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Drivers of Plant Diversity in Grassland Metacommunities: Integrating Local and Landscape-Scale Factors of Taxonomic and Functional Community Structure

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Drivers of Plant Diversity in Grassland Metacommunities: Integrating Local and Landscape-Scale Factors of Taxonomic and Functional Community Structure Wissenschaftszentrum Weihenstephan für Ernährung, Landnutzung und Umwelt Lehrstuhl für Renaturierungsökologie Drivers of plant diversity in grassland metacommunities: Integrating local and landscape-scale factors of taxonomic and functional community structure Timo Conradi Vollständiger Abdruck der von der Fakultät Wissenschaftszentrum Weihenstephan für Ernährung, Landnutzung und Umwelt der Technischen Universität München zur Erlangung des akademischen Grades eines Doktors der Naturwissenschaften genehmigten Dissertation. Vorsitzender: Univ.-Prof. Dr. Johannes Schnyder Prüfer der Dissertation: 1. Univ.-Prof. Dr. Johannes Kollmann 2. Univ.-Prof. Dr. Wolfgang W. Weisser 3. Univ.-Prof. Dr. Carl Beierkuhnlein, Universität Bayreuth (nur schriftliche Begutachtung); Univ.-Prof. Dr. Anton Fischer (nur mündlicher Prüfer) Die Dissertation wurde am 26.11.2015 bei der Technischen Universität München eingereicht und durch das Wissenschaftszentrum Weihenstephan für Ernährung, Landnutzung und Umwelt am 17.03.2016 angenommen. für Mama, Papa und Steff Content Summary ...........................................................................................................................................................i Zusammenfassung ..................................................................................................................................... iv Chapter 1 General introduction.................................................................................................................................. 1 Chapter 2 Resource availability determines the importance of niche-based vs. stochastic community assembly in grasslands .................................................................................................. 15 Chapter 3 Drivers of beta diversity in human-transformed landscapes: the key role of regional soil fertility for community assembly processes ........................................................................ 35 Chapter 4 Species pools, mass effects and environmental sorting interact to determine plant diversity and functional trait composition in secondary grasslands ................................. 59 Chapter 5 Synopsis ....................................................................................................................................................... 83 References ................................................................................................................................................... 90 Danksagung .............................................................................................................................................. 106 Intended publications and author contributions ...................................................................... 107 Appendix A1 – Rules for trait data extraction ....................................................................................................... I A2 – Curriculum vitae ............................................................................................................................. VII A3 – Publications ........................................................................................................................................IX Summary Summary Human land use is the leading driver of global losses in biodiversity. The transformation of natural ecosystems leads to widespread loss of primary habitats and divides formerly continuous landscapes into smaller and isolated fragments. Another significant characteristic of cultivated landscapes is that secondary ecosystems can re-establish after agricultural or silvicultural land use. Although their site conditions might be altered, such patches often exhibit sufficient habitat quality and thus can contribute to regional population persistence of at least some species. As a consequence, spatial dynamics of populations and communities in transformed landscapes are not only characterized by altered dispersal processes, but also by variable local interactions resulting from natural or human-induced variation in site conditions among habitat fragments. The interaction of regional dispersal and variation in site conditions, as well as the conditions under which dispersal effectively regulates biodiversity in transformed landscapes are poorly understood. Therefore, the goals of this dissertation were (1) to improve the scientific understanding of how patterns of plant diversity are generated in transformed landscapes, and (2) to identify how environmental conditions drive the degree to which dispersal influences local and regional species co-existence. To accomplish these goals, the thesis combines experimental and observational investigations in historical calcareous grassland landscapes in southern Germany with modern analytical methods in three complementary studies. The first study investigated how soil nutrient availability determines the importance of stochastic processes such as dispersal vs. niche-based selection for variation in plant species composition (i.e. beta diversity) among grasslands. For this purpose, a field experiment was established in which three levels of soil nutrient concentrations were manipulated; the successional dynamics of sown grassland communities were monitored over 3 years. Observed variation in species composition among replicate grasslands was then compared with expectations from null models. The resulting data revealed that compositional variation among oligotrophic grasslands hardly deviated from null expectations, whereas the deviation increased significantly with soil nutrient concentrations. These findings suggest that the effects of dispersal on landscape-scale biodiversity patterns decrease with increasing regional soil fertility. i SUMMARY The second study explored whether these experimental findings are transferable to real landscapes, and analyses the drivers of compositional variation among habitat fragments in transformed landscapes. Two historical calcareous grassland landscapes with contrasting regional soil fertility were compared. Using standardised methods in both landscapes, plant species composition, soil factors and – as measures of recent and past dispersal processes – contemporary and historical landscape context of multiple grassland fragments were studied. The leading drivers of species composition were identified using redundancy analysis, model selection and variation partitioning. Variation in species composition was driven by past dispersal processes in the landscape with lower regional soil fertility, whereas it was driven by variation in soil factors in the landscape with higher regional soil fertility. This confirms the experimental findings and emphasizes that regional soil nutrient concentrations are a critical environmental feature that determines the importance of dispersal vs. niche-based selection for biodiversity patterns in transformed landscapes. Secondary habitat patches can potentially contribute to the persistence and regional dynamics of plant populations. Therefore, the third study focused on the limiting factors for utilizing this potential. Using mixed effects modelling in combination with an information theoretic approach and multi-model inference, this study dissected the principal drivers of species richness, the number of habitat specialists, and the functional trait composition of secondary grasslands. To improve the mechanistic understanding of operating ecological processes, the study used traits relevant for dispersal and interactions with local site conditions. The results showed that dispersal limitation of many habitat specialists during early stages of succession is not only the main driver of plant diversity in secondary grassland, but also strongly influences their functional trait composition. The positive effects of high immigration rates of specialist species even outweighed well-known negative effects of high soil nutrient concentrations on species numbers. Besides the dominant role of dispersal, there was also a significant influence of soil moisture that selected species according to their canopy height. This shows that dispersal and certain site factors interact in determining different aspects of plant community structure in secondary grasslands. This PhD thesis demonstrates that combining complementary research approaches, large datasets and advanced analytical methods can produce novel insights into the drivers of plant diversity in human-transformed landscapes. In particular, the ii SUMMARY dissertation shows that dispersal can regulate biodiversity patterns under certain environmental conditions only, and reveals opportunities for and limits to directing successional trajectories in secondary grasslands. The results emphasize that conservation managers should actively assist dispersal and simultaneously reduce high soil nutrient concentrations. Successional trajectories in secondary grasslands can be manipulated using active species transfer measures during early stages of succession. iii Zusammenfassung Zusammenfassung Die anthropogene Transformation natürlicher oder traditionell genutzter Landschaften für eine intensive land- und forstwirtschaftliche Produktion ist weltweit der wichtigste Treiber des hohen Biodiversitätsverlusts im 20. und 21. Jahrhundert. Veränderte Landnutzung führt zu einem großflächigen Verlust von Lebensräumen vieler Arten und unterteilt ehemals zusammenhängende Habitate in kleinere und isolierte Fragmente. Ein wesentliches Merkmal solcher Landschaften ist aber auch, dass sekundäre Lebensräume
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