Natural and Human-Induced Dynamics in Plant–Animal Mutualistic Networks
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Natural and human-induced dynamics in plant–animal mutualistic networks Dissertation von Jörg Albrecht Philipps-Universität Marburg April 2014 Natural and human-induced dynamics in plant–animal mutualistic networks Dissertation zur Erlangung des Doktorgrades der Naturwissenschaften (Dr. rer. nat.) dem Fachbereich Biologie der Philipps-Universität Marburg vorgelegt von Jörg Albrecht aus Suhl Marburg an der Lahn, April 2014 Vom Fachbereich Biologie der Philipps-Universität Marburg als Dissertation am 15. April 2014 angenommen. Erstgutachterin: Junior-Prof. Dr. Nina Farwig Zweitgutachter: Prof. Dr. Roland Brandl Tag der mündlichen Prüfung am 24. Juni 2014 TABLE OF CONTENTS TABLE OF CONTENTS Chapter 1 — General introduction 1 Species interactions form the backbone of ecological communities ....................................................................... 3 Pollination and seed dispersal mutualisms are key ecological interactions............................................................. 4 Interaction networks describe community-wide patterns of interspecific interactions ............................................ 4 Frugivore-mediated competition and facilitation in plant communities .................................................................. 5 Consumer-resource dynamics and functional redundancy in plant–frugivore networks......................................... 5 Ecological perturbation of coupled mutualistic networks ....................................................................................... 6 Aims of the thesis .................................................................................................................................................... 6 Chapter 2 — Frugivore-mediated competition and facilitation in plant communities 9 Introduction ............................................................................................................................................................ 11 Summary ................................................................................................................................................................ 11 Methods ................................................................................................................................................................. 12 Results .................................................................................................................................................................... 17 Discussion .............................................................................................................................................................. 18 Chapter 3 — Consumer-resource dynamics and functional redundancy in plant–frugivore networks 23 Summary ................................................................................................................................................................ 25 Introduction ............................................................................................................................................................ 25 Methods ................................................................................................................................................................. 26 Results .................................................................................................................................................................... 30 Discussion .............................................................................................................................................................. 32 Chapter 4 — Ecological perturbation of coupled mutualistic networks 37 Summary ................................................................................................................................................................ 39 Introduction ............................................................................................................................................................ 39 Results .................................................................................................................................................................... 39 Discussion .............................................................................................................................................................. 43 Methods ................................................................................................................................................................. 44 Chapter 5 — Synthesis 49 Frugivore-mediated competition and facilitation in plant communities ................................................................ 51 Consumer-resource dynamics and functional redundancy in plant–frugivore networks....................................... 51 Ecological perturbation of coupled mutualistic networks ..................................................................................... 52 Conclusion ............................................................................................................................................................. 52 TABLE OF CONTENTS Chapter 6 — Perspectives 55 Promising avenues for future research................................................................................................................... 57 Deutsche Zusammenfassung 59 Appendix chapter 2 63 Supplementary methods 2.1 Properties of the phenological niche ...................................................................................................................... 65 Supplementary methods 2.2 Standardized Kullback-Leibler divergence as a measure of phenological niche differentiation ........................... 66 Supplementary methods 2.3 Neighbourhood effects on absolute rates of fruit removal from plants ................................................................. 68 Supplementary methods 2.4 Correlation between the co-occurrence and the phenological niche similarity of the plant species ..................... 69 Appendix chapter 3 73 Supplementary methods 3.1 Monitoring of fruit abundance along transects in 2011 ......................................................................................... 75 Supplementary methods 3.2 Estimation of consumer/resource ratios and inference about competition ............................................................ 75 Supplementary methods 3.3 Partitioning of diversity into independent richness and evenness components ..................................................... 75 Supplementary methods 3.4 Partitioning of environmental and spatial effects on composition of frugivore assemblages ............................... 76 Appendix chapter 4 83 Supplementary methods 4.1 Missing interactions ............................................................................................................................................... 85 Supplementary methods 4.2 Assessment of model fit ......................................................................................................................................... 88 References 103 Danksagung / Acknowledgements 117 CHAPTER 1 GENERAL INTRODUCTION 1 GENERAL INTRODUCTION „The history of evolution and biodiversity is fundamentally a history of the evolution of species interactions. Species in pure isolation simply do not make sense.“ John N. Thompson SPECIES INTERACTIONS FORM THE BACKBONE OF determine the outcomes of interspecific interactions in ECOLOGICAL COMMUNITIES variable biotic contexts are poorly understood. Species interactions play an eminent role in the On top of this natural variation in interspecific organization and diversification of life (Ehrlich & interactions, human land-use has a substantial and Raven 1964; Dawkins & Krebs 1979), because all increasing impact on the earth’s biosphere, and on the species require one or more coevolved interactions to persistence of species and their interactions (Vitousek et survive and reproduce (Thompson 2009). Often these al. 1997; Sala et al. 2000; Tilman et al. 2001; Tylianakis et interactions link species so strongly that they leave al. 2008). Since species are embedded in large networks a long-lasting imprint in the evolutionary history of of interdependencies, we can expect that the population species lineages (Thompson 2005; Gómez, Verdú & decline or extinction of one species either directly or Perfectti 2010). Therefore, the pervasive impact of indirectly affects populations of other species (Koh et ecological interactions on the evolution, diversification al. 2004; Dunne & Williams 2009; Säterberg, Sellman & and persistence of species is evident across the entire Ebenman 2013). This is impressively illustrated by the tree of life (e.g., Farrell 1998; Dodd, Silvertown & parallel disappearance of plants and herbivorous insects Chase 1999; Wilf et al. 2005; Grant & Grant 2006; Hu in fossil records at the end of the Cretaceous (Labandeira, et al. 2008; McKenna et al. 2009; Ramírez et al. 2011; Johnson & Wilf 2002). This example highlights that Litsios et al. 2012; Gómez & Verdú 2012). In addition, ecological interactions not only fuel the coevolution but ecological interactions form the template for a variety of also the coextinction of species. Importantly, ecological processes that are essentially linked to the functioning interactions and associated ecological functions of ecosystems and to human welfare, such as pollination often collapse long before ecologists encounter the and seed dispersal or biological control (Costanza