Phylogeny of Plecotine Bats: Reevaluation of Morphological and Chromosomal Data Author(S): Wiesław Bogdanowicz, Stephen Kasper, Robert D
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Phylogeny of Plecotine Bats: Reevaluation of Morphological and Chromosomal Data Author(s): Wiesław Bogdanowicz, Stephen Kasper, Robert D. Owen Source: Journal of Mammalogy, Vol. 79, No. 1, (Feb., 1998), pp. 78-90 Published by: American Society of Mammalogists Stable URL: http://www.jstor.org/stable/1382843 Accessed: 22/07/2008 13:35 Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at http://www.jstor.org/action/showPublisher?publisherCode=asm. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. JSTOR is a not-for-profit organization founded in 1995 to build trusted digital archives for scholarship. We work with the scholarly community to preserve their work and the materials they rely upon, and to build a common research platform that promotes the discovery and use of these resources. For more information about JSTOR, please contact [email protected]. http://www.jstor.org PHYLOGENY OF PLECOTINE BATS: REEVALUATION OF MORPHOLOGICAL AND CHROMOSOMAL DATA WIESLAWBOGDANOWICZ, STEPHEN KASPER, AND ROBERTD. OWEN Mammal Research Institute, Polish Academy of Sciences, 17-230 Bialowieta, Poland (WB) Department of Biological Sciences, Texas Tech University, Lubbock, TX 79409 (SK, RDO) Recent systematic studies of the tribe Plecotini have generated two alternative phylogenetic hierarchies: (Barbastella (Corynorhinus (Plecotus (Idionycteris Euderma)))); and (Euderma [including Idionycteris] (Barbastella (Plecotus Corynorhinus))). To test these hypotheses we examined 44-45 morphological and 11 karyological characters of 10 plecotine species, including Otonycteris hemprichii. Character states for the hypothetical ancestor were in- ferred by evaluation of selected outgroup taxa: Rhogeessa tumida, Nycticeius humeralis, Eptesicus fuscus, Myotis lucifugus, M. ciliolabrum, and Miniopterus schreibersi. The most parsimonious trees, identical in topology but different in character-state optimization, were congruent with the systematic hierarchy of Plecotini suggested by the first hypothesis. Otonycteris branched off before Corynorhinus. These results strongly support separation of Corynorhinus as an independent genus and limitation of Plecotus to Palaearctic species. The two highly derived taxa, Idionycteris phyllotis and Euderma maculatum, seem to be sufficiently different from each other to be regarded as generically distinct. It is proposed that the tribe Plecotini originated in the eastern hemisphere. Key words: Plecotini, plecotine bats, morphology, karyology, phylogenetic systematics, biogeography The tribe Plecotini (Chiroptera: Vesper- also using karyological data, proposed that tilionidae) is distributed throughout the the tribe's only short-eared bats, genus Bar- Holarctic and consists of 11 or 12 species bastella, are derived from the long-eared of mostly long-eared bats (Corbet and Hill, plecotines in the arrangement ((Euderma 1991, 1992; Koopman, 1993; Yoshiyuki, Idionycteris) (Barbastella Plecotus Cory- 1991). Numerous systematic studies have norhinus)). evaluated phylogenetic affinities within the Results from two of the most recent phy- tribe at generic, subgeneric, and specific logenetic studies of the Plecotini indicate Frost and Hand- levels (e.g., Timm, 1992; two quite different topologies. Tumlison Leniec et al., 1987; ley, 1959; Qumsiyeh and Douglas (1992) suggested the phylog- and Bickham, 1993; Tumlison and Douglas, eny (Barbastella (Corynorhinus (Plecotus 1992; Williams et al., 1970); however, no (Idionycteris Euderma)))). However, Frost consistent phylogeny has emerged. Handley and Timm (1992) proposed the cladistic re- (1959) first examined morphological rela- lationship of (Euderma [including Idionyc- tionships among plecotines and concluded teris] that the genera Plecotus (including subgen- (Barbastella (Plecotus Corynorhin- era Plecotus, Corynorhinus, and Idionycter- us))). This discrepancy might be due to the is) and Euderma shared a common ancestor fact that Tumlison and Douglas (1992) used after divergence from Barbastella in the only morphological traits, and Frost and Miocene. Based on karyological data, Wil- Timm (1992) included both morphological liams et al. (1970) suggested the phylogeny and karyological characters. As substantia- (Barbastella ((Euderma Idionycteris) (Ple- tion, one of the four alternate trees of Frost cotus Corynorhinus))). Leniec et al. (1987), and Timm (1992), based only on morpho- Journal of Mammalogy, 79(1):78-90, 1998 78 February1998 BOGDANOWICZET AL.-PHYLOGENY OF PLECOTINEBATS 79 logical characters, is identical in topology (1984). Outgroups were chosen and arranged to the tree of Tumlison and Douglas (1992). based on probable relationships among the in- Published descriptions of karyotypes of group and outgroups following the karyology- the genera of Plecotini have been inconsis- only cladogram of the Vespertilionidae (Volleth and Heller, In our were tent. Several taxa were distinguished based 1994b). study, they formed by the sister groups "Nycticeiini" (Nyc- on different chromosome banding qualities, ticeius humeralis and Rhogeessa tumida) and and of the standard num- misinterpretations Eptesicini (Eptesicus fuscus), the tribe Myotini bering system have been indicated (revised (Myotis lucifugus and M. ciliolabrum), and the by Volleth and Heller, 1994a). In addition, subfamily Miniopterinae (Miniopterus schrei- a study using G-banded chromosomes in- bersi). Chromosome banding indicates that the dicated that the large long-eared bat Oto- Miniopterinae is the first subfamily to diverge nycteris hemprichii is also a member of the from the common Vespertilionidae stem (Volleth tribe Plecotini, with Qumsiyeh and Bick- and Heller, 1994b). ham (1993) inferring the phylogeny ((Eu- Nixon and Carpenter (1993) suggested that derma Idionycteris) (Otonycteris Barbastel- characters should not be polarized prior to an la Plecotus Corynorhinus)). The karyotype analysis, rather that polarity should be deter- mined a the clado- of Otonycteris shows an extensive similar- posteriori through rooting gram between ingroup and outgroup taxa. How- ity with karyotypes of Plecotus, Corynorhi- ever, this is a reasonable procedure only when nus, and Barbastella (Qumsiyeh and Bick- there is absolutely no uncertainty concerning the Zima et Tumlison ham, 1993; al., 1992). relationship of the ingroup to the putative out- and Douglas (1992), Qumsiyeh and Bick- group taxa (i.e., certainty of ingroup monophy- ham (1993), and Bogdanowicz and Owen ly). In many systematic analyses, including this (1996) also have suggested that problems one, there may be certainty concerning most in the intepretations of some morphological taxa and uncertainty concerning a few putative characters from works such as Handley ingroup or outgroup taxa. Using the method of (1959) may have led to questionable con- Maddison et al. (1984), position of these taxa clusions. Phenetic results, based on cranial may be examined, particularly by using multiple geometric morphometrics, indicated that character sets or subdividing characters being as we have done. Otonycteris was the most divergent species examined, Characters (i.e., individual of taxic of the Plecotini (Bogdanowicz and Owen, hypotheses homology) were compiled from the literature 1996) and has undergone extensive adap- and examination of zoological material. The 61 tive changes (Horacek, 1991). skin, skull, and chromosomal characters for Our objectives were to resolve the taxo- which primitive states could be inferred, were nomic status of Idionycteris and Corynorhi- determined for each species (Table 1, Appendix nus, determine the validity of Otonycteris II). Five characters, however, were autapo- as a member of the tribe, and establish phy- morphic either for Idionycteris or Euderma, and logenetic relationships of all genera and one character (biarmed chromosome, arm com- subgenera of plecotine bats. This research bination 11/14) presented an obvious synapo- was accomplished by cladistic analysis of morphic feature of the tribe (Appendix II). Thus, morphological and karyological characters. these six characters were phylogenetically un- informative and were omitted from final analy- ses. Character states MATERIALS AND METHODS for polymorphic taxa (sensu Swofford, 1991) were intepreted as uncertain. The ingroup was composed of 10 species rep- Characters were not weighted. Because karyo- resenting the 6 genera (or subgenera) of the Ple- logical data were taken from the literature and cotini (including Otonycteris; Appendix I). A were not our direct observations, a second anal- hypothetical ancestor for the Plecotini was in- ysis was performed on a data matrix containing ferred from multiple outgroup analysis, and all only morphological characters, which we scored character states were polarized using the out- directly. group comparison method of Maddison et al. Cladograms were constructed using the 80 JOURNAL OF MAMMALOGY Vol. 79, No. I TABLE 1.-Distribution of states for 45 morphological and