World Fruit Bats Research Articles

CATATAN LAPANGAN MENGENAI SATWALIAR DI LOKASI EKOWISATA SYURGAR SAUSAPOR, KABUPATEN TAMBRAUW

This study was conducted at Syurgar ecoutourism site, subdistrict of Sausapor Tambrauw district. Data was collected from June to July 2020 using survey method, direct catching, cage trap and interview. This study aimed to list the wildlife species, especially Birds, Mammals Herpetofauna, and Butterflies in order to complete data of the government of Tambrauw District for the purpose of Syurgar ecotourism development. This study recorded that there were 15 birds’ species of 13 families, 12 mammals’ species of 6 family, 4 herpetofauna’s species of 4 families, and 25 butterflies of 4 families. Syurgar has a potential to be a birdwatching site. There are several appealing birds, including Lesser bird-of-paradise (Paradisaea minor), king bird of paradise (Cincinurus regius), blyth’s hornbill (Rhyncetores plicatus), sulphur-crested cockatoo (Cacatua galerita). Endemic species of mammals include long-beaked echidna (Zaglosus bruijnii), dorcopsis/forest’s wallaby (Dorcopsis sp.), northern common cuscus (Phalanger orientalis), common spotted cuscus (Spilocuscus maculates), old world fruit bats (Dobsonia sp.), great flying fox (Pteropus neohibernicus). Endemic species of herpetofauna include frogs (Papurana sp.), Papua wrinkled ground frog (Cornufer papuensis). Eventually, some butterflies’ species include redeye bushbrown (Mycalesis aethiops), bushbrown (Mycalesis elia), and Papuan gull (Cepora abnormis). It is strongly suspected that several birds, mammals, herpetofauna, and butterflies have not been recorded due to time constraints. Therefore, further study is recommended for data completion.

Old World Fruit Bats (Pteropodidae, Chiroptera) from the Pleistocene of Vietnam.

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.

USE OF VISCOELASTIC COAGULATION TESTING IN MEGACHIROPTERA (PTEROPUS HYPOMELANUS AND PTEROPUS VAMPYRUS) REVEALS HIGH VARIABILITY IN CLOT KINETICS.

Megachiroptera is a mammalian suborder that includes old world fruit bats. Common clinical problems among captive Megachiroptera, such as liver disease (e.g., iron storage disease), kidney disease (e.g., protein-losing nephropathy), and heart disease (e.g., dilated cardiomyopathy), carry elevated risk for hemostatic derangements. The assessment of viscoelastic coagulation assays, however, has not yet been reported in bats. The main objective of the study was to describe viscoelastography data using the Viscoelastic Coagulation Monitor (VCM) Vet in captive large flying foxes (Pteropus vampyrus) (n = 20) and variable flying foxes (Pteropus hypomelanus) (n = 10). Additional objectives were to compare viscoelastic and clotting parameters (1) between healthy P. vampyrus and P. hypomelanus bats and (2) between untreated bats and those treated with meloxicam or aspirin, and (3) to examine relationships between activated partial thromboplastin time (aPTT) and potentially homologous viscoelastic parameters clotting time (CT) and clot formation time (CFT). The results showed marked variability among clinically normal bats. The intrinsic pathway, as measured by aPTT, had prolonged times compared with most terrestrial mammals, but similar times to birds, marine mammals, and sea turtles. A search of P. vampyrus genome found stop codons present in two exons of the factor XI gene; alterations in factor XI expression would be expected to alter intrinsic coagulation. Because of the high variability, no statistically significant findings were noted in the secondary objectives. Correlation between aPTT and CT or CFT was not strong (rs = 0.406 or 0.192, respectively). The results from this study suggest that clot kinetics vary widely among Megachiroptera when using the VCM Vet with untreated blood. A prolonged intrinsic coagulation pathway, as has been found in other megachiropteran species, and activation of the extrinsic coagulation pathway during venipuncture may be responsible for the inconsistent results.

Pteropus vampyrus TRIM40 Is an Interferon-Stimulated Gene That Antagonizes RIG-I-like Receptors.

Nipah virus (NiV; genus: Henipavirus; family: Paramyxoviridae) naturally infects Old World fruit bats (family Pteropodidae) without causing overt disease. Conversely, NiV infection in humans and other mammals can be lethal. Comparing bat antiviral responses with those of humans may illuminate the mechanisms that facilitate bats' tolerance. Tripartite motif proteins (TRIMs), a large family of E3-ubiquitin ligases, fine-tune innate antiviral immune responses, and two human TRIMs interact with Henipavirus proteins. We hypothesize that NiV infection induces the expression of an immunosuppressive TRIM in bat, but not human cells, to promote tolerance. Here, we show that TRIM40 is an interferon-stimulated gene (ISG) in pteropodid but not human cells. Knockdown of bat TRIM40 increases gene expression of IFNβ, ISGs, and pro-inflammatory cytokines following poly(I:C) transfection. In Pteropus vampyrus, but not human cells, NiV induces TRIM40 expression within 16 h after infection, and knockdown of TRIM40 correlates with reduced NiV titers as compared to control cells. Bats may have evolved to express TRIM40 in response to viral infections to control immunopathogenesis.

Large Old World Fruit Bats on the Brink of Extinction: Causes and Consequences

Large Old World fruit bats (LOWFBs), species of Pteropus, Acerodon, and related genera of large bats in the pteropodid subfamily Pteropodinae, play important roles as agents of dispersal and pollination across the Paleotropics. LOWFBs are also collectively the most threatened group of bats in the world, with 71% of extant species assessed as threatened by International Union for Conservation of Nature. As highlighted here, contrary to other bats, the vast majority of LOWFBs face multiple simultaneous threats. Most importantly, biological and ecological traits, in particular life history characteristics, diet, movement, social ecology, and physiology, intensify threats and accelerate species declines. Furthermore, we demonstrate that LOWFBs are to be considered keystone species and express concern about the erosion of this role and the cascading effects expected on native ecosystems. In response to this alarming situation, we advance general recommendations and identify overarching research and conservation actions.

Phylogenetic relationships of a novel bat fly species infesting the geographically widespread Old World fruit bat, Rousettus leschenaultii, in Southern Asia.

The global epidemiological significance of bats and their blood-sucking ectoparasites is increasingly recognized. However, relevant data are scarce from Pakistan where the Palearctic and Oriental zoogeographic regions meet. In this study, 200 bats belonging to five species were examined for the presence of ectoparasites in Pakistan. Bat flies were found only on Leschenault's fruit bat (Rousettus leschenaultii). The prevalence of infestation did not correlate with habitat type and host traits including age, reproductive status, and sex. All bat flies represented the same Eucampsipoda species which was shown to be morphologically different from all species of its genus with known south Asian distribution and belonged to a separate phylogenetic group. These results highlight the existence of a hitherto undescribed bat fly species in southern Asia, which is not shared by the fruit bat species (R. leschenaultii) and insectivorous ones (e.g., Rhinopomamicrophyllum) thus probably playing a role only in intraspecific transmission of pathogens.

Comparative analyses of bat genomes identify distinct evolution of immunity in Old World fruit bats

Bats have been identified as natural reservoir hosts of several zoonotic viruses, prompting suggestions that they have unique immunological adaptations. Among bats, Old World fruit bats (Pteropodidae) have been linked to multiple spillovers. To test for lineage-specific molecular adaptations in these bats, we developed a new assembly pipeline to generate a reference-quality genome of the fruit bat Cynopterus sphinx and used this in comparative analyses of 12 bat species, including six pteropodids. Our results reveal that immunity-related genes have higher evolutionary rates in pteropodids than in other bats. Several lineage-specific genetic changes were shared across pteropodids, including the loss of NLRP1, duplications of PGLYRP1 and C5AR2, and amino acid replacements in MyD88. We introduced MyD88 transgenes containing Pteropodidae-specific residues into bat and human cell lines and found evidence of dampened inflammatory responses. By uncovering distinct immune adaptations, our results could help explain why pteropodids are frequently identified as viral hosts.

Potential Mammalian Vector-Borne Diseases in Live and Wet Markets in Indonesia and Myanmar

Vector-borne diseases spread from wild animals and their associated ectoparasites to humans and domesticated animals. Wildlife markets are recognized as important areas where this transfer can take place. We assessed the potential for spreading vector-borne diseases in two live and wet markets in Myanmar (Mong La, on the Myanmar-China border) and Indonesia (Sukahaji in Bandung on the island of Java) by making an inventory of all live and freshly killed wild mammals for sale. For eight mammal families, we quantified the number of animals on offer, and we used a heatmap cluster analysis to map vector-borne diseases that these families may carry. In Myanmar, we observed large numbers of wild pigs and deer (potentially carrying West Nile and various encephalitis viruses) whereas in Indonesia we observed Old World fruit bats (potentially carrying Chikungunya and encephalitis viruses) and squirrels (potentially carrying West Nile and encephalitis viruses). The trade in Indonesia was dominated by live mammals offered for sale as pets, and only Old World fruit bats and squirrels traded for traditional Asian medicine were killed in the markets. The trade in Myanmar was more geared towards wild meat (e.g., wild pigs, deer, primates) and traditional Asian medicine (squirrels). The combined risks of vector-borne diseases spreading from traded animals to human health highlight the need for an integrated approach protecting public health, economic interests and biodiversity.

Reproduction, seasonal morphology, and juvenile growth in three Malagasy fruit bats.

The island nation of Madagascar is home to three endemic species of Old World fruit bat in the family Pteropodidae: Pteropus rufus, Eidolon dupreanum, and Rousettus madagascariensis, all three of which are IUCN Red Listed under some category of threat. Delineation of seasonal limits in the reproductive calendar for threatened mammals can inform conservation efforts by clarifying parameters used in population viability models, as well as elucidate understanding of the mechanisms underpinning pathogen persistence in host populations. Here, we define the seasonal limits of a staggered annual birth pulse across the three species of endemic Madagascar fruit bat, known reservoirs for viruses of high zoonotic potential. Our field studies indicate that this annual birth pulse takes place in September/October for P. rufus, November for E. dupreanum, and December for R. madagascariensis in central-eastern Madagascar where the bulk of our research was concentrated. Juvenile development periods vary across the three Malagasy pteropodids, resulting in near-synchronous weaning of pups for all species in late January-February at the height of the fruiting season for this region. We here document the size range in morphological traits for the three Malagasy fruit bat species, with P. rufus and E. dupreanum among the larger of pteropodids globally and R. madagascariensis among the smaller. All three species demonstrate subtle sexual dimorphism with males being larger than females. We explore seasonal variation in adult body condition by comparing observed body mass with body mass predicted by forearm length, demonstrating that pregnant females add weight during staggered gestation periods and males lose weight during the nutritionally deficit Malagasy winter. Finally, we quantify forearm, tibia, and ear length growth rates in juvenile bats, demonstrating both faster growth and more protracted development times for P. rufus as compared with E. dupreanum and R. madagascariensis. The longer development period for the already-threatened P. rufus further undermines the conservation status of this species as human hunting is particularly detrimental to population viability during reproductive periods. Our work highlights the importance of longitudinal field studies in collecting critical data for mammalian conservation efforts and human public health alike.

First record of polydactyly for a European bat, Myotis daubentonii (Chiroptera, Vespertilionidae)

Abstract Despite being one of the most common congenital hand abnormalities in humans, polydactyly remains extremely rare in bats. To date, it has been observed only in two insectivorous families in North America (Molossidae and Vespertilionidae) and a single case for the New World fruit bats (Phyllostomidae). We report the first case of polydactyly in a European bat – a Myotis daubentonii (Vespertilionidae), with eight toes. The bat was found alive, hibernating in the Natura 2000 site (PLH080003 “Nietoperek”) in Poland.

Molecular convergence and transgenic evidence suggest a single origin of laryngeal echolocation in bats

SummaryThe laryngeal echolocation is regarded as one of the conspicuous traits that play major roles in flourishing bats. Whether the laryngeal echolocation in bats originated once, however, is still controversial. We here address this question by performing molecular convergence analyses between ancestral branches of bats and toothed whales. Compared with controls, the molecular convergences were enriched in hearing-related genes for the last common ancestor of bats (LCAB) and extant echolocating bats, but not for the LCA of Old World fruit bats (LCAP). And the convergent hearing gene prestin of the LCAB and the extant echolocating bats functionally converged. More importantly, the high-frequency hearing of the LCAP-prestin knock-in mice decreased with lower cochlear outer hair cell function compared with the LCAB-prestin knock-in mice. Together, our findings provide multiple lines of evidence suggesting a single origin of laryngeal echolocation in the LCAB and the subsequent loss in the LCAP.

Forbidden fruits? Ecosystem services from seed dispersal by fruit bats in the context of latent zoonotic risk

Old world fruit bats are important seed dispersers of forest plants as well as of commercial fruit crops. Bats scatter seeds across landscapes and also aggregate (clump) seeds under feeding and roosting trees. In agroforestry landscapes, bat frugivory and seed dispersal can result in simultaneous accrual of socio‐economic benefits and costs to humans, which may be further affected by human health risks from zoonotic spillover through human–bat interactions. In our study, we used an integrated approach to assess socio‐economic benefits (from seed dispersal) and costs (from frugivory) from bats, in relation to latent zoonotic risk. We carried out the study in five agroforestry landscapes along India's Western Ghats, for selected commercial fruit crops (cashew, areca, banana and other pulpy/fleshy fruits). Specifically, we hypothesized that people's perceptions of benefits from fruit bats would be correlated with measured extents of clumped seed dispersal in fruit plantations. We conducted ecological studies to investigate the effects of fruit crop type and plantation attributes on clump‐dispersal, and in turn, on perceived ‘net benefits' assessed from interviews with plantation‐associated people. We then explored associations between perceived socio‐economic net benefits and spatial data on disease risk factors for fruit bat‐linked henipavirus transmission. We found that the extent of bat‐mediated clump‐dispersal was highest for cashew, especially in isolated plantations. People's perceptions of benefits from fruit bats matched the measured extents of clump‐dispersal of cashew and areca. These benefits came with some costs from scatter‐dispersal and damage to other fleshy fruits from frugivory by bats. Interestingly, we did not detect tradeoffs between perceived net benefits from bats and disease risk, which is of significance for bat conservation and its implications for human well‐being. Overall, our results highlight that bat‐mediated seed dispersal needs to be sustained as an important ecosystem service, despite some latent zoonotic risk, in the Anthropocene

Geographic patterns in the diet of the Old World fruit bats: a review

Fruit bats of the family Pteropodidae are distributed in Africa, Asia, and Oceania, including many Pacific islands. Although known as good seed dispersers, there is no broader assessment of the diversity of the diet of the genera (44 to 46 recognized genera) across different localities. We analyzed the frugivorous diet of the family Pteropodidae through an extensive bibliographical review, using consumption records to document bat-fruit relations across the Old World and variation among the biogeographic realms proposed by Holt et al. (2013). Our review resulted in 239 studies and 2.597 records of 33 bat genera consuming 642 species of 320 genera from 104 angiosperms families. The most consumed fruit genus was Ficus, in almost all zoogeographic realms. However, bat-fruit relationships vary among realms with numerous links to regionally endemic plant species, suggesting a relation between the bats’ distribution and diversification of the associated flora. The genus Pteropus in particular, shows a strong geographic pattern in diet composition. Data of this nature help to visualize patterns in resource use among different regions of the globe and across diverse bat species assemblages and can direct further research on the evolution of pteropodid bats, their critical food resources, and their role in seed dispersal and ecosystem processes.

Animal‐mediated seed dispersal in India: Implications for conservation of India’s biodiversity

AbstractEffective seed dispersal is critical for forest regeneration and recruitment as well as the restoration of degraded lands. Most tropical tree species depend on frugivorous animals to disperse their seeds; frugivore‐mediated seed dispersal is thus crucial for biodiversity conservation in megadiverse tropical systems such as India. The fauna in India are increasingly threatened due to anthropogenic interventions such as deforestation and hunting. Thus, it is imperative to have an understanding of plant–frugivore interactions across the country to curb further biodiversity loss by ensuring the maintenance of these ecological processes. In this paper, I reviewed the literature on seed dispersal by animals in India to identify important groups that disperse many plant species and/or are the only recorded dispersers of certain plant species. Hornbills, bulbuls, bears, cattle, deer, civets, elephants, macaques, nilgais, Old World fruit bats, and wild pigs meet these criteria; overall 18 species were included in this list. Six of these species are currently “Threatened” on the IUCN Red List and their loss can lead to limited dispersal for at least 86 plant species. I suggest ensuring the protection of the important seed dispersers identified in this paper, irrespective of their conservation status, to prevent further biodiversity loss. The review reveals that plant–animal interactions in India are understudied and a major knowledge gap exists with regard to seed dispersal networks within the country.

The Critical Importance of Old World Fruit Bats for Healthy Ecosystems and Economies

Despite extensive documentation of the ecological and economic importance of Old World fruit bats (Chiroptera: Pteropodidae) and the many threats they face from humans, negative attitudes towards pteropodids have persisted, fuelled by perceptions of bats as being pests and undesirable neighbours. Such long-term negativity towards bats is now further exacerbated by more recent disease-related concerns, particularly associated with the current COVID-19 pandemic. There remains an urgent need to investigate and highlight the positive and beneficial aspects of bats across the Old World. While previous reviews have summarised these extensively, numerous new studies conducted over the last 36 years have provided further valuable data and insights which warrant an updated review. Here we synthesise research on pteropodid-plant interactions, comprising diet, ecological roles, and ecosystem services, conducted during 1985-2020. We uncovered a total of 311 studies covering 75 out of the known 201 pteropodid species (37%), conducted in 47 countries. The majority of studies documented diet (52% of all studies; 67 pteropodid species), followed by foraging movement (49%; 50 pteropodid species), with fewer studies directly investigating the roles played by pteropodids in seed dispersal (24%; 41 pteropodid species), pollination (14%; 19 pteropodid species), and conflict with fruit growers (12%; 11 pteropodid species). Pteropodids were recorded feeding on 1072 plant species from 493 genera and 148 families, with fruits comprising the majority of plant parts consumed, followed by flowers/nectar/pollen, leaves, and other miscellaneous parts. Sixteen pteropodid species have been confirmed to act as pollinators for a total of 21 plant species, and 29 pteropodid species have been confirmed to act as seed dispersers for a total of 311 plant species. Anthropogenic threats disrupting bat-plant interactions in the Old World include hunting, direct persecution, habitat loss/disturbance, invasive species, and climate change, leading to ecosystem-level repercussions. We identify notable research gaps and important research priorities to support conservation action for pteropodids.

Interrogating Phylogenetic Discordance Resolves Deep Splits in the Rapid Radiation of Old World Fruit Bats (Chiroptera: Pteropodidae).

The family Pteropodidae (Old World fruit bats) comprises n}{}>200 species distributed across the Old World tropics and subtropics. Most pteropodids feed on fruit, suggesting an early origin of frugivory, although several lineages have shifted to nectar-based diets. Pteropodids are of exceptional conservation concern with n}{}>50% of species considered threatened, yet the systematics of this group has long been debated, with uncertainty surrounding early splits attributed to an ancient rapid diversification. Resolving the relationships among the main pteropodid lineages is essential if we are to fully understand their evolutionary distinctiveness, and the extent to which these bats have transitioned to nectar-feeding. Here we generated orthologous sequences for n}{}>1400 nuclear protein-coding genes (2.8 million base pairs) across 114 species from 43 genera of Old World fruit bats (57% and 96% of extant species- and genus-level diversity, respectively), and combined phylogenomic inference with filtering by information content to resolve systematic relationships among the major lineages. Concatenation and coalescent-based methods recovered three distinct backbone topologies that were not able to be reconciled by filtering via phylogenetic information content. Concordance analysis and gene genealogy interrogation show that one topology is consistently the best supported, and that observed phylogenetic conflicts arise from both gene tree error and deep incomplete lineage sorting. In addition to resolving long-standing inconsistencies in the reported relationships among major lineages, we show that Old World fruit bats have likely undergone at least seven independent dietary transitions from frugivory to nectarivory. Finally, we use this phylogeny to identify and describe one new genus. [Chiroptera; coalescence; concordance; incomplete lineage sorting; nectar feeder; species tree; target enrichment.]

Explosive radiation at the origin of Old World fruit bats (Chiroptera, Pteropodidae)

Pteropodidae constitutes one of the most diverse bat families. These bats have evolved a phytophagous diet, likely lost laryngeal echolocation capability, and attained the largest body sizes among bats. Previous phylogenetic studies suggested that the family might have experienced an explosive diversification at its origin. Here, we readdress this hypothesis using a macroevolutionary approach based on Bayesian statistics (BAMM), a sampling of 139 pteropodid species, and divergence date estimates obtained in a comprehensive phylogenetic study of Chiroptera with multiple fossil calibration points. We evaluated the effect of missing data and of a reduced outgroup by repeating the analyses across simulated complete phylogenies and across a comprehensive Yinpterochiroptera phylogeny, respectively. Additionally, we performed an alternative analysis to detect diversification-rate shifts through time, the birth-death-shift method. In contrast with a previous study, we found strong statistical signals of rapid diversification at the origin of Pteropodidae. BAMM also detected diversification-rate shifts (increases) at the origin of Pteropus, as well as at crown Hipposideridae and Rhinolophidae. The birth-death-shift method detected a shift at approximately 25 million years ago, the estimated crown ages of both Pteropodidae and Hipposideridae. Our results point to a complex dynamics in the evolution of bat families, likely related to key innovations, demographic factors, and environmental opportunity enhanced by global-scale climatic and geographic changes.

Molecular adaptation and convergent evolution of frugivory in Old World and neotropical fruit bats.

Although cases of independent adaptation to the same dietary niche have been documented in mammalian ecology, the molecular correlates of such shifts are seldom known. Here, we used genomewide analyses of molecular evolution to examine two lineages of bats that, from an insectivorous ancestor, have both independently evolved obligate frugivory: the Old World family Pteropodidae and the neotropical subfamily Stenodermatinae. New genome assemblies from two neotropical fruit bats (Artibeus jamaicensis and Sturnira hondurensis) provide a framework for comparisons with Old World fruit bats. Comparative genomics of 10 bat species encompassing dietary diversity across the phylogeny revealed convergent molecular signatures of frugivory in both multigene family evolution and single-copy genes. Evidence for convergent molecular adaptations associated with frugivorous diets includes the composition of three subfamilies of olfactory receptor genes, losses of three bitter taste receptor genes, losses of two digestive enzyme genes and convergent amino acid substitutions in several metabolic genes. By identifying suites of adaptations associated with the convergent evolution of frugivory, our analyses both reveal the extent of molecular mechanisms under selection in dietary shifts and will facilitate future studies of molecular ecology in mammals.

The first complete mitochondrial genome data of Geoffroy's rousette, Rousettus amplexicaudatus originating from Malaysia

The increasing interest in understanding the evolutionary relationship between members of the Pteropodidae family has been greatly aided by genomic data from the Old World fruit bats. Here we present the complete mitogenome of Geoffroy’s rousette, Rousettus amplexicaudatus found in Peninsular Malaysia . The mitogenome constructed is 16,511bp in length containing 37 genes; 13 protein-coding genes (PCGs), 22 tRNA genes, two rRNA genes, and a D-loop region. The overall base composition is estimated to be 32.28% for A, 25.64% for T, 14.09% for G and 27.98% for C, indicating a slightly AT rich feature (57.93%). A phylogenetic and BLASTn analysis against other available mitogenomes showed Malaysian R. amplexicaudatus matched 98% similarity to the same species in Cambodia and Vietnam. However, it differed considerably (92.53% similarity) with the same species in the Philippines. This suggests flexibility in Rousettus sp. with regards to adapting to mesic and dry habitats, ability for long-distance dispersal and remarkably precise lingual echolocation thus supporting its wide-range distribution and colonization. Further taxonomical and mitogenomic comparatives are required in resolving the evolutionary relationship between Rousettus spp.

A Species-Level Phylogeny of Old World Fruit Bats with a New Higher-Level Classification of the Family Pteropodidae

Old World fruit bats (Chiroptera: Yinpterochiroptera: Pteropodidae) are a diverse radiation endemic to the tropics of Africa, Asia, Australia, and nearby island archipelagos. Recent molecular analyses have provided considerable resolution of phylogenetic relationships within this group, but many points of uncertainty have remained including the position of several enigmatic taxa (e.g., Notopteris, Eidolon), relationships among species in more diverse subfamilies and genera (e.g., Pteropodinae, Pteropus, Epomophorus), and topology of the backbone of the tree. Here we provide a new, synthetic analysis including representatives of all 45 currently recognized genera and enhanced sampling in several speciose genera. Our matrix included four nuclear genes regions (vWF, RAG1, RAG2, and BRCA1) and four mitochondrial gene loci (Cytb, tRNA valine, 12S rRNA, and 16S rRNA) for a total of >8000 bp including new sequence data for 13 species. Maximum likelihood and Bayesian analyses resulted in trees supporting recognition of six main suprageneric clades similar in content to those identified in our previous studies. We did not recover strong support for relationships among the main clades along the backbone of the tree, but identified many well-supported clades within all of the major groups. Based on these results, we propose a new classification for Pteropodidae comprising eight subfamilies and 14 tribes, and including several new and/or replacement higher-level taxonomic names for which we provide morphological diagnoses.