Ontogenetic skull variation in a shovel-headed amphisbaenian species.

Leposternon microcephalum is a species belonging to the Amphisbaenia, a group of burrowing reptiles. Amphisbaenia present various morphological and physiological adaptations that allow them to penetrate the ground and live underground, through a system of galleries and permanent chambers that they build themselves. Among the morphological adaptations in this group, those of the skull stand out as it serves as the main excavation tool. Four basic skull shapes are recognized: rounded, keeled, shovel-shaped, and spade-shaped. The skull of L. microcephalum belongs to this last type, which is considered the most specialized. The species inhabits soils that are highly compacted and difficult to penetrate. Among the species of Leposternon present in South America, L. microcephalum has the widest distribution, being found in all Brazilian biomes and neighboring countries such as Bolivia, Argentina, Paraguay, and Uruguay. The analysis of the skull of this species was carried out using three-dimensional geometric morphometrics (3D-GMM), a technique that allows comparative analysis, through robust statistical methods, of shape and its variations, using Cartesian coordinate data from a configuration of homologous landmarks. The technique allows the size and shape components of a structure to be analyzed separately. From an ontogenetic point of view, this methodology had also been used to investigate variations in Cynisca leucura, a member of the Amphisbaenidae with a rounded head. Our hypothesis is that the patterns of morphological differentiation in the skull, mainly in the intermediate and occipital regions, are similar in different Amphisbaenia species. Therefore, the objective of this study was to analyze cranial morphological variations in an ontogenetic series of L. microcephalum using 3D-GMM. Computed Tomographic scans of 13 specimens were analyzed: juveniles (N = 8) and adults (N = 5), based on 20 landmarks that characterize the skull. Principal components and regression analyses between shape (dependent variable) and size (independent variable) showed a clear difference between the cranial morphological pattern of juvenile individuals and that of adults. For instance, young specimens tend to have a dorsoventrally tall neurocranium, with the tip of the snout more anteriorly oriented and its dorsal border subtly curved. Dorsally, the parietal region is thicker and smoothly dome-shaped in juveniles. As in C. leucura, the variation was strongly correlated with the size change from juvenile to adult, indicating a dominant role for ontogenetic allometry in determining skull shape.

Scaling and mechanics of the felid calcaneus: geometric similarity without differential allometric scaling.

Six mechanically significant skeletal variables were measured on the calcanei from 60 Felidae specimens (22 species) to determine whether these variables were scaled to body mass, and to assess whether differential scaling exists. The power equation (y = a · x(b) ) was used to analyse the scaling of the six variables to body mass; we compared traditional regression methods (standardised major axis) to phylogenetically independent contrasts. In agreement with previous studies that compared these methodologies, we found no significant differences between methods in the allometric coefficients (b) obtained. Overall, the scaling pattern of the felid calcaneus conformed to the predictions of the geometric similarity hypothesis, but not entirely to those of the elastic similarity hypothesis. We found that the moment arm of the ankle extensors scaled to body mass with an exponent not significantly different from 0.40. This indicated that the tuber calcanei scaled to body mass faster than calcaneus total length. This explained why the effective mechanical advantage of the ankle extensors increased with body mass, despite the fact that limb posture does not change in felid species. Furthermore, this finding was consistent with the hypothesis of the isometric scaling of ground reaction forces. No evidence for differential scaling was found in any of the variables studied. We propose that this reflected the similar locomotor pattern of all felid species. Thus, our results suggested that the differences in allometric coefficients for 'large' and 'small' mammals were in fact caused by different types of locomotion among the species included in each category.

Asymmetry in the length of human humerus and radius during ontogeny.

Bilateral body structures usually present some kind of asymmetry. In humans, long bones are suitable to study asymmetries because they are subject to different stimulus that can influence their development and shape. This study focuses on asymmetries of humerus and radius length during ontogeny, with a sample of 1,421 people from Barcelona ranging in age from 0 to 22 years. Data were obtained from dual-energy X-ray absorptiometry images. The humerus results showed the existence of significant directional asymmetry to the left in neonates that progressively turns to right through ontogeny, being significant in people from 11 to 16 years. The radius results are not so evident, but it can be deduced directional asymmetry in children and adolescents, with little delay from humerus. Regarding fluctuating asymmetry, higher values have been seen in neonates and they decrease during ontogeny.

Lumbar ontogenetic growth and sexual dimorphism in modern humans.

To detect and differentiate between possible heterochronic processes in the ontogenetic growth pattern of the human lumbar region, in relationship with sexual dimorphism. We measured the growth trajectories of average length and width, length/width ratio, posterior projected surface area, and bone mineral density using dual energy X-ray absorptiometry, in a sample group of 1718 modern humans. These growth patterns were analyzed using the Gompertz model. In adult lumbar region, only surface area and width were significantly higher in men. Regarding the ontogenetic growth pattern leading to the dimorphic states, all values obtained for women were significantly higher than those obtained for men. Maximum initial growth rates occurred for surface area and density in women. Width scaled faster than length in both sexes. The lumbar region followed patterns similar to those of other skeletal elements when compared with a previous classification of growth patterns in the human skeleton; however, in this study, the growth rate was slower. With regard to the effect of dimorphism, sexual differences in growth rate accounted for only a small proportion of the variation in lumbar length, mineral density, and surface area. Nevertheless, these sexual differences played an important role in the increase of the length/width ratio, which was reflected in the ages at which sexual dimorphism developed. The sexual dimorphism found in the lumbar region of human adults is not caused by any heterochronic process. The lower values of bone mineral density in adult women could explain the origin of some pathologies related.

Cross-Sectional Geometry and Scaling in the Baculum of Cuban Hutias (Rodentia: Capromyidae).

Bacula from 61 individual hutia (Rodentia) from five species were studied. The purpose was to investigate cross-sectional geometry as an indicator of mechanical behavior in order to answer questions around the origin and maintenance of the mammalian baculum. From images of the apical and basal cross sections, the following variables were calculated: perimeter, cross-sectional area, maximum second moment of area, and polar moment. An allometric analysis showed that these variables were related to body size. The orientation of the maximum second moment of area was analyzed by means of circular statistics. This orientation was transverse in both the apical and basal cross sections. Values for the second moment of area and polar moment, obtained from the predicted value of the allometric equations, showed that either the bending moment or the twisting moment of the baculum must be relatively low in hutias, compared with those of the radius in the same species. The results of the second moment of area predict that the main bending stress acting on the baculum is transverse. At the same time, shear stress would not be negligible. Anat Rec, 303:1346-1353, 2020. © 2019 American Association for Anatomy.

Parametric Modeling of Human Gradient Walking for Predicting Minimum Energy Expenditure.

A mathematical model to predict the optimum gradient for a minimum energetic cost is proposed, based on previous results that showed a minimum energetic cost when gradient is -10%. The model focuses on the variation in mechanical energy during gradient walking. It is shown that kinetic energy plays a marginal role in low speed gradient walking. Therefore, the model considers only potential energy. A mathematical parameter that depends on step length was introduced, showing that the optimal gradient is a function of that parameter. Consequently, the optimal negative gradient depends on the individual step length. The model explains why recent results do not suggest a single optimal gradient but rather a range around -10%.

Another one bites the dust: bite force and ecology in three caviomorph rodents (Rodentia, Hystricognathi).

Mammals have developed sophisticated strategies adapting to particular locomotor modes, feeding habits, and social interactions. Many rodent species have acquired a fossorial, semi-fossorial, or even subterranean life-style, converging on morphological, anatomical, and ecological features but diverging in the final arrangement. These ecological variations partially depend on the functional morphology of their digging tools. Muscular and mechanical features (e.g., lever arms relationship) of the bite force were analyzed in three caviomorph rodents with similar body size but different habits and ecological demands of the jaws. In vivo forces were measured at incisors' tip using a strain gauge load cell force transducer whereas theoretical maximal performance values, mechanical advantages, and particular contribution of each adductor muscle were estimated from dissections in specimens of Ctenomys australis (subterranean, solitary), Octodon degus (semi-fossorial, social), and Chinchilla laniger (ground-dweller, colonial). Our results showed that C. australis bites stronger than expected given its small size and C. laniger exhibited the opposite outcome, while O. degus is close to the expected value based on mammalian bite force versus body mass regressions; what might be associated to the chisel-tooth digging behavior and social interactions. Our key finding was that no matter how diverse these rodents' skulls were, no difference was found in the mechanical advantage of the main adductor muscles. Therefore, interspecific differences in the bite force might be primarily due to differences in the muscular development and force, as shown for the subterranean, solitary and territorial C. australis versus the more gracile, ground-dweller, and colonial C. laniger.

Biting performance and skull biomechanics of a chisel tooth digging rodent (Ctenomys tuconax; Caviomorpha; Octodontoidea).

Biting performance is a key factor in vertebrate groups possessing particular food habits. In subterranean rodents that use the incisors as a digging tool, apart from requirements related to gnawing abrasive diets, the force exerted at the incisors tips must be sufficient to break down soils that are often exceedingly compact. The subterranean genus Ctenomys diversified in the southern portion of South America closely associated with the relatively open environments that characterize that region. This genus is considered a "claw and chisel tooth digger," that is, during the excavation of their galleries, the animals break down the soil with both the fore-claws and the incisors. We report here measurements of in vivo bite force in one of the largest species of the genus, C. tuconax, which occupies highland grasslands with compacted soils. We document the combined use of claws and incisors observed under field conditions, also providing measurements of soil compaction in the habitat occupied by this species. We report estimates of bite force at the level of the incisors and cheek teeth calculated from the physiological cross-sectional area of jaw muscles. To this aim, anatomical and biomechanical analyses of the mandibular apparatus were performed in preserved specimens. We found that C. tuconax bites with a higher force than expected for a mammal of its size. To assess anatomical correlates of biting performance, the morphology of the skull and jaw, and incisor second moment of area were compared with those of other caviomorph rodents with different lifestyle.

Scaling and adaptations of incisors and cheek teeth in caviomorph rodents (Rodentia, Hystricognathi).

The South American hystricognath rodents are one of the most diverse mammalian clades considering their occupied habitats, locomotor modes and body sizes. This might have been partly evolved by diversification of their masticatory apparatus' structure and its ecological commitment, for example, chisel-tooth digging. In this phylogeny-based comparative study, we test the relationship between ecological behavior and mechanical features of their incisors and molariforms. In 33 species of nine families of caviomorph rodents, we analyze incisor attributes related to structural stress resistance and molar features related with grinding capacity, for example, second moment of inertia and enamel index (EI) (enamel band length/occlusal surface area), respectively. Most of these variables scaled isometrically to body mass, with a strong phylogenetic effect. A principal component analysis discrimination on the EI clustered the species according to their geographic distribution. We presume that selective pressures in Andean-Patagonian regions, on particular feeding habits and chisel-tooth digging behaviors, have modeled the morphological characteristics of the teeth. Subterranean/burrower ctenomyids, coruros, and plains viscachas showed the highest bending/torsion strength and anchorage values for incisors; a simplified enamel pattern in molariforms would be associated with a better grinding of the more abrasive vegetation present in more open and drier biomes.

Sexual selection in a polygynous rodent (Ctenomys talarum): an analysis of fighting capacity.

The South American subterranean rodent genus Ctenomys (Caviomorpha: Octodontoidea), which uses both claws and teeth to dig, shows striking morphological adaptations to its specialized mode of life. Among other traits, the genus has evolved a powerful jaw musculature and procumbent incisors that are used for dento-excavation. Behavioral observations indicate that these traits are also used during male aggressive encounters, which characterize the polygynous mating system of one of the species of the genus, Ctenomys talarum. A question emerges about sexual selection: could it have induced further changes in traits primarily evolved as adaptations for digging? To address this issue, we studied functional and morphological attributes of the jaw and incisors in specimens of C. talarum. Incisor bite forces were measured on wild females and males from a local population (Mar de Cobo; Buenos Aires Province) by means of a strain gauge load cell force transducer. Museum specimens coming from the same population were studied to assess anatomical attributes of both sexes. Since this species exhibits dimorphism in body size, the possible effect of body mass on the studied traits was analyzed. Males and females showed significant differences in biting performance and mandibular width, but when size was taken into account these differences disappeared. However, other dimorphic traits can vary with a certain independence with respect to size, particularly the 2nd moment of area of the incisors and, to a lesser extent, incisor procumbency. The former geometrical parameter, which is proportional to the bending strength, was highly dimorphic. This fact suggests that, during aggressive encounters between males, biting would place large bending loads on the incisors.

Cranial ontogeny and sexual dimorphism in two new world monkeys: Alouatta caraya (Atelidae) and Cebus apella (Cebidae).

Pattern of skull development and sexual dimorphism was studied in Cebus apella and Alouatta caraya using univariate, bivariate, and multivariate statistics. In both species, sexual dimorphism develops because the common growth trajectory in males extends and because of differences in growth rates between sexes. The expectation that the ontogenetic bases of adult dimorphism vary interspecifically is well substantiated by this study. A. caraya exhibits transitional dimorphism in its subadult stage, although the condylobasal length, zygomatic breadth, and rostrum length are strongly dimorphic in the final adult stage, being greater in males. Most cranial measurements in C. apella exhibit significant dimorphism in the adult stage, being strongly influenced by a faster rate of growth in males. Sexual dimorphism is also evidenced through sex differences in growth rates in several cranial measurements. These results also indicate that different ontogenetic mechanisms are acting in C. apella and A. caraya and reveal differences in the way through which neotropical primates attain adult sexual dimorphism.