mesopithecus (Primates: Colobinae) humeri from the Turolian of Greece

The genus Mesopithecus is well represented in the late Miocene of Greece by several recognized species. The present paper investigates functional aspects of the humeri of Mesopithecus delsoni/pentelicus, M. pentelicus and M. aff. pentelicus of several Turolian sites from central and northern Greece, using multivariate approaches. For these purposes, we selected significant humeral functional features, which were represented by 23 linear dimensions and three angles on 14 fossil humeri and 104 humeri from 10 genera and 22 species of extant African and Asian Colobines. All size-adjusted measurements were examined through a principal components analysis, followed by a discriminant function analysis, and a canonical variates analysis. All analyses revealed that the selected characters were able to discriminate between extant colobine genera. Functional groups, such as arboreal walking/climbing, arboreal walking/suspensory and semi-terrestrial walking, were set apart from a central cluster formed by the arboreal walking and arboreal walking/terrestrial groups. This cluster also grouped the three studied Mesopithecus species, which were mainly classified as arboreal walkers with significant terrestrial activities. These observations match with paleoenvironmental reconstructions and the suggested opportunistic feeding habits. Moreover, the overall arboreal/terrestrial locomotor tendencies of these fossil forms are discussed in relation to their earlier migration from Africa and later dispersal to eastern and southern Asia.

Earliest colobine skeletons from Nakali, Kenya

Old World monkeys represent one of the most successful adaptive radiations of modern primates, but a sparse fossil record has limited our knowledge about the early evolution of this clade. We report the discovery of two partial skeletons of an early colobine monkey (Microcolobus) from the Nakali Formation (9.8-9.9 Ma) in Kenya that share postcranial synapomorphies with extant colobines in relation to arboreality such as mediolaterally wide distal humeral joint, globular humeral capitulum, distinctly angled zona conoidea, reduced medial trochlear keel, long medial epicondyle with weak retroflexion, narrow and tall olecranon, posteriorly dislocated fovea on the radial head, low projection of the femoral greater trochanter, wide talar head with a greater rotation, and proximodistally short cuboid and ectocuneiform. Microcolobus in Nakali clearly differs from the stem cercopithecoid Victoriapithecus regarding these features, as Victoriapithecus is postcranially similar to extant small-sized terrestrial cercopithecines. However, degeneration of the thumb, a hallmark of modern colobines, is not observed, suggesting that this was a late event in colobine evolution. This discovery contradicts the prevailing hypothesis that the forest invasion by cercopithecids first occurred in the Plio-Pleistocene, and shows that this event occurred by the late Miocene at a time when ape diversity declined.

Feeding habits of the first European colobine,Mesopithecus(Mammalia, Primates): evidence from a comparative dental microwear analysis with modern cercopithecids

This study aims to characterize the feeding habits of the first European colobines, Mesopithecus (late Miocene), through the analysis of its molar microwear pattern. Fifty-seven adult individuals of Mesopithecus (from Greece and Bulgaria) are compared to 162 wild-shot specimens representing nine modern species of African and Asian cercopithecids. Through the combination of a principal component analysis and analyses of variances, Mesopithecus displays significant differences with its extant colobine relatives. Actually, there is no significant difference between the Mesopithecus samples and Chlorocebus aethiops and Papio hamadryas. An intra-specific test dedicated to Mesopithecus pentelicus reveals no difference between the Bulgarian and the Greek samples. To sum up, the species of Mesopithecus were not leaf eaters like their present-day colobine relatives. Considering its wide paleogeographic range, its post-cranial anatomy, the paleoenvironmental data, and the present dietary reconstruction as a whole, the species of Mesopithecus can be depicted as semi-terrestrial monkeys opportunists in term of feeding preferences.

Locomotor evolution of Mesopithecus (Primates: Colobinae) from Greece: evidence from selected astragalar characters

This paper reports our investigations into functional aspects of the astragalus of four samples of the genus Mesopithecus from Greece. More particularly, it aims to infer substrate preferences of M. delsoni/pentelicus from the Middle Turolian site of Perivolaki (central Greece), M. pentelicus from the late Middle Turolian site of Pikermi (southern Greece), and M. cf. pentelicus and M. cf. monspessulanus from the Late Turolian site of Dytiko (northern Greece). For these purposes, selected astragalar functional features, such as trochlea wedging, proximal facet curvature, and head rotation were expressed as linear measurements on both fossil and selected extant colobines. The size-adjusted measurements were used for univariate comparisons as well as a multivariate principal components analysis. Both approaches revealed that the selected characters were able to discriminate between extant arboreal and semiterrestrial colobines, but all fossil forms presented mosaic morphology. Thus, the oldest representative, M. delsoni/pentelicus was reconstructed as mainly semiterrestrial. On the other hand, the astragalus of M. pentelicus appeared to reflect semiterrestrial habits with a moderate adaptation to arboreality. Similar habitus reconstruction was allocated to the more recent M. cf. pentelicus, whereas the sympatric and synchronous M. cf. monspessulanus showed semiterrestrial adaptations with a slight preference of terrestrial substrates. The results mainly conform to paleoenvironmental reconstructions of the fossiliferous localities and denote that Mesopithecus was mainly a semiterrestrial radiation throughout its evolutionary history, with differential rates of use between arboreal and terrestrial substrates. These adaptations could have promoted the dispersal of the genus throughout Eurasia during the latest Miocene and Early Pliocene.

Presence of the Maxillary Sinus in Fossil Colobinae (Cercopithecoides williamsi) from South Africa

Extant cercopithecoid monkeys, except macaques, are distinguished among primates by their lack of paranasal pneumatization, including the maxillary sinus (MS). Analysis of this structure, widespread among Eutheria, suggests that its loss occurred in the cercopithecoid common ancestor; thus, the presence of the MS in macaques is not strictly homologous to that in other primates. CT analysis of the fossil species Victoriapithecus macinnesi supports this view, demonstrating the lack of the MS in this stem cercopithecoid. Recent evidence, however, has documented the presence of the MS in extinct cercopithecoids from the late Miocene and Pliocene. This study reports on CT examination of two fossil crania attributed to Cercopithecoides williamsi from South Africa, dated in the range, 3.0-1.5 Ma. BF 42a is a complete cranium from Bolts Farm; MP113 is an intact facial skeleton, including the anterior cranial vault, from the Makapansgat Limeworks. Both demonstrate MS presence, unknown in extant colobines and unexpected in most cercopithecoid monkeys. The relative size of the MS of BF 42a is similar to that of extant tropical and subtropical macaques. The presence of sinuses in several extinct colobines suggests that our understanding of the evolutionary history of these primates, and of the MS, is incomplete, and that other fossil cercopithecoids should be examined for this feature. The developmental plasticity exhibited in this feature, indicated by multiple loss and reemergence, provides further evidence that paranasal pneumatization has undergone a complex history of suppression and expression.

Paranasal pneumatization in extant and fossil Cercopithecoidea

Unlike most primates, extant cercopithecoids lack maxillary sinuses, which are pneumatic spaces in the facial skeleton lateral of the nasal cavity proper. Character state analysis of living cercopithecoids across well-supported topologies suggests that the sinus was lost at the origin of the superfamily, only to have evolved again convergently in extant macaques. Recent work has shown that a) the ‘early loss’ hypothesis is supported by the lack of any pneumatization in Victoriapithecus, a stem cercopithecoid, b) like extant macaques, the fossil cercopithecine Paradolichopithecus shows evidence of presence of the maxillary sinus (MS), and c) unlike extant colobines, the fossil colobine Libypithecus also possesses a maxillary sinus. To more fully assess the pattern of cercopithecoid sinus evolution, fossil taxa from both subfamilies (Colobinae, Cercopithecinae) were examined both visually and by computed tomography (CT). The observations were evaluated according to standard anatomical criteria for defining sinus spaces, and compared with data from all extant Old World monkey genera. Most taxa examined conformed to the pattern already discerned from extant cercopithecoids. Maxillary sinus absence in Theropithecus oswaldi, Mesopithecus, and Rhinocolobus is typical for all extant cercopithecids except Macaca. The fossil macaque Macaca majori possesses a well-developed maxillary sinus, as do all living species of the genus. Cercopithecoides, on the other hand, differs from all extant colobines in possessing a maxillary sinus. Thus, paranasal pneumatization has reemerged a minimum of two and possibly three times in cercopithecoids. The results suggest that maxillary sinus absence in cercopithecoids is due to suppression, rather than complete loss.

Distal humerus and ulna of Parapresbytis (Colobinae) from the Pliocene of Russia and Mongolia: phylogenetic and ecological implications based on elbow morphology

Parapresbytis eohanuman is a colobine known from two middle Pliocene localities southeast of Lake Baikal, northern East Asia. This study examined the morphology of postcranial specimens, a distal humerus and an ulna, of Parapresbytis. A total of 18 and 13 linear measurements were taken from the humerus and the ulna, respectively, and compared with those of extant and European fossil colobines using principal components analysis. The distal humeral specimen of Parapresbytis is slightly larger than those of male Semnopithecus and Nasalis, while the ulnar specimen is much larger than those of extant colobines and is nearly as large as that of a male Papio ursinus. Morphologically, the distal humeri and ulnae of terrestrial colobines such as Semnopithecus, Dolichopithecus, and Mesopithecus can be distinguished from those of arboreal colobines. The morphology of the Parapresbytis elbow is within the range of the arboreal colobines, contradicting previous suggestions that this genus is terrestrially adapted and is phylogenetically close to Dolichopithecus. Because the arboreally adapted elbow is shared by most extant colobines, the morphology of the elbow does not provide evidence for phylogenetic proximity between Parapresbytis and a particular taxon of arboreal colobine such as Rhinopithecus. The elbow morphology suggests that Parapresbytis was adapted to arboreal locomotion as much as the extant arboreal colobines are. This is congruent with the paleoenvironmental evidence, which indicates the presence of forests in the Lake Baikal area during the middle Pliocene.

Victoriapithecus: The key to Old World monkey and catarrhine origins

The past ten years have witnessed major changes in reconstructions of the history of Old World monkeys, most of them driven by new material of the Miocene monkey Victoriapithecus from Maboko Island, Kenya. Before the mid-1980s, predictions about the morphological and ecological adaptations of the earliest cercopithecoids relied heavily on evidence from extant colobine and cercopithecine monkeys. It was argued that the earliest cercopithecoids were largely or at least partly folivorous, had short colobine-like faces, and were arboreal. The only studies suggesting that some of these arguments were not true were based on limited knowledge of the anatomy of Victoriapithecus. The presence of semi-terrestrial adaptations in middle Miocene monkeys hinted to some that early monkeys may not have been arboreal. Others attempted to cope with the discrepancy between neontological predictions and the fossil evidence by proposing that limb bones with stronger terrestrial adaptations within the Maboko sample were derived cercopithecine remains, while those with more arboreal features belonged in the subfamily Colobinae and should be regarded as primitive.