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). J. Morphol., 2008. © 2008 Wiley‐Liss, Inc.     

Locomotor Behavior and Body Mass of Paramys delicatus (Ischyromyidae,Rodentia) and Commentary on Other Early North American Paramyines

Paramyine ischyromyids are one of the first ancestral rodent groups to appear in North America. Studying ecological indicators of these extinct animals enables us to better understand how they integrated into North American mammalian communities. In this study we reassess the locomotor behavior of a nearly complete skeleton of a paramyine, Paramys delicatus (AMNH FM 12506), using functional limb indices and living squirrels as extant analogues. We then used the results of the functional limb index study to select an appropriate locomotor group for body mass estimations of Paramys delicatus and other early North American (Wasatchian-Bridgerian) paramyines. This was done because body mass is strongly tied to locomotor patterns and more reliable body mass estimates can be generated from an extant sample that functionally resembles the fossils being studied. Functional limb indices were calculated for three locomotor groups (arboreal, semifossorial, and gliding) of living sciurids. Comparisons among arboreal, semifossorial, and gliding sciurids show that the functional indices related to mechanical advantage of muscles and limb robusticity enable distinction among locomotor groups; however, there is considerable overlap between arboreal and semifossorial taxa. Paramys delicatus was found to have generally greater mechanical advantages and limb robusticity than most living squirrels, including semifossorial taxa. As these traits are associated with semifossorial squirrels that frequently use scratch-digging, this suggests that Paramys delicatus and perhaps other early paramyines were likely proficient scratch-diggers. However, indices reflecting limb proportions of paramyines suggest that these early rodents may have used more hind limb dominated locomotion than do living squirrels. Body mass estimations for early paramyines were therefore derived from a semifossorial squirrel sample. Statistical comparisons suggest that many of the most reliable estimators for body mass in Paramys delicatus and other paramyines are those derived from humeral dimensions, with the most reliable estimate being humeral head superoinferior breadth. Using these estimators, individual body mass estimates of early paramyines range from 3391 to 4005 g for Paramys delicatus, 1137–1329 g for Paramys copei, 1291 g for Paramys taurus, and 3357 g for Notoparamys costilloi. All body mass estimations derived from postcranial elements are substantially larger than previously published estimates derived from the dentition, which may be because postcranial elements play a larger role in supporting body weight.

     

Autopodial skeletal diversity in hystricognath rodents: Functional and phylogenetic aspects

Metapodials and phalanges of the second to fourth digital ray were measured for the hands and feet of 214 specimens belonging to 45 extant species of hystricognath rodents, encompassing members of all major clades of the radiation. Principal components analysis (PCA), the phalangeal index of the third digital ray in the hands and feet, and the relationship between second and fourth digital ray were used to investigate intrinsic autopodial proportions as well as to provide a base for comparisons between hands and feet. PCA separated cursorial Hystricognathi from arboreal ones, but lead to little distinction in other locomotory modes. Cursors have longer metapodials and shorter phalanges, particularly in their hind limb, while arboreal species have relatively longer manual and pedal phalanges. Terrestrial, scansorial, fossorial, and semi-aquatic species were not clearly distinguished, but there is a tendency towards elongated manual digits and relatively short feet in most fossorial species. Closely related species with similar locomotory habits tend to group together in PCA morphospace, and also have similar phalangeal indices. The results are in agreement with current hypotheses on locomotory adaptations of the hand and foot, and concur with many previous findings on autopodial proportions in arboreal, cursorial, and fossorial species. They also highlight the limited use of autopodial proportions for inferring systematic affinities. The lack of distinction in the majority of species is likely related to the lack of highly specialized locomotory types in Hystricognathi.     

A comparison of primate, carnivoran and rodent limb bone cross-sectional properties: are primates really unique?

The cross-sectional properties of mammalian limb bones provide an important source of information about their loading history and locomotor adaptations. It has been suggested, for instance, that the cross-sectional strength of primate limb bones differs from that of other mammals as a consequence of living in a complex arboreal environment (Kimura, 1991, 1995). In order to test this hypothesis more rigorously, we have investigated cross-sectional properties in samples of humeri and femora of 71 primate species, 30 carnivorans and 59 rodents. Primates differ from carnivorans and rodents in having limb bones with greater cross-sectional strength than mammals of similar mass. This might imply that primates have stronger bones than carnivorans and rodents. However, primates also have longer proximal limb bones than other mammals. When cross-sectional dimensions are regressed against bone length, primates appear to have more gracile bones than other mammals. These two seemingly contradictory findings can be reconciled by recognizing that most limb bones experience bending as a predominant loading regime. After regressing cross-sectional strength against the product of body mass and bone length, a product which should be proportional to the bending moments applied to the limb, primates are found to overlap considerably with carnivorans and rodents. Consequently, primate humeri and femora are similar to those of nonprimates in their resistance to bending. Comparisons between arboreal and terrestrial species within the orders show that the bones of arboreal carnivorans have greater cross-sectional properties than those of terrestrial carnivorans, thus supporting Kimura's general notion. However, no differences were found between arboreal and terrestrial rodents. Among primates, the only significant difference was in humeral bending rigidity, which is higher in the terrestrial species. In summary, arboreal and terrestrial species do not show consistent differences in long bone reinforcement, and Kimura's conclusions must be modified to take into account the interaction of bone length and cross-sectional geometry.     

Evolution of South American mammalian predators (Borhyaenoidea): anatomical and palaeobiological implications

The evolution of South American carnivorous marsupials, the borhyaenoids, has been investigated through the functional analysis of postcranial adaptive traits and palaeobiological data. There is evidence that the evolutionary history of Borhyaenoidea proceeded from a noncursorial ancestor. The locomotion and habits of the early Palaeocene Mayulestes ferox probably approached a generalized plesiomorphic pattern for marsupial locomotion, i.e. primarily terrestrial with secondary arboreal adaptations. An exceptionally rich early Miocene Patagonian fauna has yielded various morphological predator types, from scansorial ambusher to terrestrial, incipiently cursorial, taxa. The most specialized borhyaenoid was the powerful sabretooth Thylacosmilus atrox that survived until the late Pliocene. The evolution from a scansorial pattern towards a cursorial trend, illustrated by Borhyaena tuberata , and towards the probable development of postlactational parental care in Thylacosmilus , required by its dental specialization and killing strategy, suggests a modification of the selective pressures and predatory activities of the group over evolutionary time.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 140 , 487–521.     

Morphological variation in the appendicular skeleton of atlantic forest sigmodontine rodents

Rodents of the subfamily Sigmodontinae comprise a highly diversified group in the Atlantic Forest, with semifossorial, terrestrial, semiaquatic, scansorial, and arboreal forms. In this study, we analyzed morphometric variation in humerus, scapula, ulna, radius, femur, tibia, and pelvis to investigate its possible relationship with the different types of locomotion recorded in the literature. Skeletal characters were measured in 321 specimens belonging to 29 species and 19 genera either restricted to or recorded in this ecoregion. Multivariate morphometric analyses (principal component and canonical variate analyses) arranged individuals of different genera in groups congruent with the different types of locomotion. This arrangement was more clearly defined when analyses included only forelimb measurements, indicating that most of the variation in appendicular traits associated with the different locomotor modes occurs in the forelimb skeleton. Semifossorial forms exhibited the most distinct appendicular morphology, as well as the greatest frequency of endemism among analyzed species. These results suggest that this mode of locomotion led to greater differentiation in semifossorial Atlantic forest sigmodontines than in terrestrial and arboreal forms, which were found to have more subtle differentiation and fewer endemics. Scansorial species could not be set apart from terrestrial ones in terms of appendicular morphology, suggesting that these two modes of locomotion are the most similar and generalized for the group, as they occur in most lineages in the subfamily. The results of this study corroborate previous observations on the relevance of appendicular characters in the differentiation of species and genera in the subfamily Sigmodontinae. J. Morphol. 2013. © 2013 Wiley Periodicals, Inc.     

Cranial morphology and dietary habits of rodents

Rodents are important components of nearly every terrestrial ecosystem and display considerable ecological diversity. Nevertheless, a lack of data on the ecomorphology of rodents has led to them being largely overlooked in palaeoecological reconstructions. Here, geometric and linear morphometrics are used to examine how cranial and dental shapes reflect the diets of living rodent species. Although most rodents are omnivores or generalist herbivores, some species have evolved highly specialized carnivorous, insectivorous, and herbivorous diets. Results show that living rodents with similar diets display convergent morphology, despite their independent evolutionary histories. Carnivores have relatively elongate incisors, elongate and narrow incisor blades, orthodont incisor angles, reduced cheek tooth areas, and enlarged temporal fossae. Insectivores display relatively degenerate dentition, elongate rostra, narrow and thin zygomatic arches, and smaller temporal fossae. Herbivores are characterized by relatively broader incisor blades, longer molar tooth rows, larger cheek tooth areas, wider skull and rostrum, thicker and broader zygomatic arches, and larger temporal fossae. These results suggest that cranial and dental morphology can be used to accurately infer extinct rodent diets regardless of ancestry. Application to extinct beavers suggests that most had highly specialized herbivorous diets.     

Locomotor correlates of the scapholunar of living and extinct carnivorans

The relationship of carpal morphology to ecology and habitat is under studied in carnivorans and more generally in mammals. Here, we use 3D-scanning techniques to assess the usefulness of a carpal bone, the scapholunar, in carnivorans to reflect ecology and habitat, and to reconstruct the ecology of five extinct carnivorans from two fossil sites: Rancho La Brea and Natural Trap Cave. We 3D-scanned scapholunars and measured articular surface areas and angles between articular facets using GeoMagic and Rhino 3D-software. We analyzed the difference in these metrics using multivariate analysis of variance and discriminant function analysis. Results show that the scapholunar reflects ecological signal, with clear groupings of cursorial carnivorans and grappling/climbing carnivorans; however, phylogenetic signal was also present in the results with hyaenids, canids, and large felids in distinct morphospaces. Extinct species Miracinonyx trumani (American cheetah) and Smilodon fatalis (sabertooth cat) showed surprising results with M. trumani grouping with pantherines instead of Acinonyx or Puma, suggesting it runs but still retains the ability to grapple prey. S. fatalis groups with pantherines, but also shows some unique adaptations, suggesting it had a different range of wrist motion than living cats. Overall, the scapholunar is a good indicator of ecology and functional morphology and can be another tool to use in modern and fossil carnivorans to reconstruct extinct ecologies and locomotor behaviors.     

Anatomical adaptations of shrews

The family Soricidae is reviewed with special regard to anatomical structures related to habitat and life-style. Some 266 species in 20 genera are classified into six feeding and foraging categories: terrestrial, semifossorial, scansorial, semiaquatic, psammophilic and anthropophilic. Examples of corresponding anatomical adaptations are given as well as notes on adaptive radiation and convergent evolution.     

The Calcaneum—On the Heels of Marsupial Locomotion

The potential for making functional interpretations from a single postcranial element for marsupials was investigated through morphometric analysis of the calcanea of 61 extant species from Australia and New Guinea. Extant species were grouped into locomotor categories and a canonical variates analysis was carried out on measurements of their calcanea. A relationship between measurements of the calcanea and the locomotor behavior of species was found, allowing for prediction of locomotor behavior from calcaneum morphometrics. This was applied to fossil marsupial taxa, from early–late Miocene/?Pliocene deposits at Riversleigh, in an attempt to determine their locomotor behavior. Hopping (saltatorial) taxa are distinguished from quadruped terrestrial taxa and taxa capable of climbing (arboreal and scansorial) by their relatively longer tuber calcis and wider calcaneal head, by their dorso-ventrally thicker calcaneal head, and by their calcaneocuboid facet being less steeply angled antero-posteriorly. Taxa capable of climbing are distinguished from quadruped terrestrial taxa by their shorter tuber calcis relative to the calcaneal head and by their smaller calcaneo-astragalar facet. The locomotor categories distinguished in this study (arboreal/scansorial, quadruped terrestrial, and saltatorial) highlight differences between species in their use of available substrates and thus are informative with regards to the structural components of their habitat. The results of this analysis can be used, in combination with other data, to make inferences about the habitats of paleocommunities at Riversleigh through the Miocene. The calcaneum is a dense and very robust element and, therefore, has a good chance of being preserved. This method provides a quick and easy way of inferring locomotion and has a wide potential for application to many fossil deposits because it requires only a single element.     

Distal Humeral Morphology Indicates Locomotory Divergence in Extinct Giant Kangaroos

Previous studies of the morphology of the humerus in kangaroos showed that the shape of the proximal humerus could distinguish between arboreal and terrestrial taxa among living mammals, and that the extinct “giant” kangaroos (members of the extinct subfamily Sthenurinae and the extinct macropodine genus Protemnodon) had divergent humeral anatomies from extant kangaroos. Here, we use 2D geometric morphometrics to capture the shape of the distal humerus in a range of extant and extinct marsupials and obtain similar results: sthenurines have humeral morphologies more similar to arboreal mammals, while large Protemnodon species (P. brehus and P. anak) have humeral morphologies more similar to terrestrial quadrupedal mammals. Our results provide further evidence for prior hypotheses: that sthenurines did not employ a locomotor mode that involved loading the forelimbs (likely employing bipedal striding as an alternative to quadrupedal or pentapedal locomotion at slow gaits), and that large Protemnodon species were more reliant on quadrupedal locomotion than their extant relatives. This greater diversity of locomotor modes among large Pleistocene kangaroos echoes studies that show a greater diversity in other aspects of ecology, such as diet and habitat occupancy.

     

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.     

Functional-adaptive anatomy of the forelimb in the Didelphidae, and the paleobiology of the Paleocene marsupials Mayulestes ferox and Pucadelphys andinus

An attempt to determine the locomotor activities of Mayulestes ferox (Borhyaenoidea) and Pucadelphys andinus (Didelphoidea) from the early Paleocene site of Tiupampa (Bolivia) is presented. The functional anatomy of the forelimbs of these South American marsupials is compared to that of some living didelphids: Caluromys philander, Micoureus demerarae, Marmosa murina, Didelphis marsupialis, Monodelphis brevicaudata and Metachirus nudicaudatus. Deductions from bone morphology to myology and locomotor behavior in the fossils are inferred from the comparisons with living forms. Some features of the postcranial skeleton, indicative of arboreal adaptations, are found in the extinct marsupials: anteriorly projected acromion, hemispherical head of the humerus, extended humeral lateral epicondylar ridge, medially protruding humeral entepicondyle, proximal ulnar posterior convexity, and deep flexor fossa on the medial side of the ulna. But other features are related to a more terrestrial pattern: the well-developed tubercles of the humeral head, the elongated olecranon process of the ulna, and the oval shape of the radial head. Mayulestes had clear arboreal abilities, but, as a predaceous mammal, probably hunted on the ground. Pucadelphys was less specialized, close to the living Monodelphis, a terrestrial insectivorous form with some skeletal features related to arboreal locomotion that are probably plesiomorphic for marsupials.     

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.     

Correlates between calcaneal morphology and locomotion in extant and extinct carnivorous mammals

Locomotor mode is an important component of an animal's ecology, relating to both habitat and substrate choice (e.g., arboreal versus terrestrial) and in the case of carnivores, to mode of predation (e.g., ambush versus pursuit). Here, we examine how the morphology of the calcaneum, the ‘heel bone’ in the tarsus, correlates with locomotion in extant carnivores. Other studies have confirmed the correlation of calcaneal morphology with locomotion behaviour and habitat. The robust nature of the calcaneum means that it is frequently preserved in the fossil record. Here, we employ linear measurements and 2D‐geometric morphometrics on a sample of calcanea from eighty‐seven extant carnivorans and demonstrate a signal of correlation between calcaneal morphology and locomotor mode that overrides phylogeny. We used this correlation to determine the locomotor mode, and hence aspects of the palaeobiology of, 47 extinct carnivorous mammal taxa, including both Carnivora and Creodonta. We found ursids (bears), clustered together, separate from the other carnivorans. Our results support greater locomotor diversity for nimravids (the extinct ‘false sabertooths’, usually considered to be more arboreal), than previously expected. However, there are limitations to interpretation of extinct taxa because their robust morphology is not fully captured in the range of modern carnivoran morphology.     

Postcranial element shape and function: assessing locomotor mode in extant and extinct mustelid carnivorans

Assessment of locomotor modes in fossil taxa must often be made on the basis of heavily fragmented postcranial material. Previous authors have used quantitative methods to determine locomotor function from whole postcranial elements. The goals of this project were to assess the ability of element shape to discern between locomotor modes through landmark analysis, and to apply the results to assessment of fossils. Results suggest that element shape is a good predictor of function, but that different elements have different predictive capacities for each locomotor mode. Additionally, a relationship between size and shape exists that appears to drive morphological differentiation in the group. Finally, data from the extant sample were applied to fossil material of the extinct Plio-Pleistocene taxon Trigonictis . The results suggest that the locomotor mode of Trigonictis was generalized and probably an intermediate between the half-bound locomotion found in weasels and ferrets and the scansorial locomotion of martens and fishers.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 150 , 895–914.     

Locomotory adaptations in entoptychine gophers (Rodentia: Geomyidae) and the mosaic evolution of fossoriality

Pocket gophers (family Geomyidae) are the dominant burrowing rodents in North America today. Their fossil record is also incredibly rich; in particular, entoptychine gophers, a diverse extinct subfamily of the Geomyidae, are known from countless teeth and jaws from Oligocene and Miocene-aged deposits of the western United States and Mexico. Their postcranial remains, however, are much rarer and little studied. Yet, they offer the opportunity to investigate the locomotion of fossil gophers, shed light on the evolution of fossoriality, and enable ecomorphological comparisons with contemporaneous rodents. We present herein a quantitative study of the cranial and postcranial remains of eight different species of entoptychine gophers as well as many contemporary rodent species. We find a range of burrowing capabilities within Entoptychinae, including semifossorial scratch-digging animals and fossorial taxa with cranial adaptations to burrowing. Our results suggest the repeated evolution of chisel-tooth digging across genera. Comparisons between entoptychine gophers and contemporaneous rodent taxa show little ecomorphological overlap and suggest that the succession of burrowing rodent taxa on the landscape may have had more to do with habitat partitioning than competition.     

New wrist bones of the Malagasy giant subfossil lemurs

Recently discovered wrist bones of the Malagasy subfossil lemurs Babakotia radofilai, Palaeopropithecus ingens, Mesopropithecus dolichobrachion, and Megaladapis madagascariensis shed new light on the postcranial morphologies and positional behaviors that characterized these extinct primates. Wrist bones of P. ingens resemble those of certain modern hominoids in having a relatively enlarged ulnar head and dorsally extended articular surface on the hamate, features related to a large range of rotation at the inferior radioulnar and midcarpal joints. The scaphoid of P. ingens is also similar to that of the extant tree sloth Choloepus in having an elongate, palmarly directed tubercle forming a deep radial margin of the carpal tunnel for the passage of large digital flexors. In contrast, wrist remains of Megaladapis edwardsi and M. madagascariensis exhibit traits observed in the hands of extant pronograde, arboreal primates; these include a dorsopalmarly expanded pisiform and well-developed "spiral" facet on the hamate. Moreover, Megaladapis spp. and Mesopropithecus dolichobrachion possess bony tubercles (e.g., scaphoid tubercle and hamate hamulus) forming the carpal tunnel that are relatively similar in length to those of modern pronograde lemurs. Babakotia and Mesopropithecus differ from Megaladapis in exhibiting features of the midcarpal joint related to frequent supination and radioulnar deviation of the hand characteristic of animals that use vertical and quadrumanous climbing in their foraging behaviors. Comparative analysis of subfossil lemur wrist morphology complements and expands upon prior inferences based on other regions of the postcranial skeleton, and suggests a considerable degree of locomotor and postural heterogeneity among these recently extinct primates.     

Locomotor behaviour in Plio-Pleistocene sabre-tooth cats: a biomechanical analysis

The locomotor behaviour of some large extinct carnivores, including several species of Plio-Pleistocene sabre-tooth cats, is here reconstructed, based on a comparison of the cross-sectional geometric properties and linear dimensions of their femora and humeri with those of large modern carnivores. The long bones are modelled as simple beams, thereby allowing the use of basic beam theory in assessing relevant functional parameters such as second moments of area of the diaphyses when subjected to compressive and bending stresses. Three Pleistocene carnivores, Smilodon fatalis, Homotherium serum , and Panthera atrox seem to have had ecological and functional equivalents among the late Miocene-Early Pliocene genera, Barbourofelis, Machairodus , and Nimravides , respectively. Barbourofelis and Smilodon were 'cat-like'in dental morphology but some structural characteristics of their limb bones had 'bear-like'affinities. Machairodus and Homotherium were cursorial, whereas Nimravides and P. atrox display the limb morphology of true ambush predators. Various aspects of the postcranial skeleton of some of these extinct carnivores suggest that they may have employed locomotory gaits that lack modern analogs.     

More injuries in left-footed individual lizards and Sphenodon

In lizards and Sphenodon , often the fourth toes of individuals with intact tails have more subdigital lamellae on the right than on the left side, and the opposite situation frequently occurs in individuals with injured tails. The difference between intact and injured individuals in morphological directional asymmetry was statistically significant ( P <0.05) in 11.4% among 193 species from various lizard families. Lizard families varied in extent and direction of association, but no phylogenetic constraints were detected within genera. Statistical significance was greater in samples from homogenous geographic origin than from heterogeneous ones. Among gekkonid species, the difference was stronger in those with cursorial (terrestrial) habits, than in those with scansorial (rupestral or arboreal) habits. In Scincidae, loss seems more often lethal in left-footed than in right-footed individuals. Statistically significant associations between morphological left-side dominance and tail injury exist also in three independent lineages with reduced limbs (Anguidae, Scincidae and Teidae). Hence such association is probably not a result of limb function. Rather, left-side dominance seems to be the symptom of an unknown, perhaps organism-wide, detrimental trait. Polymorphism in morphological dominance existed in all species, suggesting advantages and disadvantages in different situations to both phenotypes. We propose the hypothesis that an inversion of side dominance may occur in a single trait without systematic inversion of side dominance in all traits of the body. Inversion in a single trait causes incompatibility in multiple-trait functions. Such a mechanism, rather than cultural conventions, could increase accident proneness also in left-handed Homo sapiens , and could explain increased proneness to accident and warfare mortality in left-handed men, beyond the possible involvement of cultural factors.