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BONNER ZOOLOGISCHE MONOGRAPHIEN, Nr © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at NEW WORLD NECTAR-FEEDING BATS: BIOLOGY, MORPHOLOGY AND CRANIOMETRIC APPROACH TO SYSTEMATICS by ERNST-HERMANN SOLMSEN BONNER ZOOLOGISCHE MONOGRAPHIEN, Nr. 44 1998 Herausgeber: ZOOLOGISCHES FORSCHUNGSINSTITUT UND MUSEUM ALEXANDER KOENIG BONN © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at BONNER ZOOLOGISCHE MONOGRAPHIEN Die Serie wird vom Zoologischen Forschungsinstitut und Museum Alexander Koenig herausgegeben und bringt Originalarbeiten, die für eine Unterbringung in den „Bonner zoologischen Beiträgen" zu lang sind und eine Veröffentlichung als Monographie rechtfertigen. Anfragen bezüglich der Vorlage von Manuskripten sind an die Schriftleitung zu richten; Bestellungen und Tauschangebote bitte an die Bibliothek des Instituts. This series of monographs, published by the Zoological Research Institute and Museum Alexander Koenig, has been established for original contributions too long for inclu- sion in „Bonner zoologische Beiträge". Correspondence concerning manuscripts for pubhcation should be addressed to the editor. Purchase orders and requests for exchange please address to the library of the institute. LTnstitut de Recherches Zoologiques et Museum Alexander Koenig a etabh cette serie de monographies pour pouvoir publier des travaux zoologiques trop longs pour etre inclus dans les „Bonner zoologische Beiträge". Toute correspondance concernante des manuscrits pour cette serie doit etre adressee ä I'editeur. Commandes et demandes pour echanges adresser ä la bibhotheque de I'insti- tut, s. V. p. BONNER ZOOLOGISCHE MONOGRAPHIEN, Nr. 44, 1998 Preis: 30,— DM Schriftleitung/Editor: G. Rheinwald Zoologisches Forschungsinstitut und Museum Alexander Koenig Adenauerallee 150—164, D-53113 Bonn, Germany Druck: JF.CARTHAUS, Bonn ISBN 3-925382-48-8 ISSN 0301-671 X © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at NEW WORLD NECTAR-FEEDING BATS: BIOLOGY, MORPHOLOGY AND CRANIOMETRIC APPROACH TO SYSTEMATICS BONNER ZOOLOGISCHE MONOGRAPHIEN, Nr. 44 1998 Herausgeber: ZOOLOGISCHES FORSCHUNGSINSTITUT UND MUSEUM ALEXANDER KOENIG BONN © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at Die Deutsche Bibliothek — CIP Einlieitsaufnahnie Solmsen, Ernst-Hermann: New world nectar feedmg bats: biology, morphology and craniometric approach to systematics / by Ernst- Hemiann Solnisen. Hrsg.: Zoologisches Forschungsinstitut und - Museum Alexander Koenig, Bonn. Bomi : Zoologisches Forschungsinst. und Museum Alexander Koenig, 1998 (Bonner zoologische Monographien ; Nr. 44) Zugl.: Hamburg, Univ., Diss., 1994 ISBN 3-925382-48-8 © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at CONTENTS Page Introduction 5 Acknowledgements 7 On tlie systematics, distribution and ecology of New World flower bats 8 Taxonomic position 8 Geographic distribution 10 Habitat, roosting behaviour, migration 12 Activity patterns 15 Feeding biology / Feeding ecology 16 Chiropteran adaptations to nectari- and poUenivory 16 Body size and weight 16 Body shape 17 Pelage 18 Digestive tract 18 Adaptations in host plants to chiropteropliily 20 Size and constitution 20 Blossom shape 20 Blossom exposition 20 Diet of various New World nectar feeding bat genera 21 Sensory systems / Orientation 25 Acoustic perception; echolocation 25 Optical sense 25 Olfactory sense / Olfactory perception 26 Reproductive biology 26 Cytology 31 Material and methods 31 Material 31 Methods 32 Descriptions 32 Drawings 33 Measurements 33 Calculations 34 Univariate analysis 34 Allometrics 34 Results 35 Morphology of the species examined 35 External morphology 35 Skull morphology 45 Skull descriptions of the single genera 46 © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at Morphometry 62 Univariate analysis 62 Skull proportions 63 AUometrics 66 AUometric comparison of individual genera 66 Sex dimorphism 73 Discussion 74 Morphological adaptations to nectarivory 74 Gross morphology of the head 74 Skull morphology 74 Dentition 75 Incisivi 75 Canmi 75 Premolars 76 Molar teeth 76 Rostaim 76 Forehead 77 Zygomatic arches 77 Braincase 78 Skull base 78 Mandible 80 Craniometry 80 Univariate analysis 80 Skull proportions 81 AUometrics 83 Intergeneric allometrics 83 Allometrical sex comparison 92 AUometric conclusions 92 Systematic conclusions 93 Sunmiary 97 Literature cited 98 Appendix 108 © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at 5 INTRODUCTION With more than 900 species, the Chiroptera represent the most comprehensive mammalian order next to the rodents. By developing physical ability for continuous flight bats succeeded in exploring habitats almost without vertebrate competitors - the nocturnal airspace. Tlieir morphological adaptations are just as consistent; having specialized their anatomy in flight and a hanging restmg position, most bats are hardly capable of qua- drupedal locomotion, apart from crawling over short distances. General body shape hardly varies within the Chiroptera - presumably due to strict requirements for free flight. Differences are delicate and restricted to body size, wing shape, development of the uropatagium and tail length. However, the chiropteran head is one of the most striking characters in specific detemiination. Due to echolocation many microchiropterans show an oddity of complex structures on ears and often also nose winch aid in directed emission and perception of sound. Having successfully conquered the nocturnal skies, the primarily insectivorous Cliiroptera developed a variety of new diet preferences - camivory {Megademia, Vampynim, Phyllostomus, Trachops), piscivory {Pizonyx, Noctilio) and even - unique among the manmialia - sanguivory {Desmodus, Diaemus, Diphylla). Besides, many species also take vegetable food: frugivorous and even few nectarivorous species prevail among the Megacliiroptera (Pteropodidae), but also the Microchiroptera developed a variety of fruit and flower-feeding genera within the Phyllostomidae. These various diet preferences within the Microcliiroptera have their morphological expression primarily in the shape of the head and in the dentition. Thus, skull morphology of bats developed some remarkable diversity witliin the mammalia. Fig.l: Skull shape depending on diet specialization within the Phyllostomatidae {Centurio senex = fruit eater, Choeronycteris harrisoni = blossom feeder, adapted from Koopman 1987) © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at 6 In this respect, some members of tlie phyllostomid family achieved a striking diversity, with the highly specialized flower bats representing an extreme (fig.l). Especially the Glossophaginae succeeded in extensive specialization (prolongation of the rostmm; long tongue with bristle-like papillae, weak and delicate teeth). The systematic iategration of all long-nosed Phyllostomids into a single monophyletic subfamily is still discussed controversially. As the main characters used to define glossophagine bats are derived characters influenced by fimction, they could well have developed independently within the phyllostomids. For more than 25 years, several approaches have been made to tliis problem (tab.l) and various different systematic relationships of single genera were proposed from time to tinie. Table 2: Studies on the system of New World nectaifeeding bats - morphological examinations teeth, dentition (Phillips 1971) gastrointestinal system (Forman 1971, 1979; Rouk & Glass 1970) tongue, hyoid musculature (Wille 1954; Greenbaum & Pillios 1974; Griffiths 1982) female sex apparatus (Smith & Hood 1982) cerebral anatomy (McDaniel 1976) - cx'tological examinations cromosomal analysis (Baker et al. 1967; Patton & Gardner 1971; Stock 1975; Baker & Bass 1979; Haiduk & Baker 1982) hematology (Valdivieso & Tamsit 1971) - immunological studies serum proteins (Baker et al. 1981) serum proteins (Gerber & Leone 1971) Classification of BrachyphyUa as an essentially basic form witliin its own subfamily Brachyphyllinae remained undisputed (Baker et al. 1979) as well as everyone aggreed to sunmiarize the genera Erophylla and Phyllonycteris within the Phyllonycterinae (Flower vampires), a subfamily closely related to the Brachyphyllinae (Silva Taboada & Pine 1969). Contrary conclusions primarily concern the systematic position of the genera or subfamilies having been sunmiarized so far as Glossophaginae. Some really revolutionary improvement was achieved by the works of Griffith pubhshed 1982. Analyzing the tongue and hyoid morphology and their musculature, respectively, he was tlie first to seperate three genera {Lionycteris, Lonchophylla and Platalina) from the Glossophaginae s.str., sunmiarizing tliem witliin tlieir own subfamily Lonchophyllinae. These results led to a hvely discussion between different research groups in the United States (Haiduk & Baker 1982; Warner 1983; Hood & Smiüi 1982; Griffiths 1983; Smith & Hood 1984). The point of this discussion is: do tlie New World nectarivorous bats represent a monophyletic group, or did the glossophagine bats develop twice, independently from each other? Even within
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