PRIMATE LIMB CONFORMATION: IMPLICATIONS FOR MECHANICAL ENERGY CONSERVATION

In order to understand the energy implications of primate limb conformation, the biomechanics of swing phase and the vertical movements of limb center of gravity were examined in the arboreal rhesus macaque (Macaca mulatta) and the semi-terrestrial long-tailed macaque (Macaca fascicularis). The objective was to better understand the potential variation in locomotor adaptations in these two macaque species within different ecological environments. In particular, the objective of this study was to identify the implications that these movements have on the internal energy costs of locomotion. I show that the biomechanics of swing phase for these two species have important effects on their limb center of gravity, although they may have similar limb segment mass distributions. This study suggests that mechanical energy conservation during swing phase is important in mammalian locomotion.

Splendid Innovation: The Extinct South American Native Ungulates

A remarkable diversity of plant-eating mammals known as South American native ungulates (SANUs) flourished in South America for most of the Cenozoic. Although some of these species likely filled ecological niches similar to those of modern hoofed mammals, others differed substantially from extant artiodactyls and perissodactyls in their skull and limb anatomy and probably also in their ecology. Notoungulates and litopterns were the longest-lived and most diverse SANU clades and survived into the Quaternary; astrapotheres went extinct in the late Miocene, whereas other SANU groups were restricted to the Paleogene. Neogene notoungulates were quite specialized in craniodental structure, but many were rather unspecialized postcranially; in contrast, litopterns evolved limb specializations early in their history while maintaining more conservative dentitions. In this article, we review the current understanding of SANU evolutionary relationships and paleoecology, provide an updated compilation of genus temporal ranges, and discuss possible directions for future research. ▪ South American native ungulates (SANUs) were a diverse, long-lived, and independent radiation of mammals into varied terrestrial plant-eater niches. ▪ We review origins, evolution, and paleoecology of the major SANU clades: Notoungulata, Litopterna, Astrapotheria, Xenungulata, and Pyrotheria. ▪ At their peak, during the Eocene and Oligocene, more than 40 genera of native ungulates inhabited South America at any one time. ▪ SANUs ranged from <1 kg to several tons and evolved many combinations of diet and locomotor adaptations not seen in living ungulates.

Terrestrial Locomotor Evolution in Urban Environments

The structural habitat of terrestrial urban environments can differ drastically from environments less impacted by human activities. Whether or not urban species use anthropogenic structures, they are subject to novel selection pressures to effectively locomote. Urban environments are distinctly more open than non-urban habitats, they offer few refuges, and habitat space is patchy with clustered perches. Animals must either change their behaviour to use only natural substrates or contend with manufactured substrates. Arboreal species are particularly impacted because the anthropogenic structures with which they interact, even if infrequently, differ from trees in structural, material, and surface properties. The chapter explores potential adaptive responses to the spatial structure and properties of climbing substrates in urban environments relevant to terrestrial and climbing locomotion. For each, the authors first discuss differences between urban and non-urban terrestrial habitats relevant to locomotion. They then discuss how these differences influence behaviour and locomotor demands, providing a mechanism through which natural selection shapes morphology. Lastly, they discuss the morphological traits most likely to be impacted by these altered demands and predict how natural selection may affect these traits in urban environments based on biomechanical principles. As there have been very few studies investigating urban morphological adaptation related to locomotion, the chapter draws on trait–environment relationships in natural environments. The discussion provides a starting point for developing rigorous hypotheses about functionally relevant trait shifts in urban environments and future directions for investigating locomotor adaptations in urban species.

Castor-like postcranial adaptation in an uppermost Miocene beaver from the Staniantsi Basin (NW Bulgaria)

The Staniantsi-Mazgoš Basin is one of several Neogene intramontane basins in NW Bulgaria. Recent fieldwork in the open pit coal mine yielded material of an exceptional diversity of vertebrates from the uppermost Miocene. In particular, skeletal remains of a large Castorinae are very numerous and well-preserved. Here we perform a comparative morphological description of the postcranium of the Staniantsi-beaver and compare it with the extant Castor fiber and closely related fossil taxa, noting an overall high degree of similarity in many cases. Analyses of the functional anatomy confirm similar locomotor adaptations of the large Staniantsi-beaver and the extant Castor fiber. It is shown that the hindlimb exhibits typical adaptations for swimming and the forelimb is modified for a primary fossorial movement. Further, the caudal vertebrae indicate a flattened tail. Minor osteological differences can be used in a future clarification of the taxonomic status of this fossil castorine. In conclusion, the large Staniantsi-beaver seems to be equivalent to the extant Castor fiber and several fossil castorids in his locomotor adaptations and fits perfectly into the assumed swampy to lacustrine palaeoenvironment of the Staniantsi-Mazgoš Basin. Our results highlight the importance of the postcranium for reconstructing the palaeobiology and elucidating the taxonomy of fossil rodents.

Comparative functional skeletal morphology among three genera of shrews: implications for the evolution of locomotor behavior in the Soricinae (Eulipotyphla: Soricidae)

The clade comprising the soricid tribes Blarinellini (Blarinella) and Blarinini (Blarina and Cryptotis) is notable within the Soricidae (Eulipotyphla) for the large proportion of reportedly semifossorial species. To better define locomotor modes among species in these two tribes, we quantified purported locomotor adaptations by calculating 23 functional indices from postcranial measurements obtained from museum specimens of Blarina and Blarinella and published measurements for 16 species of Cryptotis. We then analyzed relative ambulatory–fossorial function of each species using principal component analyses and mean percentile rank (MPR) analysis of the indices. Species within the Blarinellini–Blarinini clade exhibit a graded series of morphologies with four primary functional groupings that we classified as “ambulatory,” “intermediate,” “semifossorial,” and “fossorial.” To obtain a preliminary overview of evolution of locomotor modes in this group, we mapped MPRs on a composite phylogeny and examined the resulting patterns. That analysis revealed that the most recent common ancestor of the Blarinellini–Blarinini clade most likely had an intermediate or semifossorial locomotor morphology. Individual subclades subsequently evolved either more ambulatory or more fossorial morphologies. Hence, evolution of locomotor traits within this clade is complex. Multiple shifts in locomotor mode likely occurred, and no single directional tendency is apparent either among the major modes or in levels of complexity.

A tail of evolution: evaluating body length, weight and locomotion as potential drivers of tail length scaling in Australian marsupial mammals

Although mammalian tail length relative to body length is considered indicative of locomotor mode, this association has been difficult to quantify. This could be because the counterweight function of the tail might associate it more with body weight than body length. Alternatively, relative tail length might not be evolutionarily flexible owing to its integration with the remaining skeleton, particularly the spine. Using comparative analyses of morphological means and ranges in Australian marsupials, including the first co-assessment with body weight, our study supports the second hypothesis, i.e. tail length ranges within species, and tail lengths among species are explained better by body length than by body weight. However, all three variables do not differ in phylogenetic signal or rates of evolution. Associations of tail lengths with locomotion are limited, but suggest that scaling slopes, rather than intercepts, are responsible for limited divergence between relative tail lengths at different locomotor modes. This complicates (palaeo-)ecological interpretations of tail length further. We conclude that relative tail length is not a strong predictor of locomotor mode, probably owing to strong integration of tail and body length. The many well-documented bony and soft-tissue adaptations of tails are likely to be better suited to interpretations of locomotor adaptations.

Appendicular skeleton of Protoceratops andrewsi (Dinosauria, Ornithischia): comparative morphology, ontogenetic changes, and the implications for non-ceratopsid ceratopsian locomotion

Protoceratops andrewsi is a well-known ceratopsian dinosaur from the Djadokhta Formation (Upper Cretaceous, Mongolia). Since the 1920s, numerous skeletons of different ontogenetic stages from hatchlings to adults, including fully articulated specimens, have been discovered, but the postcranial anatomy of Protoceratops has not been studied in detail. A new, mostly articulated subadult individual provides an excellent opportunity for us to comprehensively describe the anatomy of the limb skeleton, to compare to other ceratopsian dinosaurs, and to study the ontogenetic and intraspecific variation in this species. New data provided by the specimen shed light on the lifestyle of P. andrewsi. The young subadult individuals present an array of morphological characters intermediate between the bipedal Psittacosaurus and fully quadrupedal adult P. andrewsi. We compare these observations with a broad range of non-ceratopsid Neoceratopsia (of various locomotor adaptations) and Psittacosauridae (obligate bipeds), which gives us insight into the evolution of the skeletal characters informative for the postural change in ceratopsian dinosaurs.