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ABOUT THIS BOOK With more than two hundred species distributed across most of mainland Mexico, Central and South America, and islands in the Caribbean Sea, the Phyllostomidae bat family (American leaf-nosed bats) is one of the world’s most diverse mammalian families in terms of its trophic, or feeding, diversity. From an insectivorous ancestry, extant species have evolved into several dietary classes, including blood-feeding, vertebrate carnivory, and the consumption of nectar, pollen, and fruit, in a period of about 30 million years. Phyllostomidae’s plant-visiting species are responsible for pollinating more than five hundred species of neotropical shrubs, trees, vines, and epiphytes—many of which are economically and ecologically important—and they also disperse the seeds of at least another five-hundred plant species. Fruit-eating and seed-dispersing members of this family thus play a crucial role in the regeneration of neotropical forests, and the fruit eaters are among the most abundant mammals in these habitats. Coauthored by leading experts in the field and synthesizing the latest advances in molecular biology and ecological methods, Phyllostomid Bats is the first overview in more than forty years of the evolution of the many morphological, behavioral, physiological, and ecological adaptations in this family. Featuring abundant illustrations as well as details on the current conservation status of phyllostomid species, it is both a comprehensive reference for these ecologically vital creatures and a fascinating exploration of the evolutionary process of adaptive radiation. AUTHOR BIOGRAPHY Theodore H. Fleming is professor emeritus of biology at the University of Miami, where he worked for thirty years. Among his recent books are Island Bats and The Ornaments of Life, both published by the University of Chicago Press, and No Species Is an Island. Liliana M. Dávalos is professor of conservation biology at Stony Brook University. She is coeditor of The Origins of Cocaine and coauthor of the 2016 World Drug Report. Marco A. R. Mello is professor of ecology at the University of São Paulo, Brazil. He served as the president of the Brazilian Bat Research Society and is the author, in Portuguese, of Sobrevivendo na Ciência: Um Pequeno Manual para a Jornada do Cientista.
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... Ancestrally, noctilionoids were predominantly insectivorous, and this dietary habit was retained in all families (Giannini and Velazco, 2020). However, the most extraordinary dietary diversification in all mammals occurred inside the Phyllostomidae; although gleaning insectivory remained as the chief dietary habit in some subfamilies, others evolved carnivory and omnivory (Phyllostominae), sanguivory (Desmodontinae), frugivory (Rhinophyllinae, Carollinae and Stenodermatinae), and nectarivory (Lonchophyllinae and Glossophaginae; Fleming, 2020). Moreover, the high diversification rate often reconstructed in phyllostomids has been related to the evolution of a plant-based diet (Rojas et al., 2012;Shi and Rabosky, 2015). ...
... Most of the diversification of the Noctilionoidea has taken place in the Neotropical region, the most diverse biogeographic region in the World, since the early-middle Eocene (Rojas et al., 2016;Amador et al., 2018). The speciation events that led to the Neotropical biodiversity have probably been promoted by several major geologic processes and interrelated climatic events (see Fleming, 2020). Orogeny in North America (Rocky Mountains and the Sierras Madre), Central America, and especially South America (the Andes) substantially modified the environment of the Neotropics, particularly during the Neogene (Graham, 2009;Hoorn et al., 2010). ...
... Other notable events are those related to the tectonics of Central America and the Caribbean, particularly the formation of the Panamanian land bridge at 3.5 Ma, which accelerated the Great American Biotic Interchange (GABI) between the North and South American continents after previous, sporadic events of biotic connections (Woodburne, 2010;Bacon et al., 2015). In the Caribbean, the Greater Antilles attained their current configuration about 25 Ma, except for volcanic Jamaica, with an age of 10 Ma (Fleming, 2020). In the case of the Lesser Antilles, the outer arc of volcanic islands dates from 40 to 45 Ma, while the inner arc dates from 25 to 20 Ma (Fleming, 2020). ...
Article
Biogeographic studies have generally relied on methods that use a few, large predefined areas, which may overlook fine‐scale patterns. Here we test previous hypotheses about the biogeographic history of a diverse bat clade regarding its association with major Neotropical geological formations, particularly the Antilles, the South American Dry Diagonal, the Andes and the Panamanian land bridge, by applying a recently available method that uses actual distributions instead of predefined areas. We compiled and curated spatially explicit, georeferenced data of 173 bat species (Mammalia: Chiroptera: Noctilionoidea) from the online database Global Biodiversity Information Facility. By taking a previous comprehensive phylogeny as an evolutionary framework, we performed computationally intensive analyses using the Geographically‐explicit Event Model. This method uses the observed species distributions to reconstruct the ancestral areas and biogeographic events at each phylogeny node. We found that sympatric speciation was the most frequently reconstructed event, and involved mainly the Panamanian Isthmus and northern South America (SA), but all sympatry reconstructions were different and specific to each node. Allopatric events were important in the Andes; vicariance caused both west/east and north/south disjunctions that went unnoticed previously. Founder events indicated bidirectional dispersal between the mainland and the Antilles since the Miocene, and across the incomplete Panamanian bridge and the SA Dry Diagonal since the early Pliocene. Overall, we found support for previous hypotheses on the influence of major Neotropical paleogeographic events in the diversification of the group, but additionally revealed multi‐scale patterns that are embedded within the mainland and were previously overlooked. Our results highlight a trans‐isthmian centre of diversification in the biogeographic history of Noctilionoidea including the Panamanian Isthmus and Northern SA.
... The Neotropical family Phyllostomidae, comprising 230 species distributed across 61 genera and 11 subfamilies (Simmons & Cirranello, 2024), represents one of the greatest adaptive radiations known among mammals (Fleming et al., 2020;Rossoni et al., 2017). This makes them the second most speciose bat family globally, with a distribution across the tropical and subtropical regions of the Americas. ...
... This makes them the second most speciose bat family globally, with a distribution across the tropical and subtropical regions of the Americas. Phyllostomid bats exhibit a wide range of feeding habits and ecomorphological characteristics, including insectivorous, carnivorous, hematophagous, nectarivorous, frugivorous, and omnivorous species, each employing a variety of strategies for food capture and showing different degrees of diet specialization (Fleming et al., 2020). The functional morphology of their wings varies significantly (Gunnell & Simmons, 2012;Norberg & Norberg, 2012), which correlates with their diverse flight methods and the habitats they utilize, facilitating their occupation of a broad range of ecological niches (Norberg & Rayner, 1987). ...
Article
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Extensive research into bat flight mechanisms has highlighted the complex functional and evolutionary dynamics of their wing structures, yet the anatomical details of certain wing muscles remain elusive. In particular, the intramembranous plagiopatagiales proprii muscles, located within the plagiopatagium—an area of the wing lacking direct joint connections—exhibit remarkable variation across bat families. These muscles, which extend anteroposteriorly in macroscopic bundles, play a crucial role in wing stiffening, modulating membrane tension, and reducing wing curvature during flight. Since larger bats tend to have higher wing loading (WL; the ratio of body mass [BMa] to wing area) and may therefore experience increased patagial curvature and resultant drag, we hypothesized that body size significantly influences the evolutionary development of the plagiopatagiales proprii muscles. This study investigates the relationship between BMa and the morphology of the plagiopatagiales proprii in New World leaf‐nosed bats (Phyllostomidae), employing bivariate allometry, multivariate analysis, and comparative phylogenetic methods across 24 species from eight phyllostomid subfamilies. Our findings reveal a significant phylogenetic signal in muscle architecture, along with positive evolutionary allometry in muscle area. This suggests an adaptive increase in muscle size in larger species, likely to counterbalance the increased WL, reduce wing curvature, and minimize drag. This research enhances our understanding of the functional and adaptive morphological evolution of intramembranous wing muscles in phyllostomid bats, underscoring their evolutionary significance.
... New Phytologist (2024) www.newphytologist.com (Rojas et al., 2016;Fleming et al., 2020). In fact, it is welldocumented that many phyllostomid bats specialize on mediumsized green and yellow fruits, including those of Neotropical solanums (Fleming, 1986;Lobova et al., 2009;Saldaña-V azquez & Fleming, 2020). ...
... Indeed, green-and yellow-fruited Solanum diversified in the mid-Miocene (c. 14 Ma), matching the evolutionary expansion of the predominantly frugivorous bats in the Carolliinae and Stenodermatinae subfamilies of Phyllostomidae (Rojas et al., 2016;Fleming et al., 2020). This time frame matching suggests that effective dispersal by these bats may have contributed to the further diversification of medium-sized green-and yellow-fruited Solanum species. ...
Article
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Mutualisms between plants and fruit‐eating animals were key to the radiation of angiosperms. Still, phylogenetic uncertainties limit our understanding of fleshy‐fruit evolution, as in the case of Solanum, a genus with remarkable fleshy‐fruit diversity, but with unresolved phylogenetic relationships. We used 1786 nuclear genes from 247 species, including 122 newly generated transcriptomes/genomes, to reconstruct the Solanum phylogeny and examine the tempo and mode of the evolution of fruit color and size. Our analysis resolved the backbone phylogeny of Solanum, providing high support for its clades. Our results pushed back the origin of Solanum to 53.1 million years ago (Ma), with most major clades diverging between 35 and 27 Ma. Evolution of Solanum fruit color and size revealed high levels of trait conservatism, where medium‐sized berries that remain green when ripe are the likely ancestral form. Our analyses revealed that fruit size and color are evolutionary correlated, where dull‐colored fruits are two times larger than black/purple and red fruits. We conclude that the strong phylogenetic conservatism shown in the color and size of Solanum fruits could limit the influences of fruit‐eating animals on fleshy‐fruit evolution. Our findings highlight the importance of phylogenetic constraints on the diversification of fleshy‐fruit functional traits.
... Habita principalmente en los claros de la vertiente caribeña, en zonas de bosque tropical, comúnmente en áreas de vegetación perennifolia, a alturas entre los 300 a 600 m.s.n.m. Ocasionalmente se observa en bosques de tipo caducifolio (Reid, 2009;Fleming et al., 2020). ...
Article
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Basados en características morfológicas específicas, presentamos el segundo registro de la especie Hylonycteris underwoodi Thomas, 1903 (Phyllostomidae: Glossophaginae) en la zona Este de la República de Panamá, presentando una nueva localidad de reporte como sitio de hábitat en el Istmo. Reporte que se obtiene por medio del proyecto de investigación sobre la biodiversidad de la Reserva Valle del Mamoní.
... Ecosystems like the tropics not only harbor a lot of species diversity but also patterns enabling species coexistence (Brown, 2014). Investigating species coexistence parameters like resource allocation, usage of similar microhabitats, prey-predator relationships, a wide range of diets, foraging strategies, and sensory abilities, etc., thus becomes interesting for morphologically similar and sympatric species (Fleming et al., 2020). ...
... Here we use developmental data to explore these questions in an ideal model clade, the bat super-family Noctilionoidea (Neotropical leaf-nosed bats and allies, more than 250 species; D avalos & Mello, 2020; Kunz & Fenton, 2003). In this hyperdiverse group, the most species-rich families are the Phyllostomidae (Neotropical leaf-nosed bats) which underwent an adaptive radiation and today exhibits great ecological richness (D avalos & Mello, 2020;Kunz & Fenton, 2003). Members of this clade have radiated into almost every mammalian ecological dietary niche (i.e., fruit, nectar and pollen, leaves, seeds, arthropods, small vertebrates, fish, and even blood). ...
Article
Sensory organs must develop alongside the skull within which they are largely encased, and this relationship can manifest as the skull constraining the organs, organs constraining the skull, or organs constraining one another in relative size. How this interplay between sensory organs and the developing skull plays out during the evolution of sensory diversity; however, remains unknown. Here, we examine the developmental sequence of the cochlea, the organ responsible for hearing and echolocation, in species with distinct diet and echolocation types within the ecologically diverse bat super‐family Noctilionoidea. We found the size and shape of the cochlea largely correlates with skull size, with exceptions of Pteronotus parnellii , whose high duty cycle echolocation (nearly constant emission of sound pulses during their echolocation process allowing for detailed information gathering, also called constant frequency echolocation) corresponds to a larger cochlear and basal turn, and Monophyllus redmani , a small‐bodied nectarivorous bat, for which interactions with other sensory organs restrict cochlea size. Our findings support the existence of developmental constraints, suggesting that both developmental and anatomical factors may act synergistically during the development of sensory systems in noctilionoid bats.
... Phyllostomidae can teach us much about the evolution of nectar consumption in bats. From insect-eating ancestors, Phyllostomidae (217 spp.) evolved to become an impressive mammalian adaptive radiation with a rich diversity in feeding morphology and behaviour (Fleming et al. 2020 Most often, we talk about nectar but ignore pollen, which as a protein source, may be closer to a proper insect food substitute than sweet nectar. Some studies even show pollen to be the most (14) important floral resource for some bat species. ...
Article
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Oceanic islands are relatively poor in insects compared to mainland areas. Therefore, insect-eating island birds and lizards may include other food sources into their diet, e.g. nectar and pollen. Here, we explore if insect-eating island bats face a similar problem and accordingly join the birds and lizards and incorporate plant resources into their diet. Thus, a priory, we assume flower visitation by bats to be more common on oceanic islands than elsewhere. To test this, we reviewed the literature to obtain information on the geographic distribution and diet of all 1,399 species of bats in the world and found that 49%, 21%, and 31% of species have a mainland, mixed mainland-island, and island distribution, respectively. Diets are known for only 65% (905 species) of the bats in the world, and 70%, 22%, and 8% of these, respectively, rely on insects, fruit, and floral resources as their major diet component. Twenty-seven species are even obligate flower visitors. This study confirms that flower-visiting bats, especially Pteropodidae, are significantly more frequent on oceanic islands, while insect eaters are more frequent on mainland and continental islands. Consequently, we argue that flower visitation and pollination by insect-eating island bats require more attention in future island ecology studies. For a start, we list known examples in the literature and report a case study from the Canary Islands. In the latter, we examined the foreheads of 34 museum specimens of the seven Canarian bat species. Half of them carried pollen from ≥ 9 taxa, but only three bat species had larger amounts. Pollen was not identified, but many Canarian and exotic plant species are candidates. Thus, flower visitation by bats may be an oceanic island phenomenon, but requires more focused research, especially night-time flower observations and examination of bats for pollen.
Article
The most significant driver of adaptive radiation in the New World leaf‐nosed bats (Phyllostomidae) is their remarkably diverse feeding habits, yet there remains a notable scarcity of studies addressing the genetic underpinnings of dietary diversification in this family. In this study, we have assembled a new genome for a representative species of phyllostomid bat, the fringe‐lipped bat ( Trachops cirrhosis ), and integrated it with eight published phyllostomid genomes, along with an additional 10 genomes of other bat species. Comparative genomic analysis across 10 200 orthologus genes has unveiled that those genes subject to divergent selection within the Phyllostomidae clade are notably enriched in metabolism‐related pathways. Furthermore, we identified molecular signatures of divergent selection in the bitter receptor gene Tas2r1 , as well as 14 genes involved in digesting key nutrients such as carbohydrates, proteins, and fats. In addition, our cell‐based functional assays conducted on Tas2r1 showed a broader spectrum of perception for bitter compounds in phyllostomids compared to nonphyllostomid bats, suggesting functional diversification of bitter taste in Phyllostomidae. Together, our genomic and functional analyses lead us to propose that divergent selection of genes associated with taste, digestion and absorption, and metabolism assumes a pivotal role in steering the extreme dietary diversification in Phyllostomidae. This study not only illuminates the genetic mechanisms underlying dietary adaptations in Phyllostomidae bats but also enhances our understanding of their extraordinary adaptive radiation.
Article
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Fossils of an insectivorous bat from the early Miocene of Panama are described as a new genus and species, Americanycteris cyrtodon (Chiroptera: Phyllostomidae: Phyllostominae). Americanycteris is a large phyllostomine bat, similar in size to the living species Chrotopterus auritus. Americanycteris cyrtodon can be distinguished from other closely related species by a posteriorly curved p4 and a thick labial cingulum on m1. Americanycteris cyrtodon occurs in two early Miocene vertebrate faunas from Panama. The holotype mandible with p4–m1 and an isolated p3 of A. cyrtodon were recovered from the early Hemingfordian (19–18 Ma) Centenario Fauna, and a mandible with p2 was found in the older late Arikareean (21 Ma) Lirio Norte Local Fauna. A similar large phyllostomine bat is known from the early Miocene Gran Barranca Fauna in Argentina. The presence of early Miocene phyllostomids in both North America and South America confirms the overwater dispersal of bats between the Americas before the late Miocene onset of the Great American Biotic Interchange. Pre-late Miocene chiropteran dispersals between the Americas were previously documented for the Emballonuridae and Molossidae. Although the five endemic New World families in the Noctilionoidea, including Phyllostomidae, were previously thought to be South American in origin, the oldest fossil records of noctilionoids (Mormoopidae and extinct Speonycteridae) are from the early Oligocene of Florida and one of the earliest records of the Phyllostomidae is from the early Miocene of Panama. The currently available fossil records from Panama and Florida suggest a possible North American origin for the Noctilionoidea.
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