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Traits in the Light of Ecology and Conservation of Ground Beetles

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Traits in the Light of Ecology and Conservation of Ground Beetles Traits in the light of ecology and conservation of ground beetles Von der Fakultät Nachhaltigkeit der Leuphana Universität Lüneburg zur Erlangung des Grades Doktorin der Naturwissenschaften - Dr. rer. nat. – genehmigte Dissertation von Dorothea Irmgard Ilse Nolte geb. Ehlers am 18.07.1987 in Bielefeld 2018 Eingereicht am: 09. November 2018 Mündliche Verteidigung am: 25. September 2019 Erstbetreuer und Erstgutachter: Prof. Dr. Thorsten Assmann Zweitgutachterin: Prof. Dr. Tamar Dayan Drittgutachter: Prof. Dr. Pietro Brandmayr Die einzelnen Beiträge des kumulativen Dissertationsvorhabens sind oder werden ggf. inkl. des Rahmenpa- piers wie folgt veröffentlicht: Nolte, D., Boutaud, E., Kotze, D. J., Schuldt, A., and Assmann, T. (2019). Habitat specialization, distribution range size and body size drive extinction risk in carabid beetles. Biodiversity and Conservation, 28, 1267-1283. Nolte, D., Schuldt, A., Gossner, M.M., Ulrich, W. and Assmann, T. (2017). Functional traits drive ground beetle community structures in Central European forests: Implications for conservation. Biological Conservation, 213, 5–12. Homburg, K., Drees, C., Boutaud, E., Nolte, D., Schuett, W., Zumstein, P., von Ruschkowski, E. and Assmann, T. (2019). Where have all the beetles gone? Long-term study reveals carabid species decline in a nature reserve in Northern Germany. Insect Conservation and Diversity, 12, 268-277. Veröffentlichungsjahr: 2019 "Look deep into nature, and then you will understand everything better." - Albert Einstein Nature awakens a great fascination in all of us and gives us a feeling of balance and peace of mind. Wherever you look, there is always something to discover. The plethora of habitats, species and various adaptation strategies is the true secret of nature’s success. But nature’s life insurance is in danger! II Copyright notice Chapters 2, 3, 4 and 5 have been either published, submitted for publication or are in preparation for publication in international peer-reviewed 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 III Contents SUMMARY 1 ZUSAMMENFASSUNG 3 1. GENERAL INTRODUCTION5 1.1. Background: Biodiversity . .5 1.2. Species traits promote extinction risks . .6 1.3. Ground beetles as model taxon . .7 1.4. Aims, research questions and methods . .9 1.4.1. Patterns and effects of ground beetle species traits across a large Central Euro- pean species pool . .9 1.4.2. Effects of ground beetle species traits on the species persistence of smaller species pools of forests . 11 2. ECOLOGICAL AND DISTRIBUTIONAL TRAITS VARY BETWEEN GROUND BEETLES OF DIFFER- ENT HABITAT TYPES IN CENTRAL EUROPE 13 2.1. Introduction . 14 2.2. Material and Methods . 16 2.2.1. Ground beetles and species traits . 16 2.2.2. Statistical analysis . 16 2.3. Results . 17 2.3.1. Analysis of phylogenetic relatedness . 17 2.3.2. Relations between traits . 18 2.3.3. Traits in different habitat types . 18 2.4. Discussion . 21 2.4.1. Relations between species traits . 21 2.4.2. Habitat-specific filtering of species traits . 22 2.5. Conclusion . 23 A. Supplementary material . 25 3. HABITAT SPECIALIZATION, DISTRIBUTION RANGE SIZE AND BODY SIZE DRIVE EXTINCTION RISK IN CARABID BEETLES 29 3.1. Introduction . 30 3.2. Methods . 32 3.2.1. Carabid beetles and extinction risk . 32 3.2.2. Species traits . 32 3.2.3. Statistical analysis . 32 3.3. Results . 33 3.4. Discussion . 36 3.4.1. Species traits as drivers of extinction risk . 36 3.4.2. Habitat preference as a drivers of extinction risk . 37 3.4.3. Implication for nature conservation . 38 IV Contents B. Supplementary material . 40 4. FUNCTIONAL TRAITS DRIVE GROUND BEETLE COMMUNITY STRUCTURES IN CENTRAL EU- ROPEAN FORESTS: IMPLICATIONS FOR CONSERVATION 41 4.1. Introduction . 42 4.2. Material and methods . 43 4.2.1. Data compilation . 43 4.2.2. Species traits . 44 4.2.3. Data analysis . 45 4.3. Results . 46 4.3.1. Patterns of richness and abundance . 46 4.3.2. Species co-occurrences . 46 4.3.3. Occurrence ranks and species traits . 48 4.4. Discussion . 49 4.4.1. Nestedness of communities . 49 4.4.2. Species occurrence ranks . 49 4.4.3. Species traits shape community composition . 50 4.4.4. Implications for habitat management of Central European forests . 51 C. Supplementary material . 53 5. WHERE HAVE ALL THE BEETLES GONE? LONG-TERM STUDY REVEALS CARABID SPECIES DE- CLINE IN A NATURE RESERVE IN NORTHERN GERMANY 59 5.1. Introduction . 60 5.2. Material and Methods . 61 5.2.1. Study site . 61 5.2.2. Trapping . 61 5.2.3. Data analysis . 62 5.3. Results . 64 5.3.1. Measures of diversity . 64 5.3.2. Trait relationships and affected traits . 64 5.4. Discussion . 67 5.4.1. Habitat . 67 5.4.2. Species traits . 68 5.4.3. Climate change . 68 5.5. Conclusions . 69 D. Supplementary material . 70 6. MAIN FINDINGS AND FUTURE PROSPECTS 73 6.1. Relations between ground beetle traits . 73 6.1.1. Impact of phylogenetic relatedness on ground beetle trait approaches . 74 6.1.2. Species traits in ground beetles and their influence on biodiversity . 75 ACKNOWLEDGEMENTS 81 Contents V REFERENCES 96 APPENDIX 97 Article overview and authors’ contribution to articles . 97 List of Publications . 99 SUMMARY 1 SUMMARY We are in a phase of an alarming biodiversity loss, by scientist already referred to as Earth’s sixth mass extinction. According to estimations, the current extinction rates are 100 to 1000 times higher than those predicted from fossil records. To counteract species loss and preserve the remaining biodiver- sity, with its important ecosystem functioning and services essential to human well-being, there is an urgent need to develop promising and long-term conservation strategies. In order to achieve these goals, extensive research to gain a better understanding of the general mechanisms underlying com- munity diversity is of greatest importance. Especially, the identification of intrinsic ecological and dis- tributional species traits is receiving increased attention in ecology and conservation biology research. Depending on the expression of their traits, species perform particular ecosystem functions and re- spond in a specific manner to environmental conditions. The identification of the effect of certain traits on community compositions can therefore significantly improve our understanding of species extinction processes and help to develop valuable and appropriate recommendations for conservation management. As trait-based analyses are applicable to different geographical, temporal and taxonom- ical scales, they may even allow for a broader generalization if similar results are found on different scales, i.e. for local species pools, the complete species pools of different habitat types or the entire species pool across several habitat types including different climatic regions. Although insects make up the largest part of animal diversity and provide essential ecosystem services in form of e.g. pollination, pest control, and decomposition, the majority of studies on extinctions have mainly focused on vertebrates. Among invertebrates either charismatic taxa or those targeted by conservation laws have been investigated until now (e.g. butterflies or saproxylic beetles). Being highly species-rich and trait-diverse, ground beetles (Coleoptera: Carabidae) should be even more suitable for conducting trait-based analyses. Thus, using ground beetles as a model taxon, four case studies focusing on the analyses of traits form the basis of this doctoral thesis. The work of this thesis was conducted with the aim of gaining general insights on the influence of species traits on ground beetle community compositions, such as habitat occupancy and species vulnerability to extinction, for instance. An important aspect when investigating species traits is the consideration of confounding factors which could influence the results, such as dependent relations between the different traits. Compil- ing a large dataset of 566 Central European species, I identified that dependent relations between the six tested traits of ground beetles (distribution range size, habitat specialization, body size, hind-wing morphology, breeding season and trophic level) are highly common. Across all identified dependent trait relations, the relation between body size and hind wing morphology or range size and hind wing morphology showed the strongest significant dependencies. Since the consideration of trait relations is necessary to provide reliable interpretations, all analyses of this thesis tested several traits simulta- neously and considered possible trait interactions. Studies on local communities found specific traits characterizing the local species pools of certain habitat types. Here, the species pools of seven different habitat types (coastal, forest, mountain, open, riparian, wetland and special habitat) were used to determine habitat-specific trait filters. The identi- fied traits, characteristic for certain habitat types, were in most cases in accordance with the previous findings on local communities. Across Germany, the.
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