Combined Effects of Climate Change and Extreme Events on Plants, Arthropods and Their Interactions
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Combined effects of climate change and extreme events on plants, arthropods and their interactions Dissertation zur Erlangung des naturwissenschaftlichen Doktorgrades der Julius-Maximilians-Universität Würzburg vorgelegt von Annette Leingärtner geboren in Benediktbeuern Würzburg 2013 Mitglieder der Promotionskommission: Vorsitzender: ___________________________ Gutachter: Prof. Dr. Ingolf Steffan-Dewenter Gutachter: Prof. Dr. Flavio Roces Eingereicht am: 11. Juli 2013 Tag des Promotionskolloquiums: 02. Oktober 2013 Doktorurkunde ausgehändigt am: ___________________________ Lilac-breasted Roller (Coracias caudatus) in the Etosha National Park (Namibia) “For the best is only bought at the cost of great pain… Or so says the legend” The Thorn Birds by Colleen McCullough 3 Table of Contents Ehrenwörtliche Erklärung .................................................................................................... 5 Curriculum vitae .................................................................................................................. 6 Publication list ..................................................................................................................... 7 Summary (English) .............................................................................................................. 8 Zusammenfassung (German) ............................................................................................. 11 I. General Introduction .................................................................................................. 15 II. Changes in the composition of butterfly assemblages and life-history traits along an altitudinal gradient ................................................................................................ 23 III. Combined effects of climate and management on plant diversity and pollination type in alpine grasslands ............................................................................................ 44 IV. Combined effects of extreme climatic events and elevation on nutritional quality and herbivory rates of alpine plants ........................................................................... 61 V. Phenological response of grassland species to manipulative snowmelt and drought along an altitudinal gradient ......................................................................... 79 VI. Elevation and experimental snowmelt manipulation affect emergence phenology and abundance of soil-hibernating arthropods ........................................................... 99 VII. General Discussion .................................................................................................. 119 References ........................................................................................................................ 124 Author Contributions ....................................................................................................... 146 Acknowledgements .......................................................................................................... 151 4 Ehrenwörtliche Erklärung Ehrenwörtliche Erklärung Hiermit erkläre ich ehrenwörtlich, dass ich die vorliegende Dissertation mit dem Titel „Combined effects of climate change and extreme events on plants, arthropods and their interactions“ selbständig, am Lehrstuhl für Tierökologie und Tropenbiologie (Zoologie III) der Julius-Maximilians-Universität Würzburg, unter der Anleitung und Betreuung durch Herrn PD Dr. Jochen Krauss und Herrn Prof. Dr. Ingolf Steffan-Dewenter angefertigt habe und dabei keine anderen als die von mir angegebenen Quellen und Hilfsmittel verwendet habe. Ich erkläre außerdem, dass die vorliegende Dissertation weder in gleicher, noch in ähnlicher Form bereits in einem Prüfungsverfahren vorgelegen hat. Des Weiteren habe ich außer den mit dem Zulassungsantrag urkundlich vorgelegten Graden keine weiteren akademischen Grade erworben oder zu erwerben versucht. Würzburg, 10.07.2013 Annette Leingärtner 5 Curriculum vitae Curriculum vitae Name Annette Leingärtner Date of birth Place of birth 2010 – 2013 PhD-student at the University of Würzburg (Animal Ecology and Tropical Ecology) 2009 – 2010 PhD-student at the University of Bayreuth (Animal Ecology I, Population Ecology) 2008 – 2009 Research assistant at the Ludwig-Maximilians-University in Munich 2007 Diploma thesis in Biology at the Ludwig-Maximilians-University in Munich; Title: “Coevolutionary arms race of the slavemaking ants Protomognathus americanus and Hapagoxenus sublaevis and their host species” 2002 – 2007 Student at the Ludwig-Maximilians-University in Munich 2002 Abitur at the Gabriel-von-Seidl-Gymnasium in Bad Tölz 6 Publication list Publication list Leingärtner, A., Krauss, J., Steffan-Dewenter, I. (subm.) Changes in the composition of butterfly assemblages and life-history traits along an altitudinal gradient. Leingärtner, A., Hoiss, B., Krauss, J., Steffan-Dewenter, I. (subm.) Comined effects of extreme events and elevation on nutritional quality and herbivory rates of alpine plants. Leingärtner, A., Krauss, J., Steffan-Dewenter, I. (subm.) Elevation and experimental snowmelt manipulation affect emergence phenology and abundance of soil- hibernating arthropods. Hoiss, B., Gaviria, J., Leingärtner, A., Krauss, J., Steffan-Dewenter, I. (2013) Combined effects of climate and management on plant diversity and pollination type in alpine grasslands. Diversity and Distributions 19, 386 – 395. Cornelius, C., Leingärtner, A., Hoiss, B., Krauss, J., Steffan-Dewenter, I., Menzel, A. (2013) Phenological response of grassland species to manipulative snowmelt and drought along an altitudinal gradient. Journal of Experimental Botany 64, 241 – 251. Pamminger, T., Leingärtner, A., Achenbach, A., Kleeberg, I., Pennings, P., Foitzik, S. Geographic distribution of the anti-parasite trait "slave rebellion". (2013) Evolutionary Ecology 27, 39-49. Germann, C., Leingärtner, A., Szallies, A. (2012) Ceutorhynchus hutchinsiae TEMPÈRE, 1975 – neu für Deutschland (Coleoptera, Curculionidae). Entomologische Nachrichten und Berichte 56, 71 – 72 Haslberger, A., Leingärtner, A. (2010) Zygaena exulans (HOHENWARTH, 1792) und Leptopterix hirsutella ([DENIS & SCHIFFERMÜLLER], 1775) in den bayerischen Alpen: aktuelle Nachweise von verschollenen alpinen Arten. Beiträge zur bayerischen Entomofaunistik 10, 19 – 22 Witte, V., Leingärtner, A., Sabas, L., Hashim, R., Foitzik, S. (2008) “Symbiont microcosm in an ant society and the diversity of interspecific interactions”. Animal Behaviour 76: 1477 - 1486. 7 Summary (English) Summary (English) I. Global climate change directly and indirectly influences biotic and abiotic components of ecosystems. Changes in abiotic ecosystem components caused by climate change comprise temperature increases, precipitation changes and more frequently occurring extreme events. Mediated by these abiotic changes, biotic ecosystem components including all living organisms will also change. Expected changes of plants and animals are advanced phenologies and range shifts towards higher latitudes and altitudes which presumably induce changes in species interactions and composition. Altitudinal gradients provide an optimal opportunity for climate change studies, because they serve as natural experiments due to fast changing climatic conditions within short distances. In this dissertation two different approaches were conducted to reveal species and community responses to climate change. First, species richness and community trait analyses along an altitudinal gradient in the Bavarian Alps (chapters II, III) and second, climate change manipulation experiments under different climatic contexts (chapters IV, V, IV). II. We performed biodiversity surveys of butterfly and diurnal moth species on 34 grassland sites along an altitudinal gradient in the National Park Berchtesgaden. Additionally, we analysed the dominance structure of life-history traits in butterfly assemblages along altitude. Species richness of butterflies and diurnal moths decreased with increasing altitude. The dominance of certain life-history-traits changed along the altitudinal gradient with a higher proportion of larger-winged species and species with higher egg numbers towards higher altitudes. However, the mean egg maturation time, population density and geographic distribution within butterfly assemblages decreased with increasing altitude. Our results indicate that butterfly assemblages were mainly shaped by environmental filtering. We conclude that butterfly assemblages at higher altitudes will presumably lack adaptive capacity to future climatic conditions, because of specific trait combinations. III. In addition to butterfly and diurnal moth species richness we also studied plant species richness in combination with pollination type analyses along the altitudinal gradient. The management type of the alpine grasslands was also integrated in the analyses to detect combined effects of climate and management on plant diversity and pollination type. Plant species richness was highest at intermediate altitudes, whereby the 8 Summary (English) management type influenced the plant diversity with more plant species at grazed compared to mown or non-managed grasslands. The pollination type was affected by both the changing climate along the gradient and the management type. These results suggest that extensive grazing can maintain high plant diversity along the whole altitudinal gradient. With ongoing climate change the diversity peak of plants may shift upwards, which