Locomotor Habits Research Articles

Humeral Cross‐Sectional Shape in Suspensory Primates and Sloths

Studies on the cross-sectional geometry of long bones in African apes have documented that shape ratios derived from second moments of area about principle axes (e.g., Imax /Imin ) are often correlated with habitual locomotor behaviors. For example, humeral cross-sections tend to appear more circular in more arboreal and forelimb suspensory chimpanzees compared with terrestrial quadrupedal gorillas. These data support the hypothesis that cross-sections that are more circular in shape are adapted for multidirectional loading regimes and bending moments encountered when using acrobatic locomotor behaviors. Whether a more circular humerus reflects greater use of forelimb suspension in other primates and nonprimate mammals is unknown. In this study, cross-sections at or near midshaft of the humerus were obtained from anthropoid primates that differ in their use of forelimb suspension, as well as from two genera of suspensory sloths. Imax /Imin ratios were compared within and between groups, and correlations were made with behavioral data. In broad comparisons, observed differences in morphology follow predicted patterns. Humeri of suspensory sloths are circular. Humeri of the more suspensory hominoids tend to be more circular than those of quadrupedal taxa. Humeri of the suspensory atelines are similar to hominoids, while those of Cebus are more like nonsuspensory cercopithecoids. There is, however, considerable overlap between taxa and within finer comparisons variation between species are not in the predicted direction. Thus, although Imax /Imin ratios of the humerus are informative for characterizing generalized locomotor modes (i.e., forelimb suspensory vs. quadrupedal), additional structural information is needed for more fine-grained assessments of locomotion.

Rauisuchia

Abstract ‘Rauisuchia’ comprises Triassic pseudosuchians that ranged greatly in body size, locomotor styles and feeding ecologies. Our concept of what constitutes a rauisuchian is changing as a result of discoveries over the last 15 years. New evidence has shown that rauisuchians are probably not a natural (monophyletic) group, but instead are a number of smaller clades (e.g. Rauisuchidae, Ctenosauriscidae, Shuvosauridae) that may not be each other's closest relatives within Pseudosuchia. Here, we acknowledge that there are still large gaps in the basic understanding in the alpha-level taxonomy and relationships of these groups, but good progress is being made. As a result of renewed interest in rauisuchians, an expanding number of recent studies have focused on the growth, locomotor habits, and biomechanics of these animals, and we review these studies here. We are clearly in the midst of a renaissance in our understanding of rauisuchian evolution and the continuation of detailed descriptions, the development of explicit phylogenetic hypotheses, and explicit palaeobiological studies are essential in advancing our knowledge of these extinct animals.

Myosin Isoform Fiber Type and Fiber Size in the Tail of the Virginia Opossum (Didelphis virginiana)

Muscle fiber type is a well studied property in limb muscles, however, much less is understood about myosin heavy chain (MHC) isoform expression in caudal muscles of mammalian tails. Didelphid marsupials are an interesting lineage in this context as all species have prehensile tails, but show a range of tail-function depending on either their arboreal or terrestrial locomotor habits. Differences in prehensility suggest that MHC isoform fiber types may also be different, in that terrestrial opossums may have a large distribution of oxidative fibers for object carrying tasks instead of faster, glycolytic fiber types expected in mammals with long tails. To test this hypothesis, MHC isoform fiber type and their regional distribution (proximal/transitional/distal) were determined in the tail of the Virginia opossum (Didelphis virginiana). Fiber types were determined by a combination of myosin-ATPase histochemistry, immunohistochemistry, and SDS-PAGE. Results indicate a predominance of the fast MHC-2A and -2X isoforms in each region of the tail. The presence of two fast isoforms, in addition to the slow MHC-1 isoform, was confirmed by SDS-PAGE analysis. The overall MHC isoform fiber type distribution for the tail was: 25% MHC-1, 71% MHC-2A/X hybrid, and 4% MHC-1/2A hybrid. Oxidative MHC-2A/X isoform fibers were found to be relatively large in cross-section compared to slow, oxidative MHC-1 and MHC-1/2A hybrid fibers. A large percentage of fast MHC-2A/X hybrids fibers may be suggestive of an evolutionary transition in MHC isoform distribution (fast-to-slow fiber type) in the tail musculature of an opossum with primarily a terrestrial locomotor habit and adaptive tail-function.

Postcranial morphology and the locomotor habits of living and extinct carnivorans

Members of the order Carnivora display a broad range of locomotor habits, including cursorial, scansorial, arboreal, semiaquatic, aquatic, and semifossorial species from multiple families. Ecomorphological analyses from osteological measurements have been used successfully in prior studies of carnivorans and rodents to accurately infer the locomotor habits of extinct species. This study uses 20 postcranial measurements that have been shown to be effective indicators of locomotor habits in rodents and incorporates an extensive sample of over 300 individuals from more than 100 living carnivoran species. We performed statistical analyses, including analysis of variance (ANOVA) and stepwise discriminant function analysis, using a set of 16 functional indices (ratios). Our ANOVA results reveal consistent differences in postcranial skeletal morphology among locomotor groups. Cursorial species display distal elongation of the limbs, gracile limb elements, and relatively narrow humeral and femoral epicondyles. Aquatic and semiaquatic species display relatively robust, shortened femora and elongate metatarsals. Semifossorial species display relatively short, robust limbs with enlarged muscular attachment sites and elongate claws. Both semiaquatic and semifossorial species have relatively elongate olecranon process of the ulna and enlarged humeral and femoral epicondyles. Terrestrial, scansorial, and arboreal species are characterized by having primarily intermediate features, but arboreal species do show relatively elongate manual digits. Morphological indices effectively discriminate locomotor groups, with cursorial and arboreal species more accurately classified than terrestrial, scansorial, or semiaquatic species. Both within and between families, species with similar locomotor habits converge toward similar postcranial morphology despite their independent evolutionary histories. The discriminant analysis worked particularly well to correctly classify members of the Canidae, but not as well for members of the Mustelidae or Ursidae. Results are used to infer the locomotor habits of extinct carnivorans, including members of several extinct families, and also 12 species from the Pleistocene of Rancho La Brea.

On Mesopithecus habitat: Insights from late Miocene fossil vertebrate localities of Bulgaria

The aim of this study is to describe the environments where the cercopithecid Mesopithecus was found during latest Miocene in Europe. For this purpose, we investigate the paleoecology of the herbivorous ungulate mesofauna of three very rich late Miocene fossil localities from southwestern Bulgaria: Hadjidimovo, Kalimantsi and Strumyani. While Mesopithecus has been found in the two first localities, no primate remains have yet been identified in Strumyani. Comparison between localities with and without primates using the herbivore mesofauna allows the cross-corroboration of paleoenvironmental conditions where this primate did and did not live. A multi-parameter statistical approach involving 117 equid and 345 bovid fossil dental and postcranial (phalanges, metapodia, astragali) remains from these three localities provides species to generic-level diet and locomotor habit information in order to characterize the environment in which Mesopithecus evolved. The analysis of dental mesowear indicates that the bovids were mainly mixed feeders, while coeval equids were more engaged in grazing. Meanwhile, postcranial remains show that the ungulate species from Hadjidimovo and Kalimantsi evolved in dry environments with a continuum of habitats ranging from slightly wooded areas to relatively open landscapes, whereas the Mesopithecus-free Strumyani locality was in comparison reflecting a rather contrasted mosaic of environments with predominant open and some more closed and wet areas. Environments in which Mesopithecus is known during the late Miocene were not contrasted landscapes combining open grassy areas and dense forested patches, but instead rather restricted to slightly wooded and homogeneous landscapes including a developed grassy herbaceous layer.

Form and function within a phylogenetic framework: locomotory habits of extant predators and some Miocene Sparassodonta (Metatheria)

In this study, we analysed locomotory habits in extant predators and Sparassodonta species through geometric morphometric techniques and discriminant analyses of the distal humerus in anterior view, proximal ulna in lateral view, and tibia in proximal view. We included a wide sample of extant predators, and considered the phylogenetic and allometric structure in the data sets. We also included some Sparassodonta, a group of carnivorous metatherians that inhabited South America during the Cenozoic, and inferred their locomotory habits. Results suggest the presence of a close relationship between shape and locomotory habits, even after removing the shape component explained by phylogeny in the three postcranial elements. Terrestrial habits were inferred for Arctodictis sinclairi, Borhyaena tuberata, ‘Lycopsis’ longirostrus, and Thylacosmilus atrox. Some degree of cursoriality was highlighted in B. tuberata and T. atrox, and climbing abilities in ‘L.’ longirostrus, and to a lesser degree in B. tuberata. Scansorial habits were inferred for Cladosictis patagonica, Sipalocyon gracilis, Prothylacynus patagonicus, and Pseudonotictis pusillus, and in the case of C. patagonica, some digging ability was also tentatively inferred.

Wired to run: exercise-induced endocannabinoid signaling in humans and cursorial mammals with implications for the ‘runner’s high’

Humans report a wide range of neurobiological rewards following moderate and intense aerobic activity, popularly referred to as the 'runner's high', which may function to encourage habitual aerobic exercise. Endocannabinoids (eCBs) are endogenous neurotransmitters that appear to play a major role in generating these rewards by activating cannabinoid receptors in brain reward regions during and after exercise. Other species also regularly engage in endurance exercise (cursorial mammals), and as humans share many morphological traits with these taxa, it is possible that exercise-induced eCB signaling motivates habitual high-intensity locomotor behaviors in cursorial mammals. If true, then neurobiological rewards may explain variation in habitual locomotor activity and performance across mammals. We measured circulating eCBs in humans, dogs (a cursorial mammal) and ferrets (a non-cursorial mammal) before and after treadmill exercise to test the hypothesis that neurobiological rewards are linked to high-intensity exercise in cursorial mammals. We show that humans and dogs share significantly increased exercise-induced eCB signaling following high-intensity endurance running. eCB signaling does not significantly increase following low-intensity walking in these taxa, and eCB signaling does not significantly increase in the non-cursorial ferrets following exercise at any intensity. This study provides the first evidence that inter-specific variation in neurotransmitter signaling may explain differences in locomotor behavior among mammals. Thus, a neurobiological reward for endurance exercise may explain why humans and other cursorial mammals habitually engage in aerobic exercise despite the higher associated energy costs and injury risks, and why non-cursorial mammals avoid such locomotor behaviors.

Hip and Thigh Anatomy of the Clouded Leopard (Neofelis nebulosa) with Comparisons to the Domestic Cat (Felis catus)

The clouded leopard (N. nebulosa) is an endangered felid native to Southeast Asia. It is considered to be the largest and most acrobatic of the arboreal cats. To date, published studies have focused on cranial morphology, particularly osteology and dentition. This study describes the hip and thigh morphology of N. nebulosa and compares it to the domestic cat (F. catus). A number of statistically significant differences were observed between N. nebulosa and F. catus that were primarily associated with muscle attachment surface areas rather than differences in overall morphology. F. catus had proportionally larger attachment areas for Mm. gluteus profundus, tensor fasciae latae, and biceps femoris, while N. nebulosa had larger attachment areas for Mm. gluteus medius, rectus femoris, and semitendinosus. By comparison, surface area analysis of nonfelid species (C. latrans and V. vulpes) showed more differences when compared to felids demonstrating that this analytical technique is useful for differentiating closely-related species from more distant ones. These results suggest that muscle map area analysis may be useful for comparing phylogenetic relationships between and within groups of different species and may also suggest variations in locomotor habits.

Estimación de Masa de las Especies de Sparassodonta (Mammalia, Metatheria) de Edad Santacrucense (Mioceno Temprano) a Partir del Tamaño del Centroide de los Elementos Apendiculares: Inferencias Paleoecológicas

BODY-MASS ESTIMATION OF SPARASSODONTA SPECIES (MAMMALIA, METATHERIA) OF SANTACRUCIAN AGE (EARLY MIOCENE) BASED ON CENTROID SIZES OF POSTCRANEAL ELEMENTS: PALAEOECOLOGICAL INFERENCES. Sparassodonts have been described as a group with a large body-mass diversity, but few works had focused on body mass estimations. In this contribution, we calculated the centroid size of the anterior view of the distal humerus, lateral view of the proximal ulna, and proximal articular view of the tibia. This data together with the known body-mass for many current predators (marsupials and placentals) allowed us to derive new allometric equations to estimate body size in the Sparassodonta. We used comparative analyses to quantify the effect of the phylogenetic structure. The relationships between body mass, centroid size, and locomotory habits were explored. The phylogenetic effect was significant for all regressions. In the humerus analysis, we found an important relationship between body mass and locomotory habit, suggesting adaptive causes. We used the ulna and tibia regressions to predict body mass, since they were less influenced by locomotory habits. Our equations produced the following masses: Arctodictis sinclari Marshall, 39.9 kg; Borhyaena tuberata Ameghino, 36.4 kg; Cladosictis patagonica Ameghino, 6.6 kg; Lycopsis longirostrus Marshall, 29.77 kg; Prothylacynus patagonicus Ameghino, 31.79 kg; Sipalocyon gracilis Ameghino, 2.11 kg; and Thylacosmilus atrox Riggs, 117.36 kg. For Borhyaena and Thylacosmilus, we obtained values higher than previously published masses, while for Pseudonotictis pusillus Ameghino, we have made the first body mass estimation of 1.17 kg. The results suggest that the guild of the santacrucian predators was structured by the body mass and other variables, such as diet and locomotion.

Relationship between humeral geometry and shoulder muscle power among suspensory, knuckle‐walking, and digitigrade/palmigrade quadrupedal primates

Shoulder morphology is functionally related to different patterns of locomotion in primates. To investigate this we performed a quantitative analysis of the relationship between cortical bone thickness (Cbt) of the muscle/tendon attachment site on the humerus and physiological cross-sectional area (PCSA) of the shoulder muscle in primates with different locomotory habits. The deltoid, subscapularis, supraspinatus, and infraspinatus were investigated. A chimpanzee, a gibbon, a baboon, two species of macaque, a lutong, a capuchin, and a squirrel monkey were included in the study. The total length of the humerus was measured and the values were converted into three-dimensional reconstructed data on a computer by computed tomography. The Cbt values were obtained from the volumes divided by the areas of the muscle/tendon attachment sites of the humerus by computer analysis. Muscle mass, muscle fascicle length, and muscle pennation angle were measured and PCSA was calculated using these parameters. A relatively high Cbt and small PCSA were characteristic of the gibbon. The gibbon's high Cbt suggests that passive tension in the muscle/tendon attachment site of suspensory primates (brachiators) may be greater than that of quadrupedal primates, whereas the relatively small PCSA indicates an association with a large amount of internal muscle fascia to endure the passive stress of brachiation. Although chimpanzees undertake some suspensory locomotion, the results for this species resemble those of the digitigrade/palmigrade quadrupedal primates rather than those of the suspensory primate. However, the deltoid and subscapularis in chimpanzee differ from those of the other primates and appear to be affected by the peculiar locomotion of knuckle-walking, i.e. the moment arm of forelimb in chimpanzees is relatively longer than that of digitigrade/palmigrade quadrupedal primates. Hence, a large PCSA in the deltoid and subscapularis may contribute to sustaining the body weight during locomotion. On the other hand, a thin cortical bone relative to a large PCSA was a feature of the cercopithecids, indicating that digitigrade/palmigrade quadrupedal locomotion produces less tension at the muscle/tendon attachment sites compared with that produced by brachiators.

Locomotor loading mechanics in the hindlimbs of tegu lizards (Tupinambis merianae): comparative and evolutionary implications

Skeletal elements are usually able to withstand several times their usual load before they yield, and this ratio is known as the bone's safety factor. Limited studies on amphibians and non-avian reptiles have shown that they have much higher limb bone safety factors than birds and mammals. It has been hypothesized that this difference is related to the difference in posture between upright birds and mammals and sprawling ectotherms; however, limb bone loading data from a wider range of sprawling species are needed in order to determine whether the higher safety factors seen in amphibians and non-avian reptiles are ancestral or derived conditions. Tegus (family Teiidae) are an ideal lineage with which to expand sampling of limb bone loading mechanics for sprawling taxa, particularly for lizards, because they are from a different clade than previously sampled iguanas and exhibit different foraging and locomotor habits (actively foraging carnivore versus burst-activity herbivore). We evaluated the mechanics of locomotor loading for the femur of the Argentine black and white tegu (Tupinambus merianae) using three-dimensional measurements of the ground reaction force and hindlimb kinematics, in vivo bone strains and femoral mechanical properties. Peak bending stresses experienced by the femur were low (tensile: 10.4 ± 1.1 MPa; compressive: -17.4 ± 0.9 MPa) and comparable to those in other reptiles, with moderate shear stresses and strains also present. Analyses of peak femoral stresses and strains led to estimated safety factor ranges of 8.8-18.6 in bending and 7.8-17.5 in torsion, both substantially higher than typical for birds and mammals but similar to other sprawling tetrapods. These results broaden the range of reptilian and amphibian taxa in which high femoral safety factors have been evaluated and further indicate a trend for the independent evolution of lower limb bone safety factors in endothermic taxa.

Neurophysiological analysis of the functional state of neuromotor apparatus in the children with cerebral palsy during treatment with Botulotoxin-A

Background A number of key clinical manifestations of infantile cerebral paralysis significantly affecting the diseases outcome can be specified. First of all, this is the time of formation of static locomotor habits reflecting “maturation” of antigravitation functional system that should be mentioned. Then the spasticity follows, significantly limiting development of voluntary motility, decreasing not only the patients’ locomotor capacity but also their vital activity and resulting in transformation of reflex contractures into the non-reflex ones, due to secondary fibro-trophic changes in muscles and capsular ligamentous complex. Spasticity triggers development of contractures. Pareses and palsies in ICP patients are less pronounced than the muscular tonicity pathology. These are contractures that serve as a leading mechanism limiting further evolution of motor sphere and significantly reducing quality of life in this category of patients (Bronnikov and Kravtsov, 2007). Though not an end in itself, muscular hypertonicity reduction in patients with spastic ICP forms is important since elimination and considerable reduction of its effect as a key factor of development of contractures would allow achieving functional improvement in the patients (Kojevnikova, 2005; Doris Valencia Valencia, 2009). Despite sufficient number of methods for the muscular tonicity reduction in ICP, they all have disadvantages, side and adverse effects and contraindications. Most modalities require prolonged management, the effect being short. Search for novel, more efficient non-invasive methods for the muscular tonicity correction with long-lasting effect, minimum side effects and contraindications is under way. In this respect local injections of botulotoxin-A seem attractive (Carla Rito et al., 2009; Doris Valencia Vlencia, 2009; Isolda Ferreira de Arajio et al., 2009). Our experience in the field, high efficiency and, ultimately, good promises of the method for rehabilitation and subsequent social adaptation of the incapacitated children were among the motives to study the aspect in detail and more profoundly. The work was initiated to study effect of botulotoxin-A injections on parameters of neuromotor apparatus’s functional condition in patients with ICP spastic forms by means of global and stimulation electromyography. Materials and methods

A possible function for an enigmatic synapomorphy of Didelphis

A conspicuous synapomorphy of the marsupial genus Didelphis is the hypertrophied spinous process of cervical vertebrae (C). The tall, thick spinous processes of C2 to C6 interlock preventing any movement of the neck. A functional interpretation of this structure has been elusive. On the basis of osteological specimens of D. albiventris and D. virginiana, we propose a possible function. These specimens show large, partially healed wounds in the spinous processes, indicating that the specimens survived major traumatisms probably caused by carnivorans, so that the spinous processes may have acted as neck armor. Didelphis exhibits several other, well-known defensive traits which, in combination with generalized feeding and locomotor habits, altogether may have helped the lineage survive the Late Tertiary extinction waves in South America and the Great American Biotic Interchange.

Ecological biogeography of Malagasy non‐volant mammals: community structure is correlated with habitat

AbstractAim To examine the relationship between ecoregions, as a proxy for regional climate and habitat type, and mammalian community structure, defined by species composition and richness (e.g. taxonomic structure) and ecological diversity (e.g. ecological structure) of non‐volant species.Location Madagascar.Methods Faunal lists of non‐volant mammal species occurring in 35 communities from five World Wildlife Fund ecoregions were collected from published and unpublished sources. Species were assigned to ecological groups defined by trophic status, locomotor habits, activity cycle and body mass. We used Mantel tests, cluster analysis and principal coordinates analysis to evaluate geographic patterning in taxonomic composition and species richness. We used stepwise multiple discriminant analysis to characterize patterns in the ecological diversity of the mammalian communities from each ecoregion. Communities from transitional habitats (e.g. representing more than one ecoregion) were used to test the predictive power of the analyses.Results Non‐volant mammal communities divided into clusters that correspond to ecoregions. There was a strong distance effect in the taxonomic structure of communities across the island and within both humid and dry forest communities, but this effect was weak within humid forest communities. Mammalian species richness was significantly lower in dry forest than in humid forest communities. The ecological structure of communities was also correlated with ecoregions. Changes in the relative percentages of omnivory, arboreal quadrupedalism, terrestrial/arboreal quadrupedalism and two body mass classes accounted for 98.1% of the variation in ecological structure. Transitional communities were projected in intermediate positions by the discriminant model.Main conclusions Our analysis demonstrates that the broad‐scale habitat and climate variables captured by the ecoregion model have shaped the assembly of non‐volant mammal communities in Madagascar over evolutionary time. The spatial pattern is consistent with ecological sorting of species ranges along environmental gradients. Historical processes, such as recent extinction and migration, may have also affected the structure of mammal communities, although these factors have played a secondary role.

Curvature Scaling in the Medial Tibial Condyle of Large Bodied Hominoids

The shape of joint articular surfaces can provide important information about the locomotor habits of animals. Understanding sources of shape variation in these surfaces is critical to correctly inferring the link between form and function. This study tests the hypothesis that increases in body size within a species result in flatter joint contours of the joint articular surface. Joint flattening is expected to regulate the magnitude and direction of transarticular stresses passing through the joint. Three-dimensional laser scans of polyvinylsiloxane molds of the medial tibial condyle of gorillas (n = 40), common chimpanzees (n = 40), and modern humans (n = 40) were evaluated for differences in articular surface topography, with all measures of joint curvature examined in relation to femoral head superoinferior diameter (as a surrogate for body size). Analyses did not detect an allometric shape change in the curvature of the medial tibial condyle with increasing body size within any of the three taxa examined here. Medial tibial condyle curvature appears to be largely independent of body size as estimated by femoral head diameter. These results do not provide direct support for the hypothesis that joints become flatter in response to increased transarticular loading.

Comparative Scaling of Humeral Cross-Sections of Felids and Canids Using Radiographic Images

The cortical thickness of long bones can be an effective indicator of locomotor modes and other stresses encountered by bone. Felids and canids are two carnivoran families that have similar levels of phylogenetic diversity and overlap in body size, but differ in their locomotor habits. Many canids and felids are cursorial, but felids also climb more frequently than canids. Felids also display a secondary use for their forelimbs not observed in any canids: they use their forelimbs to grasp and subdue prey. Large felids use their forelimbs much more extensively to subdue prey than do large canids and, therefore, should have proportionately greater forces applied to their forelimbs. This study uses a non-invasive radiographic approach to examine the differences in cortical thickness in the humerus between the Felidae and Canidae, as well as between size groups within these two families. Results show few significant differences between the two families, with a slight trend toward more positive allometry in the felids. Overall, radiographic measurements were found to be better predictors of body mass than either prey killing behavior or locomotor mode in these two carnivoran families. One canid that demonstrated exceptionally high cortical area was the bush dog, Speothos venaticus. The rarely observed bush dog has been postulated to swim and dig regularly, and it may be that the thickened cortical bone reflects these behaviors.

Apparent density of the primate calcaneo‐cuboid joint and its association with locomotor mode, foot posture, and the “midtarsal break”

Primates use a range of locomotor modes during which they incorporate various foot postures. Humans are unique compared with other primates in that humans lack a mobile fore- and midfoot. Rigidity in the human foot is often attributed to increased propulsive and stability requirements during bipedalism. Conversely, fore- and midfoot mobility in nonhuman primates facilitates locomotion in arboreal settings. Here, we evaluated apparent density (AD) in the subchondral bone of human, ape, and monkey calcanei exhibiting different types of foot loading. We used computed tomography osteoabsorptiometry and maximum intensity projection (MIP) maps to visualize AD in subchondral bone at the cuboid articular surface of calcanei. MIPs represent 3D volumes (of subchondral bone) condensed into 2D images by extracting AD maxima from columns of voxels comprising the volumes. False-color maps are assigned to MIPs by binning pixels in the 2D images according to brightness values. We compared quantities and distributions of AD pixels in the highest bin to test predictions relating AD patterns to habitual locomotor modes and foot posture categories of humans and several nonhuman primates. Nonhuman primates exhibit dorsally positioned high AD concentrations, where maximum compressive loading between the calcaneus and cuboid likely occurs during "midtarsal break" of support. Humans exhibit less widespread areas of high AD, which could reflect reduced fore- and midfoot mobility. Analysis of the internal morphology of the tarsus, such as subchondral bone AD, potentially offers new insights for evaluating primate foot function during locomotion.

EVOLUTION OF SCAPULA SIZE AND SHAPE IN DIDELPHID MARSUPIALS (DIDELPHIMORPHIA: DIDELPHIDAE)

The New World family Didelphidae, the basal lineage within marsupials, is commonly viewed as morphologically conservative, yet includes aquatic, terrestrial, scansorial, and arboreal species. Here, I quantitatively estimated the existing variability in size and shape of the Didelphidae scapula (1076 specimens from 56 species) using geometric morphometrics, and compared size and shape differences to evolutionary and ecologic distances. I found considerable variation in the scapula morphology, most of it related to size differences between species. This results in morphologic divergence between different locomotor habits in larger species (resulting from increased mechanical loads), but most smaller species present similarly shaped scapulae. The only exceptions are the water opossum and the short-tailed opossums, and the functional explanations for these differences remain unclear. Scapula size and shape were mapped onto a molecular phylogeny for 32 selected taxa and ancestral size and shapes were reconstructed using squared-changed parsimony. Results indicate that the Didelphidae evolved from a medium- to small-sized ancestor with a generalized scapula, slightly more similar to arboreal ones, but strikingly different from big-bodied present arboreal species, suggesting that the ancestral Didelphidae was a small scansorial animal with no particular adaptations for arboreal or terrestrial habits, and these specializations evolved only in larger-bodied clades.

Skeletal indicators of locomotor adaptations in living and extinct rodents

Living rodents show great diversity in their locomotor habits, including semiaquatic, arboreal, fossorial, ricochetal, and gliding species from multiple families. To assess the association between limb morphology and locomotor habits, the appendicular skeletons of 65 rodent genera from 16 families were measured. Ecomorphological analyses of various locomotor types revealed consistent differences in postcranial skeletal morphology that relate to functionally important traits. Behaviorally similar taxa showed convergent morphological characters, despite distinct evolutionary histories. Semiaquatic rodents displayed relatively robust bones, enlarged muscular attachments, short femora, and elongate hind feet. Arboreal rodents had relatively elongate humeri and digits, short olecranon processes of the ulnae, and equally proportioned fore and hind limbs. Fossorial rodents showed relatively robust bones, enlarged muscular attachments, short antebrachii and digits, elongate manual claws, and reduced hind limb elements. Ricochetal rodents displayed relatively proximal insertion of muscles, disproportionate limbs, elongate tibiae, and elongate hind feet. Gliding rodents had relatively elongate and gracile bones, short olecranon processes of the ulnae, and equally proportioned fore and hind limbs. The morphological differences observed here can readily be used to discriminate extant rodents with different locomotor strategies. This suggests that the method could be applied to extinct rodents, regardless of ancestry, to accurately infer their locomotor ecologies. When applied to an extinct group of rodents, we found two distinct ecomorphs represented in the beaver family (Castoridae), semiaquatic and semifossorial. There was also a progressive trend toward increased body size and increased aquatic specialization in the giant beaver lineage (Castoroidinae).

PARALLEL EVOLUTION OF HAND ANATOMY IN KANGAROOS AND VOMBATIFORM MARSUPIALS: FUNCTIONAL AND EVOLUTIONARY IMPLICATIONS

Abstract: The anatomy of the mammalian hand is exposed to an intriguing interplay between phylogeny and function, and provides insights on phylogenetic affinities as well as locomotory habits of extinct species. Within the marsupial order Diprotodontia, terrestrial plantigrade quadrupedalism evolved twice, in the mostly extinct vombatiforms and in extant macropodoids. To assess the influence of functional and phylogenetic signal on the manus in these two clades, manual anatomy and digital proportions in specimens of eight extinct and three extant vombatiforms were investigated and compared with extant macropodoids and extant possums. The results reveal extensive parallelisms in the carpal region of vombatiforms and macropodoids, including flattened distal metacarpal facets, reduction of the palmar process of the hamatum, reduction of mid‐wrist joint curve, extensive hamatum/scaphoid contact, and absence of a lunatum. These transformations appear to be related to stabilization of the wrist for plantigrade locomotion. Vombatiforms are apomorphic in scaphoid and triquetrum anatomy and their metacarpals are much more gracile than in other Diprotodontia. Manual diversity is greater in vombatiforms than in macropodoids, as probably was locomotor diversity. Digital proportions as well as wrist anatomy divide the extinct vombatiforms into species resembling arboreal diprotodontians, whereas others group with terrestrial quadrupedal kangaroos and wombats. The latter is suggested to be owing to plantigrade locomotion and/or large size. Carpal anatomy and digital proportions suggest that a range of earlier diverging vombatiforms may have been arboreal or scansorial. As such, we propose that the ancestor of extant vombatiforms (koalas and wombats) may have been arboreal, an option that deserves consideration in the reconstruction of vombatiform evolution.