Nacholapithecus Research Articles

New quantitative analyses of the Nacholapithecus kerioi proximal ulna confirm morphological affinities with Equatorius and large papionins.

The elbow of Nacholapithecus has been extensively described qualitatively, however its ulnar morphology has never been the focus of an in-depth quantitative analysis before. Hence, our main aim is quantifying the proximal ulnar morphology in Nacholapithecus and exploring whether it is similar to those of Equatorius and Griphopithecus as previously reported. We compared Nacholapithecus proximal ulnar morphology with a sample of extant and extinct anthropoids through principal component analysis and agglomerative hierarchical cluster analysis. Moreover, we calculated the Cophenetic Correlation Coefficient and checked for taxonomical group mean differences through MANOVA and pairwise post-hoc comparisons, as well as the phylogenetic signal in the variables used. The Nacholapithecus ulna displays a moderately long and relatively narrow olecranon, a relatively wide trochlear surface-radial notch width, and a relatively thin sigmoid notch depth. These features resemble those of large papionins and chimpanzees, and some extinct taxa, mainly Equatorius. Results presented here reinforce previous inferences on the functional morphology of the Nacholapithecus elbow, showing adaptations for general quadrupedal behaviors. However, other derived features (e.g., a relatively wide trochlear surface) might be associated with the ape-like traits described for its distal humerus (e.g., wide trochlear groove), thus displaying a combination of primitive and derived features in the proximal ulna. Finally, affinities with large papionins could suggest the presence of some terrestrial habits in Nacholapithecus. However, the lack of evidence in the rest of the skeleton prevents us from suggesting terrestrial affinities in this taxon in a conclusive manner.

The radial head of the Middle Miocene ape Nacholapithecus kerioi: Morphometric affinities, locomotor inferences, and implications for the evolution of the hominoid humeroradial joint

The radial head of the Middle Miocene ape Nacholapithecus kerioi: Morphometric affinities, locomotor inferences, and implications for the evolution of the hominoid humeroradial joint

New femoral remains of Nacholapithecus kerioi: Implications for intraspecific variation and Miocene hominoid evolution

The middle Miocene stem kenyapithecine Nacholapithecus kerioi (16–15 Ma; Nachola, Kenya) is represented by a large number of isolated fossil remains and one of the most complete skeletons in the hominoid fossil record (KNM-BG 35250). Multiple fieldwork seasons performed by Japanese–Kenyan teams during the last part of the 20th century resulted in the discovery of a large sample of Nacholapithecus fossils. Here, we describe the new femoral remains of Nacholapithecus. In well-preserved specimens, we evaluate sex differences and within-species variation using both qualitative and quantitative traits. We use these data to determine whether these specimens are morphologically similar to the species holotype KNM-BG 35250 (which shows some plastic deformation) and to compare Nacholapithecus with other Miocene hominoids and extant anthropoids to evaluate the distinctiveness of its femur. The new fossil evidence reaffirms previously reported descriptions of some distal femoral traits, namely the morphology of the patellar groove. However, results also show that relative femoral head size in Nacholapithecus is smaller, relative neck length is longer, and neck–shaft angle is lower than previously reported for KNM-BG 35250. These traits have a strong functional signal related to the hip joint kinematics, suggesting that the morphology of the proximal femur in Nacholapithecus might be functionally related to quadrupedal-like behaviors instead of more derived antipronograde locomotor modes. Results further demonstrate that other African Miocene apes (with the exception of Turkanapithecus kalakolensis) generally fall within the Nacholapithecus range of variation, whose overall femoral shape resembles that of Ekembo spp. and Equatorius africanus. Our results accord with the previously inferred locomotor repertoire of Nacholapithecus, indicating a combination of generalized arboreal quadrupedalism combined with other antipronograde behaviors (e.g., vertical climbing).

New forelimb long bone specimens of <i>Nacholapithecus kerioi</i> from the Middle Miocene of northern Kenya

This article reports eight new humeral, ulnar, and radial fragments of Nacholapithecus kerioi collected from Nachola, Kenya during the 1998/1999 field seasons. The study refines the description of its forelimb bones, which was mostly based on a single partial skeleton. The most distinctive feature of the distal humerus is a large, globular, medially tilted capitulum. The groove between the capitulum and the zona conoidea is quite deep. The medial part of the humeral trochlea is also diagnostic in showing a less salient medial border. The medial epicondyle is moderately long and more posteriorly reflected than was previously presumed. The coronoid process of the ulna is quite wide. Its medial portion is distinctly concave. The ulnar shaft is anteroposteriorly deep in its proximal half, slender, straight in frontal view, and weakly anteriorly bowing. The elbow of Nacholapithecus exhibits a primitive functional pattern as a hominoid, including lack of universal stability of the humeroulnar joint through full extension and flexion, restriction of hyperextension of the elbow, and relatively anteroposteriorly oriented loading at the proximal ulna. On the other hand, it is derived in terms of enhanced rotational mobility and stability of the radius, incipiently increased stability at the humeroulnar joint, and more frequent maximum extension of the elbow compared to proconsulids. This mosaic morphology is different from both early Miocene proconsulids and later suspensory or orthograde European fossil apes. Although Nacholapithecus was neither suspensory nor orthograde, its forelimbs may have played a greater role for body support or balance maintenance, more frequently reaching to and exploiting overhead supports than in early Miocene proconsulids.

Sexual dimorphism of body size in an African fossil ape, Nacholapithecus kerioi

Sexual size dimorphism in the African fossil ape Proconsul nyanzae (18 million years ago, 18 Ma) has been previously documented. However, additional evidence for sexual dimorphism in Miocene hominoids can provide great insight into the history of extant hominoid mating systems. The present study focused on body mass (BM) sexual dimorphism in Nacholapithecus kerioi from the Middle Miocene (16–15 Ma) in Africa. Bootstrap analysis revealed that P. nyanzae BM sexual dimorphism was lower than that in Pan troglodytes, which exhibits moderate sexual dimorphism, as reported previously. The same simulation revealed that BM sexual dimorphism of N. kerioi was comparable with that in Gorilla spp.; i.e., the males were approximately twice as large as the females. High sexual dimorphism in extant apes is usually indicative of a polygynous social structure (gorilla) or solitary/fission-fusion social system (orangutan). However, because of the high proportion of adult males in this fossil assemblage, the magnitude of dimorphism inferred here cannot be associated with a gorilla-like polygynous or oranguran-like solitary/fission-fusion social structure, and may reflect either taphonomic bias, or some other social structure. Extant hominoids have a long evolutionary history owing to their deep branching, comprising only a few existing members of the original highly successful group. Therefore, it is not surprising that the mating systems of extant hominoids fail to provide fossil apes with a perfect “model”. The mating systems of extinct hominoids may have been more diverse than those of extant apes.

Sacral vertebral remains of the Middle Miocene hominoid Nacholapithecus kerioi from northern Kenya

This study describes two new sacral specimens of Nacholapithecus kerioi, KNM-BG 42753I and KNM-BG 47687A, from the Aka Aiteputh Formation in Nachola, northern Kenya, excavated in 2002. They are of roughly equal size and are considered to belong to males. When scaled by body mass, the lumbosacral articular surface area of the better preserved specimen, KNM-BG 42753I, is smaller than that in Old World monkeys but similar to that in extant great apes and New World monkeys, as well as Proconsul nyanzae. The relatively narrow dimensions of the first sacral vertebral body in the transverse and sagittal planes are characteristics of N. kerioi and P. nyanzae and similar to those of extant great apes. In N. kerioi, lumbosacral surface area relative to body mass is small. This may simply be an extension of a trend from the previously reported small thoracolumbar vertebrae to the sacrum. ​The first sacral vertebrae of N. kerioi and Epipliopithecus vindobonensis have a higher craniocaudal vertebral body reduction (CVR; a higher CVR indicates a wider cranial width relative to a narrower caudal width), similar to that in Old World monkeys. Old World monkeys have a higher CVR, and usually have three sacral vertebrae, fewer than seen in extant great apes, which have a lower CVR and four to six (sometimes as many as eight) sacral vertebrae. New World monkeys have a lower CVR than Old World monkeys, but generally possess only three sacral vertebrae, and have a large caudal articular surface, which may be related, at least in the Atelidae, to the grasping ability of their tails. The possibility that N. kerioi had only three sacral vertebrae cannot be ruled out, because E. vindobonensis and Old World monkeys, with higher CVRs, have sacra consisting of three sacral vertebrae.

ケニア中期中新世の類人猿Nacholapithecus kerioiにおける下位胸椎―椎体サイズと棘突起について―

ケニア中期中新世の類人猿Nacholapithecus kerioiにおける下位胸椎―椎体サイズと棘突起について―

Morphology of the thoracolumbar spine of the middle Miocene hominoid Nacholapithecus kerioi from northern Kenya

A new caudal thoracic and a new lumbar vertebra of Nacholapithecus kerioi, a middle Miocene hominoid from northern Kenya, are reported. The caudal thoracic vertebral body of N. kerioi has a rounded median ventral keel and its lateral sides are moderately concave. The lumbar vertebral body has an obvious median ventral keel. Based on a comparison of vertebral body cranial articular surface size between the caudal thoracic vertebrae in the present study and one discussed in a previous study (KNM-BG 35250BO, a diaphragmatic vertebra), N. kerioi has at least two post-diaphragmatic vertebrae (rib-bearing lumbar-type thoracic vertebrae), unlike extant hominoids. It also has thick, rounded, and moderately long metapophyses on the lumbar vertebra that project dorsolaterally. The spinous process bases of its caudal thoracic and lumbar vertebrae originate caudally between the postzygapophyses, as described previously in the KNM-BG 35250 holotype specimen. In other words, the postzygapophyses of N. kerioi do not project below the caudal border of the spinous processes, similar to those of extant great apes, and unlike small apes and monkeys, which have more caudally projecting postzygapophyses. Nacholapithecus kerioi has a craniocaudally expanded spinous process in relation to vertebral body length, also similar to extant great apes. Both these spinous process features of N. kerioi differ from those of Proconsul nyanzae. The caudal thoracic vertebra of N. kerioi has a caudally-directed spinous process, whose tip is tear-drop shaped. These features resemble those of extant apes. The morphology of the spinous process tips presumably helps vertebral stability by closely stacking adjacent spinous process tips as seen in extant hominoids. The morphology of the spinous process and postzygapophyses limits the intervertebral space and contributes to the stability of the functional lumbar region as seen in extant great apes, suggesting that antipronograde activity was included in the positional behavior of N. kerioi.

New endemic platyrrhine femur from Haiti: Description and locomotor analysis

The middle Miocene stem kenyapithecine Nacholapithecus kerioi (16–15 Ma; Nachola, Kenya) is represented by a large number of isolated fossil remains and one of the most complete skeletons in the hominoid fossil record (KNM-BG 35250). Multiple fieldwork seasons performed by Japanese–Kenyan teams during the last part of the 20th century resulted in the discovery of a large sample of Nacholapithecus fossils. Here, we describe the new femoral remains of Nacholapithecus. In well-preserved specimens, we evaluate sex differences and within-species variation using both qualitative and quantitative traits. We use these data to determine whether these specimens are morphologically similar to the species holotype KNM-BG 35250 (which shows some plastic deformation) and to compare Nacholapithecus with other Miocene hominoids and extant anthropoids to evaluate the distinctiveness of its femur. The new fossil evidence reaffirms previously reported descriptions of some distal femoral traits, namely the morphology of the patellar groove. However, results also show that relative femoral head size in Nacholapithecus is smaller, relative neck length is longer, and neck–shaft angle is lower than previously reported for KNM-BG 35250. These traits have a strong functional signal related to the hip joint kinematics, suggesting that the morphology of the proximal femur in Nacholapithecus might be functionally related to quadrupedal-like behaviors instead of more derived antipronograde locomotor modes. Results further demonstrate that other African Miocene apes (with the exception of Turkanapithecus kalakolensis) generally fall within the Nacholapithecus range of variation, whose overall femoral shape resembles that of Ekembo spp. and Equatorius africanus. Our results accord with the previously inferred locomotor repertoire of Nacholapithecus, indicating a combination of generalized arboreal quadrupedalism combined with other antipronograde behaviors (e.g., vertical climbing).

Functional morphology and anatomy of cervical vertebrae in Nacholapithecus kerioi, a middle Miocene hominoid from Kenya

This paper describes the morphology of cervical vertebrae in Nacholapithecus kerioi, a middle Miocene primate species excavated from Nachola, Kenya in 1999–2002. The cervical vertebrae in Nacholapithecus are larger than those of Papio cynocephalus. They are more robust relative to more caudal vertebral bones. Since Nacholapithecus had large forelimbs, it is assumed that strong cervical vertebrae would have been required to resist muscle reaction forces during locomotion. On the other hand, the vertebral foramen of the lower cervical vertebrae in Nacholapithecus is almost the same size as or smaller than that of P. cynocephalus. Atlas specimens of Nacholapithecus resemble those of extant great apes with regard to the superior articular facet, and they have an anterior tubercle trait intermediate between that of extant apes and other primate species. Nacholapithecus has a relatively short and thick dens on the axis, similar to those of extant great apes and the axis body shape is intermediate between that of extant apes and other primates. Moreover, an intermediate trait between extant great apes and other primate species has been indicated with regard to the angle between the prezygapophyseal articular facets of the axis in Nacholapithecus. Although the atlas of Nacholapithecus is inferred as having a primitive morphology (i.e., possessing a lateral bridge), the shape of the atlas and axis leads to speculation that locomotion or posture in Nacholapithecus involved more orthograde behavior similar to that of extant apes, and, in so far as cervical vertebral morphology is concerned, it is thought that Nacholapithecus was incipiently specialized toward the characteristics of extant hominoids.

Hind limb of the <i>Nacholapithecus kerioi</i> holotype and implications for its positional behavior

We present the definitive description of the hind limb elements of the Nacholapithecus kerioi holotype (KNM-BG 35250) from the middle Miocene of Kenya. Previously, it has been noted that the postcranial (i.e. the phalanges, spine, and shoulder girdle) anatomy of N. kerioi shows greater affinity to other early/middle Miocene African hominoids, collectively called ‘non-specialized pronograde arboreal quadrupeds,’ than to extant hominoids. This was also the case for the hind limb. However, N. kerioi exhibits a unique combination of postcranial characters that distinguish this species from other early/middle Miocene African hominoids. The femoral neck has a high angle, but is relatively short, though the adaptive meaning of this form is not readily understood. In the distal femur, the shape of the patellar surface and symmetry of the femoral condylar widths suggest that the knee was not typically abducted but assumed more variable movements and/or postures. Whereas the morphologies of the talocrural and intertarsal joints are generally similar to those of the other fossil hominoids, the tibial malleolus is extremely thick and asymmetry of the talar trochlea groove is more emphasized due to a more prominent lateral trochlear rim than in Proconsul and other African fossil hominoids. The distal foot segment is more elongated. The (non-hallucal) metatarsals appear relatively gracile due to the elongation. The proximal joints of these metatarsals are, nonetheless, large. Ligamentous attachments of the tarsal/metatarsal bones are generally well developed. The distal tarsal row, which is represented only by the medial cuneiform, though, is extremely large for the presumed body mass. In terms of function, the femur, ankle, and tarsal joints are interpreted behaviorally to represent a slow-moving arboreal quadruped. However, the foot of N. kerioi appears to be more specialized for inverted grasping and subvertical support use. All of these foot features are suggestive of a greater role for antiprono-grade activities in N. kerioi relative to other Miocene ‘pronograde arboreal quadrupeds.’

Nacholapithecus and its importance for understanding hominoid evolution

Nacholapithecus kerioi is a large-sized hominoid from the Aka Aiteputh Formation (15 Ma) in Nachola, northern Kenya.1 While eight large-sized hominoid species dating to the late Early to early Middle Miocene (17-14 Ma) are known in Afro-Arabia and western Eurasia,2-6 the facial and postcranial anatomy of these apes is poorly known. However, much has been learned of the craniodental and postcranial anatomy of N. kerioi over the last ten years (A list of published specimens is available online, accompanying this article), and it plays a key role in our understanding of hominoid evolution in the Early to Middle Miocene of Africa and Eurasia. Importantly, it bears on the interpretation of the hominoid Morotopithecus bishopi from 20.6 my-old Uganda.7-10 In the article, we provide information on the anatomy and adaptations of N. kerioi as well as on the site of Nachola, and discuss how our current knowledge of N. kerioi can be incorporated into scenarios of hominoid evolution.

Comparative study of Moroto vertebral specimens

The hypodigm of Morotopithecus bishopi includes several vertebral specimens from Moroto II in addition to a scapular fragment, and femoral and craniodental specimens. The Moroto vertebral specimens include UMP 67.28, which is a well-preserved lumbar vertebra. Based on the derived morphological traits in UMP 67.28, together with evidence from other postcranial elements, it has been claimed that certain aspects of the modern hominoid body plan appeared in the hominoid lineage by as early as 20 Ma. Other vertebral specimens from the site are not well-preserved and have not been described in detail. This article provides the first detailed description of these specimen with an emphasis on a lumbar vertebral body UMP 68.06. Results confirm the existing interpretations that M. bishopi had a more dorsostable lumbar column compared to other African Miocene hominoids, such as Proconsul nyanzae/heseloni or Nacholapithecus kerioi. The vertebral body is craniocaudally short and the median ventral keel is absent through the lumbar column. However, M. bishopi might have had a similar number segments as inferred for P. nyanzae (6–7) if UMP 68.06 and UMP 67.28 are associated. Likewise, the ventral wedging of UMP 68.06 may suggest that M. bishopi had more lumbar vertebrae than extant great apes. The origin of the transverse process relative to the vertebral body is variable by level among the Moroto specimens. Thus, if these specimens derive from a single taxon, this may suggest considerable variability in this feature that would advise caution when using this feature to draw taxonomic or functional conclusions.

Postcranial bones of infant Nacholapithecus: ontogeny and positional behavioral adaptation

We describe the postcranial bones of a partial skeleton of a male infant Nacholapithecus kerioi (KNM-BG 37800) from the Middle Miocene of Kenya. The skeletal elements are associated with dental remains and its age is presumed to be 6–15 months based on a comparison with an infant Proconsul heseloni with a similar dental growth stage. The postcranial elements include the left scapula, a lumbar vertebra, left distal humerus, right proximal ulna, right radius, right and left proximal femora, a proximal fragment of metatarsal, and proximal and intermediate phalanges. Its body mass was estimated as 3.0 kg from shaft dimensions of the humerus and femur. At this early postnatal developmental stage, N. kerioi can be distinguished from P. heseloni in several postcranial features: a caudally projecting lumbar spinous process, strong dorsal divergence of the scapular spine, a coronoid fossa which is more developed than the radial fossa, higher neck shaft angle of the femur, taller trochlea of phalanges, and a lower brachial index. These features, except for the last, agree with the previously proposed positional behavioral reconstruction of N. kerioi, in which vertical climbing, orthograde clambering, transferring, or bridging are relatively important compared with P. heseloni. The apparently low brachial index results from a comparatively short radius for the predicted body mass. This might imply a relatively slow pace of growth in the radius of N. kerioi. However, it might result from body mass overestimation, incorrect length reconstruction, or individual variation. Its interpretation must await further specimens.

Vertebral morphology of Nacholapithecus kerioi based on KNM-BG 35250

This paper describes the morphology of the vertebral remains of the KNM-BG 35250 Nacholapithecus kerioi individual from the Middle Miocene of Kenya. Cervical vertebrae are generally large relative to presumed body mass, suggesting a heavy head with large jaws and well-developed neck muscles. The atlas retains the lateral and posterior bridges over the vertebral artery. The axis has a robust dens and a large angle formed by superior articular surfaces. The thoracic vertebral specimens include the diaphragmatic vertebra and one post-diaphragmatic vertebra. The thoracic vertebral bodies are much smaller that those of male Papio cynocephalus, whereas many of the dorsal elements are large and robust, exceeding those of male P. cynocephalus. Lumbar vertebral bodies are small relative to body mass, craniocaudally moderately long, and have a median ventral keel. The transverse process is craniocaudally long and arises from the widest part of the body cranially and the pedicle above the inferior vertebral notch caudally. Anapophyses are present in one of the preserved lumbar vertebrae. The postzygapophyses are thick dorsoventrally. These lumbar features are broadly shared with Proconsul. However, the base of the spinous process is longer and more caudally positioned in N. kerioi compared to Proconsul, and is more similar to the condition in Pongo. They are not dorsally (or moderately caudally) directed as is seen in P. nyanzae, Pan, and most other extant primates. A caudally directed spinous process does not permit a broad range of spinal dorsiflexion. The presumed stiff back in N. kerioi suggests a different locomotor repertoire than in Proconsul. Morotopithecus bishopi, although not possessing the same features, exhibits another morphological suite of characters for lumbar stiffness. Diverse functional adaptations of the lumbar spine were present in African hominoids during the Early to Middle Miocene.

Tail loss in Proconsul heseloni

The middle Miocene stem kenyapithecine Nacholapithecus kerioi (16–15 Ma; Nachola, Kenya) is represented by a large number of isolated fossil remains and one of the most complete skeletons in the hominoid fossil record (KNM-BG 35250). Multiple fieldwork seasons performed by Japanese–Kenyan teams during the last part of the 20th century resulted in the discovery of a large sample of Nacholapithecus fossils. Here, we describe the new femoral remains of Nacholapithecus. In well-preserved specimens, we evaluate sex differences and within-species variation using both qualitative and quantitative traits. We use these data to determine whether these specimens are morphologically similar to the species holotype KNM-BG 35250 (which shows some plastic deformation) and to compare Nacholapithecus with other Miocene hominoids and extant anthropoids to evaluate the distinctiveness of its femur. The new fossil evidence reaffirms previously reported descriptions of some distal femoral traits, namely the morphology of the patellar groove. However, results also show that relative femoral head size in Nacholapithecus is smaller, relative neck length is longer, and neck–shaft angle is lower than previously reported for KNM-BG 35250. These traits have a strong functional signal related to the hip joint kinematics, suggesting that the morphology of the proximal femur in Nacholapithecus might be functionally related to quadrupedal-like behaviors instead of more derived antipronograde locomotor modes. Results further demonstrate that other African Miocene apes (with the exception of Turkanapithecus kalakolensis) generally fall within the Nacholapithecus range of variation, whose overall femoral shape resembles that of Ekembo spp. and Equatorius africanus. Our results accord with the previously inferred locomotor repertoire of Nacholapithecus, indicating a combination of generalized arboreal quadrupedalism combined with other antipronograde behaviors (e.g., vertical climbing).

Maxillae and associated gnathodental specimens of Nacholapithecus kerioi, a large-bodied hominoid from Nachola, northern Kenya

The middle Miocene large-bodied hominoid from Nachola, initially attributed to Kenyapithecus, was recently transferred to a new genus and species Nacholapithecus kerioi. The hypodigm of N. kerioi consists of numerous maxillae, mandibles, and isolated teeth, as well as a number of postcranial bones. A detailed description of the previously discovered postcranial material has already been presented. This article aims to give a detailed description of maxillary specimens (including some mandibular fragments associated with them) of N. kerioi collected by the Japan–Kenya Joint Project team during the field seasons of 1982, 1984, and 1986. The maxillary specimens of N. kerioi retain a set of primitive catarrhine features, such as a relatively shallow palate, low position of the anterior zygomatic root, and the lack of enlarged premolars. Yet, compared to the Early Miocene Proconsul, N. kerioi is derived in having a moderately elongated subnasal clivus that appears to have overlapped the hard palate.

Nacholapithecus skeleton from the Middle Miocene of Kenya

An almost entire skeleton of a male individual of Nacholapithecus kerioi (KNM-BG 35250) was discovered from Middle Miocene (∼15 Ma) sediments at Nachola, northern Kenya. N. kerioi exhibits a shared derived subnasal morphology with living apes. In many postcranial features, such as articular shape, as well as the number of the lumbar vertebrae, N. kerioi resembles Proconsul heseloni and/or P. nyanzae, and lacks suspensory specializations characteristic of living apes. Similarly, N. kerioi shares some postcranial characters with Kenyapithecus spp. However, despite the resemblance, N. kerioi and Proconsul spp. are quite different in their body proportions and some joint morphologies. N. kerioi has proportionally large forelimb bones and long pedal digits compared to its hindlimb bones and lumbar vertebrae. Its distinctive body proportions suggest that N. kerioi was more derived for forelimb dominated arboreal activities than P. nyanzae and P. heseloni. On the other hand, it exhibits a mixture of derived and primitive cranio-dental and postcranial features relative to the contemporaneous Kenyapithecus and Early Miocene Morotopithecus. While the phylogenetic position of N. kerioi is unsettled, it seems necessary to posit parallel evolution of cranio-dental and/or postcranial features in fossil and living apes.

New Hominoid Genus from the Middle Miocene of Nachola, Kenya.

A new genus and species, Nacholapithecus kerioi have been erected for the large-bodied Miocene hominoid specimens discovered from Nachola, Kenya.

A newly discovered Kenyapithecus skeleton and its implications for the evolution of positional behavior in Miocene East African hominoids.

The middle Miocene stem kenyapithecine Nacholapithecus kerioi (16–15 Ma; Nachola, Kenya) is represented by a large number of isolated fossil remains and one of the most complete skeletons in the hominoid fossil record (KNM-BG 35250). Multiple fieldwork seasons performed by Japanese–Kenyan teams during the last part of the 20th century resulted in the discovery of a large sample of Nacholapithecus fossils. Here, we describe the new femoral remains of Nacholapithecus. In well-preserved specimens, we evaluate sex differences and within-species variation using both qualitative and quantitative traits. We use these data to determine whether these specimens are morphologically similar to the species holotype KNM-BG 35250 (which shows some plastic deformation) and to compare Nacholapithecus with other Miocene hominoids and extant anthropoids to evaluate the distinctiveness of its femur. The new fossil evidence reaffirms previously reported descriptions of some distal femoral traits, namely the morphology of the patellar groove. However, results also show that relative femoral head size in Nacholapithecus is smaller, relative neck length is longer, and neck–shaft angle is lower than previously reported for KNM-BG 35250. These traits have a strong functional signal related to the hip joint kinematics, suggesting that the morphology of the proximal femur in Nacholapithecus might be functionally related to quadrupedal-like behaviors instead of more derived antipronograde locomotor modes. Results further demonstrate that other African Miocene apes (with the exception of Turkanapithecus kalakolensis) generally fall within the Nacholapithecus range of variation, whose overall femoral shape resembles that of Ekembo spp. and Equatorius africanus. Our results accord with the previously inferred locomotor repertoire of Nacholapithecus, indicating a combination of generalized arboreal quadrupedalism combined with other antipronograde behaviors (e.g., vertical climbing).