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 (approximately 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 MioceneMorotopithecus. 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.     

Climbing,brachiation, and terrestrial quadrupedalism: Historical precursors of hominid bipedalism

The vertical-climbing account of the evolution of locomotor behavior and morphology in hominid ancestry is reexamined in light of recent behavioral, anatomical, and paleontological findings and a more firmly established phylogeny for the living apes. The behavioral record shows that African apes, when arboreal, are good vertical climbers, and that locomotion during traveling best separates the living apes into brachiators (gibbons), scrambling/climbing/brachiators (orangutans), and terrestrial quadrupeds (gorillas and chimpanzees). The paleontological record documents frequent climbing as an ancestral catarrhine ability, while a reassessment of the morphology of the torso and forelimb in living apes and Atelini suggests that their shared unique morphological pattern is best explained by brachiation and forelimb suspensory positional behavior. Further, evidence from the hand and foot points to a terrestrial quadrupedal phase in hominoid evolution prior to the adoption of bipedalism. The evolution of positional behavior from early hominoids to hominids appears to have begun with an arboreal quadrupedal-climbing phase and proceeded though an orthograde, brachiating, forelimb-suspensory phase, which was in turn followed by arboreal and terrestrial quadrupedal phases prior to the advent of hominid bipedality. The thesis that protohominids climbed down from the trees to become terrestrial bipeds needs to be reexamined in light of a potentially long history of terrestriality in the ancestral protohominid. © 1996 Wiley-Liss, Inc.     

Orang-like manual adaptations in the fossil hominoid Hispanopithecus laietanus: first steps towards great ape suspensory behaviours

Morphological and biometrical analyses of the partial hand IPS18800 of the fossil great ape Hispanopithecus laietanus (=Dryopithecus laietanus), from the Late Miocene (about 9.5Ma) of Can Llobateres (Catalonia, Spain), reveal many similarities with extant orang-utans (Pongo). These similarities are interpreted as adaptations to below-branch suspensory behaviours, including arm-swinging and clambering/postural feeding on slender arboreal supports, due to an orang-like double-locking mechanism. This is confirmed by the long and highly curved phalanges of Hispanopithecus. The short and stout metacarpals with dorsally constricted heads, together with the dorsally extended articular facets on proximal phalanges, indicate the persistence of significant degrees of palmigrady. A powerful grasping capability is indicated by the great development of basal phalangeal tubercles, the marked insertions for the flexors on phalangeal shafts and the large pits for the collateral ligaments. The morphology of the Hispanopithecus long bones of the hand indicates a unique positional repertoire, combining orthogrady with suspensory behaviours and palmigrade quadrupedalism. The retention of powerful grasping and palmigrady suggests that the last common ancestor of hominids might have been more primitive than what can be inferred on the basis of extant taxa, suggesting that pronograde behaviours are compatible with an orthograde bodyplan suitable for climbing and suspension.     

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

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

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

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

New Sivapithecus postcranial specimens from the Siwaliks of Pakistan

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

New Sivapithecus postcranial specimens from the Siwaliks of Pakistan

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

Metatarsophalangeal joint function and positional behavior in Australopithecus afarensis

Recent discussions of the pedal morphology of Australopithecus afarensis have led to conflicting interpretations of australopithecine locomotor behavior. We report the results of a study using computer aided design (CAD) software that provides a quantitative assessment of the functional morphology of australopithecine metatarsophalangeal joints. The sample includes A. afarensis, Homo sapiens, Pan troglodytes, Gorilla gorilla, and Pongo pygmaeus. Angular measurements of the articular surfaces relative to the long axes of the metatarsals and phalanges were taken to determine whether the articular surfaces are plantarly or dorsally oriented. Humans have the most dorsally oriented articular surfaces of the proximal pedal phalanges. This trait appears to be functionally associated with dorsiflexion during bipedal stride. Pongo has the most plantarly oriented articular surfaces of the proximal pedal phalanges, probably reflecting an emphasis on plantarflexion in arboreal positional behaviors, while the African hominoids are intermediate between Pongo and Homo for this characteristic. A. afarensis falls midway between the African apes and humans. Results from an analysis of metatarsal heads are inconclusive with regard to the functional morphology of A. afarensis. Overall, the results are consistent with other evidence indicating that A. afarensis was a capable climber. © 1994 Wiley-Liss, Inc.     

Reconstructing the mechanics of quadrupedalism in an extinct hominoid

Traditionally, analogising comparative anatomical approaches, working on features of individual bony elements, have led to the Miocene hominoids Proconsul heseloni and P. nyanzae being described as arboreal, with a variety of possible locomotion modes. Whilst most researchers seemingly agree that quadrupedal was one of the most frequently adopted modes, any deeper knowledge about the kinematical characteristics of such quadrupedalism is very limited. Based on the previous studies and a computer simulation technique developed in our laboratory, a set of alternative models for Proconsul quadrupedalism was created. The body measurements and initial properties for the different models were held constant, using data from published literature if available, or otherwise estimated from data for Pan. Judged by the power output of joints, the results of computer simulation indicate that the style of quadrupedal locomotion typical of living macaques fits the body proportions of Proconsul better than that of Canis domesticus, Varecia variegata, Cebus albifrons or Pan troglodytes. It may reasonably be assumed that Proconsul's quadrupedal mode was similar to that of living macaques.     

Preliminary analysis of<Emphasis Type="Italic"> Nacholapithecus</Emphasis> scapula and clavicle from Nachola,Kenya

The Miocene ape Nacholapithecus is known from rather complete skeletons; some of them preserve the shoulder joint, identified by three scapulae and one clavicle. Comparisons made with other Miocene and living apes (Proconsul, Equatorius, Ugandapithecus) suggest that the mobility of the scapulohumeral joint was important, and scapular features such as the morphology and position of the spine and the morphology of the acromion and axillary border resemble those of climbing arboreal primates except for chimpanzees, gorillas, or orang-utans. From the size of the scapula (male Nasalis size), it is clear that the animal is smaller than an adult chimpanzee, but the clavicle is almost as relatively long as those of chimpanzees. Some features closer to colobine morphology reinforce the hypothesis that Nacholapithecus was probably a good climber and was definitely adapted for an arboreal life.     

Electromyography of brachial muscles in Pan gorilla and hominoid evolution

Electromyographic studies on brachial muscles in a gorilla indicate that its elbow joint may be especially adapted for knuckle-walking and suspensory behavior. A close-packed positioning mechanism that minimizes muscular effort during full extension of the elbow joint is indicated by remarkably low levels of EMG in the brachial muscles, particularly during knuckle-walking and suspensory behavior on a trapeze. Extension of the elbow joint is facilitated by reduction of the olecranon process of the ulna, a feature that is attributable initially to aspects of an arboreal heritage in protogorilla and secondarily to selection for efficient knuckle-walking. Although notable differences exist between gorilla and man in known activity of the brachial muscles, the two species are strikingly similar in many basic features. Available evidence suggests that they share a common heritage of arboreal adaptation, including vertical climbing, hauling, hoisting, and suspensory behavior, perhaps more recently than some authors would care to admit. Knuckle-walking probably played an inconsequential role in the protohominid career. Selection for tool use, expecially involving powerful and rapid extension of the elbow joint, is the most reasonable explanation for the relatively more protuberant olecranon process in man by comparison with apes.     

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

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

Investigating the form-function interface in African apes: Relationships between principal moments of area and positional behaviors in femoral and humeral diaphyses

Investigations of cross-sectional geometry in nonhuman primate limb bones typically attribute shape ratios to qualitative behavioral characterizations, e.g., leaper, slow climber, brachiator, or terrestrial vs. arboreal quadruped. Quantitative positional behavioral data, however, have yet to be used in a rigorous evaluation of such shape-behavior connections. African apes represent an ideal population for such an investigation because their relatedness minimizes phylogenetic inertia, they exhibit diverse behavioral repertoires, and their locomotor behaviors are known from multiple studies. Cross-sectional data from femoral and humeral diaphyses were collected for 222 wild-shot specimens, encompassing Pan paniscus and all commonly recognized African ape subspecies. Digital representations of diaphyseal cross sections were acquired via computed tomography at three locations per diaphysis. Locomotor behaviors were pooled broadly into arboreal and terrestrial categories, then partitioned into quadrupedal walking, quadrumanous climbing, scrambling, and suspensory categories. Sex-specific taxonomic differences in ratios of principal moments of area (PMA) were statistically significant more often in the femoral diaphysis than the humeral diaphysis. While it appears difficult to relate a measure of shape (e.g., PMA ratio) to individual locomotor modes, general locomotor differences (e.g., percentage arboreal vs. terrestrial locomotion) are discerned more easily. As percentage of arboreal locomotion for a group increases, average cross sections appear more circular. Associations between PMA ratio and specific locomotor behaviors are less straightforward. Individual behaviors that integrate eccentric limb positions (e.g., arboreal scrambling) may not engender more circular cross sections than behaviors that incorporate repetitive sagittal movements (e.g., quadrupedal walking) in a straightforward manner.     

化石人猿超科成员指趾骨弯曲程度与位移行为

灵长类近节指趾骨的弯曲程度被认为是树栖性和悬垂位移行为的一个重要指标。几何形态测量学—多项式曲线拟合法(GM-PCF)提供了一种更加精准的指趾骨弯曲程度的定量化指标,以剔除指趾骨大小因素之后的标准化曲线高度(NPCH)作为其弯曲程度的指标,配合指趾骨的曲线长度,可以更加全面地定量分析灵长类指趾骨弯曲程度与位移行为的对应关系。尤其是涵盖灵长类大部分位移行为方式的15个类群、328个个体、5000余件指趾骨的参考样本,基本可以满足各种化石灵长类指趾骨弯曲程度分析和位移行为方式重建的需求。本文总结了发现有完整第II-V近节指趾骨化石材料的人猿超科成员的颅后骨骼形态适应及位移行为的重建,并运用GM-PCF对这些指趾骨化石的弯曲程度进行了对比分析,以通过指趾骨弯曲程度重建人猿超科成员的位移行为适应,并可为这些人猿超科成员位移行为的完整演化图景增加新的认识。     

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.     

Positional behavior in the Hominoidea

Quantitative studies on the positional behavior of members of the Hominoidea are compared in order (1) to identify consistencies across the superfamily, (2) to contrast ape positional behavior with that of Old World monkeys (forest-livingPapio anubis were chosen for study to reduce body size effects), and (3) to identify distinctive behaviors in each of the ape taxa. Differences in the way behaviors were sampled in the various studies necessitated considering posture and locomotion separately. Unimanual arm-hanging and vertical climbing were the most distinctive shared postural and locomotor modes among the apes (the gorilla excepted), constituting ≥5.0% and ≥4.9% of all behavior in each species. Arm-hanging and brachiation (sensu stricto) frequencies were the highest by far in hylobatids. Hand-foot hanging, bipedal posture, and clambering, an orthograde suspensory locomotion assisted by the hindlimbs, were more common in orangutans than in any other hominoid. Sitting and walking were observed in the highest frequencies in the African apes but were no more common than in the baboon. Relatively high frequencies of brachiation (sensu stricto) were reported for all apes except chimpanzees and gorillas. Brachiation and arm-hanging were kinematically different in apes and baboons, involving complete humeral abduction only in the former, whereas vertical climbing appeared to be kinematically similar in apes and baboons. It is concluded that the morphological specializations of the apes may be adaptations to (1) the unique physical demands of arm-hanging and (2) less kinematically distinct, but still quantitatively significant, frequencies of vertical climbing.     

The oldest ape

This paper recounts the history and significance of Morotopithecus bishopi, an early Miocene East African ape. Morotopithecus differs in intriguing ways from its contemporary Proconsul. While craniodental differences are slight, the known elements of its postcranium suggest that Morotopithecus was capable of modern ape–like positional behaviors, including vertical postures, deliberate climbing, and arm hanging. In contrast, Proconsul and other early and middle Miocene hominoids have been reconstructed as above–branch quadrupeds. Paleoanthropologists are currently divided over whether and which of the anatomical features associated with upright posture and suspension in the modern apes are due to inheritance or independent evolution. This debate has important implications for interpreting the phylogenetic positions of both Morotopithecus and Proconsul, as well as for reconstructing the pattern and timing of the emergence of modern ape adaptations.     

The locomotion of Babakotia radofilai inferred from epiphyseal and diaphyseal morphology of the humerus and femur

Palaeopropithecids, or “sloth lemurs,” are a diverse clade of large‐bodied Malagasy subfossil primates characterized by their inferred suspensory positional behavior. The most recently discovered genus of the palaeopropithecids is Babakotia, and it has been described as more arboreal than Mesopropithecus, but less than Palaeopropithecus. In this article, the within‐bone and between‐bones articular and cross‐sectional diaphyseal proportions of the humerus and femur of Babakotia were compared to extant lemurs, Mesopropithecus and Palaeopropithecus in order to further understand its arboreal adaptations. Additionally, a sample of apes and sloths (Choloepus and Bradypus) are included as functional outgroups composed of suspensory adapted primates and non‐primates. Results show that Babakotia and Mesopropithecus both have high humeral/femoral shaft strength proportions, similar to extant great apes and sloths and indicative of forelimb suspensory behavior, with Babakotia more extreme in this regard. All three subfossil taxa have relatively large femoral heads, also associated with suspension in modern taxa. However, Babakotia and Mesopropithecus (but not Palaeopropithecus) have relatively small femoral head surface area to shaft strength proportions suggesting that hind‐limb positioning in these taxa during climbing and other behaviors was different than in extant great apes, involving less mobility. Knee and humeral articular dimensions relative to shaft strengths are small in Babakotia and Mesopropithecus, similar to those found in modern sloths and divergent from those in extant great apes and lemurs, suggesting more sloth‐like use of these joints during locomotion. Mesopropithecus and Babakotia are more similar to Choloepus in humerofemoral head and length proportions while Palaeopropithecus is more similar to Bradypus. These results provide further evidence of the suspensory adaptations of Babakotia and further highlight similarities to both extant suspensory primates and non‐primate slow arboreal climbers and hangers. J. Morphol. 277:1199–1218, 2016. © 2016 Wiley Periodicals, Inc.