Morphological integration of the hominoid postcranium.

Previous research has suggested that magnitudes of integration may be distinct in the postcranium of hominoids when compared to other primate species. To test this hypothesis, we estimated and compared magnitudes of integration of eight postcranial bones from three-dimensional surface scans for 57 Hylobates lar, 58 Gorilla gorilla, 60 Pan troglodytes, 60 Homo sapiens, 60 Chlorocebus pygerythrus, and 60 Macaca fascicularis. We tested the hypotheses that 1) magnitudes of integration would be distinct in the postcranium of hominoids compared to cercopithecoids, with the explicit prediction that magnitudes of integration would be lower in hominoids than in cercopithecoids, and 2) girdle elements (scapula, os coxa) would have lower magnitudes of integration across all taxa. Integration was quantified using the integration coefficient of variation from interlandmark distances reflecting anatomical and developmental modules defined according to a priori criteria. A resampling protocol was employed to generate distributions of integration values that were then compared statistically using Mann-Whitney U tests with Bonferroni adjustment. Support for hypothesis 1 was mixed: with the exception of Gorilla, hominoid taxa were less integrated than the cercopithecoids for all anatomical modules. However, Homo, Gorilla, and, to a lesser extent, Pan showed higher integration than Hylobates and the cercopithecoids for homologous limb elements, with magnitudes of integration for both modules being lowest for Hylobates. These results generally support the hypothesis of distinct patterns of magnitudes of integration in the hominoid postcranium. The high integration of Gorilla may be explained by the effects of overall body size. The results supported the predictions of the second hypothesis. Regardless of taxon, the os coxa and scapula were generally the least integrated skeletal elements, while the femur and radius were the most integrated. The lower integration of the girdle elements suggests that the geometric complexities of particular elements may significantly influence study outcomes.

Morphological modularity in the anthropoid axial skeleton.

Previous research has found that hominoids have stronger modularity between limb elements than other anthropoids, suggesting that there is less constraint on morphological diversification (e.g., limb proportions) in hominoids in terms of evolutionary independence. However, degrees of modularity in the axial skeleton have not been investigated across a broad range of anthropoid taxa. Thus, it is unknown whether hominoids also have stronger modularity in the axial skeleton than other anthropoids, which has implications for the evolution of diverse torso morphologies in Miocene apes as well as the evolution of novel characteristics in the skull and vertebrae of fossil hominins. In this study, 12 anthropoid genera were sampled to examine degrees of modularity between axial skeletal elements (i.e., cranium, mandible, vertebrae, and sacrum). Covariance ratio coefficients were calculated using variance/covariance matrices of interlandmark distances for each axial skeletal element to evaluate degrees of modularity. The results showed that Alouatta, Hylobates, Gorilla, Pan, and Homo showed generally stronger modularity than other anthropoid taxa when considering all axial skeletal elements. When only considering the vertebral elements (i.e., vertebrae and sacrum), Alouatta, Hylobates, Gorilla, and Pan showed generally stronger modularity than other anthropoid taxa. Humans showed stronger modularity between the skull and vertebrae than other hominoids. Thus, the evolution of novel characteristics in the skull and vertebral column may have been less constrained in fossil hominins due to the dissociation of trait covariation between axial skeletal elements in hominoid ancestors, thus fostering more evolutionary independence between the skull and vertebral column.

The forearm and hand musculature of semi-terrestrial rhesus macaques (Macaca mulatta) and arboreal gibbons (fam.Hylobatidae). Part II. Quantitative analysis.

Nonhuman primates have a highly diverse locomotor repertoire defined by an equally diverse hand use. Based on how primates use their hands during locomotion, we can distinguish between terrestrial and arboreal taxa. The 'arboreal' hand is likely adapted towards high wrist mobility and grasping, whereas the 'terrestrial' hand will show adaptations to loading. While the morphology of the forearm and hand bones have been studied extensively, functional adaptations in the forearm and hand musculature to locomotor behaviour have been documented only scarcely. In this paper, we investigate the forelimb musculature of the highly arboreal gibbons (including Hylobates lar,Hylobates pileatus,Nomascus leucogenys,Nomascus concolor and Symphalangus syndactylus) and compare this with the musculature of the semi-terrestrial rhesus macaques (Macaca mulatta). Anatomical data from previous dissections on knuckle-walking bonobos (Pan paniscus) and bipedal humans (Homo sapiens) are also included to further integrate the analyses in the scope of catarrhine hand adaptation. This study indicates that the overall configuration of the arm and hand musculature of these primates is very similar but there are some apparent differences in relative size which can be linked to differences in forelimb function and which might be related to their specific locomotor behaviour. In macaques, there is a large development of wrist deviators, wrist and digital flexors, and m. triceps brachii, as these muscles are important during the different phases of palmi- and digitigrade quadrupedal walking to stabilize the wrist and elbow. In addition, their m. flexor carpi ulnaris is the most important contributor to the total force-generating capacity of the wrist flexors and deviators, and is needed to counteract the adducting torque at the elbow joint during quadrupedal walking. Gibbons show a relatively high force-generating capacity in their forearm rotators, wrist and digital flexors, which are important muscles in brachiation to actively regulate forward movement of the body. The results also stress the importance of the digital flexors in bonobos, during climbing and clambering, and in humans, which is likely linked to our advanced manipulation skills.

Mandibular shape variation in mainland and insular hylobatids.

Although hylobatids are the most speciose of the living apes, their morphological interspecies and intraspecies variation remains poorly understood. Here, we assess mandibular shape variation in two species of Hylobates, white-handed (Hylobates lar) and black-handed (Hylobates agilis) gibbons. Using 71 three-dimensional landmarks to quantify mandibular shape, interspecies and intraspecies variation and geographic patterns of mandibular shape are examined in a mixed sex sample of adult H. lar and H. agilis through generalized Procrustes analysis, Procrustes analysis of variance, and principal components analysis. We find that relative to H. agilis, H. lar exhibits a higher amount of variation in mandibular shape. Both species demonstrate similar allometric patterns in mandibular shape. We also highlight a geographic pattern in mandibular shape variation. Compared to mainland hylobatids, insular hylobatids have relatively lower, more posteriorly oriented, and anteroposteriorly wider mandibular condyles, with an increased distance between the condyles and the coronoid processes. This geographic pattern could reflect differences in functional demands on the mandible during mastication and/or could be driven by factors often associated with evolutionary pressures of island populations relative to mainland populations. The findings of this study highlight how little is known about Hylobates morphological variation and how important this is for using Hylobates to help interpret the primate fossil record. Understanding interspecific and intraspecific variation in extant primates is vital to interpreting variation in the primate fossil record.

The forearm and hand musculature of semi-terrestrial rhesus macaques (Macaca mulatta) and arboreal gibbons (Fam. Hylobatidae). Part I. Description and comparison of the muscle configuration.

Primates live in very diverse environments and, as a consequence, show an equally diverse locomotor behaviour. During locomotion, the primate hand interacts with the superstrate and/or substrate and will therefore probably show adaptive signals linked with this locomotor behaviour. Whereas the morphology of the forearm and hand bones have been studied extensively, the functional adaptations in the hand musculature have been documented only scarcely. To evaluate whether there are potential adaptations in forelimb musculature to locomotor behaviour, we investigated the forearm and hand musculature of the highly arboreal gibbons (including Hylobates lar, Hylobates pileatus, Nomascus leucogenys, Nomascus concolor, Symphalangus syndactylus) and compared this with the musculature of the semi-terrestrial rhesus macaques (Macaca mulatta) by performing complete and detailed dissections on a sample of 15 unembalmed specimens. We found that the overall configuration of the upper arm and hand musculature is highly comparable between arboreal gibbons and semi-terrestrial macaques, and follows the general primate condition. Most of the identified differences in muscle configuration are located in the forearm. In macaques, a prominent m. epitrochleoanconeus is present, which potentially helps to extend the forearm and/or stabilize the elbow joint during quadrupedal walking. The m. flexor carpi radialis shows a more radial insertion in gibbons, which might be advantageous during brachiation, as it can aid radial deviation. The fingers of macaques are controlled in pairs by the m. extensor digiti secondi et tertii proprius and the m. extensor digiti quarti et quinti proprius-a similar organization can also be found in their flexors-which might aid in efficient positioning of the hand and fingers on uneven substrates during quadrupedal walking. In contrast, extension of the little finger in gibbons is controlled by a separate m. extensor digiti minimi, whereas digits 2 to 4 are extended by the m. extensor digitorum brevis, suggesting that simultaneous extension of digits 2-4 in gibbons might be important when reaching or grasping an overhead support during brachiation. In conclusion, the overall configuration of the forelimb and hand musculature is very similar in gibbons and macaques, with some peculiarities which can be linked to differences in forelimb function and which might be related to the specific locomotor behaviour of each group.

Correlation between musculoskeletal structure of the hand and primate locomotion: Morphometric and mechanical analysis in prehension using the cross- and triple-ratios.

Biometric ratios of the relative length of the rays in the hand have been analyzed between primate species in the light of their hand function or phylogeny. However, how relative lengths among phalanges are mechanically linked to the grasping function of primates with different locomotor behaviors remains unclear. To clarify this, we calculated cross and triple-ratios, which are related to the torque distribution, and the torque generation mode at different joint angles using the lengths of the phalanges and metacarpal bones in 52 primates belonging to 25 species. The torque exerted on the finger joint and traction force of the flexor tendons necessary for a cylindrical grip and a suspensory hand posture were calculated using the moment arm of flexor tendons measured on magnetic resonance images, and were compared among Hylobates spp., Ateles sp., and Papio hamadryas. Finally, the torques calculated from the model were validated by a mechanical study detecting the force exerted on the phalanx by pulling the digital flexor muscles during suspension in these three species. Canonical discriminant analysis of cross and triple-ratios classified primates almost in accordance with their current classification based on locomotor behavior. The traction force was markedly reduced with flexion of the MCP joint parallel to the torque in brachiating primates; this was notably lower in the terrestrial quadrupedal primates than in the arboreal primates at mild flexion. Our mechanical study supported these features in the torque and traction force generation efficiencies. Our results suggest that suspensory or terrestrial quadrupedal primates have hand structures that can exert more torque at a suspensory posture, or palmigrade and digitigrade locomotion, respectively. Furthermore, our study suggests availability of the cross and triple-ratios as one of the indicators to estimate the hand function from the skeletal structure.

A Study of Selected Physicomechanical and Histological Parameters of Hair Originating from Three Primate Species.

The aim of this study was to determine selected characteristics of hair originating from adult females of three primate species: lar gibbon, crested black macaque and common chimpanzee. The research material consisted of 300 hairs collected from the top of the head. The analysis included the length, diameter, strength, breaking load, elongation and cross-section area of the hairs, and the hair index. Additionally, the scale number per 1 mm of hair length was determined using scanning electron microscope images. Lar gibbon hairs were the thinnest and the longest, while common chimpanzee hairs were the thickest and the shortest, and concurrently the least resistant. Hairs from various primate species differ in terms of their morphological structure and histological features and thus can be used for differentiation and identification of species.

The distribution, number, and certain neurochemical identities of infracortical white matter neurons in a lar gibbon (Hylobates lar) brain.

We examined the number, distribution, and immunoreactivity of the infracortical white matter neuronal population, also termed white matter interstitial cells (WMICs), in the brain of a lesser ape, the lar gibbon. Staining for neuronal nuclear marker (NeuN) revealed WMICs throughout the infracortical white matter, these cells being most numerous and dense close to cortical layer VI, decreasing significantly in density with depth in the white matter. Stereological analysis of NeuN-immunopositive cells revealed a global estimate of ~67.5 million WMICs within the infracortical white matter of the gibbon brain, indicating that the WMICs are a numerically significant population, ~2.5% of the total cortical gray matter neurons that would be estimated for a primate brain the mass of that of the lar gibbon. Immunostaining revealed subpopulations of WMICs containing neuronal nitric oxide synthase (nNOS, ~7 million in number, with both small and large soma volumes), calretinin (~8.6 million in number, all of similar soma volume), very few WMICs containing parvalbumin, and no calbindin-immunopositive neurons. These nNOS, calretinin, and parvalbumin immunopositive WMICs, presumably all inhibitory neurons, represent ~23.1% of the total WMIC population. As the white matter is affected in many cognitive conditions, such as schizophrenia, autism and also in neurodegenerative diseases, understanding these neurons across species is important for the translation of findings of neural dysfunction in animal models to humans. Furthermore, studies of WMICs in species such as apes provide a crucial phylogenetic context for understanding the evolution of these cell types in the human brain.

Muscle Spindle Density of Lateral Rotators of the Thigh in Japanese Macaques and a Gibbon.

We examined the six small lateral rotators of the hip joint, which is one of the most flexible joints and allows kinematically complex motions of the hindlimb, to elucidate the functional differentiation among these muscles and to test the hypothesis that species-specific characteristics in hindlimb use during locomotion are reflected in the muscle spindle density and in other parameters of the deep small hip joint rotators. For these purposes, we estimated the number of muscle spindles of the superior gemellus muscle (SG), inferior gemellus muscle, quadratus femoris muscle, obturator internus muscle (OI), obturator externus muscle, and piriformis muscle in three Japanese macaques and a gibbon, using 30-µm-thick serial sections throughout each muscle length after azan staining. The numbers of muscle spindles per 10,000 muscle fibers were determined to compare inter-muscle variation. The spindle density was highest in the SG and lowest in the OI in the Japanese macaques, suggesting that the SG, which is attached to the tendon of the OI, functions as a kinesiological monitor of the OI. On the other hand, SG the was missing in the gibbon, and the OI in the gibbon contained more spindles than that in the Japanese macaques. This suggests that the SG and the OI fused into one muscle in the gibbon. We postulate that the relative importance of the deep small hip rotator muscles differs between the Japanese macaques and gibbon and that the gibbon's muscles are less differentiated in terms of the spindle density, probably because this brachiating species uses its hindlimbs less frequently.

New genus of extinct Holocene gibbon associated with humans in Imperial China.

Although all extant apes are threatened with extinction, there is no evidence for human-caused extinctions of apes or other primates in postglacial continental ecosystems, despite intensive anthropogenic pressures associated with biodiversity loss for millennia in many regions. Here, we report a new, globally extinct genus and species of gibbon, Junzi imperialis, described from a partial cranium and mandible from a ~2200- to 2300-year-old tomb from Shaanxi, China. Junzi can be differentiated from extant hylobatid genera and the extinct Quaternary gibbon Bunopithecus by using univariate and multivariate analyses of craniodental morphometric data. Primates are poorly represented in the Chinese Quaternary fossil record, but historical accounts suggest that China may have contained an endemic ape radiation that has only recently disappeared.

A cerebellar substrate for cognition evolved multiple times independently in mammals.

Given that complex behavior evolved multiple times independently in different lineages, a crucial question is whether these independent evolutionary events coincided with modifications to common neural systems. To test this question in mammals, we investigate the lateral cerebellum, a neurobiological system that is novel to mammals, and is associated with higher cognitive functions. We map the evolutionary diversification of the mammalian cerebellum and find that relative volumetric changes of the lateral cerebellar hemispheres (independent of cerebellar size) are correlated with measures of domain-general cognition in primates, and are characterized by a combination of parallel and convergent shifts towards similar levels of expansion in distantly related mammalian lineages. Results suggest that multiple independent evolutionary occurrences of increased behavioral complexity in mammals may at least partly be explained by selection on a common neural system, the cerebellum, which may have been subject to multiple independent neurodevelopmental remodeling events during mammalian evolution.

Using the covariation of extant hominoid upper and lower jaws to predict dental arcades of extinct hominins.

Upper and lower jaws are well represented in the fossil record of mammals and are frequently used to diagnose species. Some hominin species are only known by either their maxillary or mandibular morphology, and in this study, we explore the possibility of predicting their complementary dental arcade shape to aid the recognition of conspecific specimens in the fossil record. To this end, we apply multiple multivariate regression to analyze 3D landmark coordinates collected on associated upper and lower dental arcades of extant Homo, Pan, Gorilla, Pongo, and Hylobates. We first study the extant patterns of variation in dental arcade shape and quantify how accurate predictions of complementary arcades are. Then we explore applications of this extant framework for interpreting the fossil record based on two fossil hominin specimens with associated upper and lower jaws, KNM-WT 15000 (Homo erectus sensu lato) and Sts 52 (Australopithecus africanus), as well as two non-associated specimens of Paranthropus boisei, the maxilla of OH 5 and the Peninj mandible. We find that the shape differences between the predictions and the original fossil specimens are in the range of variation within genera or species and therefore are consistent with their known affinity. Our approach can provide a reference against which intraspecific variation of extinct species can be assessed. We show that our method predicts arcade shapes reliably even if the target shape is not represented in the reference sample. We find that in extant hominoids, the amount of within-taxon variation in dental arcade shape often overlaps with the amount of between-taxon shape variation. This implies that whereas a large difference in dental arcade shape between two individuals typically suggests that they belong to different species or even genera, a small shape difference does not necessarily imply conspecificity.

Phenotypic integration of the cervical vertebrae in the Hominoidea (Primates).

Phenotypic integration and modularity represent important factors influencing evolutionary change. The mammalian cervical vertebral column is particularly interesting in regards to integration and modularity because it is highly constrained to seven elements, despite widely variable morphology. Previous research has found a common pattern of integration among quadrupedal mammals, but integration patterns also evolve in response to locomotor selective pressures like those associated with hominin bipedalism. Here, I test patterns of covariation in the cervical vertebrae of three hominoid primates (Hylobates, Pan, Homo) who engage in upright postures and locomotion. Patterns of integration in the hominoid cervical vertebrae correspond generally to those previously found in other mammals, suggesting that integration in this region is highly conserved, even among taxa that engage in novel positional behaviors. These integration patterns reflect underlying developmental as well as functional modules. The strong integration between vertebrae suggests that the functional morphology of the cervical vertebral column should be considered as a whole, rather than in individual vertebrae. Taxa that display highly derived morphologies in the cervical vertebrae are likely exploiting these integration patterns, rather than reorganizing them. Future work on vertebrates without cervical vertebral number constraints will further clarify the evolution of integration in this region.

Morphological self stabilization of locomotion gaits: illustration on a few examples from bio-inspired locomotion.

To a large extent, robotics locomotion can be viewed as cyclic motions, named gaits. Due to the high complexity of the locomotion dynamics, to find the control laws that ensure an expected gait and its stability with respect to external perturbations, is a challenging issue for feedback control. To address this issue, a promising way is to take inspiration from animals that intensively exploit the interactions of the passive degrees of freedom of their body with their physical surroundings, to outsource the high-level exteroceptive feedback control to low-level proprioceptive ones. In this case, passive interactions can ensure most of the expected control goals. In this article, we propose a methodological framework to study the role of morphology in the design of locomotion gaits and their stability. This framework ranges from modelling to control aspects, and is illustrated through three examples from bio-inspired locomotion: a three-dimensional micro air vehicle in hovering flight, a pendular planar climber and a bipedal planar walker. In these three cases, we will see how simple considerations based on the morphology of the body can ensure the existence of passive stable gaits without requiring any high-level control.

Development of Pelvic Sexual Dimorphism in Hylobatids: Testing the Obstetric Constraints Hypothesis.

Pelvic sexual dimorphism in primates is typically seen as the result of female-specific adaptations to obstetric constraints, which arise from the tight fit between the neonate head and the maternal pelvis. However, it remains debated to which extent pelvic dimorphism is a correlate of obstetric constraints, of body size dimorphism, and/or of other factors. Also, little is known on how pelvic dimorphism develops. Here we use methods of biomedical imaging and geometric morphometrics to address these questions in two hylobatid species, Hylobates lar and Symphalangus syndactylus. These species differ markedly in body size, but within each species, there is only little body size dimorphism. Results show that the neonates of H. lar are large compared to the maternal pelvis, resulting in high cephalopelvic proportions and substantial obstetric constraints. Pelvic sexual dimorphism is moderate but significant: During puberty, females develop a more capacious pelvic inlet than males, while overall pelvic size is similar in both sexes. S. syndactylus has substantially larger pelves than H. lar, but neonates are similar in size to those of H. lar. Cephalopelvic proportions are thus low, and there are no obstetric constraints. Pelvic sexual dimorphism is absent. Overall, these data indicate that pelvic sexual dimorphism in hylobatids reflects obstetric constraints. Anat Rec, 300:859-869, 2017. © 2017 Wiley Periodicals, Inc.

Morphological integration of the cranium in Homo, Pan, and Hylobates and the evolution of hominoid facial structures.

OBJECTIVES: Modern humans diverge from other extant hominids (chimpanzees, gorillas, and orangutans) in a series of craniofacial morphological features. Like hylobatids, they possess a face with a reduced subnasal prognathism that is associated with a globular basicranium. These traits are not independent, as the skull is a complex integrated structure. The aim of the present study is to determine relationships between the face and the basicranium in two hominid genera (Homo and Pan) and a hylobatid genus (Hylobates) to test if these taxa share common patterns of integration linking these structures. MATERIALS AND METHODS: Three dimensional (3D) geometric morphometric analyses and 3D homologous landmarks are used to compare the integration patterns between facial and basicranial structures in a comparative sample of Homo, Pan, and Hylobates. Pooled within-genus partial least squares analyses are computed to describe and quantify these patterns of integration. RESULTS: The covariation analyses show similar patterns of integration shared between the three studied taxa. These patterns correspond to the brachycephalic and dolichocephalic conditions previously defined in hominins and hominids. DISCUSSION: Results confirm that hominoids share similar patterns of integration. This is in line with the hypothesis that morphological integration is mostly conservative in hominoids. These similar patterns of integration may explain the convergent evolution of short faces in humans and hylobatids.

Primary bone microanatomy records developmental aspects of life history in catarrhine primates.

A central challenge in human origins research is to understand how evolution has shaped modern human life history. As fossilized remains of our ancestors provide the only direct evidence for life history evolution, efforts to reconstruct life history in paleontological contexts have focused on hard tissues, particularly on dental development. However, among investigators of other vertebrate groups, there is a long tradition of examining primary bone microstructure to decipher growth rates and maturational timing, based on an empirical relationship between the microanatomy of primary bone and the rate at which it is deposited. We examined ontogenetic variation in primary bone microstructure at the midshaft femur of Chlorocebus aethiops, Hylobates lar, and Pan troglodytes to test whether tissue type proportions vary in accordance with predictions based on body mass growth patterns described previously. In all taxa, younger age classes were characterized by significantly higher percent areas of fibro-lamellar and/or parallel-fibered tissues, while older age classes showed significantly higher proportions of lamellar bone. In prior experimental studies, fibro-lamellar and parallel-fibered tissue types have been associated with faster depositional rates than lamellar bone. Principal components analysis revealed differences among taxa in the timing of this transition, and in the particular tissue types observed among individuals of similar dental emergence status. Among M1 and M2 age classes, higher proportions of parallel-fibered and fibro-lamellar tissues were observed in those taxa characterized by reportedly faster body mass growth rates. Further, persistence of fibro-lamellar tissue throughout DECID, M1 and M2 age classes in chimpanzees contrasts with the pattern reported previously for modern humans. Despite the necessary limitations of our cross-sectional study design and the secondary remodeling of bone in primates, large areas of primary bone remain intact and represent a valuable and independent source of information about the evolution of growth and development in the fossil record.

Miocene small-bodied ape from Eurasia sheds light on hominoid evolution.

Miocene small-bodied anthropoid primates from Africa and Eurasia are generally considered to precede the divergence between the two groups of extant catarrhines­hominoids (apes and humans) and Old World monkeys­and are thus viewed as more primitive than the stem ape Proconsul. Here we describe Pliobates cataloniae gen. et sp. nov., a small-bodied (4 to 5 kilograms) primate from the Iberian Miocene (11.6 million years ago) that displays a mosaic of primitive characteristics coupled with multiple cranial and postcranial shared derived features of extant hominoids. Our cladistic analyses show that Pliobates is a stem hominoid that is more derived than previously described small catarrhines and Proconsul. This forces us to reevaluate the role played by small-bodied catarrhines in ape evolution and provides key insight into the last common ancestor of hylobatids (gibbons) and hominids (great apes and humans).

Reconciling the convergence of supraspinous fossa shape among hominoids in light of locomotor differences.

Differences in scapular morphology between modern humans and the African and lesser apes are associated with the distinct locomotor habits of these groups. However, several traits, particularly aspects of the supraspinous fossa, are convergent between Homo and Pongo-an unexpected result given their divergent locomotor habits. Many morphological assessments of the scapula rely on the limited number of static landmarks available, and traditional approaches like these tend to oversimplify scapular shape. Here, we present the results of two geometric morphometric (GM) analyses of hominoid supraspinous fossa shape-one employing five homologous landmarks and another with 83 sliding semilandmarks-alongside those of traditional methods to evaluate if three-dimensional considerations of fossa shape afford more comprehensive insights into scapular shape and functional morphology. Traditional measures aligned Pongo and Homo with narrow and transversely oriented supraspinous fossae, whereas African ape and Hylobates fossae are broader and more obliquely situated. However, our GM results highlight that much of the convergence between Homo and Pongo is reflective of their more medially positioned superior angles. These approaches offered a more complete assessment of supraspinous shape and revealed that the Homo fossa, with an intermediate superior angle position and moderate superoinferior expansion, is actually reminiscent of the African ape shape. Additionally, both Pongo and Hylobates were shown to have more compressed fossae, something that has not previously been identified through traditional analyses. Thus, the total morphological pattern of the Pongo supraspinous fossa is unique among hominoids, and possibly indicative of its distinctive locomotor habits.

The oldest gibbon fossil (Hylobatidae) from insular Southeast Asia: evidence from Trinil, (East Java, Indonesia), Lower/Middle Pleistocene.

A fossil femur excavated by Eugène Dubois between 1891-1900 in the Lower/Middle Pleistocene bonebed of the Trinil site (Java, Indonesia) was recognised by us as that of a Hylobatidae. The specimen, Trinil 5703 of the Dubois Collection (Leiden, The Netherlands), has the same distinctive form of fossilization that is seen in many of the bonebed fossils from Trinil in the collection. Anatomical comparison of Trinil 5703 to a sample of carnivore and primate femora, supported by morphometric analyses, lead to the attribution of the fossil to gibbon. Trinil 5703 therefore provides the oldest insular record of this clade, one of the oldest known Hylobatidae fossils from Southeast Asia. Because living Hylobatidae only inhabit evergreen rain forests, the paleoenvironment within the river drainage in the greater Trinil area evidently included forests of this kind during the Lower/Middle Pleistocene as revealed here.