SPECIAL PUBLICATIONS THE MUSEUM TEXAS TECH UNIVERSITY Phylogenetic Analyses of the Bat Subfamily Stenodermatinae (Mammalia: Chiroptera) Robert D. Owen TEXAS TECH UNIVERSITY Regents John E. Bird well (Chairman), J. Fred Bucy, Jerry Ford, Rex Fuller, Larry D. Johnson, Jean Kahle, Wm. Gordon McGee, Wesley W. Masters, and Wendell Mayes, Jr. President Lauro F. Cavazos Vice President for Academic Affairs and Research Donald R. Haragan Texas Tech University Press Editorial Committee Clyde Hendrick (Chairman), Wendell W. Broom, Jr., E. Dale Cluff, Monty E. Davenport, Oliver D. Hensley, Clyde Jones, Stephen R. Jorgensen, Richard P. McGlynn, Janet W. Perez, Marilyn E. Phelan, Rodney L. Preston, Willard B. Robinson, Charles W. Sargent, Grant T. Savage, Jeffrey R. Smitten, Idris R. Traylor, Jr., and David A. Welton. Special Publications The Museum No. 26 65 pp. 20 May 1987 Paper, $15.00 Cloth, $30.00 Special Publications of The Museum are numbered serially and published on an irregular basis under the auspices of the Vice President for Academic Affairs and Research and in cooperation with The Museum. Institutions interested in exchanging publications should address the Exchange Librarian at Texas Tech University. Copies may be purchased from: Texas Tech University Press Sales Office, Box 4139 Texas Tech University Lubbock, Texas 79409 U.S.A. ISSN 0149-1768 ISBN 0-89672-150-7 (paper) ISBN 0-89672-151-5 (cloth) Texas Tech University Press Lubbock, Texas 1987 SPECIAL PUBLICATIONS THE MUSEUM TEXAS TECH UNIVERSITY Phylogenetic Analyses of the Bat Subfamily Stenodermatinae (Mammalia: Chiroptera) Robert D. Owen No. 26 May 1987 TEXAS TECH UNIVERSITY Regents John E. Bird well (Chairman), J. Fred Bucy, Jerry Ford, Rex Fuller, Larry D. Johnson, Jean Kahle, Wm. Gordon McGee, Wesley W. Masters, and Wendell Mayes, Jr. President Lauro F. Cavazos Vice President for Academic Affairs and Research Donald R. Haragan Texas Tech University Press Editorial Committee Clyde Hendrick (Chairman), Wendell W. Broom, Jr., E. Dale Cluff, Monty E. Davenport, Oliver D. Hensley, Clyde Jones, Stephen R. Jorgensen, Richard P. McGlynn, Janet W. Perez, Marilyn E. Phelan, Rodney L. Preston, Willard B. Robinson, Charles W. Sargent, Grant T. Savage, Jeffrey R. Smitten, Idris R. Traylor, Jr., and David A. Welton. Special Publications The Museum No. 26 65 pp. 20 May 1987 Paper, $15.00 Cloth, $30.00 Special Publications of The Museum are numbered serially and published on an irregular basis under the auspices of the Vice President for Academic Affairs and Research and in cooperation with The Museum. Institutions interested in exchanging publications should address the Exchange Librarian at Texas Tech University. Copies may be purchased from: Texas Tech University Press Sales Office, Box 4139 Texas Tech University Lubbock, Texas 79409 U.S.A. ISSN 0149-1768 ISBN 0-89672-150-7 (paper) ISBN 0-89672-151-5 (cloth) Texas Tech University Press Lubbock, Texas 1987 Phylogenetic Analyses of the Bat Subfamily Stenodermatinae (Mammalia: Chiroptera) Robert D . Owen Understanding of the systematic relationships of bats has not progressed as rapidly as for many other mammalian groups. Investigators generally agree that many characters and techniques traditionally used in mammalian systematics are at best insufficient for use with bats, but few studies have been undertaken that have included an intensive quantitative examination of a chiropteran group. Exceptions are Findley’s (1972) analysis of phenetic relationships among bats of the genus Myotis, Smith’s (1972) systematic analysis of the family Mormoopidae, and Freeman’s (1981) study of the family Molossidae. Several investigators, including Dobson (1875), Winge (1892), Smith (1976, 1977, 1980), Van Valen (1979), Novacek (1980), Arnold et al. (1982), and Hood and Smith (1982), have evaluated phylogenetic relationships among bats from the generic to the ordinal level. Concerning the family Phyllostomidae, other workers have attempted to determine relationships at the subfamilial and lower levels. These studies have involved a variety of approaches, including immunology (Forman et al., 1968; Gerber and Leone, 1971), karyology (Baker, 1967, 1973; Forman et al., 1968; Davis and Baker, 1974; Greenbaum et al., 1975; Gardner, 1977; Baker et al., 1979; Johnson, 1979; Haiduk and Baker, 1982), electrophoresis (Straney et al., 1979; Koop and Baker, 1983), soft-tissue morphology (McDaniel, 1976; Griffiths, 1982; Tandler et al., 1986), classical morphologic assessment- including dentition (Miller, 1907; Andersen, 1908; Sanborn, 1955; Davis, 1958; Peterson, 1968), and morphometries (Forman et al., 1968; Davis and Baker, 1974). Although the subfamily Stenodermatinae is the most species-rich group within the Phyllostomidae, it is in several aspects the most homogeneous of the larger subfamilies. Unlike the phyllostomines, for instance, the stenodermatines all are dependent on similar foods, each being primarily or entirely frugivorous (Wilson, 1973). The subfamily as now constituted, consisting of 16 or 17 nominal genera and 55 to 59 species (Jones and Carter, 1976; Honacki et al., 1982), includes three genera that together contain over half of the species; in addition, seven to nine genera are considered to be monotypic, depending on the authority consulted. Although considered as a distinct group since 1855 (Miller, 1907), the subfamily Stenodermatinae has undergone several partial revisions, as have most of the genera within it. Miller (1907) recognized 19 genera within the group. Of these, Brachyphylla has since been placed tentatively in the Phyllonycterinae (Silva Taboada and Pine, 1969; Jones and Carter, 1976), 3 4 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Vampyriscus within Vampyressa, and Mesophylla within Ectophylla (but see Starrett and Casebeer, 1968). Of the approximately 50 species recognized by Miller (1907), at least 17 have undergone taxonomic or nominal alteration, and a number of new species have been described. Since the appearance of Jones and Carter’s (1976) checklist, the number of known species has increased from 55 to about 65. My analyses address the interspecific relationships within the subfamily. I evaluate seven problems of current interest: (1) the interspecific and subgeneric relationships within Sturnira; (2) the status and relationships of the numerous described species of Vampyrops ; (3) the relationship of Uroderma to other genera; (4) the phylogeny of the eight “short-faced” genera within the subfamily (that is, Ardops, Phyllops, Ariteus, Stenoderma, Pygoderma, Ametrida, Sphaeronycteris, and Centurio ); (5) the relationships of species and subgenera within the relatively small genus Vampyressa ; (6) the inclusion of Mesophylla within Ectophylla ; and (7) the relationships within the genus Artiheus. These questions can be addressed appropriately only by a phylogenetic analysis of the entire subfamily, because even the monophyly of a number of the genera is in question. My analyses include all known species within the subfamily, and are based upon suites of discrete-state and mensural characters. These analyses should produce robust hypotheses concerning the phylogeny of the subfamily Stenodermatinae. Species in the Subfamily Stenodermatinae I analyzed the relationships of 64 taxa considered to be species (Appendix 1) based on Jones and Carter (1976), and subsequent publications. For ease of discussion only, I used the generic names Enchisthenes and Mesophylla rather than including these bats in Artiheus and Ectophylla , respectively. I have followed Handley (1980) in subfamilial nomenclature. Considerable uncertainty exists concerning affinities of the Antillean endemic genus Brachyphylla; I have followed Dusbabek (1968), Silva Taboada and Pine (1969), Baker and Lopez (1970), Baker et al. (1979), and Sites et al. (1981) in not including this genus in the subfamily Stenodermatinae. For a review of the taxonomic history of Brachyphylla , see Swanepoel and Genoways (1978). Following Winge (1892), de la Torre (1961), Baker (1967), Gerber and Leone (1971), Gardner (1977), and Baker et ai (1979), I have included the enigmatic genus Sturnira in the subfamily. In addition to the 10 Sturnira species listed by Jones and Carter (1976), I have included S. luisi and S. bogotensis , both of which were recognized by Honacki et al. (1982). Gardner and Carter (1972a) and Carter and Rouk (1973) considered Vampyrops umbratus to be conspecific with V. dorsalis. Handley (1976), however, recognized V umbratus as distinct. Honacki et al. (1982) also recognized this separation, and I have treated these as two species. OWEN—PHYLOGENETIC ANALYSES OF STENODERMATINAE 5 One stenodermatine species ( Phyllops vetus) is known only from fossil remains. For the sake of completeness, I have included this species in the study. There is considerable disagreement concerning the taxonomy of the genus Artibeus. For many species names, it is necessary to describe exactly which populations are meant, as names have been applied in various ways by various authors. Furthermore, dispute continues over the status of names in Patten’s (1971) unpublished dissertation. The three middle-sized Artibeus (A. hirsutus , A. inopinatus, and A. concolor) are well studied. Among the large Artibeus the systematics of the Middle American taxa also is understood relatively well, and I have used Mexican and Central American specimens to represent A. jamaicensis. Artibeus lituratus was thought to be taxonomically well understood throughout its range until Davis (1984) reported a zone of overlap between two size classes of A. lituratus in Honduras. Consequently, he raised intermedius (the northern and Gulf Coast taxon) to specific status. My specimens of A. “lituratus ” were from north of this overlap zone and, therefore, are referable to the name A. intermedius. Artibeus lituratus is appropriately considered as the sister species to A. intermedius. Artibeus jamaicensis may or may not be represented south of Colombia; I have treated South American jamaicensis- like bats as distinct species. One of these, occurring on the Pacific versant of the Andes, is A. fraterculus. At least two jamaicensis- like taxa occur east of the Andes: (1) A. fuliginosus, is a relatively small, dark-colored bat; (2) a larger bat has been identified as A. planirostris (the name I have used here). See Koopman (1978) for a discussion of these large South American Artibeus, as well as the status of names used by Patten (1971). Myers and Wetzel (1983) discussed a population that they believed would prove to represent a third jamaicensis- like bat east of the Andes, and indicated that the name fimbriatus should apply to it. I have followed them in treating this population as distinct from the other species of large Artibeus; however, the status of the name fimbriatus is left to consideration by other workers. I have recognized seven species of small Artibeus. These are the six species recognized by Jones and Carter (1976) except that, following Koopman (1978) and Honacki et al. (1982), A. anderseni was considered distinct from A. cinereus. Characters and Coding When possible, I took measurements and coded multistate characters from at least 10 specimens, five of each sex (Appendix I). My objective was to derive the best estimate of stenodermatine phylogeny obtainable from external and osteological characteristics. Accordingly, suites of continuous and discrete-state characters were coded, and several analytic methods were employed. 6 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY A total of 22 discrete-state external, cranial, mandibular, and dental characters was coded for each specimen. Postcranial characters were not used because full skeletons were unavailable for 18 of the 64 species studied. The 22 characters are listed, their character states described, and an unrooted character-state tree is given for each in Appendix II. I assigned the external character states of Phyllops falcatus to P. vetus (known only from fossil remains). For five of the six characters these also were representative of P haitiensis. The 33 continuous-characters also measure aspects of the cranium, mandible, and teeth (Appendix III). Some were taken directly from, or adapted from, Freeman (1981); I developed additional characters representing consistently measurable distances between homologous points. Estimating the Ancestor—Outgroup Procedures Two steps are critical in phylogenetic analyses. The first is inclusion of a monophyletic assemblage of taxa as the ingroup, and the second is determination of the character-state transformation series. One facet of the transformation series is the order of evolutionary change, or tree topology. I specified tree topologies “based on the assumption that the most evenly graded series of changes is the evolutionarily-most-probable hypothesis” (Kluge, 1976). A second facet of the transformation series is the direction of evolutionary change, specified by the tree root. All proposed procedures for rooting character-state trees are simply indirect methods of estimating the suite of character states comprising the most recent common ancestor of the study group. Watrous and Wheeler (1981) first formalized outgroup-comparison methodology and introduced the concept and terminology of functional ingroup and functional outgroup. Maddison et al. (1984) showed the relationship of the outgroup criterion to global and local parsimony, and developed arguments for the critical importance of outgroups with known relationships to each other and to the ingroup. They presented rules for resolution of outgroup relationships, and underscored the importance of using more than one outgroup taxon simultaneously in order to determine correctly both the position and direction of character-state changes in the ingroup phylogeny. I have followed Maddison et al. (1984) in my use of outgroups in the analysis. Phyllostomid subfamilial relationships are poorly understood, especially the affinities of the Stenodermatinae. However, recent evidence (Honeycutt, 1981; Hood and Smith, 1982; Honeycutt and Sarich, 1987) indicates that the subfamily Carolliinae is the sister group to the stenodermatines. I, therefore, included the four species of Carollia among my outgroup taxa. In order to strengthen the outgroup process of character-state tree construction, I used an additional outgroup species chosen on the basis of its presumed similarity to the hypothetical primitive phyllostomid. Macrotus waterhousii has been suggested as primitive for the family, not OWEN—PHYLOGENETIC ANALYSES OF STENODERMATINAE 7 only karyotypically (Bourliere, 1955; Patton and Baker, 1978) but also on the bases of general morphology (Winge, 1892), osteology (Walton and Walton, 1968), dentition (Slaughter, 1970), and fetal membrane structure (Luckett, 1980). Analyses of Discrete-State Characters The six external and 16 cranial characters coded for each specimen are listed and described in Appendix II. The number of coded states for the discrete-state characters ranges from two (postglenoid foramen) to nine (dental formula). Some of these characters were intraspecifically variable in some species (Owen, 1987: appendix Al). To choose the character state that would most appropriately represent each species in the analyses, I used the modal, or most commonly encountered, state. If two states were equally common, I used the more plesiomorphic of the two. This is the conservative estimate in that I did not postulate a synapomorphy unless a uniquely modal state was found. After determining the representative character states for each of the outgroup species, a plesiomorphic state for the ingroup was considered as the one that is (in order of descending importance): (1) carried by all Carollia species; (2) shared by Carollia (one or more species) and Macrotus; (3) the modal state among the four Carollia species; (4) the modal state among the four Carollia species and Macrotus ; or (5) in case of a modal tie in (4), the co-mode carried by Macrotus. Criteria 2 through 5 were needed in only a few cases. Even if these criteria were invoked in some other reasonable sequence, there would be few, if any, different assignments of character states to species. Once the representative character states were determined for each species (Table 1), I transformed them into two-state (presence or absence) characters using additive binary coding (Sneath and Sokal, 1973). The 22 multistate characters thus were transformed into 72 two-state characters. Wagner analysis. —Wagner analysis is a method of estimating a phylogeny through construction of the most parsimonious (shortest), rooted branching network that reflects the character states of the taxa under study. It is an approximation of the Hennigian phylogenetic method that is used appropriately when a synapomorphy scheme (Nelson, 1979) cannot be determined. The reasons for use of the parsimony criterion have been well documented (Camin and Sokal, 1965; Kluge and Farris, 1969; Estabrook, 1978; Farris, 1982; Panchen, 1982; Maddison et al., 1984; Rohlf, 1984). Maddison et al. (1984) described a method whereby successively smaller clades of a group of taxa may be studied, thus allowing the investigator to search for locally parsimonious solutions within the previously established constraints of a global parsimony solution. I have followed their general method of: (1) determining stable portions of the cladogram; (2) determining unstable portions in need of further analysis; and (3) establishing and confirming the ancestor estimate for these successive analyses. 8 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Table 1. — Discrete-state character values for stenodermatine and outgroup species examined. See Appendix II for description of characters, character states, and character-state trees. Characters are listed here in the same order as in Appendix II. Character Species 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 Stenodermatinae Ametrida centurio 4 2 3 5 1 2 4 0 1 1 2 2 2 3 1 2 1 0 1 0 1 0 Ardops nichollsi 6 3 3 5 2 2 3 0 2 2 2 3 2 3 1 3 1 1 0 0 1 0 Ariteus flavescens 4 2 3 5 2 2 3 0 2 2 1 2 2 3 2 2 1 1 0 0 3 0 Artibeus anderseni 4 5 2 1 1 2 3 0 I 1 2 2 1 I 2 2 1 0 1 0 5 2 Artibeus aztecus 6 4 2 1 2 2 3 0 1 2 2 2 2 2 2 2 1 0 0 0 5 2 Artibeus cinereus 4 4 2 1 1 2 3 0 1 2 2 3 2 1 2 2 1 0 0 0 5 2 Artibeus concolor 4 3 3 0 1 3 3 0 ] 2 2 1 2 3 1 2 0 0 0 0 1 0 Artibeus fimbriatus 4 3 2 1 1 2 3 0 I 2 2 3 2 3 2 2 0 0 0 0 3 0 Artibeus fraterculus 4 3 2 1 1 2 3 0 1 2 2 3 3 3 2 2 0 0 0 0 3 1 Artibeus fuliginosus 4 3 2 0 1 2 3 0 0 2 2 2 2 3 1 2 0 0 0 1 1 1 Artibeus glaucus 4 5 2 1 1 2 3 0 I 1 1 2 3 2 2 2 1 0 0 0 3 1 Artibeus hirsutus 4 3 2 1 1 2 3 0 1 3 2 2 2 3 1 2 1 0 0 0 1 1 Artibeus inopinatus 4 3 2 1 1 2 3 0 1 2 2 3 2 3 1 2 1 0 0 0 1 0 Artibeus intermedius 4 3 2 1 1 2 3 0 1 2 2 3 1 3 2 2 0 0 0 0 3 1 Artibeus jamaicensis 4 3 2 1 1 2 3 0 1 2 2 2 2 3 2 2 0 0 0 0 3 1 Artibeus phaeotis 6 5 2 1 1 2 3 0 1 2 2 3 2 2 2 1 1 0 0 0 5 2 Artibeus planirostris 4 3 2 1 1 2 3 0 1 2 2 3 1 3 1 2 0 0 0 0 1 1 Artibeus toltecus 6 5 2 1 1 2 3 0 1 1 2 3 2 2 2 1 0 0 0 0 5 2 Artibeus watsoni 6 5 2 1 1 2 3 0 1 2 2 2 3 2 2 2 0 0 0 0 3 1 Centurio senex 5 1 3 6 1 2 1 0 2 1 1 2 2 3 2 2 1 1 0 0 5 2 Chiroderma doriae 2 2 3 1 0 1 3 1 0 3 3 3 1 0 2 3 1 0 2 2 5 0 Chiroderma improvisum 4 2 2 2 0 2 I 0 0 0 2 3 0 2 0 2 1 0 2 2 5 2 Chiroderma salvini 6 4 3 3 0 2 3 1 0 2 2 2 2 0 2 2 1 0 2 1 5 2 Chiroderma trinitatum 4 4 3 3 1 2 3 1 0 0 2 2 2 2 0 2 1 0 2 2 5 2 Chiroderma villosum 4 3 3 1 0 2 3 2 0 3 2 2 1 1 2 2 1 0 2 2 5 2 Ectophylla alba 3 2 1 0 1 2 1 0 1 1 2 3 4 0 2 1 1 0 1 0 5 2 Enchisthenes hartii 4 2 2 1 2 2 3 0 1 1 2 3 3 2 0 2 1 0 1 0 1 0 Mesophylla macconnelli 4 4 3 0 1 2 3 1 1 3 2 1 3 0 2 1 1 0 1 1 3 1 Phyllops falcatus 6 2 3 5 2 2 3 0 2 2 2 3 2 3 1 1 0 1 0 0 1 0 Phyllops haitiensis 6 1 3 5 2 2 1 0 2 1 2 3 1 3 1 1 0 1 0 0 1 0 Phyllops vetus 6 2 3 5 2 2 3 0 2 2 2 3 3 2 1 2 1 1 0 0 1 0 Pygoderma bilabiatum 6 2 3 5 0 2 3 0 1 2 2 3 2 3 2 3 1 0 0 1 5 2 Sphaeronycteris toxophyllum 6 1 3 5 1 2 2 0 2 1 2 2 2 3 ] 2 1 0 1 2 1 1 Stenoderma rufum 4 5 3 5 1 2 4 0 2 2 1 3 2 2 1 2 0 1 1 3 1 0 Sturnira aratathomasi 4 3 3 0 3 1 4 0 1 1 1 3 3 0 1 2 1 0 1 1 1 0 Sturnira bidens 4 3 3 0 3 1 3 0 1 2 1 1 2 0 0 2 1 0 1 0 2 0 Sturnira bogotensis 4 2 3 0 3 0 3 0 1 1 1 2 3 0 0 1 1 0 1 1 1 0 Sturnira erythromos 4 2 3 0 3 1 3 0 1 1 1 2 4 0 0 2 1 0 1 0 1 1 Sturnira lilium 3 2 3 0 3 1 3 0 1 2 1 3 3 0 1 1 1 0 1 1 1 0 Sturnira ludovici 4 2 3 0 3 1 3 1 1 1 1 3 2 0 0 1 1 0 1 1 1 0 Sturnira luisi 4 2 3 0 3 1 3 1 1 1 1 2 3 0 0 2 1 0 1 1 1 0 Sturnira magna 4 2 3 0 3 1 3 0 0 2 1 3 3 1 1 2 1 0 1 0 1 1 Sturnira mordax 4 2 3 0 3 1 3 0 1 1 1 3 3 0 0 1 1 0 1 1 1 0 OWEN—PHYLOGENETIC ANALYSES OF STENODERMATINAE 9 Table 1. —Continued. Character Species 1 2 3 4 5 6 7 8 9 10 11 12 13 H 15 16 17 18 19 20 21 22 Sturnira nana 4 3 3 0 3 2 3 0 0 2 1 2 2 0 0 1 1 0 1 0 2 0 Sturnira thomasi 4 2 3 0 3 0 3 0 0 1 1 3 4 0 0 1 I 0 1 1 8 2 Sturnira tildae 4 2 3 0 3 1 3 0 1 1 1 3 4 0 0 2 1 0 1 1 1 0 Uroderma bilobatum Uroderma 6 4 2 3 1 2 1 0 0 2 2 3 1 3 ] 2 1 0 0 0 1 1 magnirostrum 4 3 2 3 1 3 1 0 0 2 2 3 2 2 0 2 1 0 0 0 1 0 Vampyressa bidens 4 5 3 4 1 2 3 1 0 3 2 2 2 0 2 2 1 0 1 1 7 1 Vampyressa brocki 4 4 3 1 1 2 3 1 0 3 2 2 2 0 2 2 1 0 1 I 5 2 Vampyressa melissa 4 5 3 1 2 2 1 1 0 2 2 2 4 0 2 2 1 0 1 1 3 1 Vampyressa nymphaea 4 4 3 2 1 2 3 1 1 2 2 1 2 0 2 2 1 0 1 1 5 2 Vampyressa pusilla 4 5 3 1 1 2 3 1 1 2 2 2 3 0 2 2 1 0 ] 1 5 2 Vampyrodes caraccioli 4 2 2 4 2 2 3 1 0 2 2 2 3 1 2 3 1 0 1 2 3 0 Vampyrops aurarius Vampyrops 4 5 2 4 2 2 3 1 0 I 2 1 3 0 1 2 1 0 1 1 1 0 brachycephalus 6 4 2 4 2 2 3 1 1 2 2 1 3 1 1 2 1 0 1 1 1 0 Vampyrops dorsalis 4 3 3 4 2 2 3 1 0 1 2 2 2 ] 1 2 1 0 1 2 1 0 Vampyrops helleri 4 4 2 4 2 3 1 1 ] 2 2 1 3 1 1 2 1 0 1 1 1 0 Vampyrops infuscus 4 5 2 3 2 2 3 1 0 2 2 2 3 0 1 2 1 0 1 1 1 0 Vampyrops lineatus 6 3 2 4 2 2 3 1 1 2 2 2 2 0 1 2 1 0 1 1 1 0 Vampyrops nigellus 6 4 2 4 2 2 3 1 ] 1 2 I 3 1 1 2 1 0 1 1 1 0 Vampyrops recifinus 3 3 2 2 1 2 1 1 1 2 2 1 3 0 1 2 0 0 1 1 1 0 Vampyrops umbratus 3 5 2 3 2 2 3 1 1 1 2 1 2 1 1 2 1 0 1 1 1 0 Vampyrops vittatus 4 4 2 4 2 2 3 1 1 1 2 3 3 0 1 2 1 0 1 1 1 0 Carolliinae Carollia brevicauda 4 4 3 0 0 0 3 1 0 2 1 2 4 0 0 2 1 0 0 0 1 0 Carollia castanea 4 3 3 0 0 0 1 1 0 2 1 2 4 0 0 2 1 0 0 0 1 0 Carollia perspicillata 4 3 3 0 0 0 1 1 0 2 1 2 3 0 0 1 ] 0 0 0 1 0 Carollia subrufa 4 2 3 0 0 0 1 1 0 3 1 2 3 0 0 1 1 0 0 0 1 0 Phyllostominae Macrotus waterhousii 8 2 2 0 0 0 5 1 1 3 1 1 2 1 0 2 1 0 1 0 0 0 I used the WAGNER78 computer program supplied by J. S. Farris. For each analysis (all taxa together first, and then successively smaller subsets of the taxa), I ran the Wagner program 50 times, each time entering the taxa in a different order. This is especially crucial in initial analyses of all taxa because of the vast number of possible tree topologies, and concomitant low likelihood of finding the shortest in a particular run. Another important aspect of this procedure is that it allows the investigator to examine a large number of tree topologies for “stable” portions (that is, portions that do not differ among the alternate trees being examined). This procedure thus adds a robustness to the phylogenetic estimate and, simultaneously, identifies portions of the tree in need of additional analysis. For any particular analysis, I arranged the results in order of total tree length and examined the shortest 25 of the 50 trees plus any others of the 10 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY same length as those included in the group. This group is referred to as the “shortest half” of the trees. I considered as stable only those portions of the tree that did not differ among the shortest half of the trees. The initial analysis established the major clades among stenodermadne bats; successive analyses then were performed on “unstable” (thus unresolved) portions of the tree. In each case, a monophyletic portion of the tree was analyzed and, following Maddison et al. (1984), the ancestral state was estimated using those two taxa found to be the shortest distance from the hypothetical ancestor of the clade being reanalyzed. Although this method of repeated ancestor estimation for successively smaller portions of the tree might be suspected of leading to accumulated error, it should be emphasized that no locally parsimonious solution was accepted that was not concordant with one of the globally parsimonious solutions previously determined (that is, the shortest half of the trees found in the initial analysis). Analysis using Weighted Invariant Step Strategy .—In contrast to the Wagner method, the Weighted Invariant Step Strategy (WISS) is a "top- down” type of clustering algorithm (that is, the terminal taxa with the greatest degree of similarity are clustered first, on the assumption that their shared similarities are uniquely derived). Farris et al. (1970) first formalized this method and discussed its relationship to parsimony (including Wagner) methods. The procedure, simply stated, is to select a subset of species for inclusion in a monophyletic group that is “best founded” in the sense that it shares at least as many derived steps as does any other possible subset of species; delete the subset from the set of species under consideration; replace it with the Hypothetical Taxonomic Unit (HTU) that represents the most recent common ancestor of the members of the subset; and repeat the process until the tree is specified. The program used was HEN NIG, a Fortran IV program supplied by J. S. Farris. Analyses of Continuous Characters Considerable controversy exists concerning the value of and methodologies for use of continuous (mensural) data in phylogenetic analyses. Wood (1983) pointed out that there is substantial information in such characters, but the problem is to extract mathematically the information that distinguishes systematically between taxa from the overall body of information contained in the data set. I use “information” to mean “stored information” rather than the “potential information” of Weaver (1949) or the “entropy” of Shannon (1949). In his studies of cranes, storks, and grassland sparrows, Wood (1983) found that the useful information typically represented only five to 10 percent of the total variation in the data set, and that the remaining 90 to 95 percent of the variation was common to all members of the data set and, hence, not useful. Below I describe a series of transformations and other manipulations of mensural characters that I believe allow a credible phylogenetic analysis from such a data set. OWEN—PHYLOGENETIC ANALYSES OF STENODERMATINAE 11 As pointed out by Jolicoeur et al. (1984), “the statistical model generally most appropriate in biological morphometries is the lognormal distribution (in its univariate, bivariate, or multivariate versions).’’ Accordingly, all values in this study first were transformed to their natural logarithms. As biological size relationships typically are allometric, the log transformation has the effect of linearizing (but not reducing) the effect of size in the data matrix. The first principal component, therefore, will have a higher eigenvalue than it would have without the log transformation, and the remaining components will reflect less of the size relationships among taxa. The log transformation has the additional function of legitimizing arithmetic means and other linear statistics of the data (Humphries et al., 1981). As sexual dimorphism is known to exist to varying degrees among stenodermatine bats, the number of characters was doubled by using the male and female mean value of each character as separate characters (Table 2). Although this results in mostly redundant variation, the nonredundant portion (that is, the part reflecting sexual dimorphism) may have systematic usefulness (see Schnell et al., 1978). The result of the procedures described above is a matrix of appropriately transformed and averaged character-state values for each taxon. The remaining problem is to extract the useful information from the matrix and, from this reduced matrix, to construct an estimate of the phylogeny. I used two methods of data reduction to extract useful information. Size-free analysis .—One method used was a “size-out” procedure. After averaging (for each sex) the log-transformed values, a matrix of character correlations was calculated, and a principal components analysis was performed on the matrix. From this, the matrix of projections of each species on each component was calculated based on unstandardized data. The vector of projections on the first component was deleted from the matrix, and the remaining vectors of projections were taken to be character- state values for a newly created suite of characters; thus, the effect of principal component I is removed from the matrix. In the case of the transformation described above, this component mostly reflects size relationships. More importantly, the converse also is true, and the remaining matrix should contain essentially size-free information. A Wagner analysis was then performed on this matrix. Unlike the case of discrete-state information, “ties” in calculated distances (and, thus, in the order of inclusion of species in the tree) are highly improbable; therefore, the same tree is generated regardless of order of species entry into the tree-generating procedure. This single tree then represents the best phylogenetic estimate for the group based on the size-free data matrix. Common-part-removed analysis .—Wood (1983) described a transformation for phenetic analysis of continuous data. This transformation involves regression of the vector of the character values of each species studied on the character-value vector(s) of one or more closely related species. 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S 3 i iri ^ S in to ao 1 4 ao — — ■ — — CM CM 1 !2S§ i on on on SiSS ^ on cm 3 8 ' an on 1 © © t- 1 M’ M* — • in 3 3 © P^ to on CM CM CM r-t-r*Qinmo>mcM 3 3 on on on to to O cm on CM cm oi ad s 8; oi Oi ■ m m in n* O CM l CM tO id id t ■ — CM O tO o to 'tr* m — o cm — on cm to -«r on — CM CM CM — — — — -OCMCMOOOaOP^CMtO tcMt'-maotocM — ©ao minp^p^minonr^intoto Xf « xr M « O — CM © CM — C4CMCMCMCMCMCMCM — CMCM flOODiflNMnpPOOOOt htfj-qojflj Ectophylla Uroderrna -Artibeus 42 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Simodermaiinac Ectophytla /\ / Mesnphylta / / Hu. pusitla Vn. pusitla I Vampymsa’ I ChiTodrrma Pygodmna Atdops Phyllopi Arileui / I \ / | Cenlurio Sphartonyclmi \!/ I'ampyrofM* - Slumira* — Hiw/jyro/jj*- Slumira* Artibrus* Yampyrodfs B "•Pygoderma •Spharronycleris Centurio ■ChiToderma Vampyrrssa Mrsophylla Fig. 18.—Phytogenies (redrawn) from tour published arrangements of stenodermatine genera. (A) de la Torre (1961: fig. 4); (B) Baker (1973: fig. 5); (C) Smith (1976: fig. 2); (D) Gardner (1977: fig. 8). Asterisk indicates only part of genus in a particular position. Dashed line indicates alternate or uncertain phytogeny. species level for S. mordax —recognition of Sturnirops is not supported even as a subgenus. Within Vampyrops , Fig. 17 indicates at least two lineages. One contains V. nigellus, V. helleri, and V. dorsalis ; the other includes V. vittatus, V. aurarius, V. lineatus, and V. umbratus . The status of the remaining three species is unclear. Although Gardner and Carter (1972a) agreed with Sanborn (1955) in considering V. umbratus a synonym of V. dorsalis, my study indicates that Panamanian and Colombian specimens (probably OWEN—PHYLOGENETIC ANALYSES OF STENODERMATINAE 43 referable to the name umbratus) represent a species distinct from V. dorsalis of Peru. Tuttle (1970), in a review of Peruvian bats, included V. infuscus within V. vittatus, but Gardner and Carter (1972a) and Koopman (1978) recognized V. infuscus as specifically distinct. Koopman also disagreed with the description by Gardner and Carter (1972b) of V. nigellus as a distinct species; Koopman regarded this taxon as a subspecies of V. lineatus. Although my results provide little insight into the distinctness of V. infuscus, they do show V. lineatus and V. nigellus in different clades, implying full specific status for V. nigellus. The relationship of Uroderma to other stenodermatine genera has been a matter of dispute at least since Burt and Stirton (1961) suggested (without explanation) that the genus probably should be regarded as a subgenus of Artibeus. They noted that "the differences are rather subtle,” but their suggestion has not been generally accepted, particularly in light of Baker’s (1967, 1973) finding of karyotypic differences that separated Uroderma from all other stenodermatines known karyotypically at that time. Based presumably on this and other evidence, Smith (1976) arranged Uroderma as the sister taxon to Vampyressa, Vampyrodes, and Vampyrops. Gardner (1977), who also examined standard karyotypes, considered a high diploid number to be the primitive condition and placed Uroderma essentially as an outgroup to the remaining stenodermatines (including Sturnira). In studies of G- and C-banded chromosomes to determine homologous segments, Johnson (1979) and Baker et al. (1979) again found Uroderma to be quite different from Artibeus and, in fact, were unable to ascertain a close relationship between Uroderma and any other group. An analysis of electrophoretic data (Koop and Baker, 1983) also failed to find Uroderma to be closely related to any of the six Artibeus species included in the study. In my results, Uroderma appears to be a sister taxon to the species group of the large Artibeus and Ectophylla. The status of the genus certainly requires further study, but it is prudent at present to retain generic rank for Uroderma. Eight of the 17 stenodermatine genera belong to a group of short-faced bats with characteristic white epaulettes. All but one of these genera currently are regarded as containing but one species (Jones and Carter, 1976; Honacki et al., 1982). Four of the genera are Antillean endemics, and the remainder occur on the mainland from Mexico to Argentina. Investigators have not agreed on the interspecific relationships of the group, or whether the 10 living species even constitute a natural assemblage. Jones and Schwartz (1967), in their review of the genus Ardops, stated that the four Antillean genera ( Ardops , Ariteus, Phyllops , and Stenoderma) are related, with Ardops probably most closely related to Ariteus. Koopman (1968) agreed that they are all closely related, but noted that, " Ardops seems rather closely related to, and possibly congeneric with, Phyllops. ...” He also SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY 44 postulated that the derivation of the group was probably from a form of Artibeus or a close relative. Baker (1973) believed that the eight genera comprised a natural assemblage, but his chromosomal data suggested that Stenoderma was less closely related to Ardops, Phyllops, and Ariteus than to some of the mainland genera. Pygoderma was depicted (Baker, 1973: fig. 5) as being the closest mainland relative of the Antillean genera Ardops, Phyllops, and Ariteus, and the entire lineage of eight genera was postulated as having been derived (along with Enchisthenes) from the Artibeus group possessing the 30-31 diploid number. Nagorsen and Peterson (1975) published a karyotype of P. haitiensis and agreed with Baker’s (1973) assessment of the phylogeny of this group. Varona (1974), however, followed Simpson (1945) in synonomizing Ariteus, Ardops, and Phyllops under the genus Stenoderma, thus making all Antillean species congeneric. Greenbaum et al. (1975), with additional standard karyotype data, stated that the four Antillean genera represent a single invasion from the mainland and, further, that the eight short-faced genera together share a common evolutionary history exclusive of other stenodermatines. Smith (1976) agreed that the Antillean species share a common ancestor, but believed them to be more closely related to Artibeus and Enchisthenes than to the mainland short-faced taxa. Gardner (1977) agreed with Baker’s (1973) exclusion of Stenoderma from the monophyletic Antillean assemblage, indicating that this group might instead be more closely related to the mainland Ametrida. Baker and Genoways (1978) restated the belief that the four Antillean genera form a monophyletic group, and suggested that it is the second oldest bat taxon in the Caribbean region (after the endemic subfamily Phyllonycterinae). Johnson (1979) included a cladogram, based on G- and C-banded chromosomes, in which Phyllops was allied with Artibeus and Vampyrops, but not with Centurio, suggesting that the eight short-faced genera are not monophyletic. The questions of systematic relationships among these eight genera thus can be reduced to two general categories—the monophyly of Antillean genera, and the monophyly of the group overall. My results (Fig. 17) indicate that the eight short-faced genera form a monophyletic assemblage (although one possible interpretation of this tree also involves the inclusion of Enchisthenes or Artibeus concolor, or both, in the lineage). The positions of Pygoderma, Centurio, and Stenoderma are unresolved in the clade, indicating that these data at least do not support synonomy of the Antillean species under the generic name Stenoderma. The relation of Ardops and Ariteus indicated in Fig. 17 is in agreement with the statement of Jones and Schwartz (1967). The close association of these two genera with Phyllops suggests that if the number of Antillean genera of this group were to be reduced, it should be reduced to two, with Ardops and Phyllops placed in synonomy under the older name Ariteus. However, given the OWEN—PHYLOGENETIC ANALYSES OF STENODERMATINAE 45 uncertain position of Stenoderma on the tree, and the continued uncertainty concerning the phylogeny of the mainland species, I believe it best in the interest of nomenclatorial stability to retain, for the present, the generic names Ardops and Phyllops. Vampyressa and Vampyriscus, originally described as subgenera of Vampyrops (Thomas, 1900), both were raised to full generic rank by Miller in 1907. Simpson (1945) evidently disagreed, as he included both of these names (as well as Vampyrodes) within Vampyrops. Goodwin (1963), in a review of Vampyressa, did not include Vampyriscus bidens, considering it to be generically distinct. Peterson (1968), in another review, included Vaynpyriscus as a subgenus of Vampyressa, and described a new subgenus Metavampyressa to include V. ( M .) nymphaea and V. (M.) brocki. Davis (1975), in a review of variation in V. bidens, placed Metavampyressa in synonomy under Vampyriscus, which he retained as a subgenus. In 1968, Starrett and Casebeer departed from currently accepted taxonomy and returned Mesophylla to generic status, pointing out that the skull of M. macconnelli shows greater similarity to Vampyressa pusilla thyone than to Ectophylla alba. Mesophylla, described by Thomas (1901), had been placed in synonomy with Ectophylla by Laurie (1955) and Goodwin and Greenhall (1962). Starrett and Casebeer (1968) agreed that the three genera form a well-defined group, but regarded Mesophylla as sufficiently dissimilar from Ectophylla to be accorded generic rank. Based primarily on standard karyotypic data, Baker (1973) suggested that Mesophylla was derived from the Vampyressa lineage (which in turn was derived from Chiroderma). Without knowing the karyotype at the time, he assumed that Ectophylla was closely related to Mesophylla. Also, Baker et al. (1973) found that standard karyotypic data indicate that V. pusilla is more closely related to Mesophylla than to the other Vampyressa species then known karyotypically. Greenbaum et al. (1975), reporting on the karyotype of Ectophylla, found it to be unlike that of Mesophylla or any species of Vampyressa, and suggested that Ectophylla might be more distantly related to the Vampyressa-Mesophylla stock than is the genus Chiroderma. Nevertheless, Jones and Carter (1976) retained Ectophylla macconnelli in their checklist, and implied that Mesophylla might have validity as a subgenus. In the same volume, Smith’s (1976) phylogeny showed Mesophylla and Ectophylla as sister taxa, with Vampyressa in a separate clade along with Vampyrops and Vampyrodes. Gardner (1977), in a review of chromosomal variation in Vampyressa, pointed out that the subgenera Vampyressa and Vampyriscus appear to be karyotypically valid. He also commented that the karyotypes of Peruvian V. pusilla (at least three karyotypic variants are known in this species) and Mesophylla are “remarkably similar,” but declined to relegate Mesophylla to synonomy within Vampyressa. Recently, Anderson et al. (1982) recognized Ectophylla macconnelli, noting that recognition of Mesophylla as a distinct genus seems to be based more on karyology than on 46 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY morphology. Both treatments have been used in recent years and the question is still open. A suite of interrelated questions thus remains concerning these genera, the status of the two currently recognized subgenera of Vampyressa, and the relationship of Mesophylla to this genus and to Ectophylla. My results for the Vampyressa species are the least well resolved of any group, as they do not even confirm the monophyly of the genus. Nor is the subgeneric question answered here, although there is a suggestion that the distinction is not valid. The status of Mesophylla is quite clear, however, both from morphologic data and previously cited chromosomal data. Although the species of Vampyressa are not well resolved in Fig. 17, any interpretation of this tree indicates that if Vampyressa is a natural assemblage it includes Mesophylla macconnelli. Additionally, Fig. 17 does not strongly support subgeneric status for Mesophylla. I recommend therefore, that Mesophylla be placed in synonomy under Vampyressa, with the proper name of the single species being Vampyressa macconnelli. See Appendix V for additional nomenclatorial comments concerning taxa referable to this species. Ectophylla is placed in Fig. 17 in association with Uroderma and the large species of Artibeus. Neither these results nor any previous work have questioned the generic validity of Ectophylla as applied to the single species E. alba, and this combination should remain in use. Artibeus is the most species-rich genus currently recognized within the subfamily Stenodermatinae. The content of the genus was clarified by Miller (1907), who excluded from it the taxa referable to Uroderma and, agreeing with Andersen (1906, 1908), the one species referable to Enchisthenes. Miller also stated that he saw no reason to recognize subgenera within Artibeus. Simpson (1945) evidently regarded the differences between Enchisthenes and Artibeus to be minor, as he included E. hartii as a species of the latter. Gardner (1977), based on his own and other work (for example, Baker 1967, 1973), retained Enchisthenes at the generic level, showing it as the sister taxon to Artibeus. Koopman (1978) and Anderson et al. (1982), without explanation, synonomized Enchisthenes with Artibeus, as did Jones and Carter (1979) and Honacki et al. (1982), although Hall (1981) did not. The last full revision of the genus Artibeus was by Andersen (1908). Subsequent systematic work on the smaller taxa of the genus has been undertaken by (among others) Hershkovitz (1949), Dalquest (1953), Davis (1958, 1970), Baker (1973) and Koopman (1978). The larger species of Artibeus also have been studied by a number of workers, including Patten (1971), Baker (1973), Koopman (1978), Myers and Wetzel (1979), Anderson et al. (1982), Davis (1984), and Koepcke and Kraft (1984). Several recent studies have examined the relationship of the genus Artibeus to other stenodermatine taxa. Baker (1967) found Artibeus to be similar karyotypically to Vampyrops and Sturnira, and quite unlike Uroderma. Within the genus, he found A. turpis (= phaeotis) to differ from OWEN—PHYLOGENETIC ANALYSES OF STENODERMATINAE 47 A. jamaicensis, A. lituratus, and A. toltecus in the form of a chromosomal sex-determining mechanism. Straney et al. (1979) presented electrophoretic analyses of several phyllostomid bats that suggested Artibeus might be polyphyletic. Koop and Baker (1983), analyzing 22 electrophoretic loci, found apparent diphyly in Artibeus, with A. concolor and A. jamaicensis separate from the four small Artibeus species examined in the study. They also noted that A. concolor and A. jamaicensis were separated by relatively large genetic distance. The results of my morphologic analyses strongly indicate that Artibeus, as presently understood, is a polyphyletic assemblage (Fig. 17). The large species constitute a natural group, which includes the populations recognized under the names A. inopinatus, A. hirsutus, A. jamaicensis, A. planirostris, A. fraterculus, A. intermedius (and presumably A. lituratus), as well as the Paraguayan taxon referred to in this study as A. fimbriatus. These bats should continue to be referred to under the generic name Artibeus. The second natural group of Artibeus bats includes the species now recognized under the names A. watsoni, A. glaucus, A. toltecus, A. cinereus, A. phaeotis, A. aztecus, and A. anderseni, and possibly also Enchisthenes hartii and A. concolor. The oldest available generic name for this group is Dermanura Gervais 1855, and I recommend this name be applied to these bats at the generic level. Specific names ending in “us” would be changed to agree: D. glauca, D. tolteca, D. cinerea, and D. azteca. As pointed out above, the relationships of A. ( Enchisthenes ) hartii and A. concolor to the Dermanura bats are yet unclear. Koop and Baker (1983) and Tandler et al. (1986), for instance, noted that A. concolor is unlike the other species of Artibeus used in their studies. The generic validity of Enchisthenes has been in dispute since Andersen (1906) removed if from Artibeus, under which it was originally described. In the interest of stability I follow recent authors (for example, Koopman 1978; Jones and Carter, 1979; Anderson et al., 1982), who have not recognized Enchisthenes at the generic level. Thus, until additional evidence is available, I suggest applying the name Dermanura to both hartii and concolor, as well as the other species listed above. The name combinations for the affected species thus would be Dermanura hartii, Dermanura concolor, Dermanura watsoni, Dermanura glauca, Dermanura tolteca, Dermanura cinerea, Dermanura phaeotis, Dermanura azteca, and Dermanura anderseni. See Appendix V for additional nomenclatorial comments concerning taxa referable to Demanura. Within the reduced genus Artibeus, the question remains of the relationships and possible conspecific status of the South American taxa. My results (Fig. 17) suggest, but do not confirm, that neither A. planirostris nor A. fraterculus is conspecific with A. jamaicensis, and that A. hirsutus and A. inopinatus are not conspecific, as has been suggested (Honacki et al., 1982). My results strongly indicate that A. fuliginosus is specifically 48 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY distinct from A. jamaicensis, notwithstanding the comments of Anderson et al. (1982). Within the genus Dermanura, the focal question concerns D. glauca and D. watsoni, and their relationship to D. cinerea. Koopman (1978), Jones and Carter (1979), and Anderson et al. (1982) all judged glaucus and watsoni to be synonyms of A. cinereus. My results suggest that this may not be the case, but do agree with statements by the same authors that D. anderseni is specifically distinct from D. cinerea. Based on data and analyses leading to Fig. 17, I recommend that Mesophylla macconnelli be considered as congeneric with species of Vampyressa, and that the genus Artibeus (including Enchisthenes) be split into two genera, with the smaller taxa referred to under the name Dermanura . The recommended generic classification of all stenodermatine species is listed in Appendix VI, and also is indicated in Fig. 17. Acknowledgments This research was completed as part of my Ph.D. program at the University of Oklahoma. I am grateful to my committee members—Gary D. Schnell (Chair), Cluff E. Hopla, Michael A. Mares, W. Alan Nicewander, and Raymond B. Phillips—for their assistance with the completion of the dissertation. J. Knox Jones, Jr., also reviewed this manuscript and provided guidance, both editorial and systematic. I am most appreciative of June Logan for undertaking the task of typing the manuscript. Curators and other personnel of the museums listed in Appendix I were all helpful and cordial during my visits and in response to loan requests. This research was supported by National Science Foundation dissertation improvement grant DEB-7814193 and a Sigma Xi Grant-in-Aid of Research. Literature Cited Adams, E. N., III. 1972. Consensus techniques and the comparison of taxonomic trees. Syst. Zool., 21:390-397. Andersen, K. 1906. Brief diagnosis of a new genus and ten new forms of stenodermatous bats. Ann. Mag. Nat. Hist. ser. 7, 18:419-423. -. 1908. A monograph of the chiropteran genera Uroderma, Enchisthenes, and Artibeus. Proc. Zool. Soc. London, 1908:204-319. Anderson, S., K. F. Koopman, and G. K. Creighton. 1982. Bats of Bolivia: an annotated checklist. Amer. Mus. Novit., 2750:1-24. Arnold, M. L., R. L. Honeycutt, R. J. Baker, V. M. Sarich, and J. K. Jones, Jr. 1982. 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S. 1974. Catalogo de los mamiferos vivientes y extinguidos de las Antillas. Acad. Cien. Cuba, viii+139 pp. OWEN—PHYLOGENETIC ANALYSES OF STENODERMATINAE 53 Walton, D. W., and G. M. Walton. 1968. Comparative osteology of the pelvic and pectoral girdles of the Phyllostomatidae (Chiroptera:Mammalia). J. Grad. Res. Center, Southern Methodist Univ., 37:1-35. Watrous, L. E., and Q. D. Wheeler. 1981. The out-group comparison method of character analysis. Syst. Zool., 30:1-11. Weaver, W. 1949. Recent contributions to the mathematical theory of communication. Pp. 3-28, in The mathematical theory of communication (C. E. Shannon and W. Weaver, eds.), Univ. Illinois Press, Urbana, 125 pp. Wiley, E. O. 1981. Phylogenetics. The theory and practice of phylogenetic systematics. John Wiley & Sons, New York, xv+439 pp. Wilson, D. E. 1973. Bat faunas: a trophic comparison. Syst. Zool., 22:14-29. Winge, H. 1892. The interrelationships of the mammalian genera. Vol. I. Monotremata, Marsupialia, Insectivora, Chiroptera, Edentata (E. Deichmann and G. M. Allen, translators, 1941), C. A. Reitzels Forlag, Kobenhavn, 418 pp. Wood, D. S. 1983. Character transformations in phenetic studies using continuous morphometric variables. Syst. Zool., 32:125-131. 54 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Appendix I—Specimens Examined An asterisk indicates the type specimen of the species. Museum acronyms used are as follows: AMNH, American Museum of Natural History; CMNH, Carnegie Museum of Natural History; FMNH, Field Museum of Natural History; KU, Museum of Natural History, University of Kansas; LACM, Los Angeles County Museum of Natural History; LSU, Museum of Zoology, Louisiana State University; MCZ, Museum of Comparative Zoology, Harvard University; MSB, Museum of Southwestern Biology, University of New Mexico; MSU, The Museum, Michigan State University; MVZ, Museum of Vertebrate Zoology, University of California; OU, Stovall Museum of Science and History, University of Oklahoma; ROM, Royal Ontario Museum; TCWC, Texas Cooperative Wildlife Collections, Texas A&M University; TTU, The Museum, Texas Tech University; UA, Department of Biological Sciences, University of Arizona; UMMZ, Museum of Zoology, University of Michigan; USNM, United States National Museum of Natural History. Number in parentheses after localities indicates the number of specimens examined from that locality. Number in parentheses after species name indicates total number examined. Stenodermatinae Ametrida centurio (17)—Trinidad (11): CMNH 45335; TTU 5215, 8766, 8769, 8771-72, 8811, 8888, 9545, 9548, 9557. Suriname (6): CMNH 63767, 63769, 63771-73, 68418. Ardops nichollsi (14)—Montserrat (2): ROM 71463, 71467. Guadeloupe (3): TTU 20801, 20805, 20810. Dominica (7): KU 104805-10, 104813. Martinique (1): AMNH 213951. St. Lucia (1): USNM 110921. Ariteus flavescens (13)—Jamaica (13): TTU 21714-21,21763, 21772; USNM 534909-11. Artibeus anderseni (10)—Bolivia (10): AMNH 209583, 210825-26, 210828-30, 210834, 210836- 37, 210845. Artibeus aztecus (13)—Mexico (4): AMNH 203753-54, 203757; USNM 52050*. Guatemala (1): TCWC 17507. Costa Rica (7): LSU 12876; TTU 12907-08, 12910-11, 12913-14. Panama (1): USNM 323455. Artibeus cinereus (10)—Trinidad (10): TTU 5450, 5541, 5660, 5672, 5679, 8968-69, 8971, 8973, 8975. Artibeus concolor (10)—Colombia (1): TCWC 26046. Venezuela (8): USNM 387379, 387381- 82, 387386-88, 2 numbers lost. Brazil (1): AMNH 78859. Artibeus fimbriatus (10)—Paraguay (10): UMMZ 125914, 125936, 125938, 125943-44, 133721, 133724-27. Artibeus fraterculus (10)—Ecuador (10): AMNH 47248*; USNM 498940-41; 522472, 522474-75, 522478-81. Artibeus fuliginosus (10)—Peru (10): USNM 364439, 499209-14, 499216-17, 499220. Artibeus glaucus (16)—Ecuador (3): TCWC 12278, 12280-81. Peru (13): AMNH 37196-97, 214361,233750-51, 236067; LSU 16602-05, 19184, 19186, 19706. Artibeus hartii —see Enchisthenes hartii;. Artibeus hirsutus (22)—Mexico (22): LACM 9982, 9985, 9988, 13299-301, 13303, 13804, 34988, 58424; MSB 18407; TTU 8697, 8700-04, 10592-93; UA 6348-49; USNM 126449*. Artibeus inopinatus (13)—Honduras (7): TCWC 9517*, 18407-08; TTU 7685, 7688-90. Nicaragua (6): TTU 12916, 12918, 12921-24. Artibeus intermedius (7)—Mexico (3): TTU 15564-65, 15568. El Salvador (4): TTU 12990-93. Artibeus jamaicensis (13)—Mexico (12): AMNH 12038/10469; TTU 8205, 8209-10, 8212, 8214, 8217, 8344, 10043, 10045, 15557; USNM 126554. Nicaragua (1): AMNH 28335. Artibeus lituratus —see A. intermedius . Artibeus phaeotis (12)—Mexico (1): USNM 108176*. Guatemala (1): TCWC 14392. Honduras (10): TTU 12962, 12964, 12967-69, 12971-73, 12975, 12985. Artibeus planirostris (25)—Guyana (1): TCWC 28739. Peru (24): LSU 12535-36, 12538, 12540, 12542, 12545, 12547-49, 12556, 24529, 24531; MVZ 153408-09, 154900-01, 154903-05, 154909-12 157725. OWEN—PHYLOGENETIC ANALYSES OF STENODERMATINAE 55 Artibeus toltecus (11)—Mexico (11): TCWC 6499; TTU 10022-23, 10025, 10133, 10204-05, 10207-08, 10210-11. Artibeus watsoni (9)—Nicaragua (2): TTU 12968-69. Costa Rica (4): TTU 12979-82. Panama (3): USNM 503666, 503668, 503673. Centurio senex (17)—Mexico (11): LACM 13476-80; MSB 25186; MSU 16490-91; UA 10807- OS; USNM 511741. El Salvador (1): MVZ 130911. Panama (4): USNM 310238, 310242, 310244, 503837. Trinidad (1): USNM 346847. Chiroderma doriae (2)—Brazil (2): USNM 521053, 542616. Chiroderma improvisum (2)—Montserrat (1): TTU 31403. Guadeloupe (1): TTU 19900*. Chiroderma salvini (14)—Mexico (5): MSU 16502; OU 14336-14337; TTU 9996; USNM 338711. Honduras (2): TTU 12800, 12802. Panama (7): USNM 309930-34, 309946, 338044. Chiroderma trinitatum (11)—Panama (2): USNM 309903, 335295. Trinidad (9): AMNH 175325*; TTU 5223, 5265, 5336, 5382, 5675, 8989, 9014; UA 17070. Chiroderma villosum (13)—El Salvador (10): TTU 12755, 12757-59, 12762-63, 12767, 12771, 12773-74. Trinidad (1): CMNH 45357. Suriname (2): CMNH 53793-94. Ectophylla alba (13)—Costa Rica (4): LACM 14365-66; MSB 34204, 34251. Panama (9): USNM 315563, 335317-24. Ectophylla macconnelli (16)—Trinidad (13): AMNH 186433; CMNH 45358-59; TTU 5202, 5205, 5207-09, 5211-13, 5362, 5474. Suriname (3): CMNH 63829-30, 68430. Enchisthenes hartii (18)—Mexico (5): LACM 13529-30, 55911; MSU 15391; TTU 5581. Costa Rica (1): MSB 26805. Panama (6): USNM 310214-17, 323539, 539813. Trinidad (2): TTU 5243, 5371. Colombia (1): MVZ 124036. Peru (3): LSU 21534, 27251; MVZ 135599. Mesophylla macconnelli —see Ectophylla macconnelli. Phyllops falcatus (4)-Cuba (4): AMNH 176190; USNM 123187, 143844, 300504. Phyllops haitiensis (12)—Hispaniola (12): AMNH 236691-92, 236696; ROM 72797; UMMZ 123279; USNM 520534, 538339-42, 538345, 542273. Phyllops vetus (6)—Cuba (6): AMNH 41001*, 41002-06. Pygoderma bilabiatum (52)—Dutch Guiana (1): USNM 37502/14816. Bolivia (10): AMNH 246398-99, 246401-08. Brazil (2): FMNH 94716; ROM 70910. Paraguay (38): AMNH 234288, 234290-98; MCZ 28064; MVZ 144465-66; UMMZ 124373-75, 125804-12, 125824-25, 125828-29, 125833-37, 125841-43; USNM 105685. Argentina (1): CMNH 72333. Sphaeronycteris toxophyllum (12)—Colombia (1): TTU 10277. Venezuela (9): FMNH 29445; USNM 370797-99, 370801,370839-40, 405686, 455936. Bolivia (2): AMNH 209740-41. Stenoderma rufum (10)—Puerto Rico (10): TTU 8861, 8864-66, 8869, 8875-76, 8879-80, 8884. Sturnira aratathomasi (7)—Colombia (6): ROM 70874, 70876; USNM 395158*, 501064-66. Ecuador (1): ROM 46349. Sturnira bidens (19)—Colombia (3): FMNH 58719-20, 113447. Venezuela (5): USNM 386557, 386559-60, 386567, 386570. Peru (11): AMNH 214349, 216114; LSU 14197-99, 16525, 18368, 18386, 18389, 20741-42. Sturnira bogotensis (12)—Colombia (8): AMNH 207853, 207856-59, 207861-62; USNM 251989*. Bolivia (4): UMMZ 126752-53, 126836-37. Sturnira erythromos (19)—Colombia (5): FMNH 58721-25. Peru (10): FMNH 75183, 75185, 110921; LSU 14191, 18960, 18975, 18992-94; MVZ 139521. Bolivia (4): AMNH 246569, 246571- 72; UMMZ 126835. Sturnira lilium (17)—Mexico (5): OU 14423, 14430, 14433, 14436; USNM 126555. Dominica (1): USNM 361881. St. Vincent (1): USNM 361852. Trinidad (10): TTU 5367, 5408, 5415, 5648, 5651, 5669-71,5677-78. Sturnira ludovici (14)—Mexico (13): MSU 8808; TCWC 37863, 37865, 37868-69, 37874-77; TTU 5584, 7341-42, 8221. Ecuador (1): AMNH 67328*. Sturnira luisi (11)—Costa Rica (3): TCWC 9959*, 9992, 9998. Panama (1): USNM 309565. Ecuador (4): TCWC 12112, 12120-22. Peru (3): TCWC 12125-27. Sturnira magna (14)—Colombia (2): FMNH 113282-83. Peru (12): AMNH 230624; LSU 21484; MVZ 153366-69, 153371-72, 154842-43, 154856-57. 56 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Sturnira mordax (13)—Costa Rica (13): LSU 11456, 12784; MSB 28345; TCWC 9978, 10034, 10042; UA 20814; UMMZ 111332, 111415, 112038, 112040-41; USNM 250310* Sturnira nana (11)—Peru (11): AMNH 219138, 219171-73; LSU 15683*, 16519, 16521-24; TCWC 28071. Sturnira thomasi (7)—Guadeloupe (7): AMNH 234950; ROM 68118; IT U 19904-07; USNM 361883*. Sturnira tildae (19)—Trinidad (3): AMNH 149625*; CMNH 45362; TTU 5402. Suriname (6): CMNH 63842-43, 63845-46, 63848, 63851. Peru (7): AMNH 213362, 213365, 213368-69, 213371- 72, 213374. Bolivia (2): AMNH 209408-09. Brazil (1): USNM 361658. Uroderma bilobatum (11)—Mexico (1): TCWC 16603. El Salvador (10): TTU 12651-52, 12654- 57, 12659-61, 12664. Uroderma magnirostrum (18)—Honduras (1): TCWC 17189*. Peru (7): MVZ 136498-504. Bolivia (10): AMNH 210752, 210754-56, 210758, 210760-62, 210765, 210768. Vampyressa bidens (11)—Venezuela (1): USNM 387190. Ecuador (4): AMNH 67997-98, 71656, 76090. Peru (4): AMNH 208072; LSU 21536; MVZ 136506, 153381. Brazil (2): USNM 361721, 393025. Vampyressa broeki (8)—Colombia (3): TTU 8827, 8832, 9047. Guyana (2): ROM 38515*, 59745. Suriname (3): CMNH 63871-73. Vampyressa melissa (10)—Peru (10): AMNH 233769; LSU 16580-83, 19100, 19102-03; MVZ 139983,154869. Vampyressa nymphaea (13)—Nicaragua (3): TTU 12612, 12905-06. Costa Rica (1): LSU 12829. Panama (9): USNM 309882, 309885-89, 318130, 519695-96. Vampyressa pusilla (12)—Honduras (1): TTU 12894. Nicaragua (4): TTU 12897-99, 12903. Costa Rica (2): MSB 34209; TTU 12892. Panama (4): MSB 36397; USNM 173832, 503629, 503633. Ecuador (1): USNM 535095. Vampyrodes caraccioli (15)—Mexico (5): LACM 18631-33; UA 9400-01. Panama (8): UA 7892; USNM 309771, 309773-74, 309805-06, 309813-14. Peru (2): AMNH 230654-55. Vampyrops aurarius (10)—Venezuela (10): USNM 387143-44, 387146-47, 387149, 387159, 387163*, 387168, 387171-72. Vampyrops braehycephalus (11)—Venezuela (1): USNM 408411. Suriname (3): CMNH 63874- 76. Peru (5): AMNH 230639; LSU 12165-66, 12168; TCWC 12193*. Bolivia (1): MSU 28110. Vampyrops dorsalis (10)—Peru (10): AMNH 214355, 233614, 233621; LSU 20734-35, 21451- 52, 24775-77. Vampyrops helleri (12)—Trinidad (12): OU 9808-09; TTU 5259, 5365, 5423, 5481, 5483, 5540, 5647, 5674, 5830, 5840. Vampyrops infuscus (15)—Peru (15): AMNH 230645; LSU 16402, 19095-98; USNM 364397- 405. Vampyrops lineatus (16)—Brazil (6): LACM 28041, 28044-46; USNM 391085-86. Paraguay (10): UMMZ 125879-85, 125891, 125893-94. Vampyrops nigellus (13)—Peru (9): AMNH 233646, 233686; LSU 16415*, 24773, 24783-84; MVZ 154875, 154877-78. Bolivia (4): AMNH 246616-18, 246620. Vampyrops recifinus (8)—Brazil (8): FMNH 19516; USNM 27566/545001, 27959/545002, 542611,542612-15. Vampyrops umbratus (16)—Panama (2): USNM 338025, 338027. Colombia (14): AMNH 233186-87, 235778-79; LACM 18695-99; USNM 483540-41,483544-45, 483547. Vampyrops vittatus (14)—Costa Rica (7): LSU 10220, 12891; MSB 26804; TTU 12888-91. Panama (7): USNM 309720, 336031-36. Carolliinae Carollia brevicauda (10)—Mexico (10): AMNH 189732-33, 189736, 189739-42, 189745-46, 203693. Carollia castanea (10)—Panama (10): USNM 309421-22, 309424, 309439, 309450, 309452, 309461,503521,503528, 503835. OWEN—PHYLOGENETIC ANALYSES OF STENODERMATINAE 57 Carollia perspicillata (13)—Panama (4): MVZ 114402, 118779-81. Venezuela (5): MVZ 160046- 47, 160050, 160065-66. Peru (4): MVZ 157681, 157684, 157686, 157697. Carollia subrufa (11)—Guatemala (5): USNM 502216, 502223, 502234, 502238-39. Honduras (5): TCWC 19192, 19979-82. El Salvador (1): TCWC 19978. Phyllostominae Macrotus waterhousii (18)—Isle of Pines, Cuba (10): CMNH 2376, 2379, 2381, 2383-89. Cayman Islands (6): USNM 538125, 538127, 538129, 539736-38. Jamaica (1): USNM 534883. Hispaniola (1): USNM 538336. 58 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Appendix II—Discrete-State Characters Discrete-state characters, with character-state values and tree topology for each. Nonlinear (bifurcating) character-state trees are shown in Fig. 19. External Characters 1. Ear shape. —(1) Little or no posterior emargination (folding forward of latero-posterior border of pinna); pronounced indentation of medio-anterior border of pinna to point of attachment to head. (2) About one-fifth emarginated; pronounced indentation as in state 1. (3) About one-third emarginated; pronounced indentation as in state 1. (4) About one-half emarginated; pronounced indentation as in state 1. (5) About one-half emarginated; anterior lobe on pinna at medio-anterior point of attachment to head. (6) About three-fourths emarginated; pronounced indentation as in state 1. (7) About three-fourths emarginated; no medio-anterior indentation. (8) Completely emarginated; pronounced indentation as in state 1. Tree: see Fig. 19A. 2. Leaf shape (width measured at level of posterio-dorsal attachment; length measured from tip to point of posterio-dorsal attachment).—(1) Length less than width. (2) Length about equal to width. (3) Length greater than width. (4) Length about VA times width. (5) Length greater than VA times width. Tree: 1-2-3-4-5. 3. Dorsal hair bands (number of differentially colored bands on dorsal pelage).—(1) Hair uniformly colored. (2) Two bands. (3) Three bands. Tree: 1-2-3. 4. Pelage pattern. —(1) Pelage essentially uniformly colored (this state may include shoulder patches resulting from glandular secretions). (2) Paired white or pale-colored lines from nose leaf, above eye and ear; paired lines of same color from corner of mouth to base of tragus. (3) As in state 2, plus single middorsal line on posterior half of body. (4) As in state 3, with middorsal line extending nearly to neck. (5) As in state 4, with middorsal line extending to head. (6) As in state 1, with distinct white or near-white patch on shoulder near point of attachment of antebrachial membrane. (7) As in state 6, with ribbed pattern on patagium mediad to digit 4. Tree: 7-6-1-2-3-4-5. 5. Uropatagium shape (with legs extended so that uropatagium is uniformly stretched; measured along midline).—(1) Uropatagium long, extending one-half or more of distance from sacrum to base of calcar. (2) Uropatagium shorter, extending less than one-half of distance from sacrum to base of calcar. (3) Uropatagia connected at base of sacrum. (4) Uropatagia not connected to each other. Tree: 1-2-3-4. 6. Plagiopatagial attachment to leg. —(1) Attaches to tibial region. (2) Attaches to tarsal region. (3) Attaches to metatarsal region. (4) Attaches to metatarsal-phalangeal joint. Tree: 1- 2-3-4. Cranial Characters 7. Incisive foramina dex>elopment.—{ 1) Absent or minute. (2) Small, not extending past posterior margin of canines. (3) Small (as in stale 2), located farther caudad, between levels of PI and P2. (4) Medium size, not extending farther caudad than posterior margin of PI. (5) Large, extending beyond posterior margin of PI. (6) Small (as in state 2), located between levels of C and PI. Tree: see Fig. 19B. 8. Nasal septal foramen (completely penetrates nasal septum in dorsal-ventral direction).— (1) Absent. (2) One present. (3) Two (occasionally three) present. Tree: 1-2-3. 9. Hard palate posterior extent (measured from anterior border of orbit; not including any medial posterior projection of palate).—(1) Extends farther caudad than least width of palate between orbits. (2) Extends caudad to anterior border of orbits, but no farther than least width of palate. (3) Does not extend caudad to anterior border of orbit. Tree: 1-2-3. 10. Hard palate posterior border shape. —(1) Essentially square. (2) V-shaped (sharp medial notch; lateral borders straight). (3) U-shaped (lateral borders curved; no distinct medial notch). (4) As in state 3, with median posterior projection. Tree: see Fig. 19C. OWEN—PHYLOGENETIC ANALYSES OF STENODERMATINAE 59 5 3 2 1—2-3-4—6—7-8 1—2—4—5 1-3-4 A B 6 C 5 1—2-3-4 D E 3 8 Fig. 19.— Topologies of nonlinear (bifurcating) character-state trees for discrete-state characters. Sec Appendix II for descriptions of characters and character states, as well as character-state trees for all discrete-state characters not shown here. (A) 1—ear shape; (B) 7— incisive foramina development; (C) 10—hard palate posterior border shape; (D) 16—hypoglossal foramen; (E) 21—dental formula. 11. Parapterygoid foramina (just lateral to pterygoid processes),—(1) One present (passes dorso-caudally into braincase). (2) Two present (second passes dorso-cranially). (3) Three present (anterior two are smaller, appear homologous to anterior one in stale 2). Tree: 1-2-3. 12. Paraoccipital process development. —(1) No paraoccipilal process. (2) Minimal development (little disturbance of contour of the region). (3) Moderate development (less well developed than mastoid process). (4) Pronounced development (developed as much or more than mastoid process). Tree: 1-2-3-4. 13. Basisphenoid pit development .—(1) Absent. (2) Shallow, (3) Moderate. (4) Deep. (5) Extremely deep. Tree: 1-2-3-4-5. 14. Ml hypocone dei>elopment. —(1) Absent. (2) Slight protuberance. (3) Moderately developed. (4) Strongly developed. (5) Extremely strongly developed. Tree: 1-2-3-4-5. 15. M3 development. —(1) Well developed (cusps present). (2) Poorly developed (small, peglike). (3) Absent. Tree: 1-2-3. 16. Hypoglossal foramen (in occipital condyle; canal passes cranio-ventrally out of braincase).—(1) Absent. (2) Small. (3) Medium. (4) Large. (5) Two foramina present (appears as ossified division of medium-sized foramen). Tree: see Fig. 19D. 17. Secondary foramen of occipital condyle (canal passes cranio-Iaterally into braincase).— (1) Absent. (2) One present. (3) Two present. Tree: 1-2-3. 18. Postglenoid foramen (at posterior end of zygomatic arch, above mandibular condyle; canal passing ventrally from dorsal face of zygomatic arch).—(1) Absent. (2) Present. Tree: 1- 2 . 19. External nares shape (caudad extent of nares as viewed from dorsad to skull).—(1) Bony palate barely or not visible. (2) Bony palate clearly visible. (3) Nares extend caudad to level of anterior border of orbit. Tree: 1-2-3. 20. Rostral shape (dorsal border of frontal section at anterior border of orbit).—(1) Ridged or rounded convexly. (2) Flat or slightly troughed. (3) Moderately troughed. (4) Pronouncedly troughed. Tree: 1-2-3-4. 21. Dental formula.—( 1) 2/2 1/1 2/3 3/3. (2) 2/2 1/1 2/2 3/3. (3) 2/1 1/1 2/2 3/3. (4) 2/2 1/1 2/2 3/3. (5) 2/2 1/1 2/1 2/3. (6) 2/2 1/1 2/2 2/2. (7) 2/2 1/1 1/1 3/3. (8) 2/1 1/1 2/2 2/3. (9) 2/2 1/1 2/2 3/2. Tree: see Fig. 19E. 22. Development of m3. —(1) Well developed (cusps present). (2) Poorly developed (small, peglike). (3) Absent. Tree: 1-2-3. 60 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Appendix III— Continuous Characters Continuous characters (abbreviations in capitals). 1. Skull length (SKULL)—Greatest length not including incisors. 2. Condylobasal length (CONBAS)—Occipital condyle to anterior border of incisive alveolus. S. Rostral length (ROST)—Anterior border of braincase (in orbital recess) to anterior border of incisive alveolus. 4. Lacrimal width (LACR)—Greatest breadth across lacrimal protuberances. 5. Postorbital width (POST)—Least breadth of postorbital constriction. 6. Zygomatic width (ZYGO)—Greatest breadth across zygoma (all Stenodermatines have complete zygomatic arches). 7. Mastoid width (MAST)—Greatest breadth across mastoid processes. 8. Braincase width (BRAINB)—Greatest breadth of braincase, not including mastoid or paraoccipital processes. 9. Braincase height (BRAINH)—With calipers supporting antero-ventral border of foramen magnum and the hard palate, greatest cranial height excluding sagittal crest. 10. Braincase length (BRAINL)—Anterior border of braincase (in orbital recess) to posterior- most extension of occiput, excluding occipital crest. 11. Palatal length (PALAT)—Posterior palatal notch to anterior border of incisive alveolus. 12. Maxillary toothrow length (MAXIL)—Posterior border of molar alveolus to anterior border of canine alveolus. 13. Molariform toothrow length (MOLAR)—Posterior border of molar alveolus to anterior border of premolar alveolus. 14. Palate width at canines (WCAN)—Least width across palate between canines. 15. Palate width at second molar (WMOL)—Least width across palate between M2s. 16. Mandibular fossa length (MFOSL)—Greatest antero-posterior length of fossa. 17. Mandibular fossa width (MFOSW)—Greatest latero-medial length of fossa. 18. Upper second molar length (UM2L)—Greatest antero-posterior length at crown. 19. Upper second molar width (UM2W)—Greatest latero-medial width at crown. 20. Upper canine height (UCAN)—Length, on lateral aspect, from tip to dorsal edge of cingulum (not taken on noticeably worn or broken teeth). 21. Dentary length (DENTL)—Midpoint of mandibular condyle to anterior-most point of deniary. 22. Condylocanine length (CONCAN)—Midpoint of mandibular condyle to anterior border of canine alveolus. 23. Condyle to first molar length (CONM1)—Midpoint of mandibular condyle to anterior border of lower first molar alveolus. 24. Mandibular toothrow length (TOOTH)—Posterior-most border of molar alveolus to anterior border of canine alveolus. 25. Mandibular foramen to dentary anterior (MFAD)—Anterior border of mandibular foramen to anterior-most point of mandibular symphysis. 26. Condylomolar length (CONMOL)—Midpoint of mandibular condyle to posterior-most molar alveolus. 27. Temporal moment arm length (TEMMOM)—Midpoint of mandibular condyle to tip of coronoid process. 28. Masseter moment arm length (MASMOM)—Midpoint of mandibular condyle to tip of angular process. 29. Coronoid height (CORONH)—Perpendicular height from ventral mandibular border to tip of coronoid process. 30. Angular process length (ANGPRO)—Tip of angular process to nearest border of mandibular foramen. 31. Dentary thickness (DENTTH)—Vertical height of dentary at anterior base of second molar. 32. Lower canine length (LCANH)—Length, on lateral aspect, from tip to ventral edge of cingulum (not taken on noticeably worn or broken teeth). 33. Condyle length (CONDL)—Medio-lateral length of mandibular condyle. OWEN—PHYLOGENETIC ANALYSES OF STENODERMATINAE 61 Appendix IV—Linnaean Classification Annotated Linnaean classification of the species of the subfamily Stenodermatinae based on Fig. 17. Classification conventions (including sequencing) follow Wiley (1981). This classification disregards the “incertae sedis ” status of a number of taxa in Fig. 17, and is listed here for heuristic and comparative purposes only. For proposed generic classification of the Stenodermatinae, see end of text. Subfamily Stenodermatinae (all taxa sedis mutabilis) Tribe Corvirini Genus Corvira Coruira bidens Corvira nana Tribe Sturnirini Genus Sturnira (all taxa sedis mutabilis) Unnamed subgenus Sturnira luisi Unnamed subgenus Sturnira erythromos Unnamed subgenus Sturnira ludovici Unnamed subgenus Sturnira bogotensis Unnamed subgenus Sturnira thomasi Unnamed subgenus Sturnira magna Unnamed subgenus Sturnira tildae Unnamed subgenus Sturnira mordax Subgenus Sturnira Sturnira lilium Sturnira aratalhomasi Tribe Stenodermatini (all taxa sedis mutabilis) Subtribe Enchistheneini Genus Enchisthenes Enchisthenes hartii Unnamed subtribe Unnamed genus _ concolor Subtribe Dermanurini Genus Dermanura (all taxa sedis mutabilis) Unnamed subgenus Dermanura watsoni Unnamed subgenus Dermanura glauca Subgenus Dermanura (all taxa sedis mutabilis) Unnamed infragenus Dermanura tolteca Infragenus Dermanura Dermanura cinerea Unnamed infragenus (all species sedis mutabilis) Dermanura phaeotis Dermanura azteca Dermanura anderseni 62 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Subtribe Stenodermatini (all taxa sedis mutabilis) Genus Pygoderma Pygoderma bilabiatum Genus Centurio Centurio senex Genus Stenoderma Stenoderma rufum Genus Ariteus (all taxa sedis mutabilis Unnamed subgenus Ariteus vetus Subgenus Phyllops Ariteus falcatus Unnamed subgenus Ariteus haitiensis Subgenus Ariteus Ariteus nichollsi Ariteus flavescens Genus Ametrida Ametrida centurio Ametrida toxophyllum Tribe Artibeini (all taxa sedis mutabilis) Unnamed subtribe Unnamed genus _ melissa Subtribe Vampyressatini Genus Vampyressa (all species sedis mutabilis) Vampyressa pusilla Vampyressa brocki Vampyressa bidens Subtribe Mesophyllatini Genus Mesophylla Mesophylla nymphaea Mesophylla macconnelli Subtribe Chirodermini (all genera sedis mutabilis) Genus Chiroderma (all species sedis mutabilis) Chiroderma doriae Chiroderma salvini Chiroderma trinitatum Chiroderma improvisum Chiroderma villosum Genus Vampyrodes Vampyrodes caraccioli Genus Vampyrops (all taxa sedis mutabilis) Unnamed subgenus Vampyrops infuscus Unnamed subgenus Vampyrops recijinus Unnamed subgenus Vampyrops brachycephalus Subgenus Vampyrops (all species sedis mutabilis) Vampyrops vittatus Vampyrops aurarius Vampyrops lineatus Vampyrops umbratus OWEN—PHYLOGENETIC ANALYSES OF STENODERMATINAE 63 Unnamed subgenus (all species sedis mutabilis) Vampyrops nigellus Vampyrops helleri Vampyrops dorsalis Subtribe Artibeini (all genera sedis mutabilis) Genus Eclophylla Ectophylla alba Genus Uroderma Uroderma magnirostrum Uroderma bilobatum Genus Artibeus (all taxa sedis mutabilis) Unnamed subgenus Artibeus fuliginosus Unnamed subgenus Artibeus inopinatus Subgenus Artibeus (all taxa sedis mutabilis) Unnamed infragenus Artibeus hirsutus Unnamed infragenus Artibeus planirostris Unnamed infragenus Artibeus fraterculus Infragenus Artibeus (all species sedis mutabilis) Artibeus jamaicensis Artibeus intermedium Artibeus fimbriatus 64 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Appendix V—New Binomial and Trinomial Combinations New binomial and trinomial combinations resulting from taxonomic changes proposed in text. Species and subspecies listed are those generally recognized at present. Inclusion is not intended to imply validity of the several taxa; rather, this list is presented simply for the purpose of providing the appropriate name combinations to facilitate subsequent work on these bats. The list is based on those in Jones and Carter (1976), Koopman (1978), and Anderson et al. (1982). Dermanura anderseni Dermanura azteca azteca Dermanura azteca major Dermanura azteca minor Dermanura cinerea bogotensis Dermanura cinerea cinerea Dermanura cinerea pumilia Dermanura cinerea solimoesi Dermanura concolor Dermanura glauca Dermanura hartii Dermanura phaeotis nana. Dermanura phaeotis palatina Dermanura phaeotis phaeotis Dermanura tolteca hespera Dermanura tolteca rava Dermanura tolteca tolteca Dermanura watsoni Vampyressa macconnelli flavescens Vampyressa macconnelli macconnelli OWEN—PHYLOGENETIC ANALYSES OF STENODERMATINAE 65 Appendix VI—Recommended Generic Classification Recommended generic classification of species of the subfamily Stenodermatinae (taxa are listed alphabetically). This generic classification also is indicated in Fig. 17 by the names to the right of the cladogram. Name combinations are provided here to facilitate subsequent taxonomic work. Ametrida centurio. Ardops nichollsi. Ariteus flavescens. Artibeus jimbriatus, Artibeus fraterculus, Artibeus juliginosus, Artibeus hirsutus, Artibeus inopinatus, Artibeus intermedins, Artibeus jamaicensis, Artibeus lituratus, Artibeus planirostris. Centurio senex. Chiroderma doriae, Chiroderma improvisum, Chiroderma salvini, Chiroderma trinitatum, Chiroderma villosum. Dermanura anderseni, Dermanura azteca, Dermanura cinerea, Dermanura concolor, Dermanura glauca, Dermanura hartii, Dermanura phaeotis, Dermanura tolteca, Dermanura watsoni. Ectophylla alba. Phyllops falcatus, Phyllops haitiensis, Phyllops vetus. Pygoderma bilabiatum. Sphaeronycteris toxophyllum. Stenoderma rujum. Sturnira aratathomasi, Sturnira bidens, Sturnira bogotensis, Sturnira erythromos, Sturnira lilium, Sturnira ludovici, Sturnira luisi, Sturnira magna, Sturnira mordax, Sturnira nana, Sturnira thomasi, Sturnira tildae. Uroderma bilobatum, Uroderma magnirostrum. Vampyressa bidens, Vampyressa brocki, Vampyressa macconnelli, Vampyressa melissa, Vampyressa nymphaea, Vampyressa pusilla. Vampyrodes caraccioli. Vampyrops aurarius, Vampyrops brachycephalus, Vampyrops dorsalis, Vampyrops helleri, Vampyrops infuscus, Vampyrops lineatus, Vampyrops nigellus, Vampyrops recifinus, Vampyrops umbratus, Vampyrops vittatus. TEXAS TECH UNIVERSITY PRESS SELECTED TITLES IN CHIROPTERAN BIOLOGY The following titles in chiropteran biology are in print and can be purchased from the Texas Tech University Press, Sales Office, Box 4139, Lubbock, TX 79409 USA. Copies of Special Publications may be purchased on standing order or by individual titles. The Occasional Papers series is available by subscription ($16.00 per year for individuals, $19.00 for institutions) or by individual titles. Special Publications of The Museum (ISSN 0149-1768) No. 1 L. C. Watkins, J. K. Jones, Jr., and H. H. Genoways. 1972. Bats of Jalisco, Mexico. 44 pp. $2.00. No. 10 R. J. Baker, J. K. Jones, Jr., and D. C. Carter, eds. 1976. Biology of bats of the New World Phyllostomatidae. Part I. 218 pp. $8.00. No. 13 R. J. Baker, J. K. Jones, Jr., and D. C. Carter, eds. 1977. Biology of bats of the New World Phyllostomatidae. Part II. 364 pp. $16.00. No. 15 D. C. Carter, and P. G. Dolan. 1978. Catalogue of type specimens of Neotropical bats in selected European museums. 136 pp. $8.00. No. 16 R. J. Baker, J. K. Jones, Jr, and D. C. Carter, eds. 1979. Biology of bats of the New World Phyllostomatidae. Part III. 441 pp. $20.00. No. 18 C. O. Marlin, and D. J. Schmidly. 1982. Taxonomic review of the Pallid Bat, Antrozous pallidus (le Conte). 48 pp. $7.00. No. 23 M. B. Qumsiyeh. 1985. The bats of Egypt. 102 pp. $40.00. No. 24 J. O. Matson and R. H. Baker. 1986. Mammals of Zacatecas. 88 pp. $38.00 No. 26 R. Owen. 1987. Phylogenetic analyses of the bat subfamily Stenoderirtatinae (Mammalia: Chiroptera). 65 pp. $30.00. Occasional Papers of The Museum (ISSN 0149-175X) No. 10 J. K. Jones, Jr., and C. J. Phillips. 1976. Bats of the genus Sturnira in the Lesser Antilles. 16 pp. $2.00. No. 47 J. K. Jones, Jr., P. Swanepoel, and D. C. Carter. 1977. Annotated checklist of the bats of Mexico and Central America. 35 pp. $2.00. No. 70 W. B. Davis. 1980. New Sturnira (Chiroptera: Phyllostomidae) from Central and South America, with key to currently recognized species. 5 pp. $2.00. No. 77 M. L. Arnold, R. L. Honeycutt, R. J. Baker, V. M. Sarich, and J. K. Jones, Jr. 1982. Resolving a phylogeny with multiple data sets: a systematic study of phyllostomoid bats. 15 pp. $2.00. No. 83 B. E Koop, and R. J. Baker. 1983. Electrophoretic studies of relationships of six species of Artibeus (Chiroptera: Phyllostomidae). 12 pp. $2.00. No. 86 C. S. Hood, and J. D. Smith. 1983. Histomorphoiogy of the female reproductive tract in phyllostomoid bats. 38 pp. $2.00. No. 93 W. B. Davis. 1984. Review of the large fruit-eating bats of the Artibeus “lituratus” complex (Chiroptera: Phyllostomidae) in Middle America. 16 pp. $2.00. No. 106 J. K. Jones, Jr., and R. Owen. 1986. Checklist and bibliography of Nicaraguan Chiroptera. 13 pp. $2.00. Texas tech university press