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.

The extinct river shark Glyphis pagoda from the Miocene of Myanmar and a review of the fossil record of the genus Glyphis (Carcharhiniformes: Carcharhinidae).

We redescribe an extinct river shark, Glyphis pagoda (Noetling), on the basis of 20 teeth newly collected from three different Miocene localities in Myanmar. One locality is a nearshore marine deposit (Obogon Formation) whereas the other two localities represent terrestrial freshwater deposits (Irrawaddy sediments), suggesting that G. pagoda from the Irrawaddy sediments was capable of tolerating low salinity like the extant Glyphis. Glyphis pagoda likely reached up to at least 185 cm in total body length and was probably piscivorous. The fossil species occurs in rocks of Myanmar and eastern and western India and stratigraphically ranges at least from the Lower Miocene (Aquitanian) to the lower Upper Miocene (mid-Tortonian). It has been classified under at least eight other genera to date, along with numerous taxonomic synonyms largely stemming from the lack of understanding of the heterodonty in extant Glyphis in the original description. Our literature review suggests that known Miocene shark faunas, particularly those in India, are manifested with unreliable taxonomic identifications and outdated classifications that warrant the need for a comprehensive taxonomic review in order to evaluate the evolutionary history and diversity pattern of Miocene shark faunas. The genus Glyphis has a roughly 23-million-year-long history, and its success may be related to the evolution of its low salinity tolerance. While extant Glyphis spp. are considered to be particularly vulnerable to habitat degradation and overfishing, the fossil record of G. pagoda provides renewed perspective on the natural history of the genus that can be taken into further consideration for conservation biology of the extant forms.

First discovery of colobine fossils from the Late Miocene/Early Pliocene in central Myanmar.

Here we report two kinds of colobine fossils discovered from the latest Miocene/Early Pliocene Irrawaddy sediments of the Chaingzauk area, central Myanmar. A left mandibular corpus fragment preserving M1-3 is named as a new genus and species, Myanmarcolobus yawensis. Isolated upper (M(1)?) and lower (M2) molars are tentatively identified as Colobinae gen. et sp. indet. Although both forms are medium-sized colobines, they are quite different from each other in M2 morphology. The isolated teeth of the latter show typical colobine-type features, so it is difficult to identify their taxonomic position, whereas lower molars of Myanmarcolobus have unique features, such as a trapezoid-shaped long median lingual notch, a deeply concave median buccal cleft, a strongly developed mesiobuccal notch, and rather obliquely running transverse lophids. Compared with fossil and living Eurasian colobine genera, Myanmarcolobus is most similar in lower molar morphology to the Pliocene Dolichopithecus of Europe rather than to any Asian forms. In Dolichopithecus, however, the tooth size is much larger and the median lingual notch is mesiodistally much shorter than that of Myanmarcolobus. The discovery of Myanmarcolobus in central Myanmar is the oldest fossil record in Southeast Asia not only of colobine but also of cercopithecid monkeys and raises many questions regarding the evolutionary history of Asian colobine monkeys.

Prenatal ontogeny of subspecific variation in the craniofacial morphology of the Japanese macaque (Macaca fuscata).

We cross-sectionally investigated prenatal ontogeny of craniofacial shape in the two subspecies of the Japanese macaque (Macaca fuscata fuscata and Macaca fuscata yakui) using a geometric morphometric technique to explore the process of morphogenetic divergence leading to the adult morphological difference between the subspecies. The sample comprised a total of 32 formalin-fixed fetal specimens of the two subspecies, in approximately the second and third trimesters. Each fetal cranium was scanned using computed tomography to generate a three-dimensional surface model, and 68 landmarks were digitized on the external and internal surface of each cranium to trace the growth-related changes in craniofacial shape of the two subspecies. The results of our study demonstrated that the two subspecies generally shared the same craniofacial growth pattern. Both crania tend to exhibit relative contraction of the neurocranium in the mediolateral and superoinferior directions, a more superiorly positioned cranial base, a more vertically oriented occipital squama, and a more anteriorly positioned viscerocranium as the cranial size increased. However, distinctive subspecific differences, for example relatively narrower orbital breadth, higher orbit, higher position of the nuchal crest, and more protrudent snout found in Macaca fuscata yakui were already present during the prenatal period. This study demonstrated that morphological differentiation in the craniofacial shape may occur at a very early stage of the fetal period even between closely related subspecies of the Japanese macaque.

Femur length, body mass, and stature estimates of Orrorin tugenensis, a 6 Ma hominid from Kenya.

To understand the palaeobiology of extinct hominids it is useful to estimate their body mass and stature. Although many species of early hominid are poorly preserved, it is occasionally possible to calculate these characteristics by comparison with different extant groups, by use of regression analysis. Calculated body masses and stature determined using these models can then be compared. This approach has been applied to 6 Ma hominid femoral remains from the Tugen Hills, Kenya, attributed to Orrorin tugenensis. It is suggested that the best-preserved young adult individual probably weighed approximately 35-50 kg. Another fragmentary femur results in larger estimates of body mass, indicative of individual variation. The length of the femur of the young adult individual was estimated, by using anthropoid-based regression, to be a minimum of 298 mm. Because whole-femur proportions for Orrorin are unknown, this prediction is conservative and should be revised when additional specimens become available. When this predicted value was used for regression analysis of bonobos and humans it was estimated to be 1.1-1.2 m tall. This value should, however, be viewed as a lower limit.

Additional materials of Myanmarpithecus yarshensis (Amphipithecidae, Primates) from the middle Eocene Pondaung Formation.

Myanmarpithecus yarshensis is an amphipithecid primate from the middle Eocene Pondaung Formation in Myanmar. It was previously known based on maxillary fragments with P(4)-M(3) and mandibular fragments with C-P(3) and M(2-3). This study reports new materials for the genus, including a humeral head fragment, a lingual fragment of the right M(2), a lingual fragment of the right M(3), and a left I(1). These new materials were collected from approximately the same point, and likely belonged to the same individual. The upper molar morphology and size of the new materials show similarity to those of the type specimen, indicating that the new materials can be assigned to M. yarshensis. The humeral head is the first postcranial element that is associated with dental materials for amphipithecids. The morphological similarity between the previously reported larger humerus and this specimen confirms the assignment of the former specimen to Amphipithecidae and suggests common locomotor adaptations in the family. The upper central incisor is large relative to the molar fragments, but is within the variation among extant platyrrhines. The tooth is spatulate-shaped and high crowned, and lacks the mesial process, indicating similarity to I(1) of haplorhines and clear differences from that of adapoids. It has been suggested that amphipithecids, including Myanmarpithecus, have affinities with notharctine adapoids, but the morphology of I(1) does not support the notharctine hypothesis of the Amphipithecidae.

Variation in maxillary sinus anatomy among platyrrhine monkeys.

Variations in the maxillary sinus anatomy of extant and fossil catarrhine primates have been extensively examined using computed tomography (CT), and have potential utility for phylogenetic analyses. This approach has also been used to demonstrate its anatomical variation in eight of the 16 extant genera of platyrrhines and the absence of the sinus in Saimiri and Cacajao. We used this approach to evaluate the three-dimensional anatomy of the maxillary sinus in all extant platyrrhine genera, and here argue the phylogenic implications of this variation. This study confirms, for the most part, previous CT studies and augments them with the six genera not studied previously: Ateles, Lagothrix, Callithrix, Cebuella, Pithecia and Chiropotes. The entire maxilla is pneumatized by the sinus in the atelines, Cebus, and Callicebus, whereas the sinus pneumatizes only the medial part of the maxilla in the callitrichines and Aotus. Pithecia has a unique conformation in which the maxillary sinus and the expanded inferior meatus pneumatize the posteromedial and anterolateral parts of the entire maxilla, respectively. Chiropotes has no sinus, and the inferior meatus possibly expands into the area between the middle meatus and medial surface of the maxilla to disturb sinus formation, as in the case of its close relative Cacajao. Finally, we argue that the sinus that pneumatizes the entire maxilla is a primitive feature in extant platyrrhines and was probably shared by the last common ancestor of the anthropoids.

The paleobiology of Amphipithecidae, South Asian late Eocene primates.

Analysis of the teeth, orbital, and gnathic regions of the skull, and fragmentary postcranial bones provides evidence for reconstructing a behavioral profile of Amphipithecidae: Pondaungia, Amphipithecus, Myanmarpithecus (late middle Eocene, Myanmar) and Siamopithecus (late Eocene, Thailand). At 5-8 kg, Pondaungia, Amphipithecus, and Siamopithecus are perhaps the largest known Eocene primates. The dental and mandibular anatomy suggest that large-bodied amphipithecids were hard-object feeders. The shape of the mandibular corpus and stiffened symphysis suggest an ability to resist large internal loads during chewing and to recruit significant amounts of muscle forces from both the chewing and non-chewing sides of the jaw so as to increase bite force during mastication. The large spatulate upper central incisor of Pondaungia and projecting robust canines of all the larger amphipithecids suggest that incisal food preparation was important. The molars of Siamopithecus, Amphipithecus, and Pondaungia have weak shearing crests. This, and the thick molar enamel found in Pondaungia, suggests a diet of seeds and other hard objects low in fiber. In contrast, Myanmarpithecus was smaller, about 1-2 kg; its cheek teeth suggest a frugivorous diet and do not imply seed eating. Postcranial bones (humerus, ulna, and calcaneus) of a single large amphipithecid individual from Myanmar suggest an arboreal quadrupedal locomotor style like that of howler monkeys or lorises. The humeral head is rounded, proximally oriented, and the tuberosities are low indicating an extremely mobile glenohumeral joint. The great thickness of the midshaft cortical bone of the humerus implies enhanced ability to resist bending and torsion, as seen among slow moving primate quadrupeds. The elbow joint exhibits articular features for enhanced stability in habitually flexed positions, features also commonly found in slow moving arboreal quadrupeds. The short distal load arm of the calcaneus is consistent with, but not exclusive to, slow, arboreal quadrupedalism, and suggests no reliance on habitual leaping.

Endocranial cast and morphology of the olfactory bulb of Amphipithecus mogaungensis (latest middle Eocene of Myanmar).

A detailed endocranial cast of the olfactory bulb of Amphipithecus mogaungensis, a latest middle Eocene primate from the Pondaung Formation (Myanmar), was studied in comparison with some Paleogene primates, the olfactory bulb of which has been reported. The olfactory bulb of Amphipithecus is located just anterior to the postorbital constriction, that is, within the interorbital septum. It is relatively large and pedunculate, not overlapped by the frontal lobe, and consists of two parallel aligned bodies. The relative volume of the olfactory bulb shows the same pattern as in adapiforms, but the location and bilobed form are more similar to those of omomyoids than of adapiforms.