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  • Phylogeny of Western Palaearctic long-eared bats

    (Mammalia, Chiroptera, Plecotus)

    – a molecular perspective


    zur Erlangung des Grades

    Doktor der Naturwissenschaften

    am Fachbereich Biologie

    der Johannes Gutenberg-Universität in Mainz

    Andreas Kiefer

    geb. am 27.11.1965 in Bad Kreuznach

    Mainz, im Dezember 2007

  • P. auritus (links) und P. macrobullaris (rechts)

    Wir können alles schaffen

    genau wie die tollen

    dressierten Affen es schaffen

    wir müssen nur wollen…

    (Wir sind Helden)

  • Die Kapitel 3- 6 wurden in leicht abgewandelter Form veröffentlicht:

    Kapitel 3:

    KIEFER, A., MAYER, F., KOSUCH, J., VON HELVERSEN, O., VEITH, M. (2002): Conflicting

    molecular phylogenies of European long-eared bats (Plecotus) can be explained by

    cryptic diversity. – Mol. Phyl. Evol. 25: 557-566. Die ND1-Sequenzen wurden von Frieder Mayer, Erlangen, analysiert. Joachim Kosuch

    hat einen Teil der Laborarbeiten durchgeführt.

    Kapitel 4:

    KIEFER, A., VEITH, M. (2002): A new species of long-eared bat from Europe (Mammalia,

    Chiroptera, Vespertilionidae). – Myotis, 39: 5-16.

    Kapitel 5:

    MUCEDDA, M., KIEFER, A., PIDINCHEDDA, E., VEITH, M. (2002): A new species of long-eared

    bat (Chiroptera, Vespertilionidae) from Sardinia (Italy). – Acta Chiropterologica 4: 121- 135.

    Mauro Mucedda hat umfangreiche Aufsammlungen in Sardinien getätigt. Ermanno

    Pidinchedda hat bei diesen Aufsammlungen mitgeholfen.

    Kapitel 6:

    BENDA, P., KIEFER, A., HANÁK, V., VEITH, M. (2004): Systematic status of African

    populations of long-eared bats, genus Plecotus (Mammalia: Chiroptera). – Folia

    Zoologica, Monograph 1, 53: 1-47. Die umfangreichen morphologischen Analysen und Interpretationen wurden von Petr

    Benda, Museum Prag, durchgeführt, der auch zahlreiche afrikanische Fledermausbelege

    beigesteuert hat.

    Die Publikationen der Kapitel 3 – 5 wurden von mir geschrieben. Die Daten hierfür wurden

    zum größten Teil von mir erhoben und analysiert. Die oben nicht erwähnten Autoren

    waren Betreuer (Veith) oder haben neben Sammlungsmaterial auch wichtige Beiträge zur

    Diskussion geliefert. (Hanák, von Helversen). Die Publikation des Kapitels 6 wurde

    gemeinsam von Petr Benda und mir geschrieben, wobei er den morphologischen Teil und

    ich den genetisch-phylogenetischen Teil formulierte. Der Rest dieser Arbeit war eine

    echte Teamarbeit, bei der Petr Benda und ich gleiche Anteile an der Arbeit hatten. Ich war

    an der Planung, Auswertung und am Schreiben des Artikels beteiligt.

  • Contents



    1. Abstract 6

    2. General introduction 7

    3. Conflicting molecular phylogenies of European long-eared bats (Plecotus) can be

    explained by cryptic diversity 9

    4. A new species of long-eared bat (Plecotus; Vespertilionidae, Mammalia) in Europe 23

    5. A new species of long-eared bat (Chiroptera, Vespertilionidae) from Sardinia (Italy) 35

    6. Systematic status of African populations of long-eared bats, genus Plecotus (Mammalia:

    Chiroptera) 51

    7. General conclusions 91

    8. References 99

    9. Appendix 117

    10. Danksagung 129

    11. Zusammenfassung 131

  • 1 – Abstract


    1. Abstract

    Phylogeny of Western Palearctic long-eared bats (Mammalia, Chiroptera, Plecotus) – a molecular perspective

    Long-eared bats are an enigmatic group of bats that inhabit most parts of Europe up

    to the polar circle. Numerous taxa have been described in the past, but for a long time

    only two species were regarded valid. Further species were known from Northern Africa,

    the Canary Islands and Asia.

    In the present thesis I used molecular data, partial sequences of the mitochondrial

    genes for 16S rRNA, ND1 and of the mitochondrial control region to analyse the

    phylogenetic relationship within and among lineages of Western Palaearctic long-eared

    bats. I estimated the best fitting substitution models and constructed phylogenetic trees

    using four different approaches: neighbor joining (NJ), maximum likelihood (ML),

    maximum parsimony (MP) and a Bayesian approach.

    Seven lineages of long-eared bats are well differentiated at species level: Plecotus

    auritus, P. austriacus, P. balensis, P. christii, P. sardus, P. teneriffae and P. macrobullaris.

    I described three new taxa in this thesis: Plecotus sardus, P. kolombatovici gaisleri

    (= Plecotus teneriffae gaisleri, Benda et al. 2004) and P. macrobullaris alpinus [=Plecotus

    alpinus, Kiefer & Veith 2002). Morphological characteristics for field determination are

    described for the new taxa. Three of the species are polytypic: P. auritus (a western and

    eastern European lineage, and a most recently discovered Caucasian lineage), Plecotus

    kolombatovici (P. k. kolombatovici and P. k. gaisleri and P. k. ssp) and P. macrobullaris

    (P. m. macrobullaris and P. m. alpinus). A formerly fourth P. auritus subspecies, the

    Iberian P. begognae is now regarded as a distinct species (see chapter 7 and Ibanez et

    al. 2006, Mayer et al. 2007). The distribution areas of most species are refined based on

    genetically identified specimens.

    The detection of a considerable amount of cryptic diversity among Western

    Palaearctic long-eared bats will have impact on species conservation. First steps towards

    better protection of the endemic Sardinian long-eared bats have been initiated, but until

    now it did not enter national and international legislation, such as the EU-habitat directive.

  • 2 – General introduction


    2. General introduction

    In his “Systema Naturae per regna tria naturae, secundum classes, ordines, genera,

    species cum characteribus, differentiis, synonymis, locis. Editio decima, reformata” Carl

    von Linné described in 1758 Vespertilio auritus, today known as the brown long-eared bat,

    Plecotus auritus. A second variant was described by Fischer in 1829 as Vespertilio auritus

    var b austriacus (=Plecotus austriacus), the grey long-eared bat. However, P. austriacus

    was considered conspecific with P. auritus by most coeval scientists and therefore

    suffered the same fate as numerous other old forms of Plecotus that were published in the

    19th century and which never achieved scientific appreciation (Hanak 1966). Therefore, for

    most of the 20th century P. auritus was considered the only valid European species of

    long-eared bats.

    Around 1960, several scientists discovered that two forms of Plecotus occurred in

    syntopy throughout Europe (Lanza 1959, Bauer 1960). However, it was Bauer (1960) who

    affiliated one of them to P. austriacus, the form previously described by Fischer (1829).

    Consequently, he resurrected the grey long-eared bat back into species rank. His

    enumeration of diagnostic morphological characters allowed future field discrimination of

    both forms. However, there still remained regional uncertainties in the determination of

    European long-eared bats, especially in the Alps and the Balkans. Hybrid status of such

    “dubious” specimens was assumed (Bauer, in Aellen 1961), although introgression has

    never been proven (Moretti et al. 1993).

    Within Plecotus auritus, two geographically restricted subspecies were added to the

    nominotypical form: P. a. macrobullaris Kuzjakin 19651 in the Caucasus Mountains and P.

    a. begognae de Paz 1994 in the Iberian Peninsula. A single additional subspecies of the

    grey-long-eared bat, P. austriacus kolombatovici Dulic 1980, was described from the

    Mediterranean coast of former Yugoslavia.

    A first African species of long-eared bat was already described in 1838 by Gray (P.

    christii from Egypt). Offshore the African continent, Barrett-Hamilton (1907) discovered a

    comparatively large species which was regarded endemic to the Canary Islands. He

    classified it as Plecotus auritus teneriffae. After showing that it was morphologically clearly

    differentiated from both P. auritus and P. austriacus, Ibanez & Fernandez (1985a)

    considered it a full species.

    1 The description of P.a. macrobullaris is poor and published in Russian language only. It is not clear why, but all western European scientists ignored this species description, as did all Russian bat specialists (except Kuzjakin himself). Even P.P. Strelkow in his outstanding review of the genus Plecotus in the former U.S.S.R ignored Kuzjakin’s work.

  • 2 – General introduction


    Since the mid 1980s, the development and application of molecular techniques to

    phylogenetic questions accelerated the recognition of new species. Based on sequence

    analyses of mainly mitochondrial DNA, scientists started to re-examine seemingly well

    established phylogenies. Often they not only ended up in surprisingly new hypotheses on

    the phylogenetic relationships of species,