Ontogenetic shape changes in the pelvis of the Greater Rhea (Aves, Palaeognathae) and their relationships with cursorial locomotion: a geometric morphometric approach.

Knowledge of the ontogenetic pattern of morphological features is essential to improve biological interpretations. The study of morphological features of the pelvic girdle and hind limb apparatus throughout growth is an excellent approach to understand how the skeletal morphology and muscles are interrelated during growth in a bird with a specialized mode of locomotion. The Greater Rhea (Rhea americana) is a large cursorial palaeognathous bird with long legs and powerful musculature. The postnatal shape changes of the pelvis of this bird were studied with geometric morphometric techniques, using landmarks and semilandmarks. In addition, regression analyses were used to explore the association between pelvic shape changes with muscle and body mass. The pelvises of 16 specimens of Rhea americana from 1 month old to adulthood were studied in dorsal and lateral views. Noticeable differences in pelvic shape were noted between ages, particularly in lateral view. In young birds, the pre- and post-acetabular ilium was subequal in length, whereas in adults the pre-acetabular ilium became shorter. In dorsal view, the main shape changes observed were the progressive thinning of both ilium portions and the elongation of the vertex craniolateralis ilii from chicks to adulthood. In this view, the only clear differentiation was between young and adult birds. Shape differences were influenced by body mass and pelvic muscles; the post-acetabular muscle mass explained the highest percentage of the variation. The specialized locomotion of Greater Rhea is reflected in their pelvic musculoskeletal system, in which the change to a longer post-acetabular ilium correlates with the growth of the powerful post-acetabular muscles. The actions of these muscles provide the necessary strength to support the body mass, minimize the body swinging movements and propel the body forward during locomotion. Bone morphology is affected by the forces produced by body mass and the muscle activity, demonstrating the presence of common growth mechanisms, which are primordial and gave rise to a functional and properly proportioned adult.

Comparative osteology and functional morphology of the forelimb of Cyonasua (Mammalia, Procyonidae), the first South American carnivoran.

Extant procyonids only inhabit the Americas and are represented by six genera (Procyon, Nasua, Nasuella, Bassaricyon, Potos, and Bassariscus); all of them, except Bassariscus, are present in South America. The first records correspond to the early Miocene in North America (NA) and the late Miocene in South America (SA). Cyonasua was the first carnivoran to enter SA from NA, before the Great American Biotic Interchange, and went extinct in the early middle Pleistocene. This extinct procyonid is recorded in several localities of Argentina, and also in Venezuela. Paleobiological studies of procyonids are interesting from evolutionary and biogeographical viewpoints. In this study, the pectoral girdle and forelimb of 10 specimens of Cyonasua are described and compared with extant South American procyonids using a qualitative approach. Additionally, four functional morphology indexes were calculated for them and compared with an ecologically diverse sample of living carnivorans. Results indicate that Cyonasua most resembles Nasua nasua and Procyon cancrivorus, even though the extinct procyonid possessed peculiar features. Cyonasua had robust and relatively short forelimb bones, with strong stabilized joints, and movements associated with the sagittal plane, which suggest a tendency toward terrestrial habits, related to their ability to resist relatively high bending and shearing stresses. However, some features indicate a freedom in their range of movements, with moderate supination ability, compatible with climbing. When combined with previous analyses of dietary habits and estimated body mass, the morphology of Cyonasua would be well suited for digging and prey manipulation, allowing them to prey on small and relatively large-sized vertebrates, as well as to avoid some of the predators that were dominant in the Cenozoic of South America.

Carpal-metacarpal specializations for burrowing in South American octodontoid rodents.

Among the ecomorphologically diverse Octodontoidea rodents, fossorial habits are prevalent in Ctenomyidae and Octodontidae and occur in some members of Echimyidae. To detect traits linked to scratch-digging, we analyzed morpho-structural variation in the carpus and metacarpus of 27 species of extinct and living octodontoids with epigean, fossorial and subterranean habits. Within a context of relative morphological uniformity, we detected the following specialized traits in the burrowing Clyomys (Echimyidae), Spalacopus (Octodontidae), Ctenomys and †Eucelophorus (Ctenomyidae): broad shortened carpus, robust metacarpals, markedly broad and short metacarpal V, and predominance of ray III (mesaxony, incipient in Spalacopus). In addition, the specialized subterranean Ctenomys presented an enlarged scapholunar in extensive contact with the unciform, and with a complex-shaped proximal articular surface. These features are interpreted as responses to mechanical requirements of scratch-digging, providing greater carpal rigidity and resistance to direct forces exerted during the digging stroke. In Ctenomys, the radius-scapholunar joint restricts movement at wrist level. The phylogenetic distribution of traits shows that the most derived carpal and metacarpal morphologies occur among subterranean octodontoids, also possessing important craniodental adaptations, and supports the hypothesis that the acquisition of digging specializations would have been linked to increasing burrowing frequency in some lineages. Nevertheless, octodontoids with less morphological specializations have metacarpal modifications advantageous for digging, suggesting that scratch-digging specialization preceded the acquisition of tooth-digging traits, in agreement with the general claim that scratch-digging is the primary digging strategy in burrowing mammals.

Adaptive diversity of incisor enamel microstructure in South American burrowing rodents (family Ctenomyidae, Caviomorpha).

The aim of this study was to analyse the morphofunctional and adaptive significance of variation in the upper incisor enamel microstructure of South American burrowing ctenomyids and other octodontoid taxa. We studied the specialized subterranean tooth-digger daggerEucelophorus chapalmalensis (Pliocene - Middle Pleistocene), and compared it with other fossil and living ctenomyids with disparate digging adaptations, two fossorial octodontids and one arboreal echimyid. Morphofunctionally significant enamel traits were quite similar among the species studied despite their marked differences in habits, digging behaviour and substrates occupied, suggesting a possible phylogenetic constraint for the Octodontoidea. In this context of relative similarity, the inclination of Hunter-Schreger bands, relative thickness of external index (EI) and prismless enamel zone were highest in daggerEucelophorus, in agreement with its outstanding craniomandibular tooth-digging specialization. Higher inclination of Hunter-Schreger bands reinforces enamel to withstand high tension forces, while high external index provides greater resistance to wear. Results suggest increased frequency of incisor use for digging in daggerEucelophorus, which could be related to a more extreme tooth-digging strategy and/or occupancy of hard soils. Higher external index values as recurring patterns in distant clades of tooth-digging rodents support an adaptive significance of this enamel trait.