Locomotor adaptations in the Early Miocene species Diamantomys luederitzi (Rodentia, Mammalia) from Uganda (Napak).

The study of morphological adaptations to different ecological parameters among fossil vertebrates has been an important challenge in recent decades. In this paper, we focus on the link between morphological traits and locomotor behavior such as terrestriality, fossoriality and arboreality (including gliding). One of the most diverse groups in which various locomotor habits are represented is rodents, occupying a wide range of ecological niches. This work highlights morphological variations in skulls and humerus in extant rodents with varying locomotion, to predict this parameter in the extinct species Diamantomys luederitzi (Early Miocene, Napak, Uganda). Linear discriminant analysis and phylogenetic flexible discriminant analysis are used to analyze datasets obtained via traditional morphometry (measurements) and geometric morphometrics (landmarks). The results show good discrimination between locomotor groups for both structures in extant species: the skull has a wider and longer rostrum in terrestrial and fossorial taxa compared to arboreal rodents, is also higher and posteriorly wider in fossorial taxa; the distal humerus shows elongation of the trochlea and capitulum and a higher trochlea in fossorial and terrestrial species, allowing an increase of stability instead of mobility, which is more important in arboreal taxa for movement in trees. In D. luederitzi, all skull analyses except one predicted it as a terrestrial species, the other prediction as a glider was possibly linked to the diet. For the distal humerus, this species has been predicted as a terrestrial, fossorial and arboreal taxon in differing analyses, reflected by morphological traits represented in these different locomotor categories. These varying predictions could highlight the intraspecific variation in this fossil species as well as its locomotor repertoire, raising a discussion about the use of different methods in such analyses. In addition to these predictions, several issues are discussed, such as the presence of locomotor signal in the skull and its validity in locomotor studies, as well as the relevance of the use of fragmentary material in such analyses. The results obtained in this work highlight the importance of the locomotor signal in these structures, as well as the possibility of taking into account poorly preserved material, in particular the distal humerus.

Morphology and environment in some fossil Hominoids and Pedetids (Mammalia).

Linking the environment to functional anatomy is not an easy exercise, especially when dealing with fossils, which are often fragmentary and represent animals that are extinct. A holistic approach permits us to fill the gaps in reconstructing the evolutionary patterns in fossil groups. Identifying the environment where animals lived can help to interpret some anatomical structures and, vice versa, the functional morphological pattern can help to refine some fossil environments. Two examples focusing on locomotor behaviours in fossil mammals are considered in this paper: the hominoids and the origins of hominid bipedalism and the springing adaptations in fossil rodents (Pedetidae) in connection with different habitats. In the first case, the limits of the chimp-based models and the necessity to take into account detailed environmental reconstructions will be addressed. The famous 'savannah hypothesis' is no longer tenable because the palaeontological data support a more vegetated environment for the origins of bipedal hominids. Data from the environment will be considered. The earliest putative hominid fossils which preserve skeletal remains of the locomotor apparatus show mixed adaptations to terrestrial bipedalism and arboreal activities. The second example focuses on the variation in springing adaptations in Pedetidae in the Lower Miocene of East Africa and Southern Africa. In the East, the sites where Pedetidae were preserved were mainly forested, whereas in the South the region was more open and drier, with extensive grassy patches. In the first case, pedetids were robust and heavy jumpers, whereas in the South they were smaller, their skeleton more gracile and their springing was lighter. During the desertification of the southern part of Africa, the large pedetid species became extinct, but a smaller species developed. In the case of primates, as in the case of rodents, the skeletal morphology was adapted to its environment.

Reassessment of Dolichopithecus (Kanagawapithecus) leptopostorbitalis, a colobine monkey from the Late Pliocene of Japan.

In the original description of Dolichopithecus (Kanagawapithecus) leptopostorbitalis, Iwamoto, Hasegawa and Koizumi, 2005, a moderately large-sized colobine monkey from the Late Pliocene of central Japan, affinities to the European Dolichopithecus rather than to the Transbaikalian Parapresbytis were noted based on the similarities in cranial morphology. Computed tomography scans confirm the presence of the maxillary sinus in the holotype, whereas it is probably absent in specimens of the European Dolichopithecus ruscinensis, the type species of this genus. This feature is either present or absent homogeneously in any given genus of living anthropoids. Its presence or absence is unknown in Parapresbytis, but the distinct morphology of the maxillary incisors in this taxon suggests that this form had different feeding habits from the Japanese colobines. These findings suggest that the Japanese colobine should be referred to henceforth as Kanagawapithecus leptopostorbitalis. Kanagawapithecus shares many important facial and dental features with Dolichopithecus rather than with Parapresbytis, but this association depends largely on the limited availability of comparable materials for the latter. Among colobines, the presence of the maxillary sinus is recorded only in Libypithecus and Cercopithecoides. The maxillary sinus is absent in all modern Asian colobines, implying that Kanagawapithecus is an isolated form without any relationship to living forms. Nevertheless, such phylogenetic interpretations are largely dependent on the restricted fossil evidence from the Pliocene and Pleistocene of eastern Eurasia and will be reexamined when new findings are made.

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.

Preliminary analysis of Nacholapithecus 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.

From apes to humans: locomotion as a key feature for phylogeny.

If bipedalism has often been considered to be of a crucial interest for understanding hominid evolution, the acceptance of locomotor features to build phylogenies is still far from being a reality in the field. Especially for hominid evolution, it still seems to be difficult to accept that traits, other than craniodental ones, can be useful for defining the major dichotomies. The recent discovery of Australopithecus anamensis suggests a challenging view of the major dichotomy between apes and humans. Whilst it is widely accepted that Ardipithecus ramidus is ancestral to Australopithecus anamensis, which in its turn is ancestral to Australopithecus afarensis and then to later hominids, the postcranial adaptations, which should be taken into account, suggest another branching pattern. Based on the fact that by 4.0 million years two different locomotor patterns can be identified in hominids, two lineages would appear to be present: the "Australopithecine" lineage (with Australopithecus afarensis or Ardipithecus ramidus if the latter is really a hominid sensu stricto) and the "Hominine" lineage (with Australopithecus anamensis = Praeanthropus africanus).