Phyllostomid Bats A Unique Mammalian Radiation
edited by Theodore H. Fleming, Liliana M. Dávalos and Marco A. R. Mello
University of Chicago Press, 2020
Cloth: 978-0-226-69612-6 | Electronic: 978-0-226-69626-3
DOI: 10.7208/chicago/9780226696263.001.0001
ABOUT THIS BOOKAUTHOR BIOGRAPHYREVIEWSTABLE OF CONTENTS

ABOUT THIS BOOK

With more than two hundred species distributed from California through Texas and 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. From an insectivorous ancestor, species living today, over about 30 million years, have evolved a hyper-diverse range of diets, from blood or small vertebrates, to consuming nectar, pollen, and fruit. Phyllostomid 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.

REVIEWS

Phyllostomid Bats: A Unique Mammalian Radiation (edited by Theodore H. Fleming, Liliana M. Dávalos, and Marco A. R. Mello) is an outstanding book. The diversity of detailed information is exceptional. The book’s twenty-four chapters provide an in-depth look at these astonishing bats. The book will be an exceptional addition to the libraries of bat biologists, evolutionary biologists, and those who are intrigued by diversity. I am glad that I have already ordered my copy.”
— M. Brock Fenton, Department of Biology, University of Western Ontario (Canada), coeditor of “Bat Ecology” and coauthor of “Bats: A World of Science and Mystery”

Phyllostomid Bats is an excellent, comprehensive compendium of the latest research on this fascinating group. The book reflects how this incredibly diverse clade can be a model system for advancing our understanding of fundamental aspects of evolution and ecology of not just bats, but also of other organisms. The language is accessible to both budding biologists and veteran researchers looking to dip their toes into a new field, and it will surely be an indispensable volume to bat researchers everywhere.”
— Susan M. Tsang, research associate, Department of Mammalogy, American Museum of Natural History

“These are timely and eagerly anticipated reviews of the huge volume of research carried out in recent decades on this diverse and fascinating family of bats.”
— Paul Racey, Regius Professor of Natural History (Emeritus), University of Aberdeen, Scotland

Phyllostomid Bats is certainly worthwhile, as we have learned a good deal about this family in the last forty years or so (since the previous review of the whole family). From interesting speculation on the correlation between sexual dimorphism and mating systems in bats to a very nice summary of conservation threats and more importantly, perhaps, a good overview of some of the actions being taken in response, this book will serve the bat community well, and will be used by anyone studying this family in the future.”
— Don E. Wilson, curator emeritus of vertebrate zoology, National Museum of Natural History, Smithsonian Institution

"When reading a book like this magnificent new volume from the University of Chicago Press, Phyllostomid Bats: A Unique Mammalian Radiation, it’s important to understand the usage in the title: the radiation being discussed here is out-filing and endless tiny, fussy adaptations that a group of species can make to a variety of ecological and behavioral niches. As the book’s editors point out right away, the marquee example of adaptive radiation will probably always be Darwin’s famous finches, but in any case of animal adaptation, bats, specifically microchiroptera (the little bats, not the big flying foxes), some of the most successful creatures in evolutionary story, will always merit extra attention. . . . Phyllostomid Bats, although abstruse and nerdy in the extreme, is a wonderful intaglio of the sheer ferocious inventiveness of evolution. These leaf-nosed bats have radiated out to fill and exploit every conceivable niche where food and breeding might be found; in their endless forms—and in the wearily predictable ways they’re threatened in the twenty-first century—they are, among other things, fascinating bellwethers. And readers who might have a bat aficionado on their gift-recipient list? Well, their course is clear this book-season."
— Open Letters Review

Phyllostomid Bats is a well-referenced and scholarly text, coauthored by a selection of experts in the field. Well-written, generally engaging, and . . . largely accessible to more general readers, it affords an intriguing discussion of evolutionary biology and ecology, providing a fascinating glimpse into the lives of this unique group of animals. This comprehensive, up-to-date reference work will be of considerable interest to evolutionary biologists, ecologists, conservationists, and bat biologists, as well as to lay readers with more than just a passing interest in the natural history and biology of bats.” 
— NSS News

"This recently published book on phyllostomids provides a comprehensive compendium on various aspects of this family and does not disappoint. It is a well-edited volume with twenty-four chapters written by eminent specialists delivering a notable work in the field of bat biology. Most impressive is how authors integrate natural history with experimental biology thereby broadening the perspective of readers and stimulating new research hypotheses. . . . Although I could only mention a few aspects of this excellent volume in this short review, there are many more facets that readers will enjoy. This book is a worthy purchase and I commend the many authors who each provided new insights into the biology of this fascinating group of mammals."
— Quarterly Review of Biology

"Over two-hundred species of leaf-nosed bats range from the southern United States to northern Argentina and feature a wide range of diets (insects, vertebrates, blood, pollen, nectar, and fruit). This diverse family of bats demonstrates adaptive radiation patterns similar to those found among Darwin’s finches and Australian marsupials. Fleming, Dávalos, and Mello edited this nicely produced scholarly work, which features contributions from over forty contributing authors. Twenty-four chapters cover phylogeny and evolution, general biology, feeding habits, ecology, and conservation status. This volume comprehensively updates previous work on this family of bats (the most recent overview having been published during the 1970s), with new scholarship focusing especially on phylogeny. It is a fairly technical volume but will still be accessible to general readers interested in these bats. The book is illustrated by figures and tables appearing throughout the text, along with a section devoted to color plates. References at the conclusion of each chapter include literature published during the past two decades. Subject and taxonomic indexes complete the volume. This work would be a suitable addition to all college biology collections. Recommended."
— Choice

TABLE OF CONTENTS

- Theodore H. Fleming, Liliana M. Dávalos, Marco A. R. Mello
DOI: 10.7208/chicago/9780226696263.003.0001
[adaptive radiation;biological diversity;Phyllostomidae]
This book discusses in detail the adaptive radiation of American leaf-nosed bats (Phyllostomidae), the most diverse family morphologically, behaviorally, and trophically of all bats. It is divided into five major sections that cover the family’s evolution, classification, and historical biogeography; many aspects of its basic biology; its trophic, population, and community ecology; and its conservation. Because of its biological diversity, this family can serve as a model system for understanding the adaptive radiation of mammals in the last half of the Cenozoic Era. (pages 3 - 6)
This chapter is available at:
    University of Chicago Press
    https://academic.oup.com/chica...

- Theodore H. Fleming
DOI: 10.7208/chicago/9780226696263.003.0002
[climatic changes;coevolution;frugivory;gleaning insectivory;geological changes;nectarivory;sanguinivory]
I review changes in the climate, geology, and biota of the New World tropics and subtropics over the past 30 million years to understand the physical and biological opportunities and constraints phyllostomid bats faced during their adaptive radiation. This radiation occurred during a period of decreasing atmospheric CO2 levels, decreasing global air and sea temperatures, and increasing climatic seasonality that have led to the evolution of a great diversity of vegetation types, including tropical dry forests, montane forests, savannas, and deserts. Reinforcing the effects of temperature changes was the uplift of the major mountains in western North America, Central America, and western South America. Most of the extant diversity of New World noctilionoids is found on the Neotropical mainland in lowland forested habitats, which have existed in one form or another throughout the Americas since the late Cretaceous. A minority of species has evolved in more recent dry or arid habitats, in the mountains, or in the West Indies. Early phyllostomids were gleaning insectivores whose diets differed from those of other families of insectivorous bats. (pages 7 - 22)
This chapter is available at:
    University of Chicago Press
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- Norberto P. Giannini, Paúl M. Velazco
DOI: 10.7208/chicago/9780226696263.003.0003
[phylogeny;fossils;Gondwana;behavioral ecology;echolocation]
The superfamily Noctilionoidea comprises more than 230 species in 68 genera and is one of the most ecologically diverse groups of mammals, as its species consume almost the entire dietary spectrum known for terrestrial mammals, including insectivorous, omnivorous, carnivorous, piscivorous, nectarivorous, hematopagous, and frugivorous species. Here we 1) revisit the relationships of the superfamily with other groups of bats, and the family-level phylogenetic relationships within the group; 2) review the historical biogeography of the superfamily on the basis of phylogenies and the fossil record of the constituent noctilionoid families; and 3) provide an overview of the ecology of members of noctilionoid families, which collectively compose the most inclusive outgroup to the Phyllostomidae. Finally, we interpret the historical and ecological grounds and evolutionary context from which the Phyllostomidae diversified. (pages 25 - 42)
This chapter is available at:
    University of Chicago Press
    https://academic.oup.com/chica...

- Andrea L. Cirranello, Nancy B. Simmons
DOI: 10.7208/chicago/9780226696263.003.0004
[Phyllostomidae;species description;classification;taxonomic discovery]
Taxonomy is central to modern biology. Although many researchers assume that most mammal taxa are already known—and have been for decades—this is not necessarily true for small mammals including bats. Examination of patterns of taxonomic discovery over the last 200+ years reveals that we are currently in a taxonomic “Golden Age” for phyllostomid bats with more than 40 species discovered and described since 2001. Although the pace of discovery was slow throughout the 1700s, by 1850 representatives from 10 of the 11 subfamilies that we recognize today had been described, 22 genera had been named, and 39 phyllostomid species had been identified as specimen collections were amassed in European institutions. However, it was not until the early 1900s that a peak in descriptions occurred, with more than 20 species still considered valid described in the decade between 1901-1910. Most taxa described in the 19th and 20th centuries were described by scientists based in Europe, but research activity and descriptions of new taxa shifted to North American institutions between 1950 and 2000. Since 2001, the majority of published species descriptions have been authored or co-authored by researchers from Central and South American institutions. (pages 43 - 62)
This chapter is available at:
    University of Chicago Press
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- Nancy B. Simmons, Gregg F. Gunnell, Nicolas J. Czaplewski
DOI: 10.7208/chicago/9780226696263.003.0005
[diversification;extinction;fossils;Miocene;Paleogene;Phyllostomidae;Pleistocene;Pliocene;Holocene]
While modern molecular phylogenetic studies can provide a framework for understanding relationships of phyllostomids to one another and their close relatives, it is only the fossil record that can provide calibration points for reconstructing the temporal pattern of diversification. Additionally, the fossil record can facilitate synthetic understanding of patterns of morphological change over time. The fossil record of Phyllostomidae definitely extends to the early Miocene (~ 21-19 Ma) of Panama, and may extend as far back as the late middle Eocene (~41 Ma) of Peru, but it is surprisingly incomplete before the Pleistocene. Crown group members of the extant subfamilies Phyllostominae and Lonchophyllinae are known from Miocene localities in the Neotropics. Most Miocene phyllostomids exhibit dental morphology and body sizes suggesting that they were insectivores, but some may have been at least partially carnivorous or omnivorous, indicating that ecological diversification in the family was well underway in the Miocene. Large numbers of phyllostomid fossils are known from Pleistocene and Holocene deposits in Central America, South America, and the Caribbean, but no Late Miocene or Pliocene fossil phyllostomids are known, hence there is a ~10 million year gap in the fossil record. (pages 63 - 86)
This chapter is available at:
    University of Chicago Press
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- Liliana M. Dávalos, Paúl M. Velazco, Danny Rojas
DOI: 10.7208/chicago/9780226696263.003.0006
[Andes;BioGeoBEARS;dispersal;diversification;Great American Interchange;Lonchorhina;Micronycterinae;Neotropics]
Traditionally, the phylogeny of phyllostomids contained a few subfamilies sharing similar cranial and ecological traits. Family-wide genetic sequence data starting in the early 2000s, however, showed that this phylogeny obscured the taxonomy, biogeography, and evolution of ecological traits in the family. Current phylogenies strongly support the monophyly of 11 subfamilies, as well as most relationships among them. Nonetheless, the positions of several key groups are not well-supported and require further analyses, including: 1) Micronycterinae relative to Desmodontinae, 2) Lonchorhininae, and 3) the genus-level resolution of Miocene fossils. Historical biogeographic analyses have revealed a South American origin for most subfamilies; have inferred widespread and multi-continental distributions for ancient ancestors; and have rejected a special influence of Quaternary glaciations on recent speciation. These analyses have also confirmed North and Central America, as well as South America, as independent centers of diversification for some subfamilies (e.g., Macrotinae), as well as within subfamilies (e.g., Glossophaginae). These parallel centers of diversity might result from the absence of continuous dry habitat corridors between the continents, but this hypothesis has not been tested using phyloclimatic methods. (pages 87 - 104)
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    University of Chicago Press
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- Liliana M. Dávalos, Andrea L. Cirranello, Elizabeth R. Dumont, Stephen J. Rossiter, Danny Rojas
DOI: 10.7208/chicago/9780226696263.003.0007
[adaptation;dietary specialization;diversification;ecological theory of adaptive radiation;Ornstein-Uhlenbeck models;trophic level]
A diversity of cranial phenotypes and feeding ecologies characterizes phyllostomid bats. However, many subfamilies share a similar insect-feeding morphotype, apparently stable over millions of generations. Is the diversity of phyllostomid traits adaptive, and does it contribute to species diversity in the family? We evaluate predictions from Simpson’s ecological theory of adaptive radiation, applied to comparative trait analyses since 2000. The shift to higher rates of taxonomic diversification at the base of Stenodermatinae, new skull architecture capable of great bite force despite small size, and a distinctive olfactory receptor profile make this subfamily a radiation by itself. The stenodermatine skull architecture also enables making leaf tents, a new roost adaptive zone that contributes to the diversity of this subfamily. The evolution of distinct trophic adaptive zones characterized by optima in cranial musculature and mechanical advantage shows that stenodermatines are not the only phyllostomid lineage responding to ecological opportunities. Instead, nectarivores have unique cranial traits and occupy their own trophic adaptive zone, while a subset of insectivores together with sanguivores share a similarly high trophic level. There's evidence of convergent and adaptive evolution in cranial traits associated with similar trophic levels, and adaptations coupled with dietary specialization in digestive and renal function. (pages 105 - 122)
This chapter is available at:
    University of Chicago Press
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- Norberto P. Giannini, Lucila I. Amador, R. Leticia Moyers Arévalo
DOI: 10.7208/chicago/9780226696263.003.0008
[body size;continuous character optimization;phylogenetic constraints;dietary shifts;phyletic trends;past character displacement]
Body mass is the single most important factor affecting the biology of animals. Small size is generally highly favorable for flying vertebrates; in addition to this constraint, echolocating bats are also restricted by the physics of call parameters. However, we find in a single group, the Noctilionoidea, virtually the entire size range found across all echolocating bats. Here we examine the evolutionary significance of this variation. We explore how the reconstructed body mass of the ancestral phyllostomid was inherited from noctilionoid ancestors, how size changed along the branches of the phyllostomid tree, and how size co-varied with characteristic evolutionary shifts in the ecology of phyllostomid bats. We found little change along the backbone of the phylogenetic tree and across major dietary transitions, many scattered increases and decreases of variable magnitude, and most variation concentrated in phyletic change in a few groups, especially vertebrate-specialized and frugivorous phyllostomids. These trends imply sustained selection acting over millions of years in a consistent direction, principally towards an increase in size. Initial stasis may have facilitated ecological transitions, while capacity for size change may have fueled intense directional selection within highly specialized lineages. Finally, diverging trends appear to reflect past character displacement. (pages 123 - 148)
This chapter is available at:
    University of Chicago Press
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- Sharon M. Swartz, Justine J. Allen
DOI: 10.7208/chicago/9780226696263.003.0009
[aerodynamics;biomechanics;elastin;flight;musculoskeletal;sensory;skin;wings;wing;morphology]
Understanding the evolution of bat flight behavior and explaining the diversity in the structure of their wings requires examination of the functional organization and mechanical properties of wing tissues. In-depth structural analyses that consider multiple levels of organization can complement phylogenetically based comparative analyses, even if they do not sample a large number of taxa. In this chapter, we discuss recent research on the functional biology of bat flight and wing structure that has used phyllostomids as study taxa, with attention to skin morphology, muscle function kinematics, and aerodynamics. We discuss how understanding the hierarchical structure of the architecture of wing tissues, particularly intramembranous muscle, elastin fiber bundles, and collagenous structures, can provide a window into the functional dynamics of phyllostomid wings. We offer thoughts concerning differences in wing structure and flight capabilities among phyllostomids and other groups of bats, and propose future research to explore these subjects in greater depth. (pages 151 - 168)
This chapter is available at:
    University of Chicago Press
    https://academic.oup.com/chica...

- Ariovaldo P. Cruz-Neto, L. Gerardo Herrera M.
DOI: 10.7208/chicago/9780226696263.003.0010
[digestive physiology;excretory physiology;intestinal enzymes;kidney structure;metabolic physiology;oxidative metabolism;basal metabolic rate]
Dietary diversification is one of the main drivers of physiological diversification in Phyllostomidae. We analyzed the extent to which dietary diversification in phyllostomids is paralleled by diversification in whole-organism physiological traits that are essential to understanding how they interact with their environment. We focused our analysis on the interplay between diet and digestive,excretory, and metabolic physiology. We used a strict phylogenetic approach to test the relationship of these traits with diet when sufficient information for phyllostomids representative of different feeding habits was available: activity of intestinal enzymes, kidney morphology, urine concentration, and basal metabolic rate. For traits for which data are scarce or restricted to some trophic guilds, we reviewed the information available in the literature. Our analysis reveals that certain physiological traits appear to be associated with diet in an evolutionary and ecological context. Other traits seem not to be related to diet. Besides the difficulties associated with diet quantification, one potential problem with comparative analyses of the association between diet and physiological traits in phyllostomids is the paucity of data for animalivores. Future comparative work should include under-represented dietary habits and consider the use of methodological approaches currently used in the study of other vertebrates. (pages 169 - 186)
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    University of Chicago Press
    https://academic.oup.com/chica...

- Jeneni Thiagavel, Signe Brinkløv, Inga Geipel, John M. Ratcliffe
DOI: 10.7208/chicago/9780226696263.003.0011
[foraging ecology;diet selection;sensory systems;brain evolution;neuroanatomy]
Bats (order Chiroptera) exhibit wide-ranging differences in foraging ecology, morphology and behavior that often reflect the demands on their sensory systems. New World leaf-nosed bats (family Phyllostomidae) have a wide spectrum of feeding ecologies and sensory system specializations. The family consists of bats that are primarily nectarivorous (e.g., subfamily Glossophaginae), frugivorous (e.g., Stenodermatinae, Carolliinae), sanguivorous (Desmodontinae), and predatory (Phyllostominae). Phyllostomid brains typically have more balanced visual, olfactory, and auditory regions in relative size compared with other bat families. Within phyllostomid subfamilies, relative brain region volumes reflect feeding ecology and corresponding sensory specializations. For instance, phytophagous phyllostomids have larger visual and olfactory regions relative to predatory species, which in turn have larger auditory centers. This chapter uses this bat family to illustrate the influences that foraging ecology and diet selection have on the evolution of sensory systems and relative brain and brain region volumes. The diversity within this family makes it an excellent model group among bats—and mammals in general—from which to better understand sensory specializations, cognitive development, and brain evolution. (pages 187 - 204)
This chapter is available at:
    University of Chicago Press
    https://academic.oup.com/chica...

- Robert M. R. Barclay, Theodore H. Fleming
DOI: 10.7208/chicago/9780226696263.003.0012
[body size;diet;life history theory;longevity;monestrus;polyestrus;size at birth]
Using data from the primary literature for 71 species, we summarize reproductive and life history variation in this family and test predictions based on life history theory. The expectation for variation is at least partly supported. Monestry, seasonal (bimodal) polyestry, and aseasonal polyestry occur, with larger species, island species or populations, and animalivorous species being most likely to be monestrus. Litter size is universally one, however. As in other bats, phyllostomids have a “slow” life history with late maturity, one offspring per litter, and long lives for their size. Females produce relatively large offspring at birth (mean = 28% of adult mass), although relative size declines with adult mass. Young are dependent on their mother until they are almost full size at the time of weaning. Unfortunately, there is a paucity of data for many other life history traits and this precludes in-depth analyses of patterns and factors associated with variation. Many interesting evolutionary questions remain to be addressed and the variation in other aspects of the family make phyllostomids ideal subjects to test general questions regarding the evolution of reproductive and life history traits of mammals. (pages 205 - 220)
This chapter is available at:
    University of Chicago Press
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- Danielle M. Adams, Christopher Nicolay, Gerald S. Wilkinson
DOI: 10.7208/chicago/9780226696263.003.0013
[canines;female choice;male competition;sexual selection;sperm competition]
Of the 216 species of phyllostomid bats, fewer than 10% have had their mating systems studied in any great detail; however, some species exhibit mating systems ranging from apparent monogamy to extreme polygyny. Paternity studies reveal that the social mating system is generally indicative of the genetic mating system, although in some cases, subordinate males father some offspring. These findings suggest that mate selection can involve both male competition and female choice. To estimate the strength of precopulatory and postcopulatory sexual selection, we use measures of sexual dimorphism in relative body mass and canine length as indicators of direct male competition, and relative testes mass as a proxy for sperm competition. We then evaluate the influence of aggregation size and permanence of the roosting structure on the intensity of sexual selection using phylogenetically-informed analyses. Even though females are often larger than males, male-biased sexual dimorphism for relative mass and canine length is widespread and associated with large roosting aggregations. In contrast, sperm competition is greatest in species with intermediate-sized aggregations. These patterns of sexual dimorphism are largely consistent with what is known about phyllostomid mating systems, but exceptions provide potential opportunities for future study. (pages 221 - 236)
This chapter is available at:
    University of Chicago Press
    https://academic.oup.com/chica...

- Claire T. Hemingway, M. May Dixon, Rachel A. Page
DOI: 10.7208/chicago/9780226696263.003.0014
[animalivorous;foraging strategy;generalist;Phyllostominae;predator]
Phyllostomid bats are known for their spectacular radiation into distinct dietary niches. But many phyllostomid species actually have quite generalized diets. Historically, these generalist phyllostomids fell within the subfamily Phyllostominae, a diverse group that primarily hunts insects and small vertebrates such as frogs, lizards, birds, rodents, and other species of bats. This now paraphyletic group consists of over 30 species. While this group is characterized by predators with broad, generalized diets, many of these bats have distinct, specialized foraging strategies. We discuss the apparent paradox of these generalist predators: bats that can be classified as dietary generalists, broadly overlapping with one another in the prey they consume, but often possessing quite specialized foraging adaptations and distinct behavioral strategies for prey finding. To untangle the puzzle of this understudied and historically elusive group of generalist predators, we examine the influences of diet breadth on hunting behavior, spatial distribution, conservation concern, and morphology. (pages 239 - 256)
This chapter is available at:
    University of Chicago Press
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- John W. Hermanson, Gerald G. Carter
DOI: 10.7208/chicago/9780226696263.003.0015
[adaptation;cooperation;Desmodus;Diaemus;Diphylla;morphology]
The common vampire bat Desmodus rotundus, white-winged vampire bat Diaemus youngi, and hairy-legged vampire bat Diphylla ecaudata are the only mammals that feed exclusively on blood. Their blood-feeding lifestyle shapes virtually every aspect of their biology. Adaptations linked to blood-feeding include changes to morphology (e.g. dramatically reduced dentition, unique terrestrial and arboreal locomotion), physiology (e.g. anticoagulants in their saliva, an ability to rapidly eliminate water from blood meals), sensory ecology (e.g. infrared thermoperception, specialized low-frequency hearing), and social behavior (cooperative food sharing). There is even evidence for an adaptive loss of a cognitive trait: vampire bats lack the otherwise ubiquitous phenomenon of taste aversion learning, presumably because blood from live prey is never spoiled. From an evolutionary perspective, the traits that vampire bats have both gained and lost illustrate adaptive tradeoffs and the link between form and function. Vampire bats are the primary reservoir for bovine rabies virus in Central and South America, so understanding their biology has implications for agricultural development and public health. (pages 257 - 272)
This chapter is available at:
    University of Chicago Press
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- Nathan Muchhala, Marco Tschapka
DOI: 10.7208/chicago/9780226696263.003.0016
[bat pollination;chiropterophily;nectarivory;New World;specialization]
One-quarter (56) of the species of Phyllostomidae have adapted to a primarily nectarivorous diet and serve as pollinators for hundreds of species of flowering plants. Here we review the ecology and evolution of these bats and the flowers that rely on them. A suite of adaptations allows them to feed from flowers efficiently, including elongated rostrums, long, extensible tongues, reduced dentition, well-developed olfaction and spatial memory, and the ability to hover. These adaptations evolved independently in two phyllostomid subfamilies, the Lonchophyllinae and the Glossophaginae, which differ profoundly in their tongue morphology and nectar-feeding behavior: glossophagines have mop-like tongues with papillae covering the distal tip and lap nectar during flower visits, while lonchophyllines have straw-like tongues with papillae-lined grooves along the side and pull nectar through these grooves while keeping the tongue immersed in the nectar. Flowers adapted to bat pollination tend to have dull colors, strong scents, copious pollen and nectar production, and particularly well-exposed flowers. Despite heavy reliance on these flowers, nectar bats will often feed opportunistically on insects and fruit as well, particularly during times of the year when flowers are scarce. However, dietary data are still lacking for many species. (pages 273 - 294)
This chapter is available at:
    University of Chicago Press
    https://academic.oup.com/chica...

- Romeo A. Saldaña-Vázquez, Theodore H. Fleming
DOI: 10.7208/chicago/9780226696263.003.0017
[Carolliinae;climate change;foraging behavior;frugivory;seed dispersal;Stenodermatinae]
Frugivory is a hallmark of the Phyllostomidae, and frugivorous species in subfamilies Carolliinae and Stenodermatinae are among the most common mammals throughout the Neotropics. In this chapter we discuss the coevolution of fruits and phyllostomid frugivores; the morphological and other traits that these bats use to detect and process fruit; their foraging behavior and its consequences for seed dispersal; and the network structure of this coevolved mutualism and its conservation implications. Frugivorous phyllostomids have been interacting with fleshy-fruited angiosperms for about 20 Ma and currently disperse the seeds of hundreds of species of plants found throughout angiosperm phylogeny. They are especially important dispersers of early successional plants. Collectively, the feeding and foraging behavior of these bats creates strongly leptokurtic seed dispersal curves, but whose long tails can sometimes lead to the colonization of new habitats. Two foraging guilds (or modules in network parlance) have evolved: an understory guild of carolliines and Sturnira bats and a canopy guild of mostly stenodermatine bats. Ecological redundancy likely exists within these two guilds, but it is threatened whenever bat diversity is reduced by natural or anthropogenic factors. Climate change is one such factor, and its effects warrant careful monitoring. (pages 295 - 308)
This chapter is available at:
    University of Chicago Press
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- Armando Rodríguez-Durán
DOI: 10.7208/chicago/9780226696263.003.0018
[bat ecology;Noctilionoidea;Phyllostomidae;roosts;roosting behavior]
In this chapter I describe the complexity of roosts and their importance to understanding the ecology of phyllostomid bats and related noctilionoids, with an emphasis on the relationships between roosts, sociality, energetics, and species distribution. Detailed description of the roosts of phyllostomids is often neglected, due in part to the misconception that factors such as thermal stress are unimportant in tropical ecosystems. Although a number of systems for classifying roosts have been developed in the past, these schemes are in need of revision, as our knowledge about the ecology of bats has increased. I present a revised classification of roost types and identify gaps in our knowledge. In some respects this system of classification remains inevitably subjective, pending the accumulation of more detailed data. New directions for the characterization of roosts are suggested to allow an improved description of bat roosting ecology. (pages 311 - 324)
This chapter is available at:
    University of Chicago Press
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- Theodore H. Fleming, Angela M. G. Martino
DOI: 10.7208/chicago/9780226696263.003.0019
[demography;genetic subdivision;genetic variation;population sizes;relative abundance]
Many phyllostomid species, especially frugivores in subfamilies Carolliinae and Stenodermatinae, are common members of neotropical mammal communities. Population sizes of these bats reflect the size of their resource base—large for fruit and much smaller for nectar and animal prey. Colony sizes are correlated with the size of their roost structures: larger in cave or cave-like roosts—the most common roost type in these bats—and much smaller in tree or foliage roosts. Stenodermatines and animalivorous species live in smaller colonies than most other phyllostomids. From a landscape perspective, phyllostomid populations occur either in a few relatively large colonies in widely scattered caves or in many smaller, more closely spaced colonies in trees, including foliage. Colony size in many species often changes seasonally in response to changes in resource availability and reproductive activities. Because of low reproductive output, growth rates of most phyllostomid populations are low, and many species likely live at their habitat’s carrying capacity as set by resource levels and roost availability. Some species, usually plant-visitors, undergo altitudinal or latitudinal migrations. Most phyllostomid populations harbor substantial amounts of genetic variation, and the extent of genetic subdivision between populations (or metapopulations) tends to be low. (pages 325 - 346)
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    University of Chicago Press
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- Richard D. Stevens, Sergio Estrada-Villegas
DOI: 10.7208/chicago/9780226696263.003.0020
[abundance;anthrogenic effects;community properties;community structure;ecological processes;habitat size;isolation;metacommunities;species richness]
More recently, the mechanistic basis of community structure has been distilled into four higher-level fundamental processes: selection, speciation, drift and dispersal. Herein we summarize a vast literature of research on determinants of phyllostomid community structure and apply this new theoretical framework to improve our understanding phyllostomid community structure. We categorized each of 232 studies identified in the literature as focusing on phyllostomid communities as to which of these four processes was addressed. The vast majority of studies have focused on the process of selection, in particular to responses to anthropogenic habitat modification or other environmental characteristics such as food availability, elevation, latitude and seasonality. The paucity of studies focusing on drift, speciation and dispersal is striking. We provide a roadmap for future investigations that bridges the gap between a focus on selection and the other three higher-level processes. Focus on selection has not generated a unified understanding as to the selective forces that control the structure of phyllostomid bat communities, especially in the forces imposed by fragmentation versus those imposed by land-use change. Speciation rates need to be integrated in phyllostomid community structure in order to understand regional differences in species richness, abundances and trait distribution. (pages 347 - 372)
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    University of Chicago Press
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- Marco A. R. Mello, Renata L. Muylaert
DOI: 10.7208/chicago/9780226696263.003.0021
[biogeography;community ecology;ecological networks;emerging diseases;interaction networks;movement ecology;network science;species interactions]
In the past decades, bat ecology has developed considerably thanks to novel theoretical frameworks, as well as innovative tools for data collection and analysis. In this wind of change, network theory has become very useful to understand the complexity of the associations established by bats among themselves, with other organisms, and with their environment. We review how network science has been used to disentangle bats from the “web of life”, which main issues it is helping to solve, and how its application varies among studies including phyllostomids and other groups. We focus our discussion on the potential for using networks to study biological processes that shape bat systems of different kinds. Finally, we address new avenues for research, such as plant-animal interactions, movement ecology, and emerging diseases. (pages 373 - 390)
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DOI: 10.7208/chicago/9780226696263.003.0022
[biodiversity;diversity gradients;ecological niche modeling;evolutionary mechanisms]
Focus on Neotropical bats, especially Phyllostomidae, has provided many rich insights into contemporary biogeography of the Earth’s biota, in particular from perspectives of describing patterns and searching for mechanisms underlying broadscale gradients of biodiversity. Here we review this large body of research. We begin by reviewing more classical approaches to describe and explain secondary gradients of diversity related to area, latitude, and elevation but also review more contemporary analyses involving primary gradients related to climate and history. Indeed, phyllostomid bats exhibit arguably some of the strongest biodiversity gradients in the world. Moreover, gradients of phyllostomid diversity reflect responses to a complex tapestry of climatic conditions such as precipitation, temperature and their seasonality combined with historical drivers of diversification such as spatially variable speciation rates and tropical niche conservatism. We end by making explicit some of the methodological challenges that limit our understanding of phyllostomid biodiversity gradients as well as highlight some of the more exciting novel approaches that promise much in terms of improving our understanding of the vast complexity of biodiversity gradients of Phyllostomidae as well as the mechanistic basis to these patterns. (pages 391 - 410)
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    University of Chicago Press
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- Enrico Bernard, Mariana Delgado-Jaramillo, Ricardo B. Machado, Ludmilla M. S. Aguiar
DOI: 10.7208/chicago/9780226696263.003.0023
[agribusiness;bat conservation;Brazil;cave protection;forest loss;habitat loss]
Brazil is a continental-sized country that harbors at least 93 species of phyllostomids. Although large tracts of its original vegetation cover are still preserved, some of the Brazilian biomes are under severe human pressure. In this chapter, we identify and discuss major pressures and threats affecting the protection of Brazilian phyllostomids. By using species distribution modelling and GIS techniques, we estimate the impact of land conversion on some species and discuss possible future scenarios. We also identify priority areas for bat conservation in the country, as well as challenges and opportunities that must be pursued to guarantee the protection for most of Brazil’s phyllostomids. (pages 413 - 434)
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- Jafet M. Nassar, Luis F. Aguirre, Bernal Rodríguez-Herrera, Rodrigo A. Medellín
DOI: 10.7208/chicago/9780226696263.003.0024
[conservation;education;emerging threats;extrinsic threats;habitat loss;roost disturbance]
New World leaf-nosed bats stand out as the most versatile bat family within Chiroptera for their broad spectrum of ecological functions and ecosystem services. Paradoxically, phyllostomid bats, and bats in general in the region, face the negative impacts of several extrinsic threats that are mainly responsible for their population decline, in some cases driving them toward local extinction. Habitat loss and roost disturbance and destruction are the two main factors affecting them over their entire geographic range. Globally, 13 species (6% of 216 recognized taxa) have been assigned to threatened IUCN categories. At the regional level, percentage of threatened species stands below 32% (North America: 31%, Central America: 31%, South America: 26%, Caribbean: 12%). In the last decade, bat conservation activism and conservation-oriented research have increased significantly across the Americas and the Caribbean, generating measurable positive effects for many species, including many phyllostomids. These positive impacts are being achieved through the interplay between research, environmental education, and conservation applications as a function of regional and local conservation strategies. Emergence of new threats to bats in the Neotropical region, such as development of wind energy facilities and emerging and reemerging zoonotic viruses, calls for intensified action on behalf of bats. (pages 435 - 456)
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