A PATKÓSDENEVÉREK RENDSZERTANA TAXONOMY OF THE HORSESHOE BATS OF THE WORLD (CHIROPTERA: RHINOLOPHIDAE) Doktori (PhD) értekezés Csorba Gábor Debreceni Egyetem Természettudományi és Technológiai Kar Debrecen, 2008 ii Ezen értekezést a Debreceni Egyetem TTK Biológia Doktori Iskola Biodiverzitás programja keretében készítettem a Debreceni Egyetem TTK doktori (PhD) fokozatának elnyerése céljából. Debrecen, 2008 ....................................................... Tanúsítom, hogy Csorba Gábor doktorjelölt 2005-2008 között a fent megnevezett Doktori Iskola Biodiverzitás programjának keretében irányításommal végezte munkáját. Az értekezésben foglalt eredményekhez a jelölt önálló alkotó tevékenységével meghatározóan hozzájárult. Az értekezés elfogadását javasolom. Debrecen, 2008 .................................................... Dr. Varga Zoltán témavezet ı iii iv CONTENTS 1. Introduction ........................................................................................................ 1 2. Literature overview ............................................................................................ 2 3. Materials and methods ....................................................................................... 5 3.1. The family Rhinolophidae .................................................................... 5 3.2. Material investigated ............................................................................ 6 3.3. Definitions of measurements and technical terms ................................ 7 4. Results ................................................................................................................ 11 4.1. Evaluation of taxonomic characters ...................................................... 11 4.2. Taxonomic changes .............................................................................. 15 4.3. Key to the groups and species of genus Rhinolophus .......................... 16 4.4. Species accounts ................................................................................... 22 5. Összefoglaló ..................................................................................................... 154 1. Appendix: Acknowledgements ......................................................................... 157 2. Appendix: References ....................................................................................... 158 3. Appendix: Figures ............................................................................................ 168 4. Appendix: Scientific publications on the subject of the dissertation ................ 169 v vi 1. INTRODUCTION The extension and improvement of our taxonomic knowledge is rendered urgent by the biodiversity crisis which threatens to destroy much of the evidence of evolutionary history before it could be documented. This is especially emphasized in the case of mammals, a flagship group of the animal kingdom in the fields of conservation biology, phylogenetic classification, ecological and behavioural studies. The second most numerous order of mammals is that of the bats (Chiroptera), with over 1200 known living species. Due to the high number of taxa and their cryptic life-style bats are relatively poorly know. This might explain the fact that new species are described regularly from the tropical and even the temperate areas of the globe (the number of species has increased by approximately 20 per cent in the last decade) and a significant part of the bat literature is focused on their faunistics and taxonomy. The family of horseshoe bats (Rhinolophidae) is well-defined taxonomic unit with exclusively insectivorous species. The highest number of co-existing species can be found in mature tropical forests and extensive limestone cave systems of the Old World, in biotops, which are among the most threatened ones on the Earth. Specimens of many species are rarities in the collections and scattered in many countries. Our ability to differentiate the species is complicated by their uniform appearance, where differences between the species are mainly found in the complex anatomical parts, which are difficult to describe verbally (the shape of the noseleaf and the nasal swellings), and in differences in size. Moreover, in the last 100 years no serious attempt was made to provide a thorough revision and an overview of relevant literature. Neither the old species descriptions nor the new identification keys and comprehensive works give enough anatomical details or adequate quantity of measurements. Hence, the knowledge regarding the identification of these species - that forms the basis for all the other research such as ecological, physiological or conservation-oriented studies - was fairly difficult to gather. The aim of this work was therefore twofold: 1) Careful examination of specimens available in collections, information recorded as exact and repeatable measures and comparable illustrations (drawings and digital images). 2) re-organising and evaluating existing publications on various morphological characters (including external, cranial and dental features), taxonomy and systematics. By producing this database with the knowledge gained by the processes described in 1), my goal was to write a practical monography on taxonomy of this beautiful family of bats. Due to this principal task my dissertation is focusing exclusively on taxonomy , although, during the data collecting period huge amount of information were gathered on aspects of horseshoe bat biology in general, including distribution, habits, feeding, breeding, echolocation, conservation status, phylogeny and biogeography. These fields are out of the scope of this work but presented in the book of Csorba et al. (2003). 1 2. LITERATURE OVERVIEW The overview presented here is based on the compilation of Guillen et al (2003) published in the book of Csorba et al. (2003) and deals only with the systematics of the family above the species-level. The overview of the species-level taxonomy can be found in the ’species accounts’ section under the heading ’Taxonomic remarks’. These historical parts are based on, according to my best knowledge, all the information published between 1903-2002 and different taxonomic opinions related to each species are provided. This is supplemented by my critical comments and by the interpretation of taxonomic actions applied in the dissertation including lectotype designations and revisions. Beginning with the introduction of the genus name Rhinolophus by Lacépède (1799), most authors have included all extant horseshoe bats in a single genus, although some alternatives have been proposed. Leach (1816) proposed the genus name Phyllorhina for Vespertilio minutus Montagu, 1808 (now called R. hipposideros minutus ; see Hill 1963). Initially, the genus Rhinolophus also included the species of Hipposideros known at that time. Gray (1847, 1866), based on differences in the noseleaves, proposed the genus Aquias for the Indomalayan species R. trifoliatus and R. luctus and Phyllotis for the indomalayan-australasian species R. philippinensis , while keeping all other species of horseshoe bats in the genus Rhinolophus , from which he excluded the hipposiderid bats. Peters (1867) proposed the genus Coelophyllus for Rhinolophus coelophyllus . Dobson (1876) returned all Asiatic horseshoe bats to Rhinolophus and ignored generic and subgeneric partitions proposed previously. He also separated rhinolophid and hipposiderid bats in two different subfamilies, Rhinolophinae and Phyllorhininae. Matschie (1901) described Rhinolophus mehelyi under the subgenus Euryalus , together with R. euryale . Miller (1907) elevated the subfamilies erected by Dobson (1876) to family rank. Iredale and Troughton (1934) introduced Rhinophyllotis with the type being R. megaphyllus . These additional genera were obviated by subsequent authors, and Rhinolophus was kept as the only genus. Bourret (1951) described the highly characteristic R. paradoxolophus as belonging to the genus Rhinomegalophus . But Hill (1972) brought it back into Rhinolophus , noting the close similarity of this species with R. rex and other species previously included in the genus. Besides describing many new species and forms of Rhinolophus , Andersen (1905a, 1905b, 1905c, 1905d, 1905e, 1905f, 1918) reviewed the family in a classical fashion and advanced the first phylogenetic hypotheses on the evolution and biogeography of the group. Because horseshoe bats vary little in major skeletal structures, Andersen (works cited above) used few characters for establishing his systematic arrangement. Many of these characters are probably plesiomorphic or prone to homoplastic change. They included the size and degree of displacement from the toothrow of minor teeth (variable even within the same species), size and shape of noseleaves and ears, length of palate (characters involved in echolocation and prone to adaptive evolution), and relative length of finger bones of the wing (involved in adaptive flight morphology). According to the custom at the time, he also used non-objective methods of phylogenetic reconstruction and models of character evolution, regarding without clear justification some features (long palate, three mental grooves, subequal metacarpals) as primitive Andersen (1905a). In his 1905 papers, Andersen arranged the species of Rhinolophus into six groups, named after the species: R. simplex , R. lepidus , R. midas, R. philippinensis, R. macrotis and R. arcuatus , some containing a number of sections. He also sketched the 2 putative