Evaluating the muscle attachment hypothesis for sagittal cresting in Gorilla and Pongo.

Primate mandibular morphology is often associated with jaw functionality of the masticatory complex in the context of variation in diets. Recent research into the disparities between the diet and jaw functionality in male and female hominoids is inconclusive and suggests that sexual dimorphism in the mandible may be influenced by external factors such as temporalis and masseter muscle morphology, which in turn may be influenced by sexual selection. As the muscles associated with mastication (i.e., the type of chewing exhibited by primates and other mammals) encompass the mandible as well as the neurocranium, including the sagittal crest among some individuals, this study investigates sex-specific associations between regions of the mandibular ramus and neurocranium associated with mastication in a dentally mature sample of Gorilla and Pongo. A total of four cranial and mandibular variables were measured in two Gorilla taxa (Gorilla gorilla gorilla and Gorilla beringei graueri) and one Pongo taxon (Pongo pygmaeus pygmaeus) (n = 220). For all three taxa, we investigate (a) whether the degree of sexual dimorphism in cranial regions associated with sagittal cresting (sagittal crest size (SCS) and temporalis muscle attachment area (TMAA)) is proportional to the degree of mandibular ramus area (MRA) and coronoid process height (CPH) sexual dimorphism, (b) whether there are sex differences in scaling relationships between TMAA and MRA, and (c) whether there are sex differences in the strength of association between TMAA and CPH. We show that for G. g. gorilla, variables associated with sagittal cresting show higher sexual dimorphism values than our two mandibular ramus variables, which is not the case for G. b. graueri or for P. p. pygmaeus. All three taxa show similar sex-specific scaling relationships between TMAA and MRA, where for males this relationship does not diverge from isometry, and for females there is a negative allometric relationship. Our findings also show intraspecific sex differences in allometric slopes between MRA and TMAA for all three taxa. Only G. g. gorilla shows a significant association between TMAA and CPH, which is observed in both sexes. Although there are some statistical associations between the cranial and mandibular regions associated with mastication, our results show that among male gorillas and orangutans, patterns of variation in the sagittal crest, TMAA, mandibular ramus and the coronoid process cannot be explained by the muscle attachment hypothesis alone. These findings have implications surrounding the associations between social behaviour and the morphology of the craniofacial complex.

A cadaveric study of wrist-joint moments in chimpanzees and orangutans with implications for the evolution of knuckle-walking.

Understanding the mechanism underlying the evolution of knuckle-walking in African great apes but not in humans may provide important implications about the origin and evolution of human bipedal locomotion. In this study, aiming to reveal possible structural adaptations of the chimpanzee's forearm and hand musculature related to knuckle-walking, we measure the passive elastic moment of the chimpanzee's and orangutan's wrist as it was rotated into extension, immobilizing the metacarpophalangeal joint at three different positions: extended (as in knuckle-walking), flexed (as in fist-walking), and an intermediate position. Our findings demonstrate that when the metacarpophalangeal joints are extended, the rigidity of the wrist joint in the extended direction increases. This increased rigidity is attributed to the passive elongation and force generation of digital flexor muscles, which are relatively short in chimpanzees. Consequently, this enhanced wrist-joint rigidity contributes to the stability and energetically efficient transmission of propulsive force to the ground during the stance phase. Overall, our study supports the hypothesis that knuckle-walking is an adaptation to terrestrial locomotion for an ancestor characterized by the restricted capacity for wrist extension owing to the relatively shorter tendons of digital flexor muscles.

Evolution of a transcriptional regulator from a transmembrane nucleoporin.

Nuclear pore complexes (NPCs) emerged as nuclear transport channels in eukaryotic cells ∼1.5 billion years ago. While the primary role of NPCs is to regulate nucleo-cytoplasmic transport, recent research suggests that certain NPC proteins have additionally acquired the role of affecting gene expression at the nuclear periphery and in the nucleoplasm in metazoans. Here we identify a widely expressed variant of the transmembrane nucleoporin (Nup) Pom121 (named sPom121, for "soluble Pom121") that arose by genomic rearrangement before the divergence of hominoids. sPom121 lacks the nuclear membrane-anchoring domain and thus does not localize to the NPC. Instead, sPom121 colocalizes and interacts with nucleoplasmic Nup98, a previously identified transcriptional regulator, at gene promoters to control transcription of its target genes in human cells. Interestingly, sPom121 transcripts appear independently in several mammalian species, suggesting convergent innovation of Nup-mediated transcription regulation during mammalian evolution. Our findings implicate alternate transcription initiation as a mechanism to increase the functional diversity of NPC components.

Palaeobotanical evidence reveals the living conditions of Miocene Lufengpithecus in East Asia.

BACKGROUND: Understanding the relationship between human evolution and environmental changes is the key to lifting the veil on human origin. The hypothesis that environmental changes triggered the divergence of humans from apes (ca. 9.3-6.5 million years ago, Ma) has been poorly tested because of limited continuous environmental data from fossil localities. Lufengpithecus (12.5-6.0 Ma) found on the southeastern margin of the Tibetan Plateau (SEMTP) across the ape-human split provides a good chance for testing this hypothesis. RESULTS: Here, we reconstructed the habitats of L. keiyuanensis (12.5-11.6 Ma) with comprehensive vegetation, climate, and potential food web data by palaeobotanical evidence, together with other multidisciplinary data and partly tested the environment-driven hypothesis by revealing the living conditions of Lufengpithecus. CONCLUSION: A detailed comparison of hominoids on different continents reveals their behaviour and fate divergence across the ape-human split against the background of global climate change, i.e., the stable living conditions of SEMTP not only provided a so-called 'refuge' for arboreal Lufengpithecus but also acted as a 'double-edged sword', preventing their further evolution while vegetation shifts in East Africa probably stimulated the emergence of human bipedalism, and the intense climatic changes in Europe possibly prevented those hominoids from surviving that time interval. Our findings provide interesting insight into the environmental impacts on the behavioural evolution of hominoids.

The earliest hylobatid from the Late Miocene of China.

Yuanmoupithecus xiaoyuan, a small catarrhine from the Late Miocene of Yunnan in southern China, was initially suggested to be related to Miocene proconsuloids or dendropithecoids from East Africa, but subsequent reports indicated that it might be more closely related to hylobatids. Here, detailed comparisons of the material, including seven newly discovered teeth and a partial lower face of a juvenile individual, provide crucial evidence to help establish its phylogenetic relationships. Yuanmoupithecus exhibits a suite of synapomorphies that support a close phylogenetic relationship with extant hylobatids. Furthermore, based on the retention of several primitive features of the dentition, Yuanmoupithecus can be shown to be the sister taxon of crown hylobatids. The contention that Kapi ramnagarensis from the Middle Miocene of India might represent an earlier species of hylobatid is not supported here. Instead, Kapi is inferred to be a specialized pliopithecoid more closely related to Krishnapithecus krishnaii from the Late Miocene of India. Currently then, Yuanmoupithecus represents the earliest known definitively identified hylobatid and the only member of the clade predating the Pleistocene. It extends the fossil record of hylobatids back to 7-8 Ma and fills a critical gap in the evolutionary history of hominoids that has up until now remained elusive. Even so, molecular estimates of a divergence date of hylobatids from other hominoids at about 17-22 Ma signifies that there is still a substantial gap in the fossil record of more than 10 million years that needs to be filled in order to document the biogeographic origins and early evolution of hylobatids.

Exploring the functional morphology of the Gorilla shoulder through musculoskeletal modelling.

Musculoskeletal computer models allow us to quantitatively relate morphological features to biomechanical performance. In non-human apes, certain morphological features have long been linked to greater arm abduction potential and increased arm-raising performance, compared to humans. Here, we present the first musculoskeletal model of a western lowland gorilla shoulder to test some of these long-standing proposals. Estimates of moment arms and moments of the glenohumeral abductors (deltoid, supraspinatus and infraspinatus muscles) over arm abduction were conducted for the gorilla model and a previously published human shoulder model. Contrary to previous assumptions, we found that overall glenohumeral abduction potential is similar between Gorilla and Homo. However, gorillas differ by maintaining high abduction moment capacity with the arm raised above horizontal. This difference is linked to a disparity in soft tissue properties, indicating that scapular morphological features like a cranially oriented scapular spine and glenoid do not enhance the abductor function of the gorilla glenohumeral muscles. A functional enhancement due to differences in skeletal morphology was only demonstrated in the gorilla supraspinatus muscle. Contrary to earlier ideas linking a more obliquely oriented scapular spine to greater supraspinatus leverage, our results suggest that increased lateral projection of the greater tubercle of the humerus accounts for the greater biomechanical performance in Gorilla. This study enhances our understanding of the evolution of gorilla locomotion, as well as providing greater insight into the general interaction between anatomy, function and locomotor biomechanics.

Premolar enamel thickness and distribution of a Miocene hominid Lufengpithecus hudienensis compared with Pleistocene and extant hominids.

Lufengpithecus hudienensis is a Late-Miocene hominid from the Yuanmou basin of southwestern China. Previous studies link Lufengpithecus to either the Sivapithecus-orangutan clade or a derived branch from the basal stem of the Hominidae. Despite a rich fossil assemblage, the taxonomy of L. hudienensis and its phylogenetic relationship with other hominids is still unclear. Enamel thickness in fossil and modern hominids can provide insights into taxonomy, phylogeny, and dietary reconstructions. In this study, 24 upper and lower L. hudienensis premolars were imaged using high-resolution microcomputed tomography. Three-dimensional average enamel thickness and relative enamel thickness (RET) indices, as well as whole-crown enamel distribution patterns were recorded for the L. hudienensis specimens and compared with a total of 113 specimens of fossil and extant apes and recent modern humans (RMH): fossil Pongo (n = 36), Gigantopithecus blacki (n = 21), Pan troglodytes (n = 11), Gorilla gorilla (n = 7), and RMH (n = 38). RET of the premolars of L. hudienensis was found to be similar to that of fossil Pongo, P. troglodytes, and G. gorilla, but less than Gi. blacki and RMH values. The distribution of enamel thickness shows a taxon-specific pattern for L. hudienensis, generally characterized by thicker enamel on the lateral wall than on the cusp apex, on the lingual cusp of the upper and buccal cusp of the lower premolars, with some differences with respect to fossil Pongo, Gi. blacki, P. troglodytes, and G. gorilla patterns. Additional characterizations of enamel thickness distribution patterns of the molars and other dental endostructural morphologies are needed to further explore the phylogenetic relationships of L. hudienensis with other hominids.

A chimpanzee enamel-diet δ13C enrichment factor and a refined enamel sampling strategy: Implications for dietary reconstructions.

Reconstructing diets from stable carbon isotopic signals in enamel bioapatite requires the application of a δ13C enamel-diet enrichment factor, or the isotopic offset between diet and enamel, which has not been empirically determined for any primate. In this study, an enamel-diet enrichment factor (ε∗enamel-diet) of 11.8 ± 0.3‰ is calculated for chimpanzees (Pan troglodytes) at Ngogo in Kibale National Park, Uganda, based on a comprehensive isotopic assessment of previously analyzed dietary plant data and new isotopic analyses of enamel apatite. Different enamel sampling methods are evaluated to determine the potential influence of weaning on isotopic enamel values and dietary interpretations. The new chimpanzee enrichment factor and a sampling strategy that excludes teeth that formed before weaning completion are applied to all known chimpanzee δ13Cenamel data, either previously published or newly derived in this study, resulting in a dietary range of almost 6‰ across all chimpanzees sampled. This new chimpanzee enamel-diet enrichment factor is then used to reassess dietary reconstructions of 12 fossil hominin species whose isotopic enamel signatures have been determined. Results reveal hominin diets that are isotopically more positive than previously reconstructed, highlighting the widespread contribution of 13C-enriched C4/crassulacean acid metabolism (CAM) resources in fossil hominin diets and emphasizing the broad use of these resources during human evolution. These findings stress the importance of ascertaining and employing an appropriate enrichment factor for dietary reconstructions of specific taxa as well as standardizing the sampling protocol for tooth enamel in isotopic paleodietary reconstructions.

Homeotic change in segment identity derives the human vertebral formula from a chimpanzee-like one.

OBJECTIVES: One of the most contentious issues in paleoanthropology is the nature of the last common ancestor of humans and our closest living relatives, chimpanzees and bonobos (panins). The numerical composition of the vertebral column has featured prominently, with multiple models predicting distinct patterns of evolution and contexts from which bipedalism evolved. Here, we study total numbers of vertebrae from a large sample of hominoids to quantify variation in and patterns of regional and total numbers of vertebrae in hominoids. MATERIALS AND METHODS: We compile and study a large sample (N = 893) of hominoid vertebral formulae (numbers of cervical, thoracic, lumbar, sacral, caudal segments in each specimen) and analyze full vertebral formulae, total numbers of vertebrae, and super-regional numbers of vertebrae: presacral (cervical, thoracic, lumbar) vertebrae and sacrococcygeal vertebrae. We quantify within- and between-taxon variation using heterogeneity and similarity measures derived from population genetics. RESULTS: We find that humans are most similar to African apes in total and super-regional numbers of vertebrae. Additionally, our analyses demonstrate that selection for bipedalism reduced variation in numbers of vertebrae relative to other hominoids. DISCUSSION: The only proposed ancestral vertebral configuration for the last common ancestor of hominins and panins that is consistent with our results is the modal formula demonstrated by chimpanzees and bonobos (7 cervical-13 thoracic-4 lumbar-6 sacral-3 coccygeal). Hox gene expression boundaries suggest that a rostral shift in Hox10/Hox11-mediated complexes could produce the human modal formula from the proposal ancestral and panin modal formula.

Hominoid intraspecific cranial variation mirrors neutral genetic diversity.

Natural selection, developmental constraint, and plasticity have all been invoked as explanations for intraspecific cranial variation in humans and apes. However, global patterns of human cranial variation are congruent with patterns of genetic variation, demonstrating that population history has influenced cranial variation in humans. Here we show that this finding is not unique to Homo sapiens but is also broadly evident across extant ape species. Specifically, taxa that exhibit greater intraspecific cranial shape variation also exhibit greater genetic diversity at neutral autosomal loci. Thus, cranial shape variation within hominoid taxa reflects the population history of each species. Our results suggest that neutral evolutionary processes such as mutation, gene flow, and genetic drift have played an important role in generating cranial variation within species. These findings are consistent with previous work on human cranial morphology and improve our understanding of the evolutionary processes that generate intraspecific cranial shape diversity within hominoids. This work has implications for the analysis of selective and developmental pressures on the cranium and for interpreting shape variation in fossil hominin crania.

Mechanics of heel-strike plantigrady in African apes.

The initiation of a walking step with a heel strike is a defining characteristic of humans and great apes but is rarely found in other mammals. Despite the considerable importance of heel strike to an understanding of human locomotor evolution, no one has explicitly tested the fundamental mechanical question of why great apes use a heel strike. In this report, we test two hypotheses (1) that heel strike is a function of hip protraction and/or knee extension and (2) that short-legged apes with a midfoot that dorsiflexes at heel lift and long digits for whom digitigrady is not an option use heel-strike plantigrady. This strategy increases hip translation while potentially moderating the cost of redirecting the center of mass ('collisional costs') during stance via rollover along the full foot from the heel to toes. We quantified hind limb kinematics and relative hip translation in ten species of primates, including lemurs, terrestrial and arboreal monkeys, chimpanzees, and gorillas. Chimpanzees and gorillas walked with relatively extended knees but only with moderately protracted hips or hind limbs, partially rejecting the first hypothesis. Nonetheless, chimpanzees attained relative hip translations comparable with those of digitigrade primates. Heel-strike plantigrady may be a natural result of a need for increased hip translations when forelimbs are relatively long and digitigrady is morphologically restricted. In addition, foot rollover from the heel to toe in large, short-legged apes may reduce energetic costs of redirecting the center of mass at the step-to-step transition as it appears to do in humans. Heel strike appears to have been an important mechanism for increasing hip translation, and possibly reducing energetic costs, in early hominins and was fundamental to the evolution of the modern human foot and human bipedalism.

Metameric variation of upper molars in hominoids and its implications for the diversification of molar morphogenesis.

Metameric variation of molar size is in part associated with the dietary adaptations of mammals and results from slight alterations of developmental processes. Humans and great apes exhibit conspicuous variation in tooth morphology both between taxa and across tooth types. However, the manner in which metameric variation in molars emerged among apes and humans via evolutionary alterations in developmental processes remains largely unknown. In this study, we compare the enamel-dentine junction of the upper molars of humans-which closely correlates with morphology of the outer enamel surface and is less affected by wear-with that of the other extant hominoids: chimpanzees, bonobos, gorillas, orangutans, and gibbons. We used the morphometric mapping method to quantify and visualize three-dimensional morphological variation, and applied multivariate statistical analyses. Results revealed the following: 1) extant hominoids other than humans share a common pattern of metameric variation characterized by a largely linear change in morphospace; this indicates a relatively simple graded change in metameric molar shape; 2) intertaxon morphological differences become less distinct from the mesial to distal molars; and 3) humans diverge from the extant ape pattern in exhibiting a distinct metameric shape change trajectory in the morphospace. The graded shape change and lower intertaxon resolution from the mesial to distal molars are consistent with the concept of a 'key' tooth. The common metameric pattern observed among the extant nonhuman hominoids indicates that developmental patterns underlying metameric variation were largely conserved during ape evolution. Furthermore, the human-specific metameric pattern suggests considerable developmental modifications in the human lineage.

Skull reconstruction of the late Miocene ape Rudapithecus hungaricus from Rudabánya, Hungary.

We report on a computer-based reconstruction of a well-preserved ape skull from late Miocene deposits in Rudabánya, Hungary. Based on micro-computed tomographic scans of the original Rudapithecus hungaricus partial cranium RUD 200 and the associated mandible RUD 212 we realign displaced bone fragments, and reconstruct the shape of the upper and lower jaws guided by occlusal fingerprint analysis of dental wear patterns. We apply geometric morphometric methods based on several hundred landmarks and sliding semilandmarks to estimate missing data, and create multiple reconstructions of the specimen. We then compare the reconstructed overall cranial shape, as well as the volume and shape of the endocast, with extant primates. Multiple reconstructions of RUD 200 yield an average endocranial volume of 234 cc (S.D.: 9 cc; range: 221-247 cc). RUD 200 is most similar to African apes in overall cranial shape, but in a statistical analysis of endocranial shape the specimen falls closest to extant hylobatids. Our data suggest that R. hungaricus from the late Miocene in Europe displays aspects of the overall cranial geometry typical of extant African great apes, but it does not show an evolutionary reorganization of the brain evident in Pan, Gorilla, and Pongo.

Ontogeny of the distal femoral metaphyseal surface and its relationship to locomotor behavior in hominoids.

OBJECTIVE: Distal femoral metaphyseal surface morphology is highly variable in extant mammals. This variation has previously been linked to differences in locomotor behavior. We perform the first systematic survey and description of the development of this morphology in extant hominoids. MATERIALS AND METHODS: We collected 3D surface laser scans of the femora of 179 human and great ape individuals throughout all subadult stages of development. We qualitatively and quantitatively describe metaphyseal surface morphology. RESULTS: We find that the metaphysis is topographically simple in all hominoids during the fetal and infant periods relative to later developmental periods, and in apes it develops significant complexity throughout development. Humans, by contrast, retain relatively flat metaphyseal surfaces throughout ontogeny. DISCUSSION: Major shifts in morphology appear to coincide with major shifts in locomotor behavior, suggesting that metaphyseal morphology is developmentally plastic and highly dependent on the biomechanical loadings at the knee joint. This is consistent with a large body of biomedical research, which demonstrates the primacy of mechanical forces in determining growth plate ossification patterns. Additionally, specific metaphyseal morphology appears highly correlated with specific locomotor modes, suggesting that metaphyseal surface morphology will be useful for reconstructing the locomotor behavior of fossil primate taxa.

Enamel thickness and dental development in Rudapithecus hungaricus.

The fossil record of middle and late Miocene Eurasian hominoids has expanded considerably over the past few decades, particularly with the recovery of numerous isolated teeth and jaws. Scholars have turned to assessments of internal tooth structure and growth to make sense of the evolutionary radiations of these primates as well as their affinities to the living great apes (hominids). Here we characterize full-dentition enamel thickness and dental development in several juvenile Rudapithecus hungaricus individuals using multiple imaging modalities. Relative enamel thickness (RET) values for the anterior teeth and premolars of Rudapithecus are broadly akin to those of gorillas and chimpanzees and are thinner than those of orangutans. First molar RET values are most similar to chimpanzees, while posterior molar values are closer to thicker-enameled orangutans. When compared to Miocene hominoids, Rudapithecus shows an intermediate molar RET condition that is especially similar to other dryopithecines. Long-period line periodicity values are comparable to African apes and most Miocene hominoids, and lower than living and fossil orangutans. The mean cuspal daily secretion rate is similar to that of several other Miocene hominoids but is greater than extant great apes. Cusp-specific molar crown formation times generally exceed those of chimpanzees, are lower than those of orangutans, and are broadly like those of other Miocene apes. While Rudapithecus appears to have a somewhat unique pattern of enamel thickness and dental development relative to individual great ape genera, these structural and developmental features are consistent with its designation as a hominid.

New hominoid fossils from Moroto II, Uganda and their bearing on the taxonomic and adaptive status of Morotopithecus bishopi.

The early Miocene site of Moroto II, Uganda has yielded some of the oldest known hominoid fossils. A new partial mandible (UMP MORII 03'551) is notable for its long tooth row and large, narrow M2 with well-developed cristids - a morphological combination previously unknown for large bodied catarrhines of the Early Miocene and suggesting folivory. The tooth proportions are compatible with belonging to the same taxon as the maxilla UMP 62-11, the holotype of Morotopithecus bishopi; likewise, the long tooth row and vertical planum of UMP MORII 03'551 suggest that it may represent the same taxon as mandible(s) UMP 66-01 and UMP 62-10. Canine size strongly suggests UMP MORII 03'551 is a female. Comparisons of the tooth crown morphology and tooth row proportions, relative enamel thickness, enamel-dentine junction morphology, long-period line periodicity, and dental wear patterns support significant morphological, developmental, and inferred dietary differentiation, and therefore generic-level distinctiveness, among Afropithecus, Morotopithecus and the Proconsul clade. An isolated M1 (UMP MORII 03'559) is morphologically dissimilar, and much smaller than the actual or inferred size of molars in UMP MORII 03'551, UMP 66-01 and UMP 62-10, supporting the presence of two hominoid taxa at Moroto II, M. bishopi and a smaller bodied proconsulid. Given the high level of body mass dimorphism inferred for Morotopithecus and other early Miocene catarrhines, the known postcrania from Moroto II could be attributable to either taxon. However, UMP MORII 03'551 and the femora UMP MORII 94'80 derive from the same stratigraphic interval, while the isolated M1 was deposited later, increasing the likelihood that the mandible and femora are from the same individual. These new fossils expand our understanding of the taxonomic and adaptive diversity of early Miocene catarrhines.

Three-dimensional geometric morphometric analysis of the distal radius insertion sites of the palmar radiocarpal ligaments in hominoid primates.

OBJECTIVES: To identify anatomic differences in the insertion sites of the palmar radiocarpal ligaments in different species of hominoid primates that may be related to their different types of locomotion. MATERIALS AND METHODS: We have used three-dimensional geometric morphometrics (3D GM) to analyze the distal radius ligament insertion sites in 31 Homo sapiens, 25 Pan troglodytes, 31 Gorilla gorilla, and 15 Pongo pygmaeus. We have also dissected the radioscaphocapitate (RSC), long radiolunate (LRL) and short radiolunate (SRL) ligaments in six H. sapiens and five P. troglodytes to obtain quantitative values that were then compared with the results of the 3D GM analysis. RESULTS: H. sapiens had a relatively larger insertion site of the RSC + LRL ligament than the other hominoid primates. P. pygmaeus and P. troglodytes had a relatively large SRL ligament insertion site with a palmar orientation. In G. gorilla, the two ligament insertion sites were relatively smaller and the SRL insertion site had an ulnopalmar orientation. DISCUSSION: The morphological differences observed can be related to the types of locomotion used by the different species and to quantitative data obtained from the dissection of ligaments in H. sapiens and P. troglodytes. 3D GM analysis of ligament insertion sites can help in interpreting the types of locomotion used by extinct hominoid primates through the analysis of preserved fossilized fragments of the distal radius.

The gibbon's Achilles tendon revisited: consequences for the evolution of the great apes?

The well-developed Achilles tendon in humans is generally interpreted as an adaptation for mechanical energy storage and reuse during cyclic locomotion. All other extant great apes have a short tendon and long-fibred triceps surae, which is thought to be beneficial for locomotion in a complex arboreal habitat as this morphology enables a large range of motion. Surprisingly, highly arboreal gibbons show a more human-like triceps surae with a long Achilles tendon. Evidence for a spring-like function similar to humans is not conclusive. We revisit and integrate our anatomical and biomechanical data to calculate the energy that can be recovered from the recoiling Achilles tendon during ankle plantar flexion in bipedal gibbons. Only 7.5% of the required external positive work in a stride can come from tendon recoil, yet it is delivered at an instant when the whole-body energy level drops. Consequently, an additional similar amount of mechanical energy must simultaneously dissipate elsewhere in the system. Altogether, this challenges the concept of an energy-saving function in the gibbon's Achilles tendon. Cercopithecids, sister group of the apes, also have a human-like triceps surae. Therefore, a well-developed Achilles tendon, present in the last common 'Cercopithecoidea-Hominoidea' ancestor, seems plausible. If so, the gibbon's anatomy represents an evolutionary relict (no harm-no benefit), and the large Achilles tendon is not the premised key adaptation in humans (although the spring-like function may have further improved during evolution). Moreover, the triceps surae anatomy of extant non-human great apes must be a convergence, related to muscle control and range of motion. This perspective accords with the suggestions put forward in the literature that the last common hominoid ancestor was not necessarily great ape-like, but might have been more similar to the small-bodied catarrhines.

Segmental morphometrics of bonobos (Pan paniscus): are they really different from chimpanzees (Pan troglodytes)?

The inertial properties of body segments reflect performance and locomotor habits in primates. While Pan paniscus is generally described as more gracile, lighter in body mass, and as having relatively longer and heavier hindlimbs than Pan troglodytes, both species exhibit very similar patterns of (quadrupedal and bipedal) kinematics, but show slightly different locomotor repertoires. We used a geometric model to estimate the inertial properties for all body segments (i.e. head, trunk, upper and lower arms, hand, thigh, shank and foot) using external length and diameter measurements of 12 anaesthetized bonobos (eight adults and four immatures). We also calculated whole limb inertial properties. When we compared absolute and relative segment morphometric and inertial variables between bonobos and chimpanzees, we found that adult bonobos are significantly lighter than adult chimpanzees. The bonobo is also shorter in head length, upper and lower arm lengths, and foot length, and is generally lighter in most absolute segment mass values (except head and hand). In contrast, the bonobo has a longer trunk. When scaled relative to body mass, most differences disappear between the two species. Only the longer trunk and the shorter head of the bonobo remain apparent, as well as the lighter thigh compared with the chimpanzee. We found similar values of natural pendular periods of the limbs in both species, despite differences in absolute limb lengths, masses, mass centres (for the hindlimb) and moments of inertia. While our data contradict the commonly accepted view that bonobos have relatively longer and heavier hindlimbs than chimpanzees, they are consistent with the observed similarities in the quadrupedal and bipedal kinematics between these species. The morphological differences between both species are more subtle than those previously described from postcranial osteological materials.

A new species of Simiolus from the middle Miocene of the Tugen Hills, Kenya.

A new species of the "small-bodied ape" Simiolus is described here that extends the temporal range of the genus to the end of the Middle Miocene. As such, it is one of the few species of fossil primates known from East Africa during a time of significant change in which Old World monkeys and crown hominoids replaced the primitive ape-like primates that had dominated the early Miocene. The dynamics of this important event in our evolutionary history are obscured by the small number of fossil primates known from Africa between 14 and 6 million years ago, as well as persistent ambiguity regarding the phylogenetic status of the ape-like Miocene primates. The new species described here helps to fill this temporal gap, and our analysis of its phylogenetic position suggests that Simiolus and many other Miocene primates were not only ape-like, they were, indeed, stem hominoids. Judging from the available material, the new species may be the smallest known ape.