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Phylogenetic relationships of three ‘‘Nycticeiini’’ genera (Vespertilionidae, Chiroptera, Mammalia) as revealed by karyological analysis By Marianne Volleth, K.-G. Heller and J. Fahr Institut fu¨r Humangenetik, Otto-von-Guericke-Universita¨t Magdeburg, Magdeburg, Germany; Institut fu¨r Zoologie, Universita¨t Erlangen-Nu¨rnberg, Erlangen, Germany; Department of Experimental Ecology, University of Ulm, Ulm, Germany Receipt of Ms. 31.1.2005 Acceptance of Ms. 26.9.2005

Abstract GTG-banded karyotypes are presented for Scotoecus hirundo (2n ¼ 30; FN ¼ 50), Rhogeessa alleni (2n ¼ 30; FN ¼ 50), Scotophilus kuhlii (2n ¼ 36; FN ¼ 48) and Scotophilus leucogaster (2n ¼ 36; FN ¼ 50). These three genera belong to the family Vespertilionidae and have previously been placed into the tribe ‘‘Nycticeiini’’ (Tate 1942). Karyological analysis, however, points to a close relationship of Scotoecus hirundo to the tribes Pipistrellini and Vespertilionini (sensu Volleth and Heller 1994). Rhogeessa (Baeodon) alleni, a member of the karyologically diverse genus Rhogeessa, has two fusion chromosomes in common with the genus Plecotus. Together with morphological (Hill and Harrison 1987) and molecular-genetic results (Hoofer and Van Den Bussche 2003), chromosome analysis suggests a closer relationship of Plecotini and Rhogeessa. The two species examined of the genus Scotophilus show differences only in two small autosomal pairs and the Y chromosome. Chromosomal data did not reveal closer relationships of this genus to any other vespertilionid tribe. r 2005 Deutsche Gesellschaft fu¨r Sa¨ugetierkunde. Published by Elsevier GmbH. All rights reserved. Key words: Vespertilionidae, Chromosomal evolution, Phylogenetic relationships

Introduction The chiropteran family Vespertilionidae consists of about 407 species and 48 genera according to Simmons (2006). Tate (1942) placed 8 genera, amongst them Scotoecus, Scotophilus and Rhogeessa, into the tribe Nycticeiini according to a common morphological character, the missing second upper incisor. The same genera were included into Nycticeiini by Koopman (1984a, 1994) and Simmons (2006). However, Hill and Harrison (1987) classified the Nycticeiini genera ac-

cording to bacular morphology into three different tribes. They assigned Rhogeessa, Nycticeius and Otonycteris to Plecotini. Scotoecus was placed, together with Scoteanax, Scotorepens and Nycticeinops, into the Pipistrellini (sensu Hill and Harrison 1987). Together with Scotomanes, Scotophilus comprises the tribe Scotophilini of Hill and Harrison (1987). On the basis of a comprehensive study of mitochondrial DNA sequences, Hoofer and Van Den Bussche

1616-5047/$ - see front matter r 2005 Deutsche Gesellschaft fu¨r Sa¨ugetierkunde. Published by Elsevier GmbH. All rights reserved. doi:10.1016/j.mambio.2005.09.001 Mamm. biol. 71 (2006) 1  1–12

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(2001, 2003) stated that ‘‘Nycticeiini, as traditionally recognized, does not represent a natural assemblage’’. In agreement with Hill and Harrison (1987), they placed Scotoecus into Pipistrellini and Scotophilus into Scotophilini. Rhogeessa and Baeodon, however, were assigned to Antrozoini based on mt DNA results (Hoofer and Van Den Bussche 2003). In addition to similarities in the dental formula, Nycticeiini also show one striking karyological similarity: with the exception of Otonycteris, whose karyotype is similar to that of Plecotus (Zima et al. 1992), all Nycticeiini genera sensu Tate (1942) from which banded karyotypes have been published, miss at least two of the three large metacentric chromosomes (1/2, 3/4, 5/6) present in the basic karyotype (Volleth and Heller 1994). It is very likely, however, that the Robertsonian fission events leading to the acrocentric condition of arms 1–6 occurred independently several times in the family Vespertilionidae. In this study, we present detailed karyotype descriptions of four species from three genera formerly placed with the Nycticeiini. The chromosomal character states are compared with published data to elucidate the phylogenetic relationships of the species studied. Chromosomal characters belong to the category of rare genomic changes and provide an independent source of phylogenetic information (Rokas and Holland 2000). Rare genomic changes are not prone to extensive convergent or parallel evolution as found in adaptive characters such as changes in the dental formula of vespertilionid bats.

Material and methods Chromosome preparations Metaphase spreads were obtained from fibroblast cultures (see Volleth 1987 for details) and stained by differential staining procedures as GTG, QFQ, CBG, RBG and AgNOR (see Volleth et al. 2001). Chromosomal arms were numbered using Bickham’s scheme for American Myotis species (Bickham 1979). Two chromosomal arm numbers separated by a solidus (/) indicate a bi-armed chromosome resulting from a Robertsonian trans-

location. For karyotype analysis the chromosomes were compared with the ‘‘basic karyotype’’ of Vespertilionidae (2n ¼ 44, see Volleth and Heller 1994).

Specimens examined Rhogeessa alleni, 1 m, Zapotitlan, Puebla, Mexico, SMF 77908; Scotophilus leucogaster, 1 m, Ouagadougou, Burkina Faso, SMF 77907; Scotophilus kuhlii, 1 m, Ulu Gombak Field Studies Centre, Selangor, Malaysia, SMF 69317 (SMF= catalogue number of the Senckenberg Museum Frankfurt/ Main); Scotoecus hirundo, 1 f, Comoe´ National Park, Ivory Coast, field-no. 2948, currently housed at the University of Ulm. For the Scotophilus specimens, species identification follows Koch-Weser (1984) and Heller and Volleth (1989).

Results Scotoecus hirundo (de Winton, 1899) The female studied shows a karyotype with a diploid chromosome number of 30 and an autosomal fundamental number (FN) of 50. There are 11 large to small meta- to submetacentric autosomal pairs. The X chromosome is a medium-sized subtelocentric element. The three large metacentric chromosomes, composed of arms 1 and 2, 3 and 4, 5 and 6, which are present in the basic karyotype of Vespertilionidae (see Volleth and Heller 1994) and in the majority of vespertilionid species, are not present in Scotoecus hirundo. Instead, the following Robertsonian fusion products have been found: 1/12, 2/7, 3/13, 4/8, 5/14, 6/11, 9/10, 15/20, 18/19, 21/22. The small metacentric chromosome 16/17 of the basic karyotype is altered in Scotoecus, probably due to a small pericentric inversion, bringing the proximal part of arm 16 to the long arm. Further, the three small acrocentric chromosomes have been identified as arms 23, 24 and 25 (Fig. 1). Some of the chromosomal arms of the vespertilionid karyotype have been found to exist in two different states (Volleth and Heller 1994). In Scotoecus, arms 1 and 2 probably show state II, arms 11 and 23 clearly state I, arms 12 and 15 clearly state II. Due to the rearranged condition of the X

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Fig. 1. GTG-banded karyotype of a Scotoecus hirundo female. Scale bar=5 mm.

chromosome, the state could not be determined for the X. According to these characters, Scotoecus is closely related to, but not a member of the Pipistrellini (sensu Volleth and Heller 1994). In addition to the centromeric dots, CBG-banding revealed terminally located heterochromatic bands on both homologues of arm 1 and 3, and of one homologue of arm 5. Further, the distal part of the short arm of chromosome 16/17 and a small intercalary segment of the proximal part of arm 2 were CBG-positive. Due to the addition of heterochromatic material in arm 2, this arm is clearly longer than those of other vespertilionid species (Fig. 2). The Nucleolus Organizing Regions (NORs) are located in the secondary constriction (SC) of chromosomal arm 15. An unknown rearrangement led to the subtelocentric X chromo-

some of Scotoecus with three small heterochromatic bands, one of each situated proximal, central and distal on the long arm. Together with additional material (SMF 91994–97) from the same locality, this specimen represents the first record of S. hirundo for Ivory Coast. Scotophilus kuhlii Leach, 1821 The karyotype of the male studied consists of 7 meta- to submetacentric and 10 acrocentric autosomal pairs. The diploid chromosome number is 2n ¼ 36 and the fundamental number is FN ¼ 48. Due to centric fission events, the three large metacentric autosomal pairs of the basic vespertilionid karyotype are present here as acrocentric elements. Robertsonian fusions resulted in bi-armed

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heterochromatin except for a small region close to the centromere. The centromeres of the acrocentric chromosomes are very clearly stained after CBG-banding whereas the metacentric chromosomes show only weak staining in the centromeric regions. The SC on the smallest autosome, 24, was found to bear the only NOR in this species. Scotophilus leucogaster (Cretzschmar, 1826)

Fig. 2. CBG-banded metaphase spread of Scotoecus hirundo. The arrows point to telomeric heterochromatin on chromosomal arms 1, 3, 5, and 16. Intercalary heterochromatin of arm 2 is indicated by arrowheads.

autosomes 8/14, 10/20, 12/23 and 15/21. In addition to centric fusion of arms 9 and 13, a small pericentric inversion was found in the resulting metacentric chromosome, which transferred a small GTG-negative, early replicating segment from arm 9 to 13. Chromosomal arms 7, 11 and 24 are present as acrocentric elements. The composition of three chromosomes, provisionally called A, B and C, could not be determined. Chromosomal arms 1, 2 and 7 are present in state II, arms 11, 15, and 23 are present in state I (Fig. 3). The X chromosome is submetacentric with the banding pattern of state I. The acrocentric Y chromosome with a size equal to arm 21 consists of CBG-positive

This species shows a karyotype with 2n ¼ 36 and FN ¼ 50. It differs only slightly from that of S. kuhlii. The following differences were found: chromosome C is a small submetacentric chromosome whereas it is acrocentric in S. kuhlii (Fig. 3). Chromosomal arm 24 is here the short arm of a submetacentric chromosome, the long arm of which consists completely of heterochromatic material (Fig. 4). The NOR in the SC of arm 24 is separated by a small heterochromatic segment from the centromere. The Y chromosome is a very small submetacentric chromosome with a completely heterochromatic short arm. Rhogeessa (Baeodon) alleni Thomas, 1892 This species possesses a karyotype with 2n ¼ 30 and FN ¼ 50. There are 11 large to small meta- to submetacentric and three small subtelocentric autosomal pairs (Fig. 5). Out of the three large metacentric chromosomes of the basic karyotype, only chromosome 3/4 is present in Rhogeessa alleni. Arm 1 is fused with arm 5, arm 2 with arm 6. In addition, the following Robertsonian fusion products have been found: 7/8, 9/12, 10/14, 11/15, 13/ 18 and 19/21. The subtelocentric chromosomes are composed of arms 22, 23 and 24 in the long arms and a small heterochromatic short arm each. The composition of two small submetacentric chromosomes,

Fig. 3. GTG-banded and RBG-banded partial karyotypes of Scotophilus kuhlii (left chromosome) and S. leucogaster (right chromosome of each pair). Rows 1, 3 and 5 show the GTG-banding pattern, rows 2, 4 and 6 the replication pattern. The heterochromatic material of the long arm of the S. leucogaster homologue of pair 24 is clearly late replicating. The composition of chromosomes A, B and C could not be ascertained by banding analysis. The chromosomal arm numbers for rows 3 and 4 are given on top for the short arms and below for the long arms.

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Fig. 4. CBG-banded metaphase spread of Scotophilus leucogaster. The arrowheads point to the proximal region of arm 24, where the NOR is situated. The centromere and the long arm of this chromosome is clearly CBG-positive.

provisionally called A and B, could not be determined. Chromosomal arms 11, 15, and 23 are present in state I, arms 1, 7, and 12 in state II. Arm 2 is enlarged by addition of a small GTG-positive segment surrounded by two small CBG-positive heterochromatic bands (see Fig. 6). One possible explanation is the insertion of arm 25 at this position. However, in several non-related species, CBG-positive material has been found in the proximal segment of arm 2 (see Scotoecus hirundo for example). The X chromosome shows a GTG-banding pattern similar to state I. The small Y chromosome is submetacentric, late-replicating with a completely heterochromatic short arm. The NORs are situated in the SCs of the long arms of chromosomes 23 and 24. CBGbanding revealed clearly stained centromeric dots in addition to tiny heterochromatic short arms of chromosomes 22 to 24 and a heterochromatic band in the distal segment on the short arm of chromosome B.

Discussion Comparison with published chromosomal data Genus Scotoecus Nagorsen et al. (1976) presented conventionally stained chromosomes of Scotoecus hindei (Thomas, 1901), a taxon that is currently considered as a synonym of S. hirundo (Robbins 1980; Koopman 1994; but see Hill 1974; Taylor and van der Merwe 1998; Cotterill 2001). The published karyotype with 2n ¼ 30 and FN ¼ 50 seems to be very similar to the one presented here. Genus Scotophilus Several authors have published conventionally stained karyotypes of this genus. Many of them have already been cited by Zima and Hora´cˇek (1985). However, assignment to species differs due to conflicting taxonomic treatments and problems of nomenclature (e.g. Harrison and Brownlow 1978; Robbins

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Fig. 5. GTG-banded karyotype of Rhogeessa alleni.

1978; Koopman 1984b; Robbins et al. 1985). As a result, different karyotypes were assigned to the same species name by different authors (e.g. S. temmincki with 2n ¼ 36, FN ¼ 52 in Pathak and Sharma 1969 and with 2n ¼ 36, FN ¼ 48 in Harada and Kobayashi 1980; Naidu 1985). At first glance, published data on Scotophilus species lead to the impression that a remarkable number of different karyotypes, all with a 2n ¼ 36, is present in this genus. A closer examination, however, reveals that the chromosomal variation in Scotophilus seems to be restricted to the two smallest autosomal pairs (pair ‘‘C’’ and 24 in Fig. 3). Scotophilus kuhlii shows both pairs in an acrocentric condition, resulting in an FN of 48 (Harada et al. 1982; Sreepada and Gururaj 1994; Rickart et al. 1999; Lin et al. 2002). Scotophilus temminckii

(Horsfield, 1824), which has been synonymised with S. kuhlii (Hill and Thonglongya 1972), was also reported with 2n ¼ 36 and FN ¼ 48 (Harada and Kobayashi 1980; Naidu 1985). The two smallest pairs of the Scotophilus leucogaster karyotype are biarmed. However, one arm of the SC-bearing chromosome consists of heterochromatic material and is thus not counted for the FN (FN ¼ 50). The karyotype of the Asian S. heathii (Horsfield, 1831), resembles very much the S. leucogaster karyotype, showing a bi-armed chromosome with the SC in the short arm (Sharma et al. 1974; Bhatnagar and Srivastava 1974) and a long arm of polymorphic size (Sreepada and Gururaj 1994), thus being probably heterochromatic as in S. leucogaster. The confirmation of the heterochromatic nature of this arm would

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Fig. 6. CBG-banded metaphase spread of Rhogeessa alleni. Chromosomal identification was done by QFQ-staining prior to CBG. The arrowheads indicate the positions of the Nucleolus Organizing Regions on chromosomes 23 and 24, visible as secondary constrictions in the QFQ-banded chromosomes. The large arrows indicate the heterochromatic segments on the short arm of chromosome B. The small arrows point to the faint intercalary bands on chromosomal arm 2.

reduce the published FN from 52 to 50. Further remarkable is the fact that populations of S. heathii with a subtelocentric X have been reported (Sreepada and Gururaj 1994). The interpretation of the published chromosomal data for the African species S. dinganii (A. Smith, 1833) and S. viridis (Peters, 1852) is complicated by the fact that populations from South Africa have been found with an acrocentric X chromosome (Schlitter et al. 1980). Specimens of both species from Namibia and/or Somalia, however, have been published with a metacentric X-chromosome by Ruedas et al. (1990). In the third study dealing with S. dinganii (specimen originally published as S. nigrita (Schreber, 1774) by Peterson and Nagorsen 1975, but subsequently assigned to S. dinganii by Robbins 1978, see also Robbins and Baker 1978), the shape of the X-chromosome remained uncertain as only one female was studied (Peterson and Nagorsen 1975). Tak-

ing into account only the karyotype with a metacentric X chromosome, the karyotype of S. dinganii and S. viridis would be composed of 8 metacentric and 9 acrocentric autosomal pairs, with the SC located on the smallest metacentric chromosome and an acrocentric homologue to ‘‘C’’. The karyotype of the African S. nux differs from that of S. dinganii and S. viridis only in the larger SC-bearing chromosome (Ruedas et al. 1990), which resembles very much that of S. leucogaster. Banded karyotypes have only been published in two papers (Ruedas et al. 1990; Sreepada and Gururaj 1994). Unfortunately, the quality of the published figures is not sufficient for a detailed comparison. Genus Rhogeessa The karyological variation within the genus Rhogeessa has been discussed in detail by Baker et al. (1985). The results of the banding analysis (Bickham and Baker 1977; Baker et al. 1985) revealed a large number of

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Robertsonian translocation events. Therefore it is not surprising that the species studied here, Rhogeessa alleni, has only two fusion chromosomes in common with R. parvula (2n ¼ 44) and the R. tumida cytotypes (2n ¼ 30, 32, 34, 52). These are chromosomes 19/21 and B.

Phylogenetic considerations Scotoecus Formerly, the classification of the different vespertilionid tribes was based on the dental formula of the respective genera (e.g. Tate 1942). According to this character, all members of the genera treated here would belong to the same tribe, Nycticeiini. Hill and Harrison (1987), however, introduced the shape of the baculum (os penis) as an additional character. The genus Scotoecus was placed by Hill and Harrison (1987) in their tribe Pipistrellini. Previously, we showed that chromosomal characters are better suited features for classification into certain tribes, e.g. Pipistrellini and Vespertilionini (Volleth and Heller 1994). Members of both tribes show state II of chromosome 15 and the X chromosome as synapomorphic features. Scotoecus hirundo was found to have state II of chromosome 15 and a rearranged X chromosome. However, S. hirundo does neither show state II of chromosome 11 (autapomorphic feature of Vespertilionini) nor state II of chromosome 23 (autapomorphic feature of Pipistrellini), see Table 1. Therefore, it is closely related to the members of both tribes, if the unlikely event of an independent paracentric inversion leading from state I to state II of chromosome 15 is excluded. The shape of the baculum as well as the results of the mt DNA analysis (Hoofer and Van Den Bussche 2003), however, imply a closer phylogenetic relationship of Scotoecus to the Pipistrellini than to the Vespertilionini. Within the genus Scotoecus, the recognition of different taxa as distinct species or as subspecies of S. hirundo is problematical (Hill 1974; Taylor and van der Merwe 1998; Cotterill 2001) and warrants further morphometric and molecular investigations.

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Table 1. Distribution of karyological features: chromosomal arms occurring in two states (I, II). Chromosome number Taxon Ancestor1 Pipistrellini2 Vespertilionini2 Scotoecus Scotophilus Rhogeessa

11 I I II I I I

15 I II II II I I

23 I II I I I I

X I II II r3 I I

1 Presumed ancestor of the subfamily Vespertilioninae. 2 According to Volleth and Heller (1994). 3 Determination of state not possible due to rearranged X chromosome.

Scotophilus On dental and bacular reasons, the AfroOriental genus Scotophilus was placed together with the Oriental genus Scotomanes into a separate tribe, Scotophilini, by Hill and Harrison (1987). This has been criticised by Frost and Timm (1992) because Scotophilus and Eptesicus show a rather similar shape of the baculum. However, chromosomal characters clearly exclude Scotophilus from Eptesicini (sensu Volleth and Heller 1994) if the presence of the NOR on chromosomal arm 15 is used as a distinctive feature. Both genera have in common the presence of arms 1–6 as acrocentric elements instead of the three metacentrics of the basic karyotype. However, besides these two genera, fission of all three metacentrics (1/2, 3/4, and 5/6) must have occurred independently at least three times in the evolution of vespertilionid bats. Therefore, this character is clearly not suited for elucidating phylogenetic relationships. Molecular genetic studies indicate that Scotophilus is more closely related to the Plecotini than to the other vespertilionid tribes (Kawai et al. 2002; Hoofer and Van Den Bussche 2003), but is also the most divergent genus within Vespertilioninae based on mt DNA data (Hoofer and Van Den Bussche 2003: 28). Provisionally, we follow Hill and Harrison (1987) and Hoofer and Van Den Bussche (2003) in assigning Scotophilus to its own tribe, Scotophilini. Concerning the relationships within the genus Scotophilus, the results of the karyologi-

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cal and mt DNA analysis (Hoofer and Van Den Bussche 2003) are in excellent agreement. With both methods, the Asian S. heathii is more closely related to the African Scotophilus species than to the Asian S. kuhlii. Rhogeessa Due to the fact that one of the three metacentrics of the basic karyotype, i.e. 3/4, is present in R. alleni but not in the other species, R. alleni is thought to branch off early from the common Rhogeessa stem. Based on a multivariate analysis of morphological characters, this opinion was also given by LaVal (1973), justifying the classification of R. alleni as a separate subgenus, Baeodon. The basal position of R. alleni was also revealed in the molecular phylogeny of Hoofer and Van Den Bussche (2003), who treated this taxon as a full genus, i.e. Baeodon alleni, following Corbet and Hill (1991). On bacular grounds, the genus Rhogeessa has been allied to the plecotine bats together with Nycticeius humeralis (Rafinesque, 1818) and Otonycteris by Hill and Harrison (1987). Molecular genetic analyses (Hoofer and Van Den Bussche 2001, 2003) suggest closer phylogenetic relationships between traditional Plecotini, Rhogeessa, Lasiurus, Scotophilus and other New World vespertilionine genera. If further DNA studies would support these relationships, the fact that Rhogeessa alleni shares two fusion chromosomes (9/12, 13/18) with the Nearctic genus Corynorhinus and the Palaearctic plecotines could be interpreted as

a common feature and not as a mere homoplasy. The Palaearctic plecotines (Plecotus, Otonycteris, Barbastella) differ from Corynorhinus in only one autosome, i.e. a derived state of chromosome 16/17 (Zima et al. 1992). On the other hand, Corynorhinus has only one apomorphic chromosomal feature in common with the Nearctic Plecotini Euderma and Idionycteris (i.e. fusion chromosome 11/14; Stock 1983). If we assume that Rhogeessa is closely related to traditional Plecotini, the karyotype of Corynorhinus would be the most ancestral one within Plecotini. A New World origin of this tribe seems then to be very likely.

Acknowledgements We thank very much S. Koch-Weser for collecting the Scotophilus leucogaster specimen and N. Ebigbo for the specimen of Scotoecus hirundo used to prepare the respective karyotypes. For manifold support during our stay in Malaysia we would like to thank Prof. H.-S. Yong, Kuala Lumpur. Warm thanks go to Prof. O. von Helversen and C. Drews for their help with bat netting during our excursion to Mexico. JF acknowledges the funding through the BIOLOG-framework financed by the German Federal Ministry of Education and Research (BMBF), W09 BIOTA-West. This paper was written in memory of our friend and colleague Martin Rohr, who lost his life on the expedition to Mexico in 1988 and is still sadly missed.

Zusammenfassung Phylogenetische Beziehungen von drei ‘‘Nycticeiini’’ Gattungen (Vespertilionidae, Chiroptera, Mammalia) aufgrund karyologischer Analysen Fu¨r Scotoecus hirundo (2n ¼ 30; FN ¼ 50), Rhogeessa alleni (2n ¼ 30; FN ¼ 50), Scotophilus kuhlii (2n ¼ 36; FN ¼ 48) und Scotophilus leucogaster (2n ¼ 36; FN ¼ 50) werden GTGgeba¨nderte Karyotypen vorgestellt. Diese drei Gattungen geho¨ren zur Familie Vespertilionidae und wurden fru¨her dem Tribus ‘‘Nycticeiini’’ zugeordnet. Die zytogenetischen Befunde sprechen jedoch fu¨r eine nahe Verwandtschaft von Scotoecus hirundo zu den Tribus Pipistrellini und Vespertilionini. Rhogeessa (Baeodon) alleni, eine Art der karyologisch vielfa¨ltigen Gattung Rhogeessa, besitzt zwei Fusionschromosomen, die auch bei der Gattung Plecotus gefunden wurden.

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Die Ergebnisse der Chromosomenanalyse deuten also, zusammen mit morphologischen und molekulargenetischen Befunden auf eine engere Verwandtschaft zwischen Plecotini und Rhogeessa. Die beiden aus der Gattung Scotophilus untersuchten Arten unterscheiden sich lediglich in den beiden kleinsten Chromosomenpaaren und dem Y-Chromosom. Durch die Chromosomenanalyse konnte keine na¨here Verwandtschaft dieser Gattung zu einem anderen Tribus ermittelt werden. r 2005 Deutsche Gesellschaft fu¨r Sa¨ugetierkunde. Published by Elsevier GmbH. All rights reserved.

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Authors’ addresses: Marianne Volleth, Institut fu¨r Humangenetik, Ottovon-Guericke-Universita¨t Magdeburg, Leipzigerstr. 44, D-39120 Magdeburg, Germany (e-mail: [email protected]) Klaus-Gerhard Heller, Institut fu¨r Zoologie, Universita¨t Erlangen-Nu¨rnberg, Staudtstr., D-91058 Erlangen, Germany Jakob Fahr, Department of Experimental Ecology, University of Ulm, Albert-Einstein Allee 11, D-89069 Ulm, Germany

"Nycticeiini" genera

Chromosomal data did not reveal closer relationships of this genus to any other vespertilionid tribe. ..... und wurden früher dem Tribus ''Nycticeiini'' zugeordnet.

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