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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'sNotonycteris magdalenensisis 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 ofN. magdalenensisby 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 suggestN. magdalenensiswas probably an omnivore or insectivore, rather than a carnivore like its modern relativesChrotopterus auritusandVampyrum spectrum. Also, we reconstructed the body mass ofN. magdalenensisto be ~95 g, larger than most insectivorous bats, but smaller than the largest carnivorous bat (V. spectrum). Our results confirm thatN. magdalenensiswas 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 thatN. magdalenensisexhibits morphological pre-adaptations crucial for the evolution of specialized carnivory.

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.

The analysis of regional scale patterns of diversity allows insights into the processes that have shaped modern biodiversity at the macro‐scale. Previous analyses studying biogeographic regionalisation across different high‐level taxa have shown similar trends at a global scale. However, incorporating phylogenetic methods when comparing biogeographic regionalisation between subgroups facilitates identification of mechanisms leading to the biogeographic distribution of specific taxa. We analysed the spatial trends of phylogenetic diversity and phylogenetic endemism of 325 species of New World bats, using updated range maps of the modern distributions. These analyses showed phylogeographic signals that reflect the different evolutionary histories of these families. Zoogeographical zones were detected based on range‐weighted phylogenetic turnover. Values of high phylogenetic diversity and endemism were distributed differently across families, suggesting niche conservatism, but a general latitudinal trend of diversity was evident across taxa. Overall, two main bioregions were shared across New World bat taxa (Nearctic and Neotropical), with two additional subregions (Andean and La Platan). We found strong support for an additional transitional zone in the Pacific coast of South America for Emballonuridae and Molossidae. Differences in regionalisation across families indicate that niche conservatism, in situ diversification and dispersal ability are major drivers for the regionalisation of New World bats, within a dual‐centre of diversification scenario. We also found strong inter‐familial support for an independent Caribbean biogeographic region.

Diet 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 adaptations affecting different attributes of dental morphology likely facilitated the dietary diversity and specialisation found in Noctilionoidea.

Bats use their forelimbs in different ways, but flight is the most notable example of morphological adaptation. Foraging and roosting specializations beyond flight have also been described in several bat lineages. Understanding postcranial evolution during the locomotory and foraging diversification of bats is fundamental to understanding bat evolution. We investigated whether different foraging and roosting behaviors influenced humeral cross-sectional shape and biomechanical variation, following Wolff's law of bone remodeling. The effect of body size and phylogenetic relatedness was also tested, in order to evaluate multiple sources of variation. Our results suggest strong ecological signal and no phylogenetic structuring in shape and biomechanical variation in humeral phenotypes. Decoupled modes of scaling of shape and biomechanical variation were consistently indicated across foraging and roosting behaviors, suggesting divergent allometric trajectories. Terrestrial locomoting and upstand roosting species showed unique patterns of shape and biomechanical variation across all our analyses, suggesting that these rare behaviors among bats place unique functional demands on the humerus, canalizing phenotypes. Our results suggest that complex and multiple adaptive pathways interplay in the postcranium, leading to the decoupling of different features and regions of skeletal elements optimized for different functional demands. Moreover, our results shed further light on the phenotypical diversification of the wing in bats and how adaptations besides flight could have shaped the evolution of the bat postcranium.

The middle Miocene La Venta bat fauna is the most diverse bat palaeocommunity in South America, with at least 14 species recorded. They include 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’sNotonycteris magdalenensisis an extinct member of the subfamily Phyllostominae, a group of modern Neotropical animalivorous and omnivorous 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 a multivariate dental topography analysis (DTA) to reconstruct the likely diet ofN. magdalenensisby quantitatively comparing measures of molar complexity with that of 25 modern phyllostomid and noctilionid 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 suggestN. magdalenensiswas probably an omnivore or insectivore, rather than a carnivore like its modern relativesChrotopterus auritusandVampryum spectrum. Also, we reconstructed the body mass ofN. magdalenensisto be ∼50 g, which is larger than most insectivorous bats, but smaller than most carnivorous bats. Our results confirm thatNotonycteris magdalenensiswas probably not a specialised carnivore. It remains to be demonstrated that the specialised 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 thatN. magdalenensisexhibits morphological pre-adaptations crucial for the evolution of specialised carnivory.

Bats are the second-most speciose group of mammals, comprising 20% of species diversity today. Their global explosion, representing one of the greatest adaptive radiations in mammalian history, is largely attributed to their ability of laryngeal echolocation and powered flight, which enabled them to conquer the night sky, a vast and hitherto unoccupied ecological niche. While there is consensus that powered flight evolved only once in the lineage, whether laryngeal echolocation has a single origin in bats or evolved multiple times independently remains disputed. Here, we present developmental evidence in support of laryngeal echolocation having multiple origins in bats. This is consistent with a non-echolocating bat ancestor and independent gain of echolocation in Yinpterochiroptera and Yangochiroptera, as well as the gain of primitive echolocation in the bat ancestor, followed by convergent evolution of laryngeal echolocation in Yinpterochiroptera and Yangochiroptera, with loss of primitive echolocation in pteropodids. Our comparative embryological investigations found that there is no developmental difference in the hearing apparatus between non-laryngeal echolocating bats (pteropodids) and terrestrial non-bat mammals. In contrast, the echolocation system is developed heterotopically and heterochronically in the two phylogenetically distant laryngeal echolocating bats (rhinolophoids and yangochiropterans), providing the first embryological evidence that the echolocation system evolved independently in these bats.

New World bats represent over one third of global bat species and encompass the widest adaptive radiation among bats. Modern bat diversity in the Americas resulted from a mixture of migrations and colonisations of different taxa throughout the Cenozoic. Traditionally, these taxa are conceived as either South or North American, based on the location of their centres of diversification. To better understand the spatial and temporal processes behind modern biogeographic patterns of New World bat fauna, we investigated phylogenetic diversity (PD) and endemism (PE) for 325 American bat species using distribution maps and a species-level phylogenetic supertree of bats. Randomisation tests were used to evaluate the statistical significance of our results, and to derive a categorical analysis of neo- and palaeo-endemism (CANAPE) to deconstruct significant endemism into its different components. PD and PE showed different patterns than those previously reported for New World bats based on traditional measures of diversity. We found multiple centres of significant endemism across the New World for most bat families, extending the hypothesis of dual centres of diversification, previously proposed for Emballonuridae, Phyllostomidae and Mormoopidae, to Molossidae and Vespertilionidae. Our results indicate that Central America and southern North America played important roles in the diversification of New World bats, as did the Andes in the diversification of Vespertilionidae in South America. This article is protected by copyright. All rights reserved.

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 development of structures involved in flight (i.e. handwing and sternum), suggesting that flight altriciality and early ossification of pedal phalanges and sternum are common across flying vertebrates. These results indicate that the developmental modularity found in bats facilitates intramodular phenotypic diversification of the skeleton. Integration and disparity increased across developmental time in bats. We also found a delay in the ossification of highly adaptable and evolvable regions (e.g. handwing and sternum) that are directly associated with flight performance. Electronic supplementary material The online version of this article (10.1186/s12862-019-1396-1) contains supplementary material, which is available to authorized users.

Wilson, L.A.B., Hand, S.J., Lopez-Aguirre, C., Archer, M., Black, K.H., Beck, R.M.D., Armstrong, K.N. & Wroe, S., July 2016. Cranial shape variation and phylogenetic relationships of extinct and extant Old World leaf-nosed bats. Alcheringa 40, 509–524. ISSN 0311-5518The 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...

An increasing amount of research has focused on studying the drivers shaping demographics in science. As a result, we now have a better idea of the current state of gender disparity in science, which remains widespread worldwide. However, fewer studies and limited data have restricted our understanding of this phenomenon in the Neotropics, a highly important region in terms of cultural and biological diversity. Despite a civil war that lasted more than five decades and produced eight million victims (half of them women), Colombia is the fifth country with the highest scientific production in Latin America and the Caribbean, as well as the second most biodiverse country in the world. In order to evaluate the status of gender parity in science in Colombia throughout the 21st century, data of science demographics was gathered covering the 2000-2017 time period. Percentage of women in science was decomposed by research area, researcher rank level and education level. Gender disparity was also estimated for changes in average age, access to scholarships for postgraduate studies, and number of doctoral graduates. Finally, using logistic function modelling, temporal projections into the future were performed, in order to estimate how long could it take to reach gender parity. Of six research fields, medical and health science is the only one to have reached gender parity (55.99%), although it is also the only one showing a steady decrease in women representation across time. On the other hand, engineering, humanities and natural sciences had the lowest percentages of female representation (19.89%, 30.02%, and 30.21%, respectively). Female researchers were on average younger than male researchers, and they also showed a decreasing presence as they move upward to more senior levels, exemplifying the ‘leaky-pipeline phenomenon’ common in science. More men were observed both as scholarship awardees for doctoral studies, and as doctoral graduates, indicating that obtaining a doctoral degree could be a major limiting factor for women in science. Possible drivers of these results are analysed, suggesting that a combination of lack of research funding, insufficient legal framework, pre-existing biases, and poor protection of women’s rights inhibits female participation in science. Based on logistic function modelling it is estimated that, without any action to change current trends, it could take between 10 (humanities) and 175 (engineering) years to reach gender parity across all research areas.

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.

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.

Analyzing asymmetry in species associated with disturbed environments enables the evaluation of the morphological plasticity of generalistic species and the different evolutionary responses of sexes or populations to environmental or genetic stress. This report is a study of the cranial and mandibular asymmetry of Colombian Artibeus lituratus. This species has a wide distribution and high abundance, but its morphological plasticity remains uncertain. We characterized its presence, fluctuating asymmetry, directional asymmetry and antisymmetry by measuring 11 craneometric traits in 146 adults from different localities. Fluctuating asymmetry was present in all traits; directional asymmetry and antisymmetry in three; and no measurement error in any trait. Females showed more fluctuating asymmetry in the splachnocranium and males in the neurocranium. Traits with functional importance while biting had lower levels of asymmetry and higher similarity. Traits with antisymmetry did not show association while traits with directional asymmetry showed mandibular association. We discuss the relation between the presence of cranial and mandibular asymmetry, with the functional similarity of different traits.