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With 14 species recorded, the Miocene La Venta bat fauna is the most diverse bat paleocommunity in South America. It includes the oldest plant-visiting bat in the New World and some of the earliest representatives of the extant families Phyllostomidae, Thyropteridae, and Noctilionidae. La Venta's Notonycteris magdalenensis is an extinct member of the subfamily Phyllostominae, a group of modern Neotropical animalivorous bats, and is commonly included in studies of the evolution of Neotropical bats, but aspects of its biology remain unclear. In this study, we used multivariate dental topography analysis (DTA) to reconstruct the diet of N. magdalenensis by quantitatively comparing measures of molar complexity with those of 25 modern noctilionoid species representing all major dietary habits in bats. We found clear differences in molar complexity between dietary guilds, indicating that DTA is potentially an informative tool to study bat ecomorphology. Our results suggest N. magdalenensis was probably an omnivore or insectivore, rather than a carnivore like its modern relatives Chrotopterus auritus and Vampyrum spectrum. Also, we reconstructed the body mass of N. magdalenensis to be ∼95 g, larger than most insectivorous bats, but smaller than the largest carnivorous bat (V. spectrum). Our results confirm that N. magdalenensis was not a specialized carnivore. It remains to be demonstrated that the specialized carnivory ecological niche was occupied by the same lineage of phyllostomines from at least the middle Miocene. Combining our diet and body-mass reconstructions, we suggest that N. magdalenensis exhibits morphological pre-adaptations crucial for the evolution of specialized carnivory.

Fluctuating asymmetry (random fluctuations between the left and right sides of the body) has been interpreted as an index to quantify both the developmental instabilities and homeostatic capabilities of organisms, linking the phenotypic and genotypic aspects of morphogenesis. However, studying the ontogenesis of fluctuating asymmetry has been limited to mostly model organisms in postnatal stages, missing prenatal trajectories of asymmetry that could better elucidate decoupled developmental pathways controlling symmetric bone elongation and thickening. In this study, we quantified the presence and magnitude of asymmetry during the prenatal development of bats, focusing on the humerus, a highly specialized bone adapted in bats to perform under multiple functional demands. We deconstructed levels of asymmetry by measuring the longitudinal and cross-sectional asymmetry of the humerus using a combination of linear measurements and geometric morphometrics. We tested the presence of different types of asymmetry and calculated the magnitude of size-controlled fluctuating asymmetry to assess developmental instability. Statistical support for the presence of fluctuating asymmetry was found for both longitudinal and cross-sectional asymmetry, explaining on average 16% of asymmetric variation. Significant directional asymmetry accounted for less than 6.6% of asymmetric variation. Both measures of fluctuating asymmetry remained relatively stable throughout ontogeny, but cross-sectional asymmetry was significantly different across developmental stages. Finally, we did not find a correspondence between developmental patterns of longitudinal and cross-sectional asymmetry, indicating that processes promoting symmetrical bone elongation and thickening work independently. We suggest various functional pressures linked to newborn bats’ ecology associated with longitudinal (altricial flight capabilities) and cross-sectional (precocial clinging ability) developmental asymmetry differenti

Bats show a remarkable ecological diversity that is reflected both in dietary and foraging guilds (FGs). Cranial ecomorphological adaptations linked to diet have been widely studied in bats, using a variety of anatomical, computational and mathematical approaches. However, foraging-related ecomorphological adaptations and the concordance between cranial and postcranial morphological adaptations remain unexamined in bats and limited to the interpretation of traditional aerodynamic properties of the wing (e.g. wing loading [WL] and aspect ratio [AR]). For this reason, the postcranial ecomorphological diversity in bats and its drivers remain understudied. Using 3D virtual modelling and geometric morphometrics (GMM), we explored the phylogenetic, ecological and biological drivers of humeral morphology in bats, evaluating the presence and magnitude of modularity and integration. To explore decoupled patterns of variation across the bone, we analysed whole-bone shape, diaphyseal and epiphyseal shape. We also tested whether traditional aerodynamic wing traits correlate with humeral shape. By studying 37 species from 20 families (covering all FGs and 85% of dietary guilds), we found similar patterns of variation in whole-bone and diaphyseal shape and unique variation patterns in epiphyseal shape. Phylogeny, diet and FG significantly correlated with shape variation at all levels, whereas size only had a significant effect on epiphyseal morphology. We found a significant phylogenetic signal in all levels of humeral shape. Epiphyseal shape significantly correlated with wing AR. Statistical support for a diaphyseal-epiphyseal modular partition of the humerus suggests a functional partition of shape variability. Our study is the first to show within-structure modular morphological variation in the appendicular skeleton of any living tetrapod. Our results suggest that diaphyseal shape correlates more with phylogeny, whereas epiphyseal shape correlates with diet and FG.

Most morphological and physiological adaptations associated with bat flight are concentrated in the postcranium, reflecting strong functional demands for flight performance. Despite an association between locomotory diversity and trophic differentiation, postcranial morphological diversity in bats remains largely unexplored. Evolutionary developmental biology is a novel approach providing a link between the analysis of genotypic and phenotypic variation resulting from selective pressures. To quantify the morphological diversity of the postcranium in bats and to explore its developmental basis, we reconstructed the postcranial allometric trajectories of nine bat species from different prenatal developmental series, representing five families and both suborders. We tested for allometric growth in Chiroptera and also quantified levels of allometric disparity and inter-trajectory distances. Using a phylogenetic scaffold, we assessed whether ontogenetic differences reflect evolutionary relationships. We found significant allometric growth trajectories in almost all species. Interspecific trajectory distances showed lower variance within Yinpterochiroptera than within Yangochiroptera and between suborders. Each suborder occupied nonoverlapping sections of allometric space, showing changes in the growth rates of specific bones for each suborder. The allometry-corrected disparity was significantly higher in larger species. Statistically significant phylogenetic signal in our results suggests that there is an ontogenetic basis for the postcranial morphological diversity in modern bats. Ancestral state reconstruction also showed an increase in the amount of change in shape with size in the larger species studied. We hypothesize that differences in allometric patterns among bat taxa may reflect a size-dependent evolutionary constraint, whereby variability in body size and allometric patterns are associated.

Background: Self-powered flight is one of the most energy-intensive types of locomotion found in vertebrates. It is also associated with a range of extreme morpho-physiological adaptations that evolved independently in three different vertebrate groups. Considering that development acts as a bridge between the genotype and phenotype on which selection acts, studying the ossification of the postcranium can potentially illuminate our understanding of bat flight evolution. However, the ontogenetic basis of vertebrate flight remains largely understudied. Advances in quantitative analysis of sequence heterochrony and morphogenetic growth have created novel approaches to study the developmental basis of diversification and the evolvability of skeletal morphogenesis. Assessing the presence of ontogenetic disparity, integration and modularity from an evolutionary approach allows assessing whether flight may have resulted in evolutionary differences in the magnitude and mode of development in bats. Results: We quantitatively compared the prenatal ossification of the postcranium (24 bones) between bats (14 species), non-volant mammals (11 species) and birds (14 species), combining for the first time prenatal sequence heterochrony and developmental growth data. Sequence heterochrony was found across groups, showing that bat postcranial development shares patterns found in other flying vertebrates but also those in non-volant mammals. In bats, modularity was found as an axial-appendicular partition, resembling a mammalian pattern of developmental modularity and suggesting flight did not repattern prenatal postcranial covariance in bats. Conclusions: Combining prenatal data from 14 bat species, this study represents the most comprehensive quantitative analysis of chiropteran ossification to date. Heterochrony between the wing and leg in bats could reflect functional needs of the newborn, rather than ecological aspects of the adult. Bats share similarities with birds in the devel

Sparassodontans are a diverse but now extinct group of metatherians that were apex predators in South America during most of the Cenozoic. Studying their decline has been controversial mainly due to the scarcity of the fossil record, and different methodological approaches have led to contradictory hypotheses. In an effort to explore questions about their extinction, we developed a novel multi-model statistical approach to analyse all of the currently available data at a continental scale. Using multiple regression analysis and new advances in beta diversity analysis, we used all currently available fossil data at a continental scale to test four competing hypotheses to account for the decline of sparassodontans: competition with placental carnivorans, competition with avian phorusrhacids, non-competitive ecological interactions, and environmental fluctuations. Our results show that the sparassodontan extinction was a gradual process with species disappearing throughout the Cenozoic. Multiple regression analysis supported non-competitive ecological interactions as the best extinction model. Native South American ungulates, African migrants (caviomorph rodents and platyrrhine primates) and didelphimorphians were the groups with the highest statistical significance. Sparassodontan beta diversity increased between South American Land Mammal Ages after the Paleocene–Eocene boundary. Our results demonstrate that ecological modelling techniques illuminate aspects of extinction processes whilst mitigating the limitations of the fossil record. Our study suggests that non-competitive ecological interactions could have been the main driver for sparassodontan extinction rather than, as commonly assumed, a result of competition and/or abiotic fluctuations.

The leaf-nosed bats in Hipposideridae and Rhinonycteridae currently have an Old World tropical to subtropical distribution, with a fossil record extending back to the middle Eocene of Europe. The Riversleigh World Heritage fossil site in northwestern Queensland constitutes a particularly rich archive of faunal diversity for Old World leaf-nosed bats, having yielded more than 20 species. We used 2D geometric morphometrics to quantify cranial shape in hipposiderids and rhinonycterids, with the aim of referring unallocated fossil species, particularly from Riversleigh, to each family within a phylogenetic framework, and using a quantitative approach to reconstruct cranial shape for key clades in these Old World radiations. Our sample comprised 21 extant hipposiderids and rhinonycterids, 1 megadermatid and 1 rhinolophid, in which 31 landmarks were placed in lateral and ventral views, and five measurements were taken in dorsal view. The phylogeny used as the framework for this study was based on an analysis of 64 discrete morphological characters from the dentition, cranium and postcranium scored for 42 extant and fossil hipposiderids and rhinonycterids and five outgroup taxa (rhinolophids and megadermatids). The phylogenetic analysis was conducted using maximum parsimony, with relationships among selected extant taxa constrained to match the results of recent comprehensive molecular studies. Our phylogenetic results suggest that the Riversleigh leaf-nosed bats probably do not represent an endemic Australian radiation, with fossil species spread throughout the tree and several with sister-group relationships with non-Australian taxa. Discriminant analyses (DA) conducted separately on each dataset resulted in cross-validated classification success ranging from 61.9% for ventral landmarks to 71.4% for lateral landmarks. Classification of the original grouped cases resulted in success of 81% for each dataset. Of the eight fossil taxa included as unknowns in the DA, six were

Neotropical bats of the genus Carollia are widely studied due to their abundance, distribution and relevance for ecosystems. However, the ecomorphological boundaries of these species are poorly differentiated, and consequently correspondence between their geographic distribution, ecological plasticity and morphological variation remains unclear. In this study, patterns of cranial and mandibular morphological variation were assessed for Carollia brevicauda, C. castanea and C. perspicillata from Colombia. Using geometric morphometrics, morphological variation was examined with respect to: differences in intraspecific variation, morphological modularity and integration, and biogeographic patterns. Patterns of intraspecific variation were different for each species in both cranial and mandibular morphology, with functional differences apparent according to diet. Cranial modularity varied between species whereas mandibular modularity did not. High cranial and mandibular correlation reflects Cranium-Mandible integration as a functional unit. Similarity between the biogeographic patterns in C. brevicauda and C. perspicillata indicates that the Andes do not act as a barrier but rather as an independent region, isolating the morphology of Andean populations of larger-bodied species. The biogeographic pattern for C. castanea was not associated with the physiography of the Andes, suggesting that large body size does not benefit C. brevicauda and C. perspicillata in maintaining homogeneous morphologies among populations.

AbstractDiet has been linked to the diversification of the bat superfamily Noctilionoidea, a group that underwent an impressive ecological adaptive radiation within Mammalia. For decades, studies have explored morphological adaptations and diversity of noctilionoid bats to reveal traits associated with their ecological diversity. Surprisingly, despite such interest and recent application of novel techniques, ecomorphological studies have failed to fully resolve the link between diet and a critical component of the feeding apparatus: dental morphology. Using multivariate dental topographic analysis and phylogenetic comparative methods, we examined the phylogenetic, biological and ecological signal in the dental morphology of noctilionoid bats. Analysing the lower first molars of 110 species, we explored relationships between diet and dental morphology, accounting for three different dimensions of diet (guild, composition and breadth). Phylogenetic and size-dependent structuring of the dental topography data shows it does not correlate only to diet, highlighting the need to account for multiple sources of variation. Frugivorous noctilionoids have sharper molars than other previously reported frugivorous mammals. Nectarivorous noctilionoids showed reduced lower molar crown height and steepness, whereas animalivorous species had larger molars. Dietary composition suggested that the intensity of exploitation of a resource is also linked to different dimensions of dental morphology. Increasing carnivory positively correlated with MA, explaining the highest proportion of its variation, and increasing frugivory explained the highest proportion of variation in all other variables. Dietary breadth showed generalist species have sharper, more topographically-complex molars, whereas specialist herbivores and specialist animalivores fell at opposite ends in the range of tooth steepness and crown height. Together, the results suggest that adaptati