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Sperm morphology is considered the best indicator of male fertility. In Neotropical bats, important aspects of sperm morphology have been scantly studied. The aim of the present study was to characterize and compare the sperm morphology and morphometry of Artibeus planirostris and Sturnira erythromos. A total of 11 specimens were analyzed from the Colección de Mamíferos Lillo: five A. planirostris and six S. erythromos. The fixed epididymis were extracted and macerated in Farmer's solution, followed by the routine cytological procedure with different stains. To carry out the description and morphometric analysis, microphotographs were taken under an optical, epifluorescence and scanning electron microscope. A total of 50 sperm from each individual were measured for morphometric analysis. The percentage of normal/abnormal spermatozoa was estimated and the sperm abnormalities were classified. Both species showed morphologically simple spermatozoa with a spatulate head, a short neck, a helical midpiece and a tail that tapers at the final end, similar to other species of Phyllostomidae. The differences observed were: apex of the head was conical in A. planirostris and was oval in S. erythromos; longer head and midpiece in S. erythomos and longer sperm in A. planirostris. Both species showed a high percentage of sperm with normal appearance: 65% for A. planirostris and 72% for S. erythromos. The main sperm abnormalities were: scattered tails and heads, coiled tails, folded midpieces and presence of cytoplasmic droplets. The present work will improve the understanding of their reproductive biology. RESEARCH HIGHLIGHTS: Morphological descriptions and morphometric analyses of the sperm of Artibeus planirostris and Sturnira erythromos were carried out with optical, epifluorescence and scanning electron microscopy., (© 2024 Wiley Periodicals LLC.)

Fluctuating asymmetry, the non-directional deviation from bilateral symmetry resulting from developmental instability, can indicate early-life environmental stress. While fluctuating asymmetry can affect individual survival and reproductive success, its effect on fitness differs between species. Here, we analyzed up to 27 years of mark-recapture data from 894 RFID tagged individuals of three forest-living bat species in southern Germany to investigate the degree of fluctuating asymmetry in forearm length. In Bechstein's bats, Myotis bechsteinii, the species with the highest sample size, we furthermore investigated if fluctuating asymmetry has become more frequent over the study period, a time when juvenile bats have grown larger forearms in response to warmer summers. We also investigated whether fluctuating asymmetry affects individual lifespan and lifetime reproductive success in female Myotis bechsteinii. The degree of fluctuating asymmetry clearly exceeding the measurement error estimated on recaptured individuals was similar in all three species (1.8%). In female Myotis bechsteinii, the frequency of fluctuating asymmetry did not increase over the course of the study and even strong asymmetry had no effect on individual reproductive success and life expectancy. Our data suggest that fluctuating asymmetry is a poor predictor of fitness in the female Myotis bechsteinii studied, and is so far unaffected by the warming environment which is leading to larger individuals in our study population., Competing Interests: Competing interests: The authors declare no competing interests., (© 2024. The Author(s).)

The external ear in eutherian mammals is composed of the annular, auricular (pinna), and scutellar cartilages. The latter extends between the pinnae, across the top of the head, and lies at the intersection of numerous auricular muscles and is thought to be a sesamoid element. In bats, this scutulum consists of two distinct regions, (1) a thin squama that is in contact with the underlying temporalis fascia and (2) a lateral bossed portion that is lightly tethered to the medial surface of the pinna. The planar size, shape, and proportions of the squama vary by taxa, as does the relative size and thickness of the boss. The origins, insertions, and relative functions of the auricular muscles are complicated. Here, 30 muscles were tallied as to their primary attachment to the pinnae, scutula, or a pre-auricular musculo-aponeurotic plate that is derived from the epicranius. In contrast to Yangochiroptera, the origins and insertions of many auricular muscles have shifted from the scutulum to this aponeurotic plate, in both the Rhinolophidae and Hipposideridae. We propose that this functional shift is a derived character related primarily to the rapid translations and rotations of the pinna in high-duty-cycle rhinolophid and hipposiderid bats., (© 2024 The Author(s). Journal of Morphology published by Wiley Periodicals LLC.)

Background: Eonycteris spelaea ( E. spelaea ) is a sizable nectar-feeding bat that falls within the taxonomic classification of order Chiroptera and family Pteropododae. The form and structure of the tongue play a crucial role for bats in the intake and digestion of food in their mouth. Each papilla's morphology, dimensions, spatial arrangement, and physiological role exhibit variations among different animal species, contingent upon their respective lifestyles., Aim: This research attempts to examine the morphology and microstructure of the E. spelaea tongue papillae, collected from Timor Island, East Nusa Tenggara, Indonesia., Methods: This study aimed to achieve a scanning electron microscope and a light microscope in the presence of hematoxylin-eosin staining and employed a sample of 6 sexually indiscriminate adult E. spelaea bats., Results: The tongue of E. spelaea is separated into three distinct parts: the apex, corpus, and radix. The structure's apex contains filiform papillae, which come in many varieties, such as scale-like filiform papilla, enormous trifid papilla, and little crown-like papilla. Additionally, there is a cluster of fungiform papillae on the outside edge of the highest point and transitional papillae connecting the large trifid papillae with the smaller crown-like papillae. The corpus section comprises two papilla types: filiform papilla (leaf-shaped filiform papilla and big crown-like papilla) and fungiform papilla. The radix comprises the elongated conical papilla, rosette-shaped filiform papilla, short conical papilla, transitional papilla, and three circumvallate papillae at the back of the tongue., Conclusion: The tongue papillae of E. spelaea comprise a wide variety of mechanic papillae and also sensory papillae which have specific dietary regimens in their living habitat., Competing Interests: The authors declare that there is no conflict of interest.

Bats rely on their hand-wings to execute agile flight maneuvers, to grasp objects, and cradle young. Embedded in the dorsal and ventral membranes of bat wings are microscopic hairs. Past research findings implicate dorsal wing hairs in airflow sensing for flight control, but the function of ventral wing hairs has not been previously investigated. Here, we test the hypothesis that ventral wing hairs carry mechanosensory signals for flight control, prey capture, and handling. To test this hypothesis, we used synchronized high-speed stereo video and audio recordings to quantify flight and echolocation behaviors of big brown bats (Eptesicus fuscus) engaged in an aerial insect capture task. We analyzed prey-capture strategy and performance, along with flight kinematics, before and after depilation of microscopic hairs from the bat's ventral wing and tail membranes. We found that ventral wing hair depilation significantly impaired the bat's prey-capture performance. Interestingly, ventral wing hair depilation also produced increases in the bat's flight speed, an effect previously attributed exclusively to airflow sensing along the dorsal wing surface. These findings demonstrate that microscopic hairs embedded in the ventral wing and tail membranes of insectivorous bats provide mechanosensory feedback for prey handling and flight control., (© 2023. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

The Hoary Bat Chalinolobus nigrogriseus is the only species of the genus known from the island of New Guinea. A new species of Chalinolobus from Papua New Guinea is described based on DNA sequence and morphological criteria using material previously assigned to C. nigrogriseus. The new species most resembles the eastern Australian subspecies of the Hoary Bat C. n. nigrogriseus in general size and appearance but is easily distinguished by an enlarged, rather than rudimentary lobe at the terminal outer ear margin. The new species might also be confused on external characters with smaller individuals of Australian Gould's Wattle Bat C. gouldii, from which it differs in having bifid first upper incisors and uniform dark dorsal fur. The inclusion of C. nigrogriseus in the bat fauna of New Guinea is now in doubt, pending a re-assessment of the identity of Chalinolobus specimens in world museum collections. Locality records of all Chalinolobus spp. from Papua New Guinea are reviewed. Most localities are below 60 m elevation in coastal savannah and woodlands. The identity of specimens of C. nigrogriseus and C. gouldii from northern Australia should be reviewed to determine whether the new species also occurs in Australia.

Isolated teeth of a lesser short-nosed fruit bat Cynopterus brachyotis (Müller, 1838), a fulvous fruit bat Rousettus leschenaultii (Desmarest, 1820), and a dawn bat Eonycteris spelaea (Dobson, 1871) are described from the Middle Pleistocene Tham Hai cave locality in northern Vietnam (Lang Son Province). These are the first fossil findings of the Old World fruit bats in Vietnam. The Middle Pleistocene association of Pteropodidae from the Tham Hai locality may largely reflect the composition of species that roosted in local caves., (© 2024. Pleiades Publishing, Ltd.)

Biological variation in the mammalian skull is the product of a series of factors including changes in gene expression, developmental timing, and environmental pressures. When considering the diversity of extant mammalian crania, it is important to understand these mechanisms that contribute to cranial growth and in turn, how differences in cranial morphology have been attained. Various researchers, including Dr. Sue Herring, have proposed a variety of mechanisms to explain the process of cranial growth. This work has set the foundation on which modern analysis of craniofacial morphology happens today. This study focused on the analysis of modularity in three mammalian taxa, all of which exhibit facial reduction. Specifically, we examined facial reduction as a morphological phenomenon through the use of two-module and six-module modularity hypotheses. We recorded three-dimensional coordinate data for 55 cranial landmarks that allowed us to analyze differences in cranial shape in these three taxa (primates n = 88, bats n = 64, dogs n = 81). When assessing modularity within the two-module modularity hypothesis specifically, dogs exhibited the lowest levels of modularity, while bats and primates both showed a slightly more modular covariance structure. We further assessed modularity in the same sample using the Goswami six-module model, where again dogs exhibited a low degree of modularity, with bats and primates being more moderate. We then broke the sample into subsets by analyzing each morphotype separately. We hypothesized that the modularity would be more pronounced in the brachycephalic morphotype. Surprisingly, we found that in brachycephalic dogs, normocephalic dogs, brachycephalic primates, and normocephalic primates, there was a moderate degree of modularity. Brachycephalic bats had a low degree of modularity, while normocephalic bats were the most modular group observed in this study. Based on these results, it is evident that facial reduction is a complex and multifaceted phenomenon with unique morphological changes observed in each of the three taxa studied., (© 2024 Wiley Periodicals LLC.)

Males of the bat Eptesicus furinalis show at least one process of testicular regression, in which the testes regress and temporarily interrupt the production of sperm, during its annual reproductive cycle. As the process of spermatogenesis is under hormonal control, mainly of pituitary and androgen hormones, our aim was to analyze the morphological variations and the hormonal control of the testes of E. furinalis during the four phases of its reproductive cycle. Testes of 18 adult males, divided into four sample groups (active, regressing, regressed, and recrudescence phases), were submitted to morphological, morphometric, and immunohistochemical analyzes. The results demonstrate that the processes of testicular regression and recrudescence of E. furinalis are under the control of pituitary, androgen and estrogen hormones. The regulation is exerted mainly through the activation and cross signaling of AR and FSHR in Sertoli cells and of LHR in Leydig cells. The testicular regression appears to be activated by an inhibition/reduction of AR expression in Sertoli cells, which inhibits the proliferation and differentiation of new spermatogonia and causes the deactivation of spermatogenesis. Conversely, the testicular recrudescence occurs by the increasing of the expression of LHR in Leydig cells, and AR and FSHR in Sertoli cells, which reactivates the testicular production of androgens and estrogens, the proliferation of spermatogonia and restarts the spermatogenesis., (© 2023 American Association for Anatomy.)

Bats are the only mammals capable of powered flight and have correspondingly specialized body plans, particularly in their limb morphology. The origin of bat flight is still not fully understood due to an uninformative fossil record but, from the perspective of a functional transition, it is widely hypothesized that bats evolved from gliding ancestors. Here, we test predictions of the gliding-to-flying hypothesis of the origin of bat flight by using phylogenetic comparative methods to model the evolution of forelimb and hindlimb traits on a dataset spanning four extinct bats and 231 extant mammals with diverse locomotor modes. Our results reveal that gliders exhibit adaptive trait optima (1) toward relatively elongate forelimbs that are intermediate between those of bats and non-gliding arborealists, and (2) toward relatively narrower but not longer hindlimbs that are intermediate between those of non-gliders and bats. We propose an adaptive landscape based on limb length and width optimal trends derived from our modeling analyses. Our results support a hypothetical evolutionary pathway wherein glider-like postcranial morphology precedes a bat-like morphology adapted to powered-flight, setting a foundation for future developmental, biomechanical, and evolutionary research to test this idea., Competing Interests: The authors declare that they have no competing interests., (© 2024 Burtner et al.)

Birds, bats and insects have evolved unique wing structures to achieve a wide range of flight capabilities. Insects have relatively stiff and passive wings, birds have a complex and hierarchical feathered structure and bats have an articulated skeletal system integrated with a highly stretchable skin. The compliant skin of the wing distinguishes bats from all other flying animals and contributes to bats' remarkable, highly manoeuvrable flight performance and high energetic efficiency. The structural and functional complexity of the bat wing skin is one of the least understood although important elements of the bat flight anatomy. The wing skin has two unusual features: a discrete array of very soft elastin fibres and a discrete array of skeletal muscle fibres. The latter is intriguing because skeletal muscle is typically attached to bone, so the arrangement of intramembranous muscle in soft skin raises questions about its role in flight. In this paper, we develop a multi-scale chemo-mechanical constitutive model for bat wing skin. The chemo-mechanical model links cross-bridge cycling to a structure-based continuum model that describes the active viscoelastic behaviour of the soft anisotropic skin tissue. Continuum models at the tissue length-scale are valuable as they are easily implemented in commercial finite element codes to solve problems involving complex geometries, loading and boundary conditions. The constitutive model presented in this paper will be used in detailed finite element simulations to improve our understanding of the mechanics of bat flight in the context of wing kinematics and aerodynamic performance.

Bats have the ability to fly without eye application in the darkness. In this study, we aimed to characterize the functional and structural acclimations of the lenses of two common bats with a various lifestyle in the Egyptian environment: the insectivorous bat (IB) (Pipistrellus kuhlii) and Egyptian fruit bat (FB) (Rousettus aegyptiacus). From each species, seven lenses were extracted from adult eyes. The scanning electron microscopic (SEM) and light microscopic examination of the lens were carried out. FB lenses were made up primarily of fiber cells and sheets, which were encapsulated by a thin collagenous capsule and covered by single epithelial layer anteriorly. On the other hand, the IB lens had two poles and was visibly oval shaped. Both lenses had epithelial cells of the same cuboidal form that were subjected to continuous division and differentiation into new fiber cells at the center. SEM revealed that the normal FB lens had regularly organized shells of fiber cells of intact lens fibers which were connected by membrane interdigitations with different shapes mainly ball-and-socket junctions through the superficial cortical fiber cells. The IB lens was composed of parallel, evenly spaced fibers with various types of interdigitations between fibers that can be seen and increased close to the middle region revealing tiny bumps along the scrubby portions and sockets and balls in the center of the wide portions. Near the center of both lenses, there were large interlocking paddles with little and lengthy protrusions along their short sides. In conclusion, our study discovered several ultrastructural and structural variations among the investigated species. The detection of specialized membrane interdigitations with different shapes protruding from the lens fiber sheets is considered the most characteristic of the FB lens. RESEARCH HIGHLIGHTS: FB lens has more organized sheets of fibers parallel to each other than IB lens. Different shapes of interdigitations protruded from the FB lens have been detected. Interlocking paddles, balls, and sockets with tongue-like fiber flabs are characteristic to FB lens., (© 2024 Wiley Periodicals LLC.)

As bones age in most mammals, they typically become more fragile. This state of bone fragility is often associated with more homogenous collagen fiber orientations (CFO). Unlike most mammals, bats maintain mechanically competent bone throughout their lifespans, but little is known of positional and age-related changes in CFO within wing bones. This study tests the hypothesis that age-related changes in CFO in big brown bats (Eptesicus fuscus) differ from those of the standard mammalian model for skeletal aging, the C57BL/6 laboratory mouse. We used data from quantitative polarized light microscopy (qPLM) to compare CFO across the lifespan of long-lived big brown bats and age matched C57BL/6 mice. Eptesicus and C57BL/6 mice displayed idiosyncratic patterns of CFO. Consistent age-related changes were only apparent in the outer cortical bone of Eptesicus, where bone tissue is more longitudinally arranged and more anisotropic in older individuals. Both taxa displayed a ring of more transversely oriented bone tissue surrounding the medullary cavity. In Eptesicus, this tissue represents a greater proportion of the overall cross-section, and is more clearly helically aligned (arranged at 45° to the bone long axis) than similar bone tissue in mice. Bat wing bones displayed a proximodistal gradient in CFO anisotropy and longitudinal orientation in both outer and inner cortical bone compartments. This study lays a methodological foundation for the quantitative evaluation of bone tissue architecture in volant and non-volant mammals that may be expanded in the future., (© 2023 The Authors. The Anatomical Record published by Wiley Periodicals LLC on behalf of American Association for Anatomy.)

Whether the endocrine pancreas exhibits structural features to couple with dietary patterns is not fully explored. Considering the lack of data comparing endocrine pancreas and islet cell distribution among different bat species in the same study, we considered this an opportunity to explore the topic, including five species within three different predominant diets. For this, we applied morphometric techniques to compare the islets of frugivorous Artibeus lituratus and Carollia perspicillata, insectivorous Molossus molossus and Myotis nigricans, and nectarivorous Glossophaga soricina bats. Data for islet size, cellularity, and mass were equivalent between frugivorous A. lituratus and nectarivorous G. soricina, which differed from insectivorous bats. The frugivorous C. perspicillata bat exhibited morphometric islet values between A. lituratus and the insectivorous species. A. lituratus and G. soricina but not C. perspicillata bats had higher islet mass than insectivorous species due to larger size, instead of a higher number of islets per area. Insectivorous bats, on the other hand, had a higher proportion of α-cells per islet. These differences in the endocrine pancreas across species with different eating habits indicate the occurrence of species-specific adjustments along the years of evolution, with the demand for α-cells higher in bats with higher protein intake., Competing Interests: Declaration of Competing Interest The authors declare that there are no conflicts of interest., (Copyright © 2024 Elsevier Ltd. All rights reserved.)

The extinct mountain long-eared bat Plecotus macrobullaris sarmaticus subsp. nov. is described on the base of jaw remains from the Lower Pleistocene of the Taurida cave in central Crimea. This is the first record of the genus Plecotus in the Lower Pleistocene of Russia and the first fossil find of P. macrobullaris Kuzyakin, 1965, documenting the early stage of the evolutionary history of the species., (© 2024. Pleiades Publishing, Ltd.)

Body size is a key morphological attribute, often used to delimit species boundaries among closely related taxa. But body size can evolve in parallel, reaching similar final states despite independent evolutionary and geographic origins, leading to faulty assumptions of evolutionary history. Here, we document parallel evolution in body size in the widely distributed leaf-nosed bat genus Hipposideros, which has misled both taxonomic and evolutionary inference. We sequenced reduced representation genomic loci and measured external morphological characters from three closely related species from the Solomon Islands archipelago, delimited by body size. Species tree reconstruction confirms the paraphyly of two morphologically designated species. The nonsister relationship between large-bodied H. dinops lineages found on different islands indicates that large-bodied ecomorphs have evolved independently at least twice in the history of this radiation. A lack of evidence for gene flow between sympatric, closely related taxa suggests the rapid evolution of strong reproductive isolating barriers between morphologically distinct populations. Our results position Solomon Islands Hipposideros as a novel vertebrate system for studying the repeatability of parallel evolution under natural conditions. We conclude by offering testable hypotheses for how geography and ecology could be mediating the repeated evolution of large-bodied Hipposideros lineages in the Solomon Islands., (© The Author(s) 2024. Published by Oxford University Press on behalf of The Society for the Study of Evolution (SSE).)

Molossus molossus is an insectivorous molossid bat that is important in the control of nocturnal insects. It is the nominal and the most representative species of the family. However, there are few studies about its reproduction. Thus, this study aimed to evaluate variations of its female reproductive organs during the different reproductive phases. Twenty adult females, divided into four sample groups (non-reproductive, early and advanced pregnancy and lactation), were submitted to morphological and morphometric analyses. Results show that the female reproductive system of M. molossus is composed of ovaries, a short bicornuate uterus, slightly convoluted uterine tubes and vagina. The system presents a distinct morphofunctional asymmetry, with a marked dextro-dominance. The right ovaries of all analyzed groups (NON, P1, P2, and LAC) showed follicles at different stages of development, a large number of interstitial glands and a small, but persistent corpus luteum. Ovulation is simple, unilateral and preferential, occurring exclusively in the right ovary. Follicular development in the left ovary usually does not pass the secondary stage. Implantation is fundic and preferential, occurring exclusively in the right uterine horn. The placenta is formed with two distinct chorioallantoic portions, one diffuse endotheliochorial, which covers the entire uterine cavity and regresses in the final stages of pregnancy, and the principal discoidal hemochorial portion, formed in the implantation site. The uterine cervix presents a pseudostratified epithelium, while the vagina has a little keratinized stratified epithelium, which does not accentually vary in the different reproductive stages, but can disrupt and shed in some cases., Competing Interests: Declaration of Competing Interest The authors declare that they have no conflict of interest., (Copyright © 2023 Elsevier Ltd. All rights reserved.)

Sperm morphology is considered a species-specific character and has been used as a tool in the classification of numerous mammalian taxa. Neotropical bats have been poorly studied, and important aspects on sperm morphology have not been elucidated. The aim of the present study was to describe and compare the sperm morphology and morphometry of Molossus molossus and Molossops temminckii. A total of 14 adults specimens were analyzed from the Colección Mamíferos Lillo, Universidad Nacional de Tucumán: five M. molossus and nine M. temminckii. The epididymis were extracted and macerated in Farmer's solution, followed by a coloration with different stains. To carry out the description and morphometric analysis, microphotographs were taken under an optical, epifluorescence, and scanning electron microscope (SEM). A total of 50 sperm from each individual were measured for morphometric analysis. The length and width of the head, midpiece and tail were taken as variables. Sperm from M. molossus and M. temminckii were practically identical, both morphologically and morphometrically. In both species, a distal bulge was observed at the end of the intermediate piece in a percentage greater than 85%. The main characteristics shared between the species were: presence of acrosomal blebs in the upper half of the head of the spermatozoa; cephalic equatorial segment with filiform ornamentations; intermembrane space of head apex wedge-shaped; helical middle piece and annulus at the end of middle piece. In the present study, SEM allowed us to visualize structures, such as acrosomal vesicles, that were not detected with other types of microscopy. RESEARCH HIGHLIGHTS: The similarities in the sperm morphology between M. molossus and M. temminckii were observed with three types of microscopy: optical, epifluorescence and scanning electron, and supported by morphometric and statistical analyses., (© 2023 Wiley Periodicals LLC.)

Echolocation is the primary sense used by most bats to navigate their environment. However, the influence of echolocating behaviors upon the morphology of the auditory apparatus remains largely uninvestigated. While it is known that middle ear ossicle size scales positively with body mass across mammals, and that peak call frequency scales negatively with body mass among bats, there are still large gaps in our understanding of the degree to which allometry or ecology influences the morphology of the chiropteran auditory apparatus. To investigate this, we used μCT datasets to quantify three morphological components of the inner and middle ear: ossicle size, ossicle shape, and cochlear spirality. These data were collected across 27 phyllostomid species, spanning a broad range of body sizes, habitats, and dietary categories, and the relationships between these variables and ear morphology were assessed using a comparative phylogenetic approach. Ossicle size consistently scaled with strong negative allometry relative to body mass. Cochlear spirality was significantly (p = .025) associated with wing aspect ratio (a proxy for habitat use) but was not associated with body mass. From a morphological perspective, the malleus and incus exhibited some variation in kind with diet and call frequency, while stapes morphology is more closely tied to body size. Future work will assess these relationships within other chiropteran lineages, and investigate potential morphological differences in the middle and inner ear of echolocating-vs-non-echolocating taxa., (© 2023 American Association for Anatomy.)

Despite the remarkable morphological modifications that occurred in the thoracic limbs of bats, information about the brachial plexus in this group is still scarce. The present study aimed to describe the origin, structure, and distribution of these peripheral nerves in four Phyllostomidae species. Both antimeres of six Artibeus lituratus, five Desmodus rotundus, seven Glossophaga soricina, and five Phyllostomus hastatus-all adult males from the Adriano Lúcio Peracchi Collection (UFRRJ)-were dissected. After complete exposure of the structure, we found that the brachial plexus of D. rotundus and P. hastatus is formed by the same roots (C5-T1), whereas the fourth cervical spinal nerve and the second thoracic spinal nerve are present in G. soricina (C4-T1) and A. lituratus (C5-T2), respectively. There was intraspecific variation and asymmetry in the origin of the structure and the combinations of nerve segments forming terminal branches. The distribution to the target muscles and patagium, however, was not subject to significant variation in our sample. Data presented here support the presence of two prevailing conditions in distribution of nerves to the bat muscles, and the innervation of the membranes seems to be explained by embryogenesis. Although the brachial plexus in phyllostomid bats is similar to that of other terrestrial Laurasiatheria, aspects identified in these bats, apparently unique to Chiroptera, may be related to anatomical changes in the thoracic limbs functionally linked to flight., (© 2022 American Association for Anatomy.)

The extensive diversity observed in bat nasal chemosensory systems has been well-documented at the histological level. Understanding how this diversity evolved and developing hypotheses as to why particular patterns exist require a phylogenetic perspective, which was first outlined in the work of anatomist Kunwar Bhatnagar. With the onset of genetics and genomics, it might be assumed that the puzzling patterns observed in the morphological data have been clarified. However, there is still a widespread mismatch of genetic and morphological correlations among bat chemosensory systems. Novel genomic evidence has set up new avenues to explore that demand more evidence from anatomical structures. Here, we outline the progress that has been made in both morphological and molecular studies on the olfactory and vomeronasal systems in bats since the work of Bhatnagar. Genomic data of olfactory and vomeronasal receptors demonstrate the strong need for further morphological sampling, with a particular focus on receiving brain regions, glands, and ducts., (© 2023 The Authors. The Anatomical Record published by Wiley Periodicals LLC on behalf of American Association for Anatomy.)

Bite force is a performance metric commonly used to link cranial morphology with dietary ecology, as the strength of forces produced by the feeding apparatus largely constrains the foods an individual can consume. At a macroevolutionary scale, there is evidence that evolutionary changes in the anatomical elements involved in producing bite force have contributed to dietary diversification in mammals. Much less is known about how these elements change over postnatal ontogeny. Mammalian diets drastically shift over ontogeny-from drinking mother's milk to feeding on adult foods-presumably with equally drastic changes in the morphology of the feeding apparatus and bite performance. Here, we investigate ontogenetic morphological changes in the insectivorous big brown bat (Eptesicus fuscus), which has an extreme, positive allometric increase in bite force during development. Using contrast-enhanced micro-computed tomography scans of a developmental series from birth to adult morphology, we quantified skull shape and measured skeletal and muscular parameters directly related to bite force production. We found pronounced changes in the skull over ontogeny, including a large increase in the volume of the temporalis and masseter muscles, and an expansion of the skull dome and sagittal crest that would serve to increase the temporalis attachment area. These changes indicate that development of the jaw adductors play an important role in the development of biting performance of these bats. Notably, static bite force increases with positive allometry with respect to all anatomical measures examined, suggesting that modifications in biting dynamics and/or improved motor coordination also contribute to increases in biting performance., (© 2023 American Association for Anatomy.)

The gular gland is a skin gland located in the suprasternal region of adult males of some bat families. Knowledge of the morphology and functional aspects of these gland types is often limited. This study aimed to describe the structure and composition of the gular glands of three molossid species (Eumops patagonicus, Molossus fluminensis and Molossus molossus) with respect to their reproductive activity and to define the mechanism involved in secretion release. Different histological, histochemical and immunohistochemical techniques were used to achieve these goals. The results revealed that the size and composition of this gland are variable and are mostly related to the lipid content during the reproductive season. The results also documented, for the first time, the occurrence of mechanoreceptors associated with the surface of the glandular duct by detecting an S100 protein, indicating that an external stimulus activates secretion. Previous studies on other species have classified the gland using obsolete criteria; hence, we adopted a new classification of adenomeres in this study. Moreover, we investigated the gland secretion mechanism previously proposed. This study defines the implications of this gland in the reproduction of this species. Our preliminary interpretation of the function of the gular gland is that it is a cutaneous exocrine gland activated by mechanoreceptors involved in the reproductive behaviour of the Molossidae family., (© 2023 American Association for Anatomy.)

Dedicated nectarivory is a derived feeding habit that requires specialized cranial and soft-tissue morphologies to extract nectar from flowers. Nectarivory has evolved many times in terrestrial vertebrates, and in four bat families (Pteropodidae, Phyllostomidae, Vespertilionidae, and Mystacinidae). Within phyllostomids, specializations to nectarivory have been well documented in two subfamilies, Glossophaginae and Lonchophyllinae. However, nectarivory has also evolved independently in the genus Phyllostomus (subfamily Phyllostominae). Since Phyllostomus species have an omnivorous diet with a high consumption of nectar, they can be used to explore the basic morphological modifications linked to evolving a nectarivorous habit. Here, we focused on describing and comparing the morphological features potentially associated with nectarivory in Phyllostomus discolor. We present the first detailed tongue and palate morphological descriptions for P. discolor and perform skull morphometric analysis including 10 species. We found hair-like papillae on the tongue of P. discolor, a convergent feature with Glossophaginae and nectarivorous Pteropodids; these papillae likely confer an advantage when feeding on nectar. P. discolor does not show skull morphological features characteristic of nectarivorous bats, such as a long and narrow snout. We pose that the consumption of a variety of food, such as hard insects and fruits, and the large size of P. discolor relative to specialized nectarivores may create trade-offs against morphological specialization of the skull towards nectarivory. In contrast, a long and mobile tongue with hair-like papillae may be an evolutionary solution for nectar extraction that does not have a major impact on this species' ability to feed on other resources., (© 2022 American Association for Anatomy.)

Despite the high number of species and wide geographic dispersion, reproductive accessory glands (RAGs) of bats have traditionally received little attention in the literature, with some species not even having a basic description of their composition and structure. Thus, this study aimed to analyze and compare the composition, anatomy, and histology of male RAGs of bat species belonging to three of the largest (cosmopolitan) bat families: Vespertilionidae (Histiotus velatus), Molossidae (Molossus rufus), and Emballonuridae (Peropteryx leucoptera), in order to understand the variations in the bat RAGs. The results showed that the RAGs of H. velatus, M. rufus, and P. leucoptera are composed of an intra-abdominal prostatic complex, associated with the urethra, urethral glands, and a pair of inguinal bulbourethral glands; without ampullary glands or seminal vesicles. The prostatic complex can be composed of two (M. rufus and P. leucoptera) or three (H. velatus) prostatic regions, and can be compact (P. leucoptera), semi-lobed (M. rufus), or multilobed (H. velatus). Each prostatic region has unique and distinct characteristics, with the ventral region presenting a holocrine nature, exclusive to bats; while the dorsal and/or dorsolateral regions have similar characteristics to the ventral prostate of rats and to the human peripheral zone., (© 2022 American Association for Anatomy.)

Although evolvability of genes and traits may promote specialization during species diversification, how ecology subsequently restricts such variation remains unclear. Chemosensation requires animals to decipher a complex chemical background to locate fitness-related resources, and thus the underlying genomic architecture and morphology must cope with constant exposure to a changing odorant landscape; detecting adaptation amidst extensive chemosensory diversity is an open challenge. In phyllostomid bats, an ecologically diverse clade that evolved plant visiting from a presumed insectivorous ancestor, the evolution of novel food detection mechanisms is suggested to be a key innovation, as plant-visiting species rely strongly on olfaction, supplementarily using echolocation. If this is true, exceptional variation in underlying olfactory genes and phenotypes may have preceded dietary diversification. We compared olfactory receptor (OR) genes sequenced from olfactory epithelium transcriptomes and olfactory epithelium surface area of bats with differing diets. Surprisingly, although OR evolution rates were quite variable and generally high, they are largely independent of diet. Olfactory epithelial surface area, however, is relatively larger in plant-visiting bats and there is an inverse relationship between OR evolution rates and surface area. Relatively larger surface areas suggest greater reliance on olfactory detection and stronger constraint on maintaining an already diverse OR repertoire. Instead of the typical case in which specialization and elaboration are coupled with rapid diversification of associated genes, here the relevant genes are already evolving so quickly that increased reliance on smell has led to stabilizing selection, presumably to maintain the ability to consistently discriminate among specific odorants-a potential ecological constraint on sensory evolution., (© 2022 The Authors. Evolution published by Wiley Periodicals LLC on behalf of The Society for the Study of Evolution.)

Background: Mammal evolution has generated diverse craniofacial morphologies, including remarkable movable rostral appendages. However, the muscular and skeletal architecture providing the mobility of these appendages remains largely unexplored. Here, we focus on chiropteran noseleaves and compare the three-dimensional internal morphology of late-stage embryos between the greater horseshoe bat Rhinolophus ferrumequinum, which possesses a noseleaf, and the Asian bent-winged bat Miniopterus fuliginosus and Egyptian fruit bat Rousettus aegyptiacus, which do not. We also assess earlier stage cell proliferation within the rostrum to elucidate cellular mechanisms underlying noseleaf-associated morphological modifications., Results: The musculus maxillolabialis inserted into proximal vibrissae follicles in M fuliginosus and R aegyptiacus embryos but instead inserted into the horseshoe plate in R ferrumequinum. This modification suggests that the M maxillolabialis has adapted to controlling the noseleaf rather than vibrissae in rhinolophid bats. Our cellular analysis showed higher cell proliferation within the maxillary and frontonasal processes of St. 14 embryos in R ferrumequinum compared to M fuliginosus and R aegyptiacus, suggesting that the spatial alteration of noseleaf-associated muscle is derived from changes in facial morphogenesis that occur by St. 14., Conclusion: This is the first study clarifying the morphological and cellular bases underlying the development of mammalian rostral appendages., (© 2022 American Association for Anatomy.)

Our knowledge of nasal cavity anatomy has grown considerably with the advent of micro-computed tomography (CT). More recently, a technique called diffusible iodine-based contrast-enhanced CT (diceCT) has rendered it possible to study nasal soft tissues. Using diceCT and histology, we aim to (a) explore the utility of these techniques for inferring the presence of venous sinuses that typify respiratory mucosa and (b) inquire whether distribution of vascular mucosa may relate to specialization for derived functions of the nasal cavity (i.e., nasal-emission of echolocation sounds) in bats. Matching histology and diceCT data indicate that diceCT can detect venous sinuses as either darkened, "empty" spaces, or radio-opaque islands when blood cells are present. Thus, we show that diceCT provides reliable information on vascular distribution in the mucosa of the nasal airways. Among the bats studied, a nonecholocating pteropodid (Cynopterus sphinx) and an oral-emitter of echolocation sounds (Eptesicus fuscus) possess venous sinus networks that drain into the sphenopalatine vein rostral to the nasopharynx. In contrast, nasopharyngeal passageways of nasal-emitting hipposiderids are notably packed with venous sinuses. The mucosae of the nasopharyngeal passageways are far less vascular in nasal-emitting phyllostomids, in which vascular mucosae are more widely distributed in the nasal cavity, and in some nectar-feeding species, a particularly large venous sinus is adjacent to the vomeronasal organ. Therefore, we do not find a common pattern of venous sinus distribution associated with nasal emission of sounds in phyllostomids and hipposiderids. Instead, vascular mucosa is more likely critical for air-conditioning and sometimes vomeronasal function in all bats., (© 2021 American Association for Anatomy.)

Cryptic species that coexist in sympatry are likely to simultaneously experience strong competition and hybridization. The first phenomenon would lead to character displacement, whereas the second can potentially promote morphological similarity through adaptive introgression. The main goal of this work was to investigate the effect of introgressive hybridization on the morphology of cryptic Iberian Eptesicus bats when facing counteracting evolutionary forces from interspecific competition. We found substantial overlap both in dentition and in wing morphology traits, though mainly in individuals in sympatry. The presence of hybrids contributes to a fifth of this overlap, with hybrids showing traits with intermediate morphometry. Thus, introgressive hybridization may contribute to species adaptation to trophic and ecological space responding directly to the macro-habitats characteristics of the sympatric zone and to local prey availability. On the other hand, fur shade tended to be browner and brighter in hybrids than parental species. Colour differences could result from partitioning of resources as an adaptation to environmental factors such as roost and microhabitats. We argue that a balance between adaptive introgression and niche partitioning shapes species interactions with the environment through affecting morphological traits under selection., (© 2022. The Author(s).)

The shape and size of a skull provides insight into the age, breed and gender of the animal. Skull shape variations have been reported in different animals, with some theories linking these variations to evolution and/or migration. This study assessed the variations observed in the skull shape, size and gross morphometrics of two groups of the Eidolon helvum obtained from two geographical regions in Nigeria (south and north). All skulls were rostro-caudally elongated, having a dolichocephalic appearance. The skulls from the north had a distinct dome shape, with a more prominent zygomatic process, absence of a 'diastema' and an extra upper molar, while the southern skulls showed a more dorsally flattened skull and a less prominent zygomatic process. The shape of the sagittal crest was different in the two groups, while there was the presence of an accessory infraorbital foramen in some of the southern skulls. The southern skulls lacked the palatine foramen. The lacrimal foramen was observed to be more caudally placed in the southern skulls. Values for most linear measurements were higher in the northern skulls, although statistically significant difference was not present in all. The value for the neurocranial volume was considerably higher in the northern skulls (4.41 ± 0.28 mls) relative to the southern skulls (2.0 ± 0.27 mls). Statistically significant differences were not observed between males and females (within regions). Data obtained from this study may find application in evolution and migration studies, wildlife medicine and surgery and comparative and forensic anatomy.

The morphology of the stylohyal-tympanic bone articulation found in laryngeally echolocating bats is highly indicative of a function associated with signal production. One untested hypothesis is that this morphology allows the transfer of a sound signal from the larynx to the tympanic bones (auditory bulla) via the hyoid apparatus during signal production by the larynx. We used µCT data and finite element analysis to model the propagation of sound through the hyoid chain into the tympanic bones to test this hypothesis. We modeled sound pressure (dB) wave propagation from the basihyal to the tympanic bones, vibratory behavior (m) of the stylohyal-tympanic bone unit, and the stylohyal and tympanic bones when the stylohyal bone is allowed to pivot on the tympanic bone. Sound pressure wave propagation was modeled using the harmonic acoustics solver in ANSYS and vibratory behavior was modeled using coupled modal and harmonic response analyses in ANSYS. For both analyses (harmonic acoustics and harmonic response), the input excitation on the basihyal and thyrohyals was modeled as the estimated pressure (Pa) imposed by the collision of the vibrating thyroid cartilage of the larynx against these bones during signal production. Our models support the hypothesis that this stereotypical hyoid morphology found in laryngeally echolocating bats can transfer sound to the auditory bullae at an amplitude that is likely heard for the species Artibeus jamaicensis and Rhinolophus pusillus., (© 2021 Anatomical Society.)

Body size and shape fundamentally determine organismal energy requirements by modulating heat and mass exchange with the environment and the costs of locomotion, thermoregulation, and maintenance. Ecologists have long used the physical linkage between morphology and energy balance to explain why the body size and shape of many organisms vary across climatic gradients, e.g., why larger endotherms are more common in colder regions. However, few modeling exercises have aimed at investigating this link from first principles. Body size evolution in bats contrasts with the patterns observed in other endotherms, probably because physical constraints on flight limit morphological adaptations. Here, we develop a biophysical model based on heat transfer and aerodynamic principles to investigate energy constraints on morphological evolution in bats. Our biophysical model predicts that the energy costs of thermoregulation and flight, respectively, impose upper and lower limits on the relationship of wing surface area to body mass (S-MR), giving rise to an optimal S-MR at which both energy costs are minimized. A comparative analysis of 278 species of bats supports the model’s prediction that S-MR evolves toward an optimal shape and that the strength of selection is higher among species experiencing greater energy demands for thermoregulation in cold climates. Our study suggests that energy costs modulate the mode of morphological evolution in bats—hence shedding light on a long-standing debate over bats’ conformity to ecogeographical patterns observed in other mammals—and offers a procedure for investigating complex macroecological patterns from first principles.

Phylogenomics of bats suggests that their echolocation either evolved separately in the bat suborders Yinpterochiroptera and Yangochiroptera, or had a single origin in bat ancestors and was later lost in some yinpterochiropterans 1-6 . Hearing for echolocation behaviour depends on the inner ear, of which the spiral ganglion is an essential structure. Here we report the observation of highly derived structures of the spiral ganglion in yangochiropteran bats: a trans-otic ganglion with a wall-less Rosenthal's canal. This neuroanatomical arrangement permits a larger ganglion with more neurons, higher innervation density of neurons and denser clustering of cochlear nerve fascicles 7-13 . This differs from the plesiomorphic neuroanatomy of Yinpterochiroptera and non-chiropteran mammals. The osteological correlates of these derived ganglion features can now be traced into bat phylogeny, providing direct evidence of how Yangochiroptera differentiated from Yinpterochiroptera in spiral ganglion neuroanatomy. These features are highly variable across major clades and between species of Yangochiroptera, and in morphospace, exhibit much greater disparity in Yangochiroptera than Yinpterochiroptera. These highly variable ganglion features may be a neuroanatomical evolutionary driver for their diverse echolocating strategies 4,14-17 and are associated with the explosive diversification of yangochiropterans, which include most bat families, genera and species., (© 2022. The Author(s), under exclusive licence to Springer Nature Limited.)

Haeckel's recapitulation theory has been a controversial topic in evolutionary biology. However, we have seen some recent cases applying Haeckel's view to interpret the interspecific variation of prenatal ontogeny. To revisit the validity of Haeckel's recapitulation theory, we take bats that have undergone drastic morphological changes and possess a characteristic ecology as a case study. All members of Rhinolophoidea and Yangochiroptera can generate an ultrasonic pulse from the larynx to interpret surrounding objects (laryngeal echolocation) whereas Pteropodidae lacks such ability. It is known that the petrosal bone is particularly derived in shape and expanded in laryngeal echolocators. If Haeckel's recapitulation theory holds, the formation of this derived trait should occur later than those of other bones. Therefore, we compared the prenatal ossification timing of the petrosal in 15 bat species and five outgroup species. We found that the ossification of the petrosal is accelerated in laryngeal echolocators while it is the last bone to ossify in non-laryngeal echolocating bats and non-volant mammals, which runs counter to the prediction generated by Haeckel's recapitulation theory. We point out the evolutionarily labile nature of trait developmental timing and emphasize that Haeckel's recapitulation theory does not hold in many cases. We caution that generating predictions on ancestral conditions and evolutionary history leading from Haeckel's recapitulation theory is not well supported., (© 2021 Wiley Periodicals LLC.)

Background: This work was designed to provide a morphologic, morphometric and histochemical description of the eye of the African straw-coloured fruit bat (Eidolon helvum). An explanation of the optical role of the choroidal papillae in the vision of megachiropteran bats was provided., Materials and Methods: Enucleated eyes of captured fruit bats were measured and processed for light microscopy., Results: Typical gross features of the mammalian eye including an anterior transparent cornea, posterior whitish sclera and a golden-brown iris surrounding a round pupil were observed in the eye. Presence of undulating retina typically found in megachiropterans was also seen. The ratio of mean corneal diameter to mean axial eye diameter was 0.58 ± 0.08. The histochemical investigation of the eye indicated the presence of mucins, proteoglycans, hyaluronic acid, glycogen and/or glycoproteins in the corneal, scleral, choroidal and retinal tissues., Conclusions: The presence of reflective materials of the tapetum lucidum on the undulating retina was shown to be a morphological adaptation for increased light sensitivity as each parabolic surface of the choroidal papillae served as a convex mirror, reflecting the light rays to the adjacent parabolic surface, thus sensitising photoreceptors in affected regions. This phenomenon thus empowers megachiropteran bats with improved scotopic visual capability and could explain why most of them are reliant on their vison without the need for echolocation.

The most superficial epidermal layer in endotherms is the stratum corneum (SC), which is composed of dead corneocytes embedded in a lipid matrix with free fatty acids, cholesterol, ceramides, and cerebrosides; the lipid composition of the SC determines its permeability to water vapor. Lipids that are more polar, have longer hydrocarbon chains, and are less bulky are often packed in more ordered phase states to slow cutaneous evaporative water loss (CEWL); these lipids also resist transitions to more disordered phases at high ambient temperatures (T a ). In bats, wing and tail membranes (wing patagia and tail uropatagium, respectively) allow powered flight, but increase surface area, and hence CEWL, with implications for survival in arid environments. We captured Pipistrellus kuhlii from an arid habitat and measured the lipid composition of the SC of the plagiopatagium in the wing, the uropatagium, and the non-membranous region (NMR) of the body using thin layer chromatography and reversed phase high performance liquid chromatography coupled with atmospheric pressure photoionization mass spectrometry. The patagia contained more cholesterol and shorter-chained ceramides, and fewer cerebrosides than the NMR, indicating that the lipid phase transition temperature in the patagia is lower than in the NMR. Thus, at moderate T a the lipids in the SC in all body regions will remain in an ordered phase state, allowing water conservation; but as T a increases, the lipids in the SC of the patagia will more easily transition into a disordered phase, resulting in increased CEWL from the patagia facilitating efficient heat dissipation in hot environments., (Copyright © 2021 Elsevier Inc. All rights reserved.)

Species richness exhibits well-known patterns across elevational gradients in various taxa, but represents only one aspect of quantifying biodiversity patterns. Functional and phylogenetic diversity have received much less attention, particularly for vertebrate taxa. There is still a limited understanding of how functional, phylogenetic and taxonomic diversity change in concert across large gradients of elevation. Here, we focused on the Himalaya-representing the largest elevational gradients in the world-to investigate the patterns of taxonomic, functional and phylogenetic diversity in a bat assemblage. Combining field data on species occurrence, relative abundance, and functional traits with measures of phylogenetic diversity, we found that bat species richness and functional diversity declined at high elevation but phylogenetic diversity remained unchanged. At the lowest elevation, we observed low functional dispersion despite high species and functional richness, suggesting a niche packing mechanism. The decline in functional richness, dispersion, and divergence at the highest elevation is consistent with patterns observed due to environmental filtering. These patterns are driven by the absence of rhinolophid bats, four congeners with extreme trait values. Our data, some of the first on mammals from the Himalayan region, suggest that in bat assemblages with relatively high species diversity, phylogenetic diversity may not be a substitute to measure functional diversity., (© 2021. The Author(s).)

Echolocation is the use of reflected sound to sense features of the environment. Here, we show that soft-furred tree mice ( Typhlomys ) echolocate based on multiple independent lines of evidence. Behavioral experiments show that these mice can locate and avoid obstacles in darkness using hearing and ultrasonic pulses. The proximal portion of their stylohyal bone fuses with the tympanic bone, a form previously only seen in laryngeally echolocating bats. Further, we found convergence of hearing-related genes across the genome and of the echolocation-related gene prestin between soft-furred tree mice and echolocating mammals. Together, our findings suggest that soft-furred tree mice are capable of echolocation, and thus are a new lineage of echolocating mammals., (Copyright © 2021 The Authors, some rights reserved; exclusive licensee American Association for the Advancement of Science. No claim to original U.S. Government Works.)

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., (© 2020 Anatomical Society.)

The gain or loss of anatomical features is an important mechanism of morphological evolution and ecological adaptation. Dental anomalies-the loss or gain of teeth-are widespread and a potential source of craniodental specialization among mammals, yet their macroevolutionary patterns have been rarely explored. We present the first phylogenetic comparative study of dental anomalies across the second largest mammal Order, Chiroptera (bats). We conducted an extensive literature review and surveyed a large sample of museum specimens to analyze the types and prevalence of dental anomalies across bats, and performed phylogenetic comparative analyses to investigate the role of phylogenetic history and dietary specialization on incidence of dental anomalies. We found dental anomalies have a significant phylogenetic signal, suggesting they are not simply the result of idiosyncratic mutations or random developmental disorders, but may have ancestral genetic origins or result from shared developmental pathways among closely related species. The incidence of dental anomalies was not associated with diet categories, suggesting no effect of craniodental specialization on dental anomalies across bats. Our results give insight into the macroevolutionary patterns of dental anomalies in bats, and provide a foundation for investigating new hypotheses underlying the evolution of dental variation and diversity in mammals., (© 2021 The Authors. Evolution © 2021 The Society for the Study of Evolution.)

Living primates show a complex trend in reduction of nasal cavity spaces and structures due to moderate to severe constraint on interorbital breadth. Here we describe the ontogeny of the posterior end of the primate cartilaginous nasal capsule, the thimble shaped posterior nasal cupula (PNC), which surrounds the hind end of the olfactory region. We used a histologically sectioned sample of strepsirrhine primates and two non-primates (Tupaia belangeri, Rousettus leschenaulti), and histochemical and immunohistochemical methods to study the PNC in a perinatal sample. At birth, most strepsirrhines possess only fragments of PNC, and these lack a perichondrium. Fetal specimens of several species reveal a more complete PNC, but the cartilage exhibits uneven or weak reactivity to type II collagen antibodies. Moreover, there is relatively less matrix than in the septal cartilage, resulting in clustering of chondrocytes, some of which are in direct contact with adjacent connective tissues. In one primate (Varecia spp.) and both non-primates, the PNC has a perichondrium at birth. In older, infant Varecia and Rousettus, the perichondrium of the PNC is absent, and PNC fragmentation at its posterior pole has occurred in the former. Loss of the perichondrium for the PNC appears to precede resorption of the posterior end of the nasal capsule. These results suggest that the consolidation of the basicranial and facial skeletons happens ontogenetically earlier in primates than other mammals. We hypothesize that early loss of cartilage at the sphenoethmoidal articulation limits chondral mechanisms for nasal complexity, such as interstitial expansion or endochondral ossification., (© 2021 American Association for Anatomy.)

Mammalian nasal capsule development has been described in only a few cross-sectional age series, rendering it difficult to infer developmental mechanisms that influence adult morphology. Here we examined a sample of Leschenault's rousette fruit bats (Rousettus leschenaultii) ranging in age from embryonic to adult (n = 13). We examined serially sectioned coronal histological specimens and used micro-computed tomography scans to visualize morphology in two older specimens. We found that the development of the nasal capsule in Rousettus proceeds similarly to many previously described mammals, following a general theme in which the central (i.e., septal) region matures into capsular cartilage before peripheral regions, and rostral parts of the septum and paries nasi mature before caudal parts. The ossification of turbinals also generally follows a rostral to the caudal pattern. Our results suggest discrete mechanisms for increasing complexity of the nasal capsule, some of which are restricted to the late embryonic and early fetal timeframe, including fissuration and mesenchymal proliferation. During fetal and early postnatal ontogeny, appositional and interstitial chondral growth of cartilage modifies the capsular template. Postnatally, appositional bone growth and pneumatization render greater complexity to individual structures and spaces. Future studies that focus on the relative contribution of each mechanism during development may draw critical inferences how nasal morphology is reflective of, or deviates from the original fetal template. A comparison of other chiropterans to nasal development in Rousettus could reveal phylogenetic patterns (whether ancestral or derived) or the developmental basis for specializations relating to respiration, olfaction, or laryngeal echolocation., (© 2020 American Association for Anatomy.)

Morphological evolution is often assumed to be causally related to underlying patterns of ecological trait evolution. However, few studies have directly tested whether evolutionary dynamics of-and major shifts in-ecological resource use are coupled with morphological shifts that may facilitate trophic innovation. Using diet and multivariate cranial (microCT) data, we tested whether rates of trophic and cranial evolution are coupled in the radiation of New World bats. We developed a generalizable information-theoretic method for describing evolutionary rate heterogeneity across large candidate sets of multirate evolutionary models, without relying on a single best-fitting model. We found considerable variation in trophic evolutionary dynamics, in sharp contrast to a largely homogeneous cranial evolutionary process. This dichotomy is surprising given established functional associations between overall skull morphology and trophic ecology. We suggest that assigning discrete trophic states may underestimate trophic generalism and opportunism, and that this radiation could be characterized by labile crania and a homogeneous dynamic of generally high morphological rates. Overall, we discuss how trophic classifications could substantively impact our interpretation of how these dynamics covary in adaptive radiations., (© 2021 The Authors. Evolution © 2021 The Society for the Study of Evolution.)

Bats have adapted to many different feeding habits, which are known to induce morphophysiological adaptations in several tissues, especially those particularly involved with absorption, metabolism and excretion. The common vampire bat (Desmodus rotundus) has a very unique diet (blood), which, among other challenges, seems to pose a risk to their kidneys, due to the increased nitrogen excretion imposed by their remarkably high protein meal. Fruit-eating bats (Artibeus lituratus) consume a high carbohydrate diet and may be taken as a suitable species for this dietary comparative study. Here we aimed at investigating the renal morphology and stereology, kidneys antioxidant capacity, and plasma antidiuretic hormone (ADH) concentrations in adult fruit-eating and vampire bats. Sixteen animals were captured and used in this study, being 8 adult males from each species. Our results showed higher morphological standards of glomerular area, volumetric density of glomeruli, and renal somatic index for vampire bats, as well as higher reactive species of oxygen (ROS) production, such as nitric oxide (NO), higher plasma iron reduction ability (FRAP), higher activity of the antioxidant enzyme glutathione-S-transferase (GST) and a higher malondialdehyde production (MDA) in vampires' kidneys, compared to the fruit-eating species. The activities of the antioxidant enzymes superoxide dismutase (SOD) and catalase (CAT) were higher in fruit-eating bats. Plasma ADH concentrations were not different between species. Taken together, the renal morphophysiology conditions presented by vampire bats might be associated with a high demand for nitrogenous products excretion imposed by protein and iron overload. These features may play an important role on preventing protein-overload nephropathy, allowing vampires to survive under such a unique diet., (Copyright © 2020 Elsevier GmbH. All rights reserved.)

The following mammal assemblage was identified among the materials collected in 2020 from the Pleistocene of the Lang Trang cave (northern Vietnam): the primates Pongo sp., Trachypithecus sp., Macaca cf. nemestrina (Linnaeus, 1766), and Macaca sp.; the carnivorans Arctonyx collaris rostratus Matthew et Granger, 1923 and Panthera sp.; the chiropteran Ia io Thomas, 1902; the rodent Hystrix kiangsenensis Wang, 1931; the proboscidean Elephas sp.; the perissodactyls Tapirus indicus (Desmarest, 1819) and Dicerorhinus sumatrensis (Fischer, 1814); the artiodactyls Sus scrofa Linnaeus, 1758, S. barbatus Müller, 1838, Tragulus kanchil (Raffles, 1821), Hydropotes inermis Swinhoe, 1870, Muntiacus muntjak (Zimmermann, 1780), Axis porcinus (Zimmermann, 1780), Rusa unicolor (Kerr, 1792), and Capricornis sumatraensis (Bechstein, 1799). I. io, S. barbatus, T. kanchil, H. inermis, and A. porcinus were detected in the Lang Trang fauna for the first time. The mammal assemblage is dominated by inhabitants of tropical forests (from lowlands to mountains), subtropical forms are also present.

This study was aimed to characterize the spermatogenic process and its seasonal variation in Desmodus rotundus, in the Caatinga biome, a water-limited ecosystem, with marked water restriction during most of the year. Collections of adult animals were performed during the dry and rainy seasons, and after euthanasia, their testes were processed histologically to perform morphological, morphometric, ultrastructural and immunohistochemical analyzes. The percentage of seminiferous epithelium, number of Leydig cells per gram of testis, and population of Sertoli cells and A-type spermatogonia presented by D. rotundus were significantly higher in the rainy season, while the percentage of lumen, mitotic index, support capacity performed by Sertoli cells, and overall yield of spermatogenesis were higher in the dry season. The ultrastructure of spermatogenesis was similar to that described in other mammals, and the immunohistochemical analysis revealed activity of the aromatase enzyme in Sertoli cells, Leydig cells, spermatocytes and spermatids, as well as the presence of androgen receptors in Sertoli cells and Leydig cells. FGF2 activity was detected in primary spermatocytes in zygotene and pachytene, as well as secondary spermatocytes and rounded and elongated spermatids, while the BCL-2 protein was expressed in primary spermatocytes in zygotene and pachytene, secondary spermatocytes, and rounded spermatids. The activity of these molecules was similar in both seasons, and associated with the morphometric findings, indicates maintenance in the integrity of the seminiferous epithelium throughout the year. The seasonal study of D. rotundus spermatogenesis indicates a continuous spermatogenesis pattern and suggests a greater production of spermatozoa in the rainy season in the Caatinga biome., Competing Interests: The authors have declared that no competing interests exist.

Small echolocating bats are set apart from most other mammals by their relatively large cerebellum, a feature that has been associated to echolocation, as it is presumed to indicate a relatively enlarged number of neurons in the cerebellum in comparison to other brain structures. Here we quantify the neuronal composition of the cerebral cortex, cerebellum and remaining brain structures of seven species of large Pteropodid bats (formerly classified as megachiropterans), one of which echolocates, and six species of small bats (formerly classified as microchiropterans), all of which echolocate. This chiropteran data is compared to 60 mammalian species in our dataset to determine whether the relatively large cerebellum of the small echolocating bats, and possibly that of the echolocating Pteropodid, contains a relatively enlarged number of neurons. We find no evidence that the distribution of neurons differs between microchiropterans and megachiropterans, but our data indicate that microchiropterans, like the smallest shrew in our dataset, have diminutive cerebral cortices, which makes the cerebellum appear relatively large. We propose that, in agreement with the diminutive brain size of the earliest fossil mammals, this is a plesiomorphic, not a derived, feature of microchiropteran brains. The results of this study also reveal important neural characteristics related to the phylogenetic affinities and relationships of the chiropterans., (© 2020 Wiley Periodicals LLC.)

Centurio senex is an iconic bat characterized by a facial morphology deviating far from all other New World Leaf Nosed Bats (Phyllostomidae). The species has a bizarrely wrinkled face and lacks the characteristic nose leaf. Throughout its distribution from Mexico to Northern South America the species is most of the time rarely captured and only scarce information on its behavior and natural history is available. Centurio senex is frugivorous and one of the few bats documented to consume also hard seeds. Interestingly, the species shows a distinct sexual dimorphism: Adult males have more pronounced facial wrinkles than females and a fold of skin under the chin that can be raised in style of a face mask. We report the first observations on echolocation and mating behavior of Centurio senex, including synchronized audio and video recordings from an aggregation of males in Costa Rica. Over a period of 6 weeks we located a total of 53 perches, where during the first half of the night males were hanging with raised facial masks at a mean height of 2.35 m. Most of the time, the males moved just their wing tips, and spontaneously vocalized in the ultrasound range. Approaches of other individuals resulted in the perching male beating its wings and emitting a very loud, low frequency whistling call. Following such an encounter we recorded a copulation event. The observed aggregation of adult C. senex males is consistent with lek courtship, a behavior described from only few other bat species., Competing Interests: The authors have declared that no competing interests exist. The owner of the study site, Hotel Villa Blanca, provided support in the form of salaries for authors J.V. and E.H. This does not alter our adherence to PLOS ONE policies on sharing data and materials.

Bats possess wings comprised of a flexible membrane and a jointed skeletal structure allowing them to execute complex flight maneuvers such as rapid tight turns. The extent of a bat's tight turning capability can be explored by analyzing a 180-degree U-turn. Prior studies have investigated more subtle flight maneuvers, but the kinematic and aerodynamic mechanisms of a U-turn have not been characterized. In this work, we use 3D optical motion capture and aerodynamic simulations to investigate a U-turn maneuver executed by a great roundleaf bat (Hipposideros armiger: mass = 55 g, span = 51 cm). The bat was observed to decrease its flight velocity and gain approximately 20 cm of altitude entering the U-turn. By lowering its velocity from 2.0 m/s to 0.5 m/s, the centripetal force requirement to execute a tight turn was substantially reduced. Centripetal force was generated by tilting the lift force vector laterally through banking. During the initiation of the U-turn, the bank angle increased from 20 degrees to 40 degrees. During the initiation and persisting throughout the U-turn, the flap amplitude of the right wing (inside of the turn) increased relative to the left wing. In addition, the right wing moved more laterally closer to the centerline of the body during the end of the downstroke and the beginning of the upstroke compared to the left wing. Reorientation of the body into the turn happened prior to a change in the flight path of the bat. Once the bat entered the U-turn and the bank angle increased, the change in flight path of the bat began to change rapidly as the bat negotiated the apex of the turn. During this phase of the turn, the minimum radius of curvature of the bat was 5.5 cm. During the egress of the turn, the bat accelerated and expended stored potential energy by descending. The cycle averaged total power expenditure by the bat during the six wingbeat cycle U-turn maneuver was 0.51 W which was approximately 40% above the power expenditure calculated for a nominally straight flight by the same bat. Future work on the topic of bat maneuverability may investigate a broader array of maneuvering flights characterizing the commonalities and differences across flights. In addition, the interplay between aerodynamic moments and inertial moments are of interest in order to more robustly characterize maneuvering mechanisms., Competing Interests: The authors have declared that no competing interests exist.