amnb 00189 Mp 1 File # 01TQ PHYLOGENETIC RELATIONSHIPS OF MORMOOPID BATS (CHIROPTERA: MORMOOPIDAE) BASED ON MORPHOLOGICAL DATA NANCY B. SIMMONS Associate Curator Division of Vertebrate Zoology (Mammalogy) American Museum of Natural History TENLEY M. CONWAY Scientific Assistant Division of Vertebrate Zoology (Mammalogy) American Museum of Natural History BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY Number 258, 97 pp., 12 figures, 4 tables Issued February 15, 2001 Price: $10.10 a copy Copyright ᭧ American Museum of Natural History 2001 ISSN 0003-0090 2 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 258 CONTENTS Abstract ....................................................................... 3 Introduction .................................................................... 3 Historical Background ......................................................... 3 Mormoopid Species: A Synopsis ............................................... 8 Goals of the Present Study .................................................... 12 Materials and Methods ......................................................... 12 Taxonomic Sampling, Outgroups, and Tree Rooting .............................. 12 Sources of Data ............................................................. 13 Definition of Characters and Ordering of Character States ........................ 13 Polarity ..................................................................... 15 Completeness ............................................................... 15 Methods of Phylogenetic Analysis ............................................. 16 Character Descriptions .......................................................... 17 Skull ....................................................................... 17 Dentition ................................................................... 24 Vomeronasal Complex and Brain .............................................. 29 Trachea and Hyoid Apparatus ................................................. 30 Tongue ..................................................................... 35 Face, Ears, and Vibrissae ..................................................... 36 Pelage and Patagia ........................................................... 42 Postcranial Skeleton .......................................................... 45 Postcranial Myology ......................................................... 58 Reproductive Tract ........................................................... 65 Digestive Tract .............................................................. 66 Results ....................................................................... 68 Discussion and Conclusions ..................................................... 73 Intrafamilial Relationships of Mormoopids ...................................... 73 Classification and Taxonomic Diagnoses of Mormoopid Clades ................... 74 Monophyly and Diagnoses of Noctilionoid Families ............................. 81 Interfamilial Relationships .................................................... 83 Directions for Future Research ................................................ 84 Acknowledgments ............................................................. 85 References .................................................................... 85 Appendix 1: Specimens Examined ............................................... 93 Appendix 2: Taxon-Character Matrix ............................................. 96 2001 SIMMONS AND CONWAY: PHYLOGENY OF MORMOOPID BATS 3 ABSTRACT Mormoopidae is a small family of Neotropical microchiropteran bats that includes two genera (Mormoops and Pteronotus) and ten species, two of which are known only from fossils. Mormoopidae is typically classified as a member of Noctilionoidea, a group that minimally includes two other Neotropical families (Phyllostomidae and Noctilionidae) and may also include Mystacinidae, a taxon endemic to New Zealand and Australia. Phylogenetic relation- ships of extant mormoopid species and one extinct taxon, Pteronotus pristinus, were investi- gated in a series of parsimony analyses of 209 morphological characters including features of the skull, dentition, vomeronasal organ complex and brain, trachea and hyoid apparatus, tongue, face, ears, pelage, patagia, postcranial skeleton, postcranial myology, reproductive tract, and digestive tract. Three extant phyllostomid species, two noctilionids, two mystacinids, and one emballonurid species were included as outgroups to test monophyly of Mormoopidae and to provide a context for determining the sister group of the family. Results of parsimony analyses under a variety of different assumption sets indicate that Mormoopidae is monophy- letic, and that Pteronotus and Mormoops are monophyletic sister taxa. Within the genus Pter- onotus, several clades were repeatedly recovered: (1) P. davyi ϩ P. gymnonotus (ϭ subgenus Pteronotus); (2) P. macleayi ϩ P. quadridens; (3) P. personatus ϩ P. macleayi ϩ P. quad- ridens (ϭ subgenus Chilonycteris); (4) P. parnellii ϩ P. pristinus ϩ P. personatus ϩ P. macleayi ϩ P. quadridens; and (5) P. parnellii ϩ P. pristinus (ϭ subgenus Phyllodia). These results support monophyly of all subgenera of Pteronotus previously recognized, and addi- tionally indicate that the subgenera Pteronotus and Chilonycteris are sister taxa. Comprehen- sive diagnoses for each species and clade of Mormoopidae are provided based on character optimizations and ancillary morphometric data from the literature. Results of our parsimony analyses also have implications for understanding higher-level phy- logeny of noctilionoid bats. Monophyly of each of the traditionally recognized families (i.e., Noctilionidae, Mystacinidae, Mormoopidae, and Phyllostomidae) was strongly supported. In con- gruence with recent analyses of mitochondrial gene-sequence data and DNA hybridization ex- periments, we found strong support for inclusion of Mystacinidae in Noctilionoidea. Using an emballonurid species to root the tree, we found the following interfamilial relationships of noc- tilionoids: (Noctilionidae (Mystacinidae (Phyllostomidae, Mormoopidae))). Lists of morpholog- ical synapomorphies of each of these groups are provided based on character optimizations. INTRODUCTION tropical forest to semiarid and arid subtropical forest and scrubland (Handley, 1976; Em- Mormoopidae is a small family of Neo- mons, 1997; Reid, 1997; Smith, 1972). The tropical microchiropteran bats that currently most recent revision of Mormoopidae was includes two genera (Mormoops and Pteron- that of Smith (1972), which has formed the otus) and eight extant species; two additional basis for most modern treatments of the fam- species are known only from Quaternary fos- ily. sils (Smith, 1972; Silva-Taboada, 1974, 1979; The history of classification and nomen- Koopman, 1993, 1994). Commonly known as clature of mormoopids is complex and often mustached, ghost-faced, or naked-backed confusing. Smith (1972) provided a compre- bats, these taxa are characterized by the pres- hensive systematic literature review for the ence of flaplike outgrowths below the lower family, which we will not reproduce here. A lip and funnel-shaped ears. Mormoopids are few critical points from the pre-1970 litera- small to medium in size (e.g., forearm length ture, together with more recent taxonomic 35–66 mm) and are thought to be exclusively changes and phylogenetic inferences, are insectivorous (Koopman, 1984, 1994). Mem- summarized below. bers of the family presently range from the southwestern United States to southern Brazil, HISTORICAL BACKGROUND and also occur in the Greater Antilles, Lesser Antilles, Dutch West Indes, and Trinidad and CLASSIFICATION Tobago (Koopman, 1993, 1994). They live in The taxonomic history of Mormoopidae a wide variety of habitats ranging from humid began with Leach (1821a, 1821b), who de- 4 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 258 scribed Mormoops and Aello. Pteronotus and sification did not entirely match his phylo- Chilonycteris were subsequently described genetic tree (fig. 1), which depicted Pteron- by Gray (1838, 1839), and Lobostoma was otus personatus as more closely related to P. named by Gundlach (1840). Gray (1843) davyi and P. suapurensis than to P. macleayii named a sixth genus, Phyllodia, followed al- and P. fuliginosus. However, Smith’s (1972) most 60 years later by Gill’s (1901) propo- revision was completed in a precladistic con- sition of Dermonotus as a replacement name text, so this discrepancy was not widely rec- for Pteronotus. Various authors have pro- ognized. Subsequent subgeneric classifica- posed different synonomies for these taxa tions (e.g., Corbet and Hill, 1980; Herd, over the last 150 years, and misplaced ho- 1983; Koopman, 1994) retained Smith’s lotypes, overlooked names, and arguments (1972) subgeneric usage. over nomenclatural priority have served to Although Smith’s (1972) classification of further complicate matters (see review in Mormoopidae survived more-or-less intact Smith, 1972). For most of the 20th century, into the 1990s, several significant nomencla- classifications of Mormoopidae were based tural changes have occurred since that pub- on that of Miller (1907), who recognized lication. Silva-Taboada (1976) showed that three genera (Chilonycteris, Pteronotus, and Lobostoma quadridens Gundlach, 1840 is a Mormoops), which he placed in the subfam- senior synonym of Chilonycteris fuliginosus ily Chilonycterinae in the family Phyllostom- Gray, 1843, so Pteronotus quadridens is now idae.