Phylogeny of western palaearctic long-eared bats (Mammalia, Chiroptera, Plecotus) [Elektronische Ressource] : a molecular perspective / Andreas Kiefer

Kiw

Informations

• Publié par: johannes_gutenberg-universitat_mainz
• Publié le: 01 janvier 2008
• Nombre de lectures: 32
• Poids de l'ouvrage: 2 Mo

Extrait

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






Dissertation
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

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


5 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.
6 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
th19 century and which never achieved scientific appreciation (Hanak 1966). Therefore, for
thmost of the 20 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
1nominotypical form: P. a. macrobullaris Kuzjakin 1965 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.
7 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,