Cercopithecoid Monkeys Research Articles

What can monkey fossils tell us about early human evolution? New insights from the analysis of molar cusp proportions in fossil cercopithecoids from Laetoli, Tanzania

IntroductionLaetoli, Tanzania preserves the earliest fossils attributed to Australopithecus afarensis (3.7 mya). A diversity of non‐human primates including cercopithecoid monkeys have also been recovered. Cercopithecoids provide valuable insights into the paleoenvironments they shared with early humans and, by extension, early hominin paleobiology. Although cercopithecoid fossils at Laetoli represent both more omnivorous cercopihecines and more folivorous and arboreal colobines, the former is more abundant. The diversity and proportion of cercopithecoids figures prominently in debate over the prevalence of closed woodland/forested environments vs more open scrub/bushland. This study uses dental morphometrics to establish the morphological affinity of molar crowns from additional cercopithecoid craniodental fossils recovered from Laetoli localities 1, 2, 6 and 7 to clarify the paleoenvironmental context of our early human ancestors.MethodsStandard linear metric data (mesiodistal length, buccolingual breadth) were collected using dental calipers and total occlusal and individual cusp areas (mm2) were obtained for all molar crowns in an extant cercopithecoid sample (n=90; 18 genera) using high‐resolution 2D images and ImageJ. These data are a comparative baseline for a fossil cercopithecoid sample including a complete (LP) mandible (M1‐M3), maxillary fragment (M1‐M3) and four isolated maxillary molars (n=3 M1’s, n=1 M2). Discriminant function analysis (DFA) using subfamily (n=2 groups) and tribe (n=5 groups) was used to predict group membership and morphological affinity of the fossil sample.ResultsDFA results indicate that predictive accuracy is highest when subfamily is used as the grouping variable and ranges from 62.8% (M3) to 87.2% (M2). Mandibular molars more reliably segregate subfamilies (77.1% ‐ 87.2%) than do maxillary molars (64.4% ‐ 70.2%). The associated mandibular molars group are most closely associated with African colobines. All maxillary specimens cluster with cercopithecines and, more specifically, the papioni. Colobines have expanded hypoconids and metacones but reduced metaconids, entoconids and paracones relative to cercopithecoids.ConclusionMolar cusp proportions discriminate colobines from cercopithecines. This study provides additional evidence for the continued presence of both groups in the upper beds at Laetoli and confirms that the complete LP mandible is most closely aligned with the Colobinae.The presence of a large colobine is consistent with palaeoecological reconstructions supporting the significant presence of closed woodland/forested habitats.SignificanceDespite obligate bipedalism, early hominins retain numerous upper limb and torso adaptations consistent with the use of arboreal substrates. The continued presence of colobines at Laetoli confirms the presence of closed woodland/forested environments and suggests that Au. afarensis at Laetoli may have potentially exploited a variety of habitats including arboreal substrates.

A geometric morphometric approach to investigate primate proximal phalanx diaphysis shape.

Current approaches to quantify phalangeal curvature assume that the long axis of the bone's diaphysis approximates the shape of a portion of a circle (included angle method) or a parabola (second-degree polynomial method). Here we developed, tested, and employed an alternative geometric morphometrics-based approach to quantify diaphysis shape of proximal phalanges in humans, apes and monkeys with diverse locomotor behaviors. 100 landmarks of the central longitudinal axis were extracted from 3D surface models and analyzed using 2DGM methods, including Generalized Procrustes Analyses. Principal components analyses were performed and PC1 scores (>80% of variation) represented the dorsopalmar shape of the bone's central longitudinal axis and separated taxa consistently and in accord with known locomotor behavioral profiles. The most suspensory taxa, including orangutans, hylobatids and spider monkeys, had significantly lower PC1 scores reflecting the greatest amounts of phalangeal curvature. In contrast, bipedal humans and the quadrupedal cercopithecoid monkeys sampled (baboons, proboscis monkeys) exhibited significantly higher PC1 scores reflecting flatter phalanges. African ape (gorillas, chimpanzees and bonobos) phalanges fell between these two extremes and were not significantly different from each other. PC1 scores were significantly correlated with both included angle and the a coefficient of a second-degree polynomial calculated from the same landmark dataset, but had a significantly higher correlation with included angles. Our alternative approach for quantifying diaphysis shape of proximal phalanges to investigate dorsopalmar curvature is replicable and does not assume a priori either a circle or parabola model of shape, making it an attractive alternative compared with existing methodologies.

Redefining varicose projection astrocytes in primates.

Varicose projection astrocytes (VP-As) are found in the cerebral cortex and have been described to be specific to humans and chimpanzees. To further examine the phylogenetic distribution of this cell type, we analyzed cortical tissue from several primates ranging from primitive primates to primates evolutionary closer to human such as apes. We specifically analyzed tissue from four strepsirrhine species, one tarsier, six species of platyrrhine monkeys, ten species of cercopithecoid monkeys, two hylobatid ape species, four to six cases each of chimpanzee, bonobo, gorilla, and orangutan, and thirteen human. We found that VP-As were present only in human and other apes (hominoids) and were absent in all other species. We showed that VP-As are localized to layer VI and the superficial white matter of the cortex. The presence of VP-As co-occured with interlaminar astrocytes that also had varicosities in their processes. Due to their location, their long tangential processes, and their irregular presence within species, we propose that VP-As are astrocytes that develop varicosities under specific conditions and that are not a distinct astrocyte type.

Homo naledi pollical metacarpal shaft morphology is distinctive and intermediate between that of australopiths and other members of the genus Homo

Homo naledi fossils from the Rising Star cave system provide important insights into the diversity of hand morphology within the genus Homo. Notably, the pollical (thumb) metacarpal (Mc1) displays an unusual suite of characteristics including a median longitudinal crest, a narrow proximal base, and broad flaring intrinsic muscle flanges. The present study evaluates the affinities of H. naledi Mc1 morphology via 3D geometric morphometric analysis of shaft shape using a broader comparative sample (n = 337) of fossil hominins, recent humans, apes, and cercopithecoid monkeys than in prior work. Results confirm that the H. naledi Mc1 is distinctive from most other hominins in being narrow at the proximal end but surmounted by flaring muscle flanges distally. Only StW 418 (Australopithecus cf. africanus) is similar in these aspects of shape. The gracile proximal shaft is most similar to cercopithecoids, Pan, Pongo, Australopithecus afarensis, and Australopithecus sediba, suggesting that H. naledi retains the condition primitive for the genus Homo. In contrast, Neandertal Mc1s are characterized by wide proximal bases and shafts, pinched midshafts, and broad distal flanges, while those of recent humans generally have straight shafts, less robust muscle flanges, and wide proximal shafts/bases. Although uncertainties remain regarding character polarity, the morphology of the H. naledi thumb might be interpreted as a retained intermediate state in a transformation series between the overall gracility of the shaft and the robust shafts of later hominins. Such a model suggests that the addition of broad medial and lateral muscle flanges to a primitively slender shaft was the first modification in transforming the Mc1 into the overall more robust structure exhibited by other Homo taxa including Neandertals and recent Homo sapiens in whose shared lineage the bases and proximal shafts became expanded, possibly as an adaptation to the repeated recruitment of powerful intrinsic pollical muscles.

Reconstruction of the Early Miocene Critical Zone at Loperot, Southwestern Turkana, Kenya

The Hominoidea (apes and, eventually, humans) and Cercopithecoidea (Old World monkeys) diverged from a common ancestor during the late Oligocene (~25 Ma) in East Africa. Subsequently, both catarrhine groups diversified in the early Miocene (~23-16 Ma), making this time period an extremely important one for understanding our (deep) ancestral roots and the link between our hominoid ancestors and the environment. A remote region of west Turkana known as Loperot provides an exciting opportunity to study early Miocene primate paleoecology and has the potential to reveal the phylogenetic relationship between cercopithecoid monkeys and stem hominoids. The site of LpM4 is particularly fossiliferous and has produced a diverse fauna that includes several catarrhine fossils that await description. Radioisotopic dating indicates that the Loperot fauna are older than ~17 Ma (early Miocene). Using geochemical data from paleosols and stable isotope values from rhizolith calcite, coupled with detailed sedimentology and stratigraphy, we reconstruct the early Miocene Critical Zone of Loperot to reveal a large perennial river system fed by ephemeral streams that created localized riparian forest microhabitats within a larger open ecosystem under semi-arid climate conditions, making Loperot’s environmental reconstruction unlike many other early Miocene catarrhine-bearing localities. The perennial river system supported corridors of C3 forests; however, because these forests were restricted, they could not support a high diversity of forest indicator taxa. This is the most comprehensive analysis of the environmental conditions at Loperot to date, and provides new empirical evidence to suggest that African climate varied significantly by region and that arid habitats certainly existed during the early Miocene.

Foraging Profile, Activity Budget and Spatial Ecology of Exclusively Natural-Foraging Chacma Baboons (Papio ursinus) on the Cape Peninsula, South Africa

Exploitation of anthropogenic foods has profound effects on various aspects of the behavior and ecology of nonhuman primates. On the Cape Peninsula, South Africa, incentives to exploit novel resources are high and most local chacma baboons (Papio ursinus) feed on anthropogenic foods. Here we investigate the foraging profile, activity budget, and ranging behavior of a Peninsula group that feeds exclusively on natural foods. We predicted that the group would exploit the marine foods available to them and feed more extensively on subterranean foods, spend more time feeding and traveling, and occupy a larger area and travel further than groups that feed on anthropogenic foods, and groups elsewhere that occupy highly productive habitats. We tracked the group using handheld GPS units, and recorded behavioral data using instantaneous scan sampling in four seasons through 13 months. Our predictions were supported with the study group feeding on marine foods during all for seasons, but allocating a small percentage (<5%) of their total feeding time to exploiting these foods. Also, the group used a greater area (45.3 km2) and traveled further (5.9 km/day), and allocated more time to feeding and traveling, than previously studied groups on the Peninsula that exploit anthropogenic foods. These findings highlight behavioral variation in baboons associated with anthropogenic food use. Comparison of our results with those reported elsewhere should allow management authorities to more accurately predict changes in behavior of cercopithecoid monkeys as their access to anthropogenic foods is gradually restricted.

Arteriviruses, Pegiviruses, and Lentiviruses Are Common among Wild African Monkeys.

Nonhuman primates (NHPs) are a historically important source of zoonotic viruses and are a gold-standard model for research on many human pathogens. However, with the exception of simian immunodeficiency virus (SIV) (family Retroviridae), the blood-borne viruses harbored by these animals in the wild remain incompletely characterized. Here, we report the discovery and characterization of two novel simian pegiviruses (family Flaviviridae) and two novel simian arteriviruses (family Arteriviridae) in wild African green monkeys from Zambia (malbroucks [Chlorocebus cynosuros]) and South Africa (vervet monkeys [Chlorocebus pygerythrus]). We examine several aspects of infection, including viral load, genetic diversity, evolution, and geographic distribution, as well as host factors such as age, sex, and plasma cytokines. In combination with previous efforts to characterize blood-borne RNA viruses in wild primates across sub-Saharan Africa, these discoveries demonstrate that in addition to SIV, simian pegiviruses and simian arteriviruses are widespread and prevalent among many African cercopithecoid (i.e., Old World) monkeys. Primates are an important source of viruses that infect humans and serve as an important laboratory model of human virus infection. Here, we discover two new viruses in African green monkeys from Zambia and South Africa. In combination with previous virus discovery efforts, this finding suggests that these virus types are widespread among African monkeys. Our analysis suggests that one of these virus types, the simian arteriviruses, may have the potential to jump between different primate species and cause disease. In contrast, the other virus type, the pegiviruses, are thought to reduce the disease caused by human immunodeficiency virus (HIV) in humans. However, we did not observe a similar protective effect in SIV-infected African monkeys coinfected with pegiviruses, possibly because SIV causes little to no disease in these hosts.

Mandibular bone mineral density variation in three West African Cercopithecoid monkey species: Associations with diet and feeding behavior

ObjectiveProcolobus badius, Colobus polykomos, and Cercocebus atys are three West African primate species known to have distinctive feeding behaviors. Our objectives were (1) to test whether intra-taxon variation in bone mineral content exists between anterior and posterior regions of the mandible and (2) to determine if interspecific differences are interpretable via feeding and dietary idiosyncracies among the three taxa. DesignTwenty-one mandibles from naturally deceased C. polykomos (n=7), P. badius (n=9), and C. atys (n=5) were scanned using cone beam computerized tomography. Alveolar bone (AB) and basal cortical bone (CB) of incisor (I1) and second molar (M2) regions were digitally isolated. Degree of bone mineralization (DBM) distribution was obtained using histograms of CT attenuation values. Mean, standard deviation (SD), the 5th (Low5) and the 95th (High5) percentiles of the DBM histogram were compared between the jaw regions in species. ResultsThe mean and Low5 of DBM were significantly lower for AB than CB of all species (p<0.001). The AB DBM parameters were not significantly different between I1 and M2 of all species (p>0.056) except the mean of C. polykomos (p<0.05). The mean, SD, High5 of CB DBM at M2 of C. atys was significantly higher than those of C. polykomos and P. badius (p<0.006). ConclusionsThe durophagous C. atys had higher CB DBM value at the M2 region than C. polykomos and P. badius, which supports the hypothesis that materially stiffer mandibular bone in C. atys can develop in response to the generation of high bite forces during hard object feeding.

Giant pandas (Carnivora: Ailuropoda melanoleuca) and living hominoids converge on lumbar vertebral adaptations to orthograde trunk posture

Living hominoids share a common body plan characterized by a gradient of derived postcranial features that distinguish them from their closest living relatives, cercopithecoid monkeys. However, the evolutionary scenario(s) that led to the derived postcranial features of hominoids are uncertain. Explanations are complicated by the fact that living hominoids vary considerably in positional behaviors, and some Miocene hominoids are morphologically, and therefore probably behaviorally, distinct from modern hominoids. Comparative studies that aim to identify morphologies associated with specific components of positional behavioral repertoires are an important avenue of research that can improve our understanding of the evolution and adaptive significance of the hominoid postcranium. Here, we employ a comparative approach to offer additional insight into the evolution of the hominoid lumbar vertebral column. Specifically, we tested whether giant pandas (Carnivora: Ailuropoda melanoleuca) converge with living hominoids on lumbar vertebral adaptations to the single component of their respective positional behavioral repertoires that they share—orthograde (i.e., upright) trunk posture. We compare lumbar vertebral morphologies of Ailuropoda to those of other living ursids and caniform outgroups (northern raccoons and gray wolves). Mirroring known differences between living hominoids and cercopithecoids, Ailuropoda generally exhibits fewer, craniocaudally shorter lumbar vertebrae with more dorsally positioned transverse processes that are more dorsally oriented and laterally directed, and taller, more caudally directed spinous processes than other caniforms in the sample. Our comparative evidence lends support to a potential evolutionary scenario in which the acquisition of hominoid-like lumbar vertebral morphologies may have evolved for generalized orthograde behaviors and could have been exapted for suspensory behavior in crown hominoids and for other locomotor specializations (e.g., brachiation) in extant lineages.

The Middle Miocene Ape Pierolapithecus catalaunicus Exhibits Extant Great Ape-Like Morphometric Affinities on Its Patella: Inferences on Knee Function and Evolution

The mosaic nature of the Miocene ape postcranium hinders the reconstruction of the positional behavior and locomotion of these taxa based on isolated elements only. The fossil great ape Pierolapithecus catalaunicus (IPS 21350 skeleton; 11.9 Ma) exhibits a relatively wide and shallow thorax with moderate hand length and phalangeal curvature, dorsally-oriented metacarpophalangeal joints, and loss of ulnocarpal articulation. This evidence reveals enhanced orthograde postures without modern ape-like below-branch suspensory adaptations. Therefore, it has been proposed that natural selection enhanced vertical climbing (and not suspension per se) in Pierolapithecus catalaunicus. Although limb long bones are not available for this species, its patella (IPS 21350.37) can potentially provide insights into its knee function and thus on the complexity of its total morphological pattern. Here we provide a detailed description and morphometric analyses of IPS 21350.37, which are based on four external dimensions intended to capture the overall patellar shape. Our results reveal that the patella of Pierolapithecus is similar to that of extant great apes: proximodistally short, mediolaterally broad and anteroposteriorly thin. Previous biomechanical studies of the anthropoid knee based on the same measurements proposed that the modern great ape patella reflects a mobile knee joint while the long, narrow and thick patella of platyrrhine and especially cercopithecoid monkeys would increase the quadriceps moment arm in knee extension during walking, galloping, climbing and leaping. The patella of Pierolapithecus differs not only from that of monkeys and hylobatids, but also from that of basal hominoids (e.g., Proconsul and Nacholapithecus), which display slightly thinner patellae than extant great apes (the previously-inferred plesiomorphic hominoid condition). If patellar shape in Pierolapithecus is related to modern great ape-like knee function, our results suggest that increased knee mobility might have originally evolved in relation to enhanced climbing capabilities in great apes (such as specialized vertical climbing).

The relationship between bending stress and the shape of maxillary canines in cercopithecoid monkeys

Primate canines function in social displays but are also recruited for biting in agonistic encounters. Although the precise nature of the loads such behavior places on the canine crown is unknown, it is thought that bending is a major component of such loads. To date, modeling of canine bending strength has relied on idealized geometric representations. Accounting for the tapering of the crown as well as shape changes along an apical-basal axis provides a more realistic model for assessment of bending stress in canines. We provide such an accounting and evaluate the hypothesis that the morphology of the cercopithecoid canine represents a structural solution for maintaining constant maximum bending stress under apical or distributed loading in a parasagittal plane. This isostress hypothesis is analogous to a design criterion of minimum mass for a given structural requirement. Examining permanent maxillary canines from males and females representing eight West African cercopithecoid species, we reconstructed crown geometry from apex to base using microcomputed tomography. From reconstructed cross-sections, we determined section moduli about a buccolingual centroidal axis. We then determined what the taper should be for a variety of parasagittal loading distributions if the isostress hypothesis were true and compared these theoretical tapers to actual crown geometry. We found that a variety of loading distributions can be accommodated by the canines, particularly among males. These results suggest that our sample of canines are not optimized for resisting particular biting loads, but effectively limit stress gradients associated with a range of behaviors.

Partial skeleton of Theropithecus brumpti (Primates, Cercopithecidae) from the Chemeron Formation of the Tugen Hills, Kenya

Here we describe a complete skull and partial skeleton of a large cercopithecoid monkey (KNM-TH 46700) discovered in the Chemeron Formation of the Tugen Hills at BPRP Site #152 (2.63 Ma). Associated with the skeleton was a mandible of an infant cercopithecoid (KNM-TH 48364), also described here. KNM-TH 46700 represents an aged adult female of Theropithecus brumpti, a successful Pliocene papionin taxon better known from the Omo Shungura Formation in Ethiopia and sites east and west of Lake Turkana, Kenya. While the morphology of male T. brumpti is well-documented, including a partial skeleton with both cranial and postcranial material, the female T. brumpti morphotype is not well-known. This skeleton represents some of the first associated evidence of cranial and postcranial female T. brumpti remains. In addition to the complete skull, postcranial material includes elements of the axial skeleton and lower limb. While aspects of the skeleton conform to those of specimens previously assigned to T. brumpti, other features on the femur and tibia appear to differ from those previously described for this species. It is unclear whether these differences represent general variation within the T. brumpti population, variation between the sexes in T. brumpti, or the incorrect assignment of previous isolated hindlimb specimens. In total, the observable morphological features of the hindlimb suggest that KNM-TH 46700 was a terrestrial quadruped similar to modern savannah baboons ( Papio). From the available evidence, it is difficult to assess whether or not KNM-TH 46700 frequently engaged in the specialized squatting and shuffling behavior observed in extant geladas ( Theropithecus gelada).

The dangers of multi‐male groupings: trauma and healing in cercopithecoid monkeys from Cameroon

This study examines the potential linkage between social organization and trauma in a sample of cercopithecids from Cameroon. Skeletal trauma is described in a museum collection of eight sympatric monkey species. Macroscopic analysis was carried out on a total of 139 complete skeletons of mangabeys, colobines and guenons. Species in multi-male groups were found to have higher fracture frequencies than those in uni-male groups. These higher frequencies may be related to intra-specific male-male aggression; however, similarities in fracture patterning between males and females in multi-male groups suggest that other factors may be involved. Although fracture etiology may not be identified with certainty, this study suggests that predation may indirectly be a cause of traumatic injuries in those species of cercopithecid monkeys displaying multi-male social organizations. The data presented also highlight the utility of museum collections as an additional resource in analyses of primate behavior, demonstrating that behavioral information does not die when the animal does.

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.

Masticatory stress and the mechanics of “wishboning” in colobine jaws

Cercopithecoid monkeys experience relatively high strains along the lingual aspect of the mandibular symphysis because of lateral transverse bending of the mandibular corpora ("wishboning") during mastication. Hylander (Am J Phys Anthropol 64 (1984) 1-46; Am Zool 25 (1985) 315-330) demonstrated that the distribution of strains arising from wishboning loads is comprehensible with reference to the mechanics of curved beams. Theory of curved beams suggests that lingual tensile strains are some multiple of labial compressive strains, yet limitations of experimental methods and uncertainty in estimating parameters needed for theoretical calculations have confounded attempts to characterize the magnitude of this disparity of normal strains. We evaluate the theoretical disparity of normal strains in wishboning in comparison to in vitro strains collected under controlled loads for a sample of mandibles representing two colobine species (N = 6). These data suggest that in colobine monkeys, maximum normal lingual strains should be at least twice maximum labial strains. In addition, we reexamine the distribution of symphyseal stress under an assumption of asymmetric bending, a general approach for calculation of stress appropriate for members that lack a plane of symmetry and are bent along an axis that is not coincident with the member's principal axes. Under asymmetric bending in colobine mandibles, the effect of symphyseal inclination on lingual strain is mitigating at the superior transverse torus and exacerbating at the inferior transverse torus. Relative compliance of colobine mandibular bone further supports the hypothesis that the structural and material properties of the colobine mandibular symphysis do not represent a morphological strategy for minimizing masticatory strain.

Precision and Consistency of the Taphonomic Signature of Predation by Crowned Hawk-Eagles (Stephanoaetus coronatus) in Kibale National Park, Uganda

Abstract Prey material, which was collected from beneath the nests of two pairs of crowned hawk-eagles (Stephanoaetus coronatus) in the Ngogo study area, Kibale National Park, Uganda, was analyzed to identify the taphonomic features diagnostic of eagle predation. Material from a recent three-year collection interval was compared to an earlier three-year collection subsample to determine signal consistency over time. The bulk of the assemblage is comprised of cercopithecoid monkey remains, reflecting the taxonomic composition of the Ngogo area. Taxonomic composition of the assemblage remains highly consistent over the collection period. Bone survivability and fragmentation profiles, as well as damage patterns, also were investigated. Crowned hawk-eagles inflict much less damage to prey skeletons than do mammalian carnivores. Crania, scapulae, and hindlimb elements are most likely to survive predation by crowned hawk-eagles and to be concentrated at nest sites, but these elements (especially crania) show ag...

Red-tailed monkeys (Cercopithecus ascanius) hunt green pigeons (Treron calva) in the Kalinzu Forest in Uganda

Red-tailed monkeys (Cercopithecus ascanius) were observed hunting green pigeons (Treron calva) in the Kalinzu Forest in Uganda. During 2 h 39 min, I observed two cases of successful hunting and one case of unsuccessful hunting in a Ficus saussureana tree. Red-tailed monkeys stalked the pigeons until they were within 2-3 m, and then jumped and caught them. In both successful cases, blue monkeys (C. mitis) ran to the hunting site from adjacent trees in order to poach the prey, and the red-tailed monkeys fled. One of these red-tailed monkeys dropped the pigeon while fleeing, and the blue monkey climbed down from the tree to search for it. This is the first record of cercopithecoid monkeys hunting birds that are outside of the nest and moving freely, and also the first record of red-tailed monkeys hunting vertebrates. However rare it may be, the repeated hunting attempts using similar techniques and the immediate attempt of blue monkeys to poach the prey suggest that this type of hunting was not a one-time event that happened by chance. Blue monkeys and an adult chimpanzee (Pan troglodytes schweinfurthii) in and around the fig tree did not attempt to hunt. The hunting of volant birds may be enabled by the small body size and the quick movements of red-tailed monkeys.

Paleoenvironmental basis of cognitive evolution in great apes.

A bias favoring tree-dominated habitats and ripe-fruit frugivory has persisted in great ape evolution since the early Miocene. This bias is indicated by fossil ape paleoenvironments, molar morphology, dental microwear, the geographic pattern of extinctions, and extant apes' reliance on wooded settings. The ephemeral aspect of high-quality fruit has placed a premium on cognitive and social means of finding and defending food sources, and appears related to great apes' affinity since the Miocene for wooded, fruit-rich environments. These habitats have, however, undergone a severe withdrawal toward the low latitudes of Africa and Southeast Asia since the late Miocene, corresponding to a decline in the diversity of great apes beginning 9.5 million years ago. Plio-Pleistocene records imply that wooded settings of Africa and SE Asia were prone to substantial fragmentation and coalescence. Once apes were confined to equatorial settings, therefore, habitat instability heightened the spatial/temporal uncertainty of ripe-fruit sources. Prolonged learning, the assignment of attributes to distant places, mental representation, and reliance on fallback foods were all favored in this dynamic environmental context. These abilities helped sustain forest frugivory in most lineages. Fluid social grouping afforded the animals opportunities to locate ephemeral foods in continuous and fragmented forests. Fission-fusion grouping also magnified the problems of object impermanence (of individuals) and dispersion manifested by food sources in the ecological realm. Thus the spatial and temporal dynamics of fruit and wooded habitats since the Miocene are reflected in important components of great ape cognition, foraging, and sociality. In contrast to great apes, cercopithecoid monkeys have increased their plant dietary options and diversified in seasonal environments since the late Miocene. Early hominins eventually severed the habitat bias that characterized the evolution of great apes, and later expanded into diverse environments.

Taphonomic aspects of crowned hawk-eagle predation on monkeys.

This study provides a taphonomic analysis of prey accumulations of crowned hawk-eagles (Stephanoaetus coronatus) from Ngogo, Kibale National Park, Uganda, collected over 37 months from below nests of two eagle pairs. Crowned hawk-eagles are powerful predators capable of killing animals much larger than themselves, and are significant predators of cercopithecoid monkeys in forest habitats throughout sub-Saharan Africa. At Ngogo, 81% of the individuals in the kill sample are monkeys. Redtail monkeys (Cercopithecus ascanius) are particularly well represented in the sample, making up 66% of monkeys identified to species. Despite an impressive killing apparatus, crowned hawk-eagles are fastidious eaters that inflict far less damage to bone than mammalian predators. Examination of skeletal material from the Ngogo kill sample reveals that crania, hindlimb elements, and scapulae survive predation better than do other bones. Crania of adults are typically complete and accompanied by mandibles, while crania of young individuals are usually dissociated from mandibles and lack basicrania and faces. Long bones are often whole or show minimal damage. Thin bones, such as crania and innominates, are marked by numerous nicks, punctures, and "can-opener" perforations. Scapular blades are heavily raked and shattered. Along with the strong preference for cercopithecoids, these distinct patterns of bone survival and damage indicate the feasibility of recognizing specific taphonomic signatures of large raptors in fossil assemblages. Berger and Clarke (1995) hypothesized that crowned hawk-eagles or similar large raptors were principally responsible for the accumulation of the late Pliocene fossil fauna from Taung, South Africa, including the type infant skull of Australopithecus africanus. The results of our study suggest that the faunal composition and type of damage to the hominid skull and other bone from Taung are consistent with the predatory activities of large raptors. More rigorous assessment of their hypothesis will require sorting the Taung fauna by locality and further detailed analysis of species composition and bone damage and survivability patterns.

Paleosols and paleoenvironments of the middle Miocene, Maboko Formation, Kenya.

The middle Miocene (15 Ma) Maboko Formation of Maboko Island and Majiwa Bluffs, southwestern Kenya, has yielded abundant fossils of the earliest known cercopithecoid monkey (Victoriapithecus macinnesi), and of a kenyapithecine hominoid (Kenyapithecus africanus), as well as rare proconsuline (Simiolus leakeyorum, cf. Limnopithecus evansi) and oreopithecine apes (Mabokopithecus clarki, M. pickfordi), and galagids (Komba winamensis). Specific habitat preferences can be interpreted from large collections of primate fossils in different kinds of paleosols (pedotypes). Fossiliferous drab-colored paleosols with iron-manganese nodules (Yom pedotype) are like modern soils of seasonally waterlogged depressions (dambo). Their crumb structure and abundant fine root-traces, as well as scattered large calcareous rhizoconcretions indicate former vegetation of seasonally wet, wooded grassland. Other fossiliferous paleosols are evidence of nyika bushland (Ratong), and early-successional riparian woodland (Dhero). No fossils were found in Mogo paleosols interpreted as saline scrub soils. Very shallow calcic horizons (in Yom, Ratong, and Mogo paleosols) and Na-montmorillonite (in Mogo) are evidence of dry paleoclimate (300-500 mm MAP=mean annual precipitation). This is the driest paleoclimate and most open vegetation yet inferred as a habitat for any Kenyan Miocene apes or monkeys. Victoriapithecus was abundant in dambo wooded grassland (Yom) and riparian woodland (Dhero), a distribution like that of modern vervet monkeys. Kenyapithecus ranged through all these paleosols, but was the most common primate in nyika bushland paleosols (Ratong), comparable to baboons and macaques today. Mabokopithecus was virtually restricted to riparian woodland paleosols (Dhero), and Simiolus had a similar, but marginally wider, distribution. Habitat preferences of Mabokopithecus and Simiolus were like those of modern colobus monkeys and mangabeys. A single specimen of Komba was found in dambo wooded grassland paleosol (Yom), a habitat more like that of the living Senegal bushbaby than of rainforest galagids. A shift to non-forest habitats may explain the terrestrial adaptations of Victoriapithecus, basal to the cercopithecid radiation, and of Kenyapithecus, basal to the hominoid radiation. Both taxa are distinct from earlier Miocene arboreal proconsulines, oreopithecines and galagids.