Masticatory myology of the llama (Lama glama, Camelidae) and comparisons with other camelids and euungulates.

Camelids are the only living representatives of the Suborder Tylopoda, and present a unique set of osteo-myological masticatory features, differing from all other extant euungulates. They combine selenodont dentition and rumination with a fused symphysis, and roughly plesiomorphic muscle proportions. Despite its potential relevance as an euungulate model in comparative anatomy studies, the available data is strikingly scarce. The present study represents the first description of the masticatory muscles of a Lamini, analyzing the functional morphology of Lama glama and other camelids in a comparative framework. Both sides of the head of three adult specimens from Argentinean Puna were dissected. Descriptions, illustrations, muscular maps, and weighing of all masticatory muscles were performed. Some facial muscles are also described. The myology of llamas confirms that camelids possess relatively large temporalis muscles, with Lama being less extreme than Camelus. This plesiomorphic feature is also recorded in suines and some basal euungulates. Conversely, the direction of the fibers of the M. temporalis is mainly horizontal, resembling grinding euungulates such as equids, pecorans, and some derived suines. Although the M. masseter of camelids and equids do not reach the particularly modified configuration of pecorans, in which it is rostrally extended and arranged horizontally, the posterior sectors of Mm. masseter superficialis and pterygoideus medialis have acquired relatively horizontal disposition in the former lineages, suitable for protraction. The pterygoidei complex presents several bundles, and its relative size is intermediate between suines and derived grinding euungulates. The whole masticatory muscles are relatively light when compared to jaw weight. The evolution of the masticatory muscles and chewing of camelids implied that grinding abilities were reached with less extreme modifications of the topography and/or proportions than pecoran ruminants and equids. A relatively large M. temporalis recruited as a powerful retractor during the power stroke is a key feature of camelids. The relaxed pressure on chewing derived from the acquisition of rumination explains the slenderer build masticatory musculature of camelids compared to other euungulates except ruminants.

Functional and phylogenetic interpretation of the forelimb myology of two South American carnivorans, the ring-tailed coati (Nasua nasua) and crab-eating raccoon (Procyon cancrivorus).

A comparative analysis of the forelimb myology of two neotropical procyonids (Nasua nasua and Procyon cancrivorus) was performed to assess how observed differences in their myological configuration would be related to their diverse ecological behaviors and phylogeny. Although both species are associated with the arboreal substrate, N. nasua is a more agile climber that usually digs; whereas P. cancrivorus spends most of its time on the ground foraging, climbing on the trees as shelter and is a good swimmer. Here, myological descriptions, muscle maps, phylogenetic optimizations, and muscle mass data of the forelimb of these two procyonids are presented. The main functional muscular groups are discussed in a comparative framework with other carnivorans that present a wide ecological diversity. Also, muscular characters were mapped onto a phylogeny to explore their evolution and to obtain ancestral state reconstructions. Results indicate clear myological differences among the two neotropical procyonids associated with their ecological preferences. One of the most remarkable anatomical differences is the arrangement and relative mass of the extrinsic musculature, mainly the musculus rhomboideus and the delto-pectoral complexes. In Nasua nasua, these suggested a greater stability in their shoulder girdle for climbing and digging and probably would provide stronger neck and head movements when they use them for foraging on the ground. Conversely, P. cancrivorus has a different extrinsic muscular configuration, which would allow an increment on the stride length and faster movements of the forelimb associated with more frequent terrestrial gaits. Also, significant differences are observed in the distal musculature, associated with strong movements of forepaws when climbing and digging in N. nasua; whereas, P. cancrivorus configuration suggested precise forearm and digits movements, related to manipulation of food items when they are catching prey or feeding. Most of the codified features of P. cancrivorus would reflect retention of plesiomorphies acquired in the common ancestor of caniforms or arctoids, whereas N. nasua shows derived traits, particularly in the proximal forelimb region. The present work increases the information available on the myology of these particular taxa and extant generalized arctoid models in general. The analyses presented here will be useful both for other comparative myological studies (morpho-functional and phylogenetic) and for muscular reconstruction in extinct procyonids, as well as other carnivorans.

Differential locomotor and predatory strategies of Gondwanan and derived Laurasian dromaeosaurids (Dinosauria, Theropoda, Paraves): Inferences from morphometric and comparative anatomical studies.

Tetrapod limbs morphology is a reliable proxy of locomotor capacities. Beyond this, other aspects of life habits, such as predation abilities, can also be relevant to determine main morphofunctional appendicular properties, which ultimately reflect a compromise between different factors of the biological role. Dromaeosauridae is a dinosaur clade belonging to Theropoda, a group of bipedal predators. Dromaeosaurids represent an interesting study case, in which the hindlimbs have been proposed to be involved in both locomotion and predation activity. A peculiar feature characterizing all dromaeosaurids is a modified second pedal digit, which is typically related to predation. This theropod group is closely related to birds and diversified during the Cretaceous Period, mainly in the Northern Hemisphere (Laurasia). However, a subclade of dromaeosaurids, the Unenlagiinae, was recently recognized for Gondwana. Nevertheless, there are morphological differences between derived Laurasian dromaeosaurids (eudromaeosaurs) and unenlagiines. Such differences are observed in the proportions between hindlimb bones and in the presence of a subarctometatarsalian condition in unenlagiines, which is mainly characterized by a proximally constricted metatarsal III. To evaluate the function of these divergent morphologies, we conducted morphometric analyses and comparisons of qualitative morphological aspects, encompassing unenlagiines, other dromaeosaurids, as well as taxa from other theropod groups, including extant birds. The former approach consisted of two phylogenetic principal component analyses, one based on the main measurements of the hindlimb, and the other focused on the lengths of the pedal phalanges. The first analysis drew the unenlagiines close to taxa with long tibiae, as well as long and slender metatarsi. Instead, eudromaeosaurs are closer to taxa with shorter tibiae, and shorter and wider metatarsi. The second analysis showed that eudromaeosaurs and unenlagiines have similar phalangeal proportions, including the elongation of distal phalanges. However, the shorter second phalanx of the pedal digit II of eudromaeosaurs could have increased the force generated by this digit, which was the main predatory tool of the autopodium. This, together with a shorter and wider metatarsus, and a marked hinge-like morphology of the articular surfaces of metatarsals and phalanges, possibly allowed eudromaeosaurs to exert a great gripping strength and hunt large prey. Conversely, the longer and slender subarctometatarsus, and less well-marked hinge joints of unenlagiines possibly gave them greater cursorial capacities. Additionally, the longer second phalanx of digit II allowed unenlagiines fast movements of this digit to hunt smaller and elusive prey. Thus, the distinctive morphological evolutionary pathways of these two dromaeosaurid clades seem to have been influenced by the particular locomotor and predatory specializations that characterized each of these lineages.

Sciuromorphy outside rodents reveals an ecomorphological convergence between squirrels and extinct South American ungulates.

Notoungulates were a diverse group of South American ungulates that included the rodent-like typotherians. However, they are typically compared with other ungulates and interpreted as grazers. Here we present the first detailed reconstruction of the masticatory muscles of the pachyrukhine typotherians Paedotherium and Tremacyllus. An outstanding feature is the presence of a true sciuromorph condition, defined by an anterior portion of the deep masseter muscle originating from a wide zygomatic plate that reaches the rostrum, a trait traceable since the Oligocene pachyrukhines. Consequently, pachyrukhines are the first case of sciuromorph non-rodent mammals. This morphology would have allowed them to explore ecological niches unavailable for the exclusively hystricomorph coexisting rodents. This innovative acquisition seems to be synchronous in Pachyrukhinae and sciuromorph rodents and related to hard-food consumption. We postulate the expansion of nut and cone trees during the major environmental changes at Eocene-Oligocene transition as a potential trigger for this convergence.

Comparative myology of the ankle of Leopardus wiedii and L. geoffroyi (Carnivora: Felidae): functional consistency with osteology, locomotor habits and hunting in captivity.

Leopardus wiedii (margay) is the only arboreal Neotropical felid able to climb head-first down trees, due to its ability to rotate its tarsal joint 180°. A closely related, similar-sized species, L. geoffroyi (Geoffroy's cat) exhibits more typical terrestrial habits and lacks the arboreal capabilities of L. wiedii. There is osteological evidence that supports a mechanical specialization of L. wiedii's tarsal joint for inversion, but there have been no studies on the myology of this specialization. Based on comparative gross-anatomy dissections of zeugo- and autopodial muscles related to the ankle joint of one margay specimen and two Geoffroýs cats, we identified myological specializations of L. wiedii that support its arboreal abilities. In addition, we documented both species hunting the same prey (domestic pigeon Columba livia, Aves: Columbidae) in captivity, to complement. We report differences in the origin, insertion and belly in 8 of the 10 dissected muscles. At least 3 of these interspecific variations can be associated with strengthening of the main muscles that command inversion/eversion movements of the tarsal joint and support the body weight in the head-down climbing position typical of L. wiedii. Frame-by-frame video reconstructions depict the sequence of movements in these species while hunting and highlight the advantages of the arboreal abilities of L. wiedii.

Tracing the origin of the panda's thumb.

We investigate the relative development of the carnivoran radial sesamoids to untangle the evolution of this iconic structure. In the pandas (both giant and red), this 'false thumb' is known to perform a grasping role during bamboo feeding in both the red and giant pandas. An original locomotor role has been inferred for ailurids, but this remains to be ascertained for ursids. A large sample of radial sesamoids of Indarctos arctoides from the Miocene of Batallones-3 (Spain) indicates that this early ailuropodine bear displayed a relatively hypertrophied radial sesamoid, with a configuration more similar to that of the red panda and other carnivorans than to that of giant pandas. This false thumb is the first evidence of this feature in the Ursidae, which can be linked to a more herbivorous diet. Moreover, in the two extant pandas, the false thumb should not be interpreted as an anatomical convergence, but as an exaptive convergence regarding its use during the bamboo feeding, which changes the evolutionary view of this singular structure.

The more, the better: the use of multiple landmark configurations to solve the phylogenetic relationships in musteloids.

Although the use of landmark data to study shape changes along a phylogenetic tree has become a common practice in evolutionary studies, the role of this sort of data for the inference of phylogenetic relationships remains under debate. Theoretical issues aside, the very existence of historical information in landmark data has been challenged, since phylogenetic analyses have often shown little congruence with alternative sources of evidence. However, most analyses conducted in the past were based upon a single landmark configuration, leaving it unsettled whether the incorporation of multiple configurations may improve the rather poor performance of this data source in most previous phylogenetic analyses. In the present study, we present a phylogenetic analysis of landmark data that combines information derived from several skeletal structures to derive a phylogenetic tree for musteloids. The analysis includes nine configurations representing different skeletal structures for 24 species. The resulting tree presents several notable concordances with phylogenetic hypotheses derived from molecular data. In particular, Mephitidae, Procyonidae, and Lutrinae plus the genera Martes, Mustela, Galictis, and Procyon were retrieved as monophyletic. In addition, other groupings were in agreement with molecular phylogenies or presented only minor discordances. Complementary analyses have also indicated that the results improve substantially when an increasing number of landmark configurations are included in the analysis. The results presented here thus highlight the importance of combining information from multiple structures to derive phylogenetic hypotheses from landmark data.

Locomotion in some small to medium-sized mammals: a geometric morphometric analysis of the penultimate lumbar vertebra, pelvis and hindlimbs.

We assessed the influence of a variety of aspects of locomotion and ecology including gait and locomotor types, maximal running speed, home range, and body size on postcranial shape variation in small to medium-sized mammals, employing geometric morphometric analysis and phylogenetic comparative methods. The four views analyzed, i.e., dorsal view of the penultimate lumbar vertebra, lateral view of the pelvis, posterior view of the proximal femur and proximal view of the tibia, showed clear phylogenetic signal and interesting patterns of association with movement. Variation in home range size was related to some tibia shape changes, while speed was associated with lumbar vertebra, pelvis and tibia shape changes. Femur shape was not related to any locomotor variables. In both locomotor type and high-speed gait analyses, locomotor groups were distinguished in both pelvis and tibia shape analyses. These results suggest that adaptations to both typical and high-speed gaits could explain a considerable portion of the shape of those elements. In addition, lumbar vertebra and tibia showed non-significant relationships with body mass, which suggests that they might be used in morpho-functional analyses and locomotor inferences on fossil taxa, with little or no bias for body size. Lastly, we observed morpho-functional convergences among several mammalian taxa and detected some taxa that achieve similar locomotor features following different morphological paths.

Mandible shape in marsupial and placental carnivorous mammals a morphological comparative study using geometric morphometrics

We analysed mandible shape of the orders Dasyuromorpha, Didelphimorphia, and Carnivora using twodimensional geometric morphometrics, in order to explore the relationship between shape, size, and phylogeny. We studied 541 specimens, covering most of the genera of the terrestrial Carnivora (115 species) and a wide sample of marsupials (36 species). The observed shape variation had an ecological component. As an example, omnivorous carnivores have thick mandibles and large talonids in the carnassials, while hypercarnivores possess short mandibles and reduced talonids. There is also a discrimination between different taxonomic groups (i.e. marsupials and Carnivora), indicating some kind of constraint. Size explains a large percentage of total variance (large species had shorter and stronger mandibles, with anteriorly displaced carnassials), was significant when phylogeny was taken into account with a comparative method, but not when size and shape were optimized on the phylogeny. Carnivora presents a larger disparity and variation in body size, which could be related to the difference in teeth replacement. The optimization of mandible shape on the phylogenetic tree indicates that functional aspects, such as diet, are a key factor in the evolution of the carnivore mandible, but also that there is a phylogenetic pattern that cannot be explained by differences in diet alone.