Commonalities and evolutionary divergences of mandible shape ontogenies in rodents.

In mammals, significant changes take place during postnatal growth, linked to changes in diet (from sucking to gnawing). During this period, mandible development is highly interconnected with muscle growth and the epigenetic interactions between muscle and bone control the spatialization of bone formation and remodelling in response to biomechanical strain. This mechanism contributes to postnatal developmental plasticity and may have influenced the course of evolutionary divergences between species and clades. We sought to model postnatal changes at a macroevolutionary scale by analysing ontogenetic trajectories of mandible shape across 16 species belonging mainly to two suborders of Rodents, Myomorpha and Hystricomorpha, which differ in muscle attachments, tooth growth and life-history traits. Myomorpha species present a much stronger magnitude of changes over a shorter growth period. Among Hystricomorpha, part of the observed adult shape is set up prenatally, and most postnatal trajectories are genus-specific, which agrees with nonlinear developmental trajectories over longer gestational periods. Beside divergence at large scale, we find some collinearities between evolutionary and developmental trajectories. A common developmental trend was also observed, leading to enlargement of the masseter fossa during postnatal growth. The tooth growth, especially hypselodonty, seems to be a major driver of divergences of postnatal trajectories. These muscle- and tooth-related effects on postnatal trajectories suggest opportunities for developmental plasticity in the evolution of the mandible shape, opportunities that may have differed across Rodent clades.

Middle ear pneumatization in nonhuman primates: A comparative analysis.

OBJECTIVES: We test the effects of body mass and phylogeny on middle ear cavity pneumatization, and the role of pneumatization in hearing function, spanning the anatomical, ecological, and behavioral diversity of nonhuman primates. MATERIALS AND METHODS: All cavities were segmented in middle ear scans of 96 specimens, from 12 strepsirrhine and 15 haplorhine extant species. We measured the tympanic cavity (TC) separately, and all other middle ear spaces together (MES), calculating the degree of pneumatization with the surface area-to-volume ratio. We tested body mass effect with linear regression; we evaluated the phylogenetic signal and selection patterns, using a Kappa statistic test, and Ornstein-Uhlenbeck models (OU). We investigated the link between pneumatization and hearing sensitivity using phylogenetic regression. RESULTS: Testing body mass reveals an allometric pattern for both TC and MES dimensions. Degree of pneumatization in MES is dependent on body mass in haplorhines: larger animals have more pneumatized MES. Differences at various taxonomic ranks were observed for MES, while no phylogenetic influence was observed for TC. Infraorder selection patterns are different. Auditory performance is significantly related to degree of pneumatization, indicating that a pneumatized middle ear is associated with better perception of low frequencies. DISCUSSION: Pneumatization in MES is under differential selective pressure, indicating several optima for this trait. Pneumatization in MES probably modifies hearing sensitivity through pressure regulation mechanisms, auditory bulla size reduction, and frequency modulation. This could explain strepsirrhine adaptation to high-frequency perception, while haplorhine auditory perception is adapted to a broader sound range, including high and low frequencies.

Developmental integration in a functional unit: deciphering processes from adult dental morphology.

The evolution of mammalian dentition is constrained by functional necessity and by the non-independence of morphological structures. Efficient chewing implies coherent tooth coordination from development to motion, involving covariation patterns (integration) within dental parts. Using geometric morphometrics, we investigate the modular organization of the highly derived vole dentition. Integration patterns between and within the upper and lower molar rows are analyzed to identify potential modules and their origins (functional and developmental). Results support an integrated adult dentition pattern for both developmental and functional aspects. The integration patterns between opposing molar pairs suggest a transient role for the second upper and lower molars during the chewing motion. Upper and lower molar rows form coherent units but the relative integration of molar pairs is in contradiction with existing developmental models. Emphasis on the first three cusps to grow leads to a very different integration pattern, which would be congruent with developmental models. The early developmental architecture of traits is masked by later stages of growth, but may still be deciphered from the adult phenotype, if careful attention is paid to relevant features.

Functional constraints channel mandible shape ontogenies in rodents.

In mammals, postnatal growth plays an essential role in the acquisition of the adult shape. During this period, the mandible undergoes many changing functional constraints, leading to spatialization of bone formation and remodelling to accommodate various dietary and behavioural changes. The interactions between the bone, muscles and teeth drive this developmental plasticity, which, in turn, could lead to convergences in the developmental processes constraining the directionality of ontogenies, their evolution and thus the adult shape variation. To test the importance of the interactions between tissues in shaping the ontogenetic trajectories, we compared the mandible shape at five postnatal stages on three rodents: the house mouse, the Mongolian gerbil and the golden hamster, using geometric morphometrics. After an early shape differentiation, by both longer gestation and allometric scaling in gerbils or early divergence of postnatal ontogeny in hamsters in comparison with the mouse, the ontogenetic trajectories appear more similar around weaning. The changes in muscle load associated with new food processing and new behaviours at weaning seem to impose similar physical constraints on the mandible, driving the convergences of the ontogeny at that stage despite an early anatomical differentiation. Nonetheless, mice present a rather different timing compared with gerbils or hamsters.

Variations in the carapace shape of whip spiders (Arachnida: Amblypygi).

Morphological studies often need to reference body size to correctly characterise the shape of organisms. In arthropods, the most commonly used reference for this is the length or width of the carapace, thorax, or the prosoma in the case of chelicerates. However, in the case of animals with unlimited growth, such as whip spiders, this measure could be irrelevant if growth is allometric. In this study, we analyse the ontogenetic modifications in prosoma outline shape in whip spiders during growth and compare the differences in shape between species. Differences are important for the relative prosoma width between species and, in the case of Damon medius, during growth in the juvenile stages, whereas the shape remains stable in mature stages. We conclude that a one-dimensional measure (i.e., length or width) suffices for mature specimens of a single species or family, but for larger studies, or when including immature specimens, at least the prosoma area (within the outline shape) should be used as a size estimator.

Cryptic diversity within three South American whip spider species (Arachnida, Amblypygi).

Cryptic diversity (CD), the presence of highly divergent phylogenetic lineages within closed morphological species, has been documented for many taxa. Great arachnid orders such as Araneae or Scorpiones are well studied and many cases of CD have been described therein; to date, however, related research on smaller arachnid orders, such as whip spiders (Amblypygi), remains lacking. In the current study, we investigated CD based on cytochrome oxidase 1 ( COI) in three nominal species of the genus Heterophrynus ( H. alces, H. batesii, and H. longicornis), represented by 65 specimens. The sequences were compared using three different methods. All three species showed geographically structured CD. Thus, given its existence in this genus, it is important that CD and its spatial distribution be considered in future studies and possible conservation projects.

Morphological modularity and assessment of developmental processes within the vole dental row (Microtus arvalis, Arvicolinae, Rodentia).

Knowledge of mammalian tooth formation is increasing, through numerous genetic and developmental studies. The prevalence of teeth in fossil remains has led to an intensive description of evolutionary patterns within and among lineages based on tooth morphology. The extent to which developmental processes have influenced tooth morphologies and therefore the role of these processes in these evolutionary patterns are nonetheless challenging. Recent methodological advances have been proposed allowing the inference of developmental processes from adult morphologies and the characterization of the degree of developmental integration/modularity of morphological traits by studying the patterns of variation within and among individuals. This study focuses on the geometric shape of the lower molars of the vole species Microtus arvalis. Our results suggest (i) quasi-independence of each molar at the developmental level (developmental modules), even slightly stronger for the third molar supporting some genetic and developmental hypotheses and (ii) more pervasive integration processes among molars at the morphological level.

Investigating the Influence of Climate Changes on Rodent Communities at a Regional-Scale (MIS 1-3, Southwestern France).

Terrestrial ecosystems have continuously evolved throughout the Late Pleistocene and the Holocene, deeply affected by both progressive environmental and climatic modifications, as well as by abrupt and large climatic changes such as the Heinrich or Dansgaard-Oeschger events. Yet, the impacts of these different events on terrestrial mammalian communities are poorly known, as is the role played by potential refugia on geographical species distributions. This study examines community changes in rodents of southwestern France between 50 and 10 ky BP by integrating 94 dated faunal assemblages coming from 37 archaeological sites. This work reveals that faunal distributions were modified in response to abrupt and brief climatic events, such as Heinrich events, without actually modifying the rodent community on a regional scale. However, the succession of events which operated between the Late Pleistocene and the Holocene gradually led to establishing a new rodent community at the regional scale, with intermediate communities occurring between the Bølling and the Allerød.

Tooth wear as a means to quantify intra-specific variations in diet and chewing movements.

In mammals, tooth function, and its efficiency, depends both on the mechanical properties of the food and on chewing dynamics. These aspects have rarely been studied in combination and/or at the intra-specific level. Here we applied 3D dental surface texture analysis to a sample of field voles (Microtus agrestis) trapped from Finnish Lapland at different seasons and localities to test for inter-population variations. We also explored intra-individual variation in chewing dynamics by analysing two facets on the second upper molars. Our results confirm that the two localities have similar environments and that the voles feed on the same items there. On the other hand, the texture data suggest that diets are seasonally variable, probably due to varying concentrations of abrasives. Lastly, the textures on the buccal facets are more isotropic and their direction deviates more from the mesial chewing direction than the lingual facets. We interpret these results as reflecting food, rather than chewing, movements, where food particles are more guided on the lingual side of the molars. This has implications for the application of dental microwear analysis to fossils: only homologous facets can be compared, even when the molar row seems to constitute a functional unit.

Isotopic partitioning by small mammals in the subnivium.

In the Arctic, food limitation is one of the driving factors behind small mammal population fluctuations. Active throughout the year, voles and lemmings (arvicoline rodents) are central prey in arctic food webs. Snow cover, however, makes the estimation of their winter diet challenging. We analyzed the isotopic composition of ever-growing incisors from species of voles and lemmings in northern Finland trapped in the spring and autumn. We found that resources appear to be reasonably partitioned and largely congruent with phylogeny. Our results reveal that winter resource use can be inferred from the tooth isotopic composition of rodents sampled in the spring, when trapping can be conducted, and that resources appear to be partitioned via competition under the snow.

Evolution of mammal tooth patterns: new insights from a developmental prediction model.

The study of mammalian evolution is often based on insights into the evolution of teeth. Developmental studies may attempt to address the mechanisms that guide evolutionary changes. One example is the new developmental model proposed by Kavanagh et al. (2007), which provides a high-level testable model to predict mammalian tooth evolution. It is constructed on an inhibitory cascade model based on a dynamic balance of activators and inhibitors, regulating differences in molar size along the lower dental row. Nevertheless, molar sizes in some mammals differ from this inhibitory cascade model, in particular in voles. The aim of this study is to point out arvicoline and murine differences within this model and to suggest an alternative model. Here we demonstrate that the inhibitory cascade is not followed, due to the arvicoline's greatly elongated first lower molar. We broaden the scope of the macroevolutionary model by projecting a time scale onto the developmental model. We demonstrate that arvicoline evolution is rather characterized by a large gap from the oldest vole to more recent genera, with the rapid acquisition of a large first lower molar contemporaneous to their radiation. Our study provides alternative evolutionary hypotheses for mammals with different trajectories of development.