EFFECTS OF CLIMATE ON DIVERSITY PATTERNS IN GROUND BEETLES: CASE STUDIES IN MACROECOLOGY, PHYLOGEOGRAPHY AND GLOBAL CHANGE BIOLOGY Der Fakultät Nachhaltigkeit der Leuphana Universität Lüneburg zur Erlangung des Grades Doktorin der Naturwissenschaften – Dr. rer. nat. – vorgelegte Dissertation von Katharina Homburg geb. Schäfer am 16.03.1984 in Stadthagen 2013 Eingereicht am: 15.01.2013 Betreuer und Erstgutachter: Prof. Dr. Thorsten Assmann Zweitgutachter: Prof. Dr. Tamar Dayan Drittgutachter: Prof. Dr. Achille Casale Tag der Disputation: 20.12.2013 COPYRIGHT NOTICE: Cover picture: Modified photograph of Carabus irregularis. Copyright of the underlying photograph is with O. Bleich, www.eurocarabidae.de. Chapters 2, 3, 4 and 5 have been either published, submitted for publication or are in preparation for publication in international ecological journals. Copyright of the text and the figures is with the authors. However, the publishers own the exclusive right to publish or use the material for their purposes. Reprint of any of the materials presented in this thesis requires permission of the publishers and the author of this thesis. CONTENTS CONTENTS Zusammenfassung ...................................................................................................................... 1 Summary .................................................................................................................................... 4 1 General introduction and conclusions ................................................................................ 6 1.1 Background: Biodiversity studies and climate change ................................................ 6 1.1.1 Macroecology ....................................................................................................... 6 1.1.2 Phylogeography .................................................................................................... 7 1.1.3 Global change biology ......................................................................................... 8 1.2 Ground beetles as a study taxon .................................................................................. 9 1.3 Aims and hypotheses of this thesis ............................................................................ 11 1.3.1 Large-scale patterns in carabid species’ traits .................................................... 11 1.3.2 Glacial refugia of a cold-adapted, flightless ground beetle species ................... 12 1.3.3 Future distribution and conservation of a flightless ground beetle species ........ 13 1.4 Main findings and prospects ...................................................................................... 14 1.4.1 Insights from a species-rich taxon and future research perspectives ................. 14 1.4.2 Insights from a specialised species and future research perspectives ................ 16 1.5 Appendix: Authors’ contributions to manuscripts ..................................................... 19 2 Broad-scale geographic patterns in body size and hind wing development of western Palaearctic carabid beetles (Coleoptera: Carabidae) ................................................................ 21 2.1 Abstract ...................................................................................................................... 22 2.2 Introduction ............................................................................................................... 22 2.3 Material and methods ................................................................................................ 24 2.3.1 Study area ........................................................................................................... 24 2.3.2 Species data ........................................................................................................ 25 2.3.3 Environmental data ............................................................................................ 26 2.3.4 Statistical analyses .............................................................................................. 27 2.4 Results ....................................................................................................................... 29 2.4.1 Body size ............................................................................................................ 29 2.4.2 Hind wing development ..................................................................................... 31 2.4.3 Relationships between body size and hind wing development .......................... 34 2.5 Discussion .................................................................................................................. 35 CONTENTS 2.5.1 Contemporary climate ........................................................................................ 35 2.5.2 Dispersal and historical climate ......................................................................... 37 2.5.3 Phylogeny ........................................................................................................... 38 2.5.4 Country size ........................................................................................................ 39 2.6 Conclusions ............................................................................................................... 40 2.7 Acknowledgements ................................................................................................... 40 2.8 Supplementary Material ............................................................................................ 41 3 Carabids.org – A dynamic online database of ground beetle species traits (Coleoptera, Carabidae) ................................................................................................................................ 71 3.1 Abstract ...................................................................................................................... 72 3.2 Introduction ............................................................................................................... 72 3.3 Content of carabids.org .............................................................................................. 74 3.3.1 Taxonomic and systematic affiliation ................................................................ 74 3.3.2 Distributional range ............................................................................................ 74 3.3.3 Trait data ............................................................................................................ 79 3.3.4 Body size and dispersal ...................................................................................... 79 3.3.5 Food and habitat preferences .............................................................................. 80 3.3.6 Activity and reproduction ................................................................................... 81 3.4 Structure and application of carabids.org .................................................................. 81 3.4.1 Technical structure ............................................................................................. 81 3.4.2 User registration ................................................................................................. 82 3.4.3 Data search and export ....................................................................................... 82 3.4.4 Data contributions .............................................................................................. 82 3.4.5 Relationship between body size and taxon description ...................................... 82 3.5 Prospects .................................................................................................................... 84 3.5.1 Extension of the database ................................................................................... 84 3.6 Acknowledgements ................................................................................................... 84 3.7 Supplementary Material ............................................................................................ 85 4 Multiple glacial refugia of the low-dispersal ground beetle Carabus irregularis: Molecular data support predictions of species distribution models ......................................... 87 4.1 Abstract ...................................................................................................................... 88 4.2 Introduction ............................................................................................................... 88 CONTENTS 4.3 Material and methods ................................................................................................ 90 4.3.1 Study species ...................................................................................................... 90 4.3.2 Species distribution modelling ........................................................................... 91 4.3.3 Sample collection and molecular methods ......................................................... 94 4.3.4 Sequence alignment and phylogenetic analysis ................................................. 95 4.3.5 Intraspecific divergence and genetic diversity ................................................... 96 4.4 Results ....................................................................................................................... 96 4.4.1 Current and past predicted distribution .............................................................. 96 4.4.2 Sequence characteristics ....................................................................................