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.

Skull morphology of bottlenose dolphins from different ocean populations with emphasis on South America.

The bottlenose dolphin, genus Tursiops, is cosmopolitan occurring in tropical and temperate regions, with morphological variation between and within different oceans. Since the genus' taxonomy has been under discussion for a long time, this work aimed at analyzing the cranial variability of T. truncatus from different regions of the world. Geometric Morphometrics analyses were performed in 201 skulls of adult specimens, on dorsal, ventral, and lateral views, from the Eastern North Pacific, Eastern North Atlantic, Eastern South Atlantic, and Western South Atlantic oceans. The results indicate differences between individuals that inhabit the Atlantic and Pacific oceans. Within the Atlantic Ocean, there is an evident longitudinal differentiation of specimens from the eastern and western regions. A latitudinal separation was also observed, considering specimens from the North and South Atlantic Ocean. In the Western South Atlantic statistical differences were found between two morphological groups, identified as T. gephyreus (sensu Lahille, 1908) and T. truncatus, and the cross-validation presented 98% as minimum confidence for correct classification of these two groups. The present study provides strong morphological support to consider these two lineages as separate species.

Variation in the skull morphometry of four taxonomic units of Thrichomys (Rodentia: Echimyidae), from different Neotropical biomes.

The echimyid rodents of the genus Thrichomys vary considerably in their behavior and feeding ecology, reflecting their occurrence in environments as different as the Caatinga, Cerrado, Pantanal, and Chaco biomes. While the genus was originally classified as monospecific, a number of Thrichomys species have been recognized in recent decades, based on morphometric, cytogenetic, and molecular analyses. While Thrichomys is well studied, the variation found in its cranial morphology is poorly understood, given the taxonomic and ecological complexities of the genus. Using a geometric morphometric approach, we characterized the differences found in the cranial morphology of four Thrichomys taxonomic units, including three established species, Thrichomys apereoides, Thrichomys fosteri, and Thrichomys laurentius, and one operational taxonomic unit (OTU), Thrichomys aff. laurentius. No significant differences were found among these units in cranium size, but significant variation was found in skull shape. The Procrustes distances provided a quantification of the differences in the shape of the skull, with the largest distances being found between T. aff. laurentius and T. fosteri in the dorsal view, and between T. aff. laurentius and T. apereoides in the ventral view. A Discriminant Function Analysis (DFA) with cross-validation determined that the pairings with the highest correct classification were T. aff. laurentius vs. T. apereoides and T. aff. laurentius vs. T. fosteri, in both views. The principal variation in skull shape was found in the posterior region and the zygomatic arch, which may be related to differences in diet.

Skull variation in a shovel-headed amphisbaenian genus, inferred from the geometric morphometric analysis of five South American Leposternon species.

Amphisabenia is a group of squamates adapted for a fossorial lifestyle. The skull is the animal's the main digging tool, and can present one of four principal shapes. The shovel-headed shape is considered to be the most specialized for digging. The South American genus Leposternon presents a shovel-headed morphotype, and is widely distributed on this continent. The general shovel-headed skull pattern may vary considerably, even within the same genus, and we hypothesized that this variation may be influenced primarily by body size and geographical factors. This study investigated the variation in skull size and shape among five Leposternon species, and examined the potential relationship between this variation and the size of the specimens and bioclimatic variables, through a geometric morphometric approach. Significant morphological variation was found among the species, and was also related systematically to body size and the geographical distribution of the specimens. As even subtle differences in the skull size or shape may represent significant modification in bite force and digging capacity and digging speed, the cranial variation found among the Leposternon species and specimens may have a direct influence on their diet and locomotor performance. Our results, together with direct observations of some of these species, suggest that shovel-headed amphisbaenians may be able to penetrate different soil types under a range of climatic conditions, especially considering the ample, but often sympatric distribution of the species studied here.

Body and skull morphometric variations between two shovel-headed species of Amphisbaenia (Reptilia: Squamata) with morphofunctional inferences on burrowing.

BACKGROUND: Morphological descriptions comparing Leposternon microcephalum and L. scutigerum have been made previously. However, these taxa lack a formal quantitative morphological characterization, and comparative studies suggest that morphology and burrowing performance are be related. The excavatory movements of L. microcephalum have been described in detail. However, there is a lack of studies comparing locomotor patterns and/or performance among different amphisbaenids sharing the same skull shape. This paper presents the first study of comparative morphometric variations between two closely related amphisbaenid species, L. microcephalum and L. scutigerum, with functional inferences on fossorial locomotion efficiency. METHODS: Inter-specific morphometric variations were verified through statistical analyses of body and cranial measures of L. microcephalum and L. scutigerum specimens. Their burrowing activity was assessed through X-ray videofluoroscopy and then compared. The influence of morphological variation on the speed of digging was tested among Leposternon individuals. RESULTS: Leposternon microcephalum and L. scutigerum are morphometrically distinct species. The first is shorter and robust with a wider head while the other is more elongated and slim with a narrower head. They share the same excavatory movements. The animals analyzed reached relatively high speeds, but individuals with narrower skulls dug faster. A negative correlation between the speed and the width of skull was determined, but not with total length or diameter of the body. DISCUSSION: The morphometric differences between L. microcephalum and L. scutigerum are in accord with morphological variations previously described. Since these species performed the same excavation pattern, we may infer that closely related amphisbaenids with the same skull type would exhibit the same excavatory pattern. The negative correlation between head width and excavation speed is also observed in others fossorial squamates. The robustness of the skull is also related to compression force in L. microcephalum. Individuals with wider heads are stronger. Thus, we suggest trade-offs between excavation speed and compression force during burrowing in this species.

Age and sex-related geometrical variation of ramus mandibulae of Sus scrofa's (Mammalia: Artiodactyla) jaws / Variaciones geométricas del ramus mandibulae en mandíbulas de Sus scrofa (Mammalia: Artiodactyla) según edad y sexo

Morphological variation of the ramus mandibulae of wild Sus scrofa was described and correlated to sex and age differences. Young forms (n=27, 15 males, 12 females) and adults (n=23, 13 males, 10 females) were analyzed through Geometric Morphometrics (2-D) to assess differences between groups. We found significant differences between age- and sex-related shape variations at ramus mandibulae level. Young wild boars present bulky angular process with backward oriented coronoid process, while adults show slender jaw vertical ramus, and orthogonally oriented coronoid process. The ramus mandibulae in female wild boars is more robust and bulky than in males. The morphological changes between young and adult forms are related to alterations in the osteo-muscular jaw complex along wild boars ontogeny. The jaw morphological variations observed in adult wild boars could also be related to dietary differences between sexes.

Se describen las variaciones morfológicas a nivel de la rama mandibular del jabali (Sus scrofa) en función del sexo y la edad. Se analizó material procedente de jabalíes juveniles (n=27, 15 machos y 12 hembras) y adultos (n=23, 13 machos y 10 hembras) a través de la morfometría geométrica (2-D) para valorar diferencias entre grupos. Se encontraron diferencias a nivel de la forma del ángulo mandibular entre edades y sexos, de manera que los individuos juveniles presentaron un ángulo más abultado y un proceso coronoides orientado hacia atrás, mientras que los adultos muestran una rama mandibular más estrecha y vertical, así como, un proceso coronoides orientado ortogonalmente. En las hembras la parte posterior de la mandíbula es más abultada y compacta que en los machos. Los cambios morfológicos entre juveniles y adultos están relacionados con los procesos ontogenéticos que afectan al complejo osteomuscular durante el crecimiento. Por otra parte, las variaciones morfológicas observadas en los adultos podrían tener relación con diferencias en la dieta entre ambos sexos.

Microsatellite-based characterization of the Castanea sativa cultivar heritage of southern Switzerland.

Southern Switzerland has a long tradition of chestnut cultivation as a staple food. Local inhabitants constantly selected varieties according to the ripening period, the type of use, and the adaptability to the territory. As a result, the panorama of chestnut varieties is very complex, as reflected by more than 120 different variety names in an area of 26,000 ha. Since 1994, 47 varieties have been conserved in the chestnut germplasm of southern Switzerland (CSS), including Marroni, Euro-Japanese, and French varieties. A selection of 164 individuals from the CSS was analysed by 8 SSR markers (4 of which were developed in this study). Microsatellite analysis indicated that the CSS was accurately established, as 86% of the individuals grafted were correctly labeled. The identification of 98 genotypes, 10 clonal chestnut groups, 4 synonym groups, and 12 homonym groups reflected the complex ethnogeographical structure of the chestnut distribution. The 17 Marroni individuals considered clustered in 2 differentiated genetic groups instead of only 1 as expected. The fundamental problem of the frequent cases of homonymy and synonymy is discussed, as is the need for criteria for discriminating between polyclonal varieties and distinct homonymous varieties.

Excavatory cycle of Leposternon microcephalum Wagler, 1824 (Reptilia, Amphisbaenia) / Ciclo excavatorio de Leposternon microcephalum Wagler, 1824 (Reptilia, Amphisbaenia)

The excavatory movements of the spade-snouted amphisbaenid Leposternon microcephalum (Reptilia, Squamata) was studied with the aid of videofluorscopy (X-ray) techniques. This allows the observation of skull and column movements along tunneling, and a more detailed motion observation, being so a novel approach for amphisbaenian excavatory rescarches. A single specimen of Leposternon microcephalum was kept in a glass terrarium filled with semoline, and filmed with a scopy (X-ray) machine. Fixed anatomical marks on the head of the specimen were put in drawings from the framed recordings. Selected sequences of the recordings were fragmented in isolated frames for motion observation. The analysis of the recordings revealed a repetitive pattern of excavatory cycles, with retreating and downward bending of the head before its upstroke to compact the substrate tunnel roof. Follows a dropping of the head, which lays over the substrate giving support for the next retreating and downward head bending. This is an essential step that was neglected in earlier cycle descriptions. The initial downward head bending was not previously properly described for spade-snouted amphisbaenians. The excavatory movements of spade-snouted amphisbaenians are usually treated as a two-stepped cycle, but the evidence that this excavatory cycle has three steps is given here.

Los movimientos excavatorios de la Amphisbaenia (culebrilla ciega), con hocico en forma de pala, Leposternon microcephalum (Reptilia: Squamata) se estudiaron con la ayuda de técnicas de videofluoroscopía (rayos X). Esto permite la observación de los movimientos del cráneo y columna, a lo largo de túneles, y una propuesta más detallada, siendo un enfoque novedoso para investigaciones de la culebrilla ciega excavadora. Un solo espécimen de Leposternon microcephalum se mantuvo en un terrario de vidrio lleno de semolina, y fue filmado con una máquina de escopía (rayos X). Se fijaron marcas anatómicas en la cabeza de éste y luego fueron dibujadas desde las grabaciones. Las secuencias seleccionadas de las grabaciones fueron fragmentadas en fotogramas aislados de la observación del movimiento. Los análisis de las grabaciones revelaron un patrón repetitivo de ciclos excavatorios, con la retirada y doblamiento hacia abajo de la cabeza, antes de su movimiento hacia arriba para compactar el sustrato del techo del túnel. Sigue una caída de la cabeza, que se pone sobre el apoyo del sustrato para la próxima retirada y doblamiento hacia abajo de la cabeza. Esto es un paso esencial que es descuidado en las descripciones anteriores del ciclo. La primera flexión baja de la cabeza no fue debidamente descrita anteriormente para Amphisbaenia con hocico en forma de pala. Los movimientos excavatorios de estos animales suelen ser tratados como un período de dos ciclos intensificados, pero la evidencia que este ciclo excavatorio consta de tres pasos, se dan aquí.