Chromosome differentiation and the radiation of the butterfly subgenus Agrodiaetus (Lepidoptera: Lycaenidae: Polyommatus) [Elektronische Ressource] : a molecular phylogenetic approach / vorgelegt von Martin Wiemers

Informations

• Publié par: rheinische_friedrich-wilhelms-universitat_bonn
• Publié le: 01 janvier 2003
• Nombre de lectures: 47
• Poids de l'ouvrage: 11 Mo

Extrait

Chromosome differentiation and the radiation
of the butterfly subgenus Agrodiaetus
(Lepidoptera: Lycaenidae: Polyommatus)
– a molecular phylogenetic approach




Dissertation
zur Erlangung des Doktorgrades (Dr. rer. nat.)
der
Mathematisch-Naturwissenschaftlichen Fakultät
der
Rheinischen Friedrich-Wilhelms-Universität Bonn


vorgelegt von
Martin Wiemers
aus
Münster (Westf.)


Bonn – September 2003
Angefertigt mit Genehmigung der Mathematisch-Naturwissenschaftlichen Fakultät der
Rheinischen Friedrich-Wilhelms-Universität Bonn












































1. Referent: Prof. Dr. Clas M. Naumann
2. Referent: Priv.-Doz. Dr. Bernhard Misof

Tag der Promotion: _________________
2 Contents

CONTENTS

CHAPTER 1: INTRODUCTION........................................................................................................................ 5
SCOPE AND AIMS................................................................................................................................................. 7
CHAPTER 2: NEW KARYOLOGICAL RESULTS IN AGRODIAETUS...................................................... 9
INTRODUCTION.................... 9
MATERIAL AND METHODS.... 9
RESULTS............................ 10
Karyotype photographs of Agrodiaetus....................................................................................................... 11
Overview of karyotypes in the subgenus Agrodiaetus Hübner, 1822 .......................................................... 15
DISCUSSION....................... 29
CHAPTER 3: A MOLECULAR PHYLOGENY OF AGRODIAETUS ......................................................... 33
MATERIAL AND METHODS................................................................................................................................. 33
Material........................ 33
Selection of taxa........... 36
DNA extraction............. 37
Selection of genes......... 37
PCR and sequencing..... 38
Sequence alignment...... 39
Choice of methods for phylogenetic inference............................................................................................. 40
Sequence comparisons and phylogenetic inference..................................................................................... 41
RESULTS............................ 43
Sequences and alignment............................................................................................................................. 43
Base composition and sequence divergence................................................................................................ 43
Bayesian inference of phylogeny ................................................................................................................. 48
Maximum parsimony analysis ..................................................................................................................... 60
Statistical parsimony networks.................................................................................................................... 74
DISCUSSION ...................................................................................................................................................... 87
Comparisons with traditional systematics................................................................................................... 87
Congruence between gene trees and species trees in outgroups................................................................. 90
Comparison with allozyme results............................................................................................................... 94
Hybridization in Agrodiaetus ...................................................................................................................... 94
Radiation of Agrodiaetus and the colonization of Europe........................................................................... 96
CHAPTER 4: SYSTEMATICS OF AGRODIAETUS BASED ON MOLECULAR EVIDENCE ............... 97
CHAPTER 5: EVOLUTION OF MORPHOLOGICAL TRAITS IN AGRODIAETUS............................. 107
INTRODUCTION................ 107
MATERIAL AND METHODS 107
RESULTS.......................... 108
DISCUSSION .................................................................................................................................................... 110
SUMMARY........................ 112
CHAPTER 6: THE ROLE OF CHROMOSOME EVOLUTION IN THE RADIATION OF
AGRODIAETUS................. 113
INTRODUCTION................ 113
MATERIAL AND METHODS 113
RESULTS ......................................................................................................................................................... 114
DISCUSSION..................... 116
SUMMARY........................ 119
SUMMARY 120
ZUSAMMENFASSUNG... 120
ACKNOWLEDGEMENTS.............................................................................................................................. 121
REFERENCES.................................................................................................................................................. 123
HILFSMITTEL.................. 143
3Contents

APPENDIX 1. KARYOLOGICAL RESULTS IN AGRODIAETUS....................................................... 145
A2. LIST OF MATERIAL............................................................................................. 149
APPENDIX 3. SEQUENCE DATA ............................................................................................... 165
PLATES. AGRODIAETUS WINGS 201

4 Introduction

Chapter 1: Introduction

The Holarctic butterfly genus Polyommatus Latreille, 1804 consists of approximately 120-180
species (HÄUSER & ECKWEILER 1997), many of which have been placed into subgroups
which are considered separate genera according to numerous authors (e.g. the subgenera
Meleageria, Cyaniris, Plebicula, Sublysandra, Neolysandra). The subgenus Agrodiaetus
Hübner, 1822, which is distributed in the southern Palaearctic, is the most rich in species.
ECKWEILER & HÄUSER (1997) recognize between 56 and 84 species whereas BÁLINT &
JOHNSON (1997) list 102 species (including 8 species placed under Paragrodiaetus Rose &
Schurian, 1977). Since 1997 alone, 23 further species rank taxa have been described from
Anatolia and Iran. The distribution of this subgenus ranges from the Iberian Peninsula to
northern Pakistan and Central Mongolia, but the majority of species is found in Anatolia,
Transcaucasia and Iran. The characteristic morphological feature of this subgenus is the
almost complete reduction of the submarginal elements of the complex wing pattern on the
underside of both pairs of wings, and a prominent white streak on the underside of the hind
wings (although this can be missing in some cases). Secondly the larval foodplants of all
Agrodiaetus species as far as they are known belong to only two closely related genera of
Fabaceae, Onobrychis and Hedysarum which are only used by few other Polyommatus
species. However, the relevance of these two features for a presumed monophylum
Agrodiaetus has never been checked systematically, and therefore the delimitation of this
taxon in literature remains contentious. HIGGINS (1975) and TOLMAN & LEWINGTON (1997)
also include e.g. the taxa thersites Cantener, 1834 (larval foodplant Onobrychis), escheri
Hübner, 1823 (larval foodplant Astragalus) and amanda Schneider, 1792 (larval foodplants
Vicia and Lathyrus) within Agrodiaetus. BÁLINT & JOHNSON (1997) also place thersites with
Agrodiaetus, but split off eight taxa (incl. glaucias (Lederer, 1871), erschoffii (Lederer, 1869)
and dagmara (Grum-Grshimailo, 1888)) from Agrodiaetus and place them into
Paragrodiaetus Rose & Schurian, 1977. This genus was synonymized with Agrodiaetus by
HÄUSER & ECKWEILER (1997). Despite a rather up-to-date species inventory (ECKWEILER &
HÄUSER 1997) the exact number of species is uncertain.
The reasons are threefold. First, this is the only Palaearctic group of butterflies in which a
high number of new species is still being discovered, especially in remote parts of Iran. 15%
of the almost 250 nominal taxa described in Agrodiaetus (Fig. 1) have been named the last
five years, 2/3 of them of species rank.
Taxa descriptions in Agrodiaetus
250
200
150
100 Taxa descriptions
Total50
0
1763 1803 1843 1883 1923 1963 2003
Period

Fig. 1. Taxa descriptions in Agrodiaetus within periods of 20 years
5Chapter 1


Secondly, many taxa represent allopatric populations which differ only slightly in
morphology or karyology (morphospecies and karyospecies) and a decision on their status as
distinct species or subspecies is controversial and subjective. Thirdly and most importantly
many Agrodiaetus taxa are extremely similar in phenotype (HESSEL