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Higher Level Classification of Phyllostomid Bats with a Summary of DNA Synapomorphies

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Higher Level Classification of Phyllostomid Bats with a Summary of DNA Synapomorphies Acta Chiropterologica, 18(1): 1–38, 2016 PL ISSN 1508-1109 © Museum and Institute of Zoology PAS doi: 10.3161/15081109ACC2016.18.1.001 Higher level classification of phyllostomid bats with a summary of DNA synapomorphies ROBERT J. BAKER1, SERGIO SOLARI2, ANDREA CIRRANELLO3, and NANCY B. SIMMONS4 1Department of Biological Sciences and Museum, Texas Tech University, Lubbock, TX 79409, USA 2Instituto de Biología, Universidad de Antioquia, Medellín, Colombia 3Department of Anatomical Sciences, Stony Brook University, Stony Brook, NY 11794, USA 4Division of Vertebrate Zoology, American Museum of Natural History, New York, NY 10024, USA 5Corresponding author: E-mail: [email protected] The family Phyllostomidae is recognized as representing the most extensive radiation known in any mammalian family. Creating a Linnaean classification for this clade has been difficult and controversial. In two companion papers, we here propose a revised classification drawing on the strengths of genetic and morphological data and reflecting current ideas regarding phylogenetic relationships within this monophyletic clade. We recognize 11 subfamilies (Macrotinae, Micronycterinae, Desmodontinae, Phyllostominae, Glossophaginae, Lonchorhininae, Lonchophyllinae, Glyphonycterinae, Carolliinae, Rhinophyllinae, and Stenodermatinae), 12 tribes (Diphyllini, Desmodontini, Macrophyllini, Phyllostomini, Vampyrini, Glossophagnini, Brachyphyllini, Choeronycterini, Lonchophyllini, Hsunycterini, Sturnirini, and Stenodermatini), and nine subtribes (Brachyphyllina, Phyllonycterina, Anourina, Choeronycterina, Vampyressina, Enchisthenina, Ectophyllina, Artibeina, and Stenodermatina). The proposed arrangement avoids non-monophyletic associations, only keeping those detected based on analyses of DNA sequence data. We propose that a classification based on the strengths of the most complete morphological and genetic data sets will provide the most robust classification for multiple uses by science and society. Key words: Phyllostomidae, higher-level classification, DNA sequence data INTRODUCTION Dávalos et al., 2014). Because this ecomorphologi- cal diversity has fascinated scientists for over a cen- The family Phyllostomidae (New World leaf- tury, Phyllostomidae is one of the best-known and nosed bats) comprises more than 200 species in 60 well-studied chiropteran groups (Jones et al., 2002). genera (see Solari and Martínez-Arias, 2014; Hur - Although comparative studies abound in the litera- tado and Pacheco, 2014), and is the second most ture, remarkably different systematic and phyloge- speciose chiropteran family (Simmons, 2005). The netic analyses have been proposed based on differ- family has undergone a radiation unparalleled in ent data sets and analysis methods (see reviews in other mammalian families in terms of ecological and Wetterer et al., 2000, and Baker et al., 2003). Until morphological diversity (Freeman, 2000; Dumont et recently, there has been little agreement regarding al., 2012). Phyllostomidae encompasses a range of the deep branching patterns and relationships in this dietary diversity larger than that seen in any other remarkable radiation, leading to considerable insta- monophyletic mammal family, including omnivo- bility in classifications (Simmons, 2005; Baker et rous, insectivorous, carnivorous, nectarivorous, fru- al., 2012; Dávalos et al., 2012, 2014). Because of givorous and even hematophagous species (Gard - their diversity, abundance, and ubiquity across the ner, 1977a; Ferrarezzi and Gimenez, 1996; Dumont Neotropics, phyllostomid bats are the main focus of et al., 2012). Ecological variation in diets is associ- a number of research efforts; therefore, a well-sup- ated with extensive morphological diversity that ported and stable classification is highly desirable to involves skeletal, muscle, digestive, kidney, sensory communicate information among those studying systems, and behavior (Phillips, 2000; Wetterer et this family as well as those in other fields who ap- al., 2000; Dumont, 2004; Monteiro and Nogueira, preciate the biodiversity within this complex. The 2011; Baker et al., 2012; Dumont et al., 2012; categories of phyllostomid bats showing the least Acta Chiropterologica, 18(1): 39–71, 2016 PL ISSN 1508-1109 © Museum and Institute of Zoology PAS doi: 10.3161/15081109ACC2016.18.1.002 Morphological diagnoses of higher-level phyllostomid taxa (Chiroptera: Phyllostomidae) ANDREA CIRRANELLO1, 2, 5, NANCY B. SIMMONS1, SERGIO SOLARI3, and ROBERT J. BAKER4 1Division of Vertebrate Zoology, American Museum of Natural History, New York, NY 10024, USA 2Department of Anatomical Sciences, Stony Brook University, Stony Brook, NY 11794, USA 3Instituto de Biología, Universidad de Antioquia, Medellín, Colombia 4Department of Biological Sciences and Museum, Texas Tech University, Lubbock, TX 79409, USA 5Corresponding author: E-mail: [email protected] Phyllostomidae (New World leaf-nosed bats), the second most speciose chiropteran family, is one of the best-known and well- studied chiropteran groups. Due to the ecological and morphological diversity of this family, comparative studies of phyllostomids abound in the literature, and numerous systematic and phylogenetic analyses have been published. Unfortunately, many of these studies have reached different conclusions concerning phyllostomid relationships, and have proposed different classification schemes. This has led to confusion, and highlighted the need for a well-supported and stable classification of the family, particularly at the level of subfamilies and tribes, areas of the greatest controversy. The goal of this paper is to provide morphological diagnoses of higher-level taxa (subtribes, tribes, and subfamilies). Herein we provide morphological diagnoses for 11 subfamilies (Macrotinae, Micronycterinae, Desmodontinae, Lonchorhininae, Phyllostominae, Glyphonycterinae, Glossophaginae, Lonchophyllinae, Carollinae, Rhinophyllinae, and Stenodermatinae), 12 tribes (Desmodontini, Diphyllini, Macrophyllini, Phyllostomini, Vampyrini, Choeronycterini, Glossophagini, Brachyphyllini, Lonchophyllini, Hsunycterini, Sturnirini, and Stenodermatini), and nine subtribes (Anourina, Choeronycterina, Brachyphyllina, Phyllonycterina, Vampyressina, Enchisthenina, Ectophyllina, Artibeina, and Stenodermatina). Key words: morphology, taxonomy, Phyllostomidae INTRODUCTION I have founded the fourth great school of classifica- tion, the It-Doesn’t-Matter-Very-Much school.” Classification and taxonomy are often regarded Although we respect Felsenstein’s (2004) view as boring exercises in bookkeeping, but scientific that it is the phylogeny that matters, referencing names provide a critical framework for storage and nodes on a phylogeny by numbers or letters — or re- communication of scientific knowledge. Herein, we producing large phylogenies simply to be able to define taxonomy as the description, naming, and talk about them — is cumbersome and can hinder classification of organisms. At alpha taxonomic lev- effective scientific communication. To us, higher els (i.e., the level of the species), the importance of level taxonomy is important because it provides taxonomy is clear — it is difficult to imagine setting names, increasingly applied to clades that have been conservation priorities or conducting most field re- supported by data and tested through systematic re- search without first identifying study organisms to search. This shorthand approach to discussion of species. However, at higher taxonomic levels, this phylogenetic groups is both satisfying and neces- clarity of purpose may not be as obvious. To quote sary. Names of phylogenetic groups are used in Felsenstein (2004: 145): studies concerned with evolution (e.g., adaptive “A phylogenetic systematist and an evolutionary radiation, or biogeography), ecology (e.g., behav- systematist may make very different classifications, ioral ecology), and conservation biology (e.g., while inferring much the same phylogeny. If it is hotspot location, identification of unique lineages). the phylogeny that gets used by other biologists, We believe that having a well-supported and stable their differences about how to classify may not be classification for any group reduces confusion that important. I have consequently announced that among non-experts, enabling productive discussion, Acta Chiropterologica, 18(1): 73–90, 2016 PL ISSN 1508-1109 © Museum and Institute of Zoology PAS doi: 10.3161/15081109ACC2016.18.1.003 The evolutionary history of the African fruit bats (Chiroptera: Pteropodidae) FRANCISCA CUNHA ALMEIDA1, 3, 4, NORBERTO PEDRO GIANNINI1, 2, and NANCY B. SIMMONS1 1Department of Mammalogy, Division of Vertebrate Zoology, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024, USA 2Facultad de Ciencias Naturales e Instituto Miguel Lillo, Universidad Nacional de Tucumán, Consejo Nacional de Investigaciones Científicas y Tecnológicas (CONICET),Unidad Ejecutora Lillo, Miguel Lillo, 205, San Miguel de Tucumán, 4000, Argentina 3Current address: Departamento de Ecología, Genética y Evolución, Universidad de Buenos Aires, Consejo Nacional de Investigaciones Científicas y Tecnológicas (CONICET), Intendente Güiraldes y Costanera Norte s/n, Pabellón II - Ciudad Universitaria, 1428, Capital Federal, Argentina 4Corresponding author: E-mail: [email protected] Bats of the family Pteropodidae, also known as megabats or Old World fruit bats, are widely distributed in tropical areas of Africa, Asia, and Oceania. Of 45 genera in
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