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Plants, herbivores, and their interactions in human-modified landscapes

  • Human forest modification is among the largest global drivers of terrestrial degradation of biodiversity, species interactions, and ecosystem functioning. One of the most pertinent components, forest fragmentation, has a long history in ecological research across the globe, particularly in lower latitudes. However, we still know little how fragmentation shapes temperate ecosystems, irrespective of the ancient status quo of European deforestation. Furthermore, its interaction with another pivotal component of European forests, silvicultural management, are practically unexplored. Hence, answering the question how anthropogenic modification of temperate forests affects fundamental components of forest ecosystems is essential basic research that has been neglected thus far. Most basal ecosystem elements are plants and their insect herbivores, as they form the energetic basis of the tropic pyramid. Furthermore, their respective biodiversity, functional traits, and the networks of interactions they establish are key for a multitude of ecosystem functions, not least ecosystem stability. Hence, the thesis at hand aimed to disentangle this complex system of interdependencies of human impacts, biodiversity, species traits and inter-species interactions. The first step lay in understanding how woody plant assemblages are shaped by human forest modification. For this purpose, field investigations in 57 plots in the hyperfragmented cultural landscape of the Northern Palatinate highlands (SW Germany) were conducted, censusing > 4,000 tree/shrub individuals from 34 species. Use of novel, integrative indices for different types of land-use allowed an accurate quantification of biotic responses. Intriguingly, woody tree/shrub communities reacted strikingly positive to forest fragmentation, with increases in alpha and beta diversity, as well as proliferation of heat/drought/light adapted pioneer species. Contrarily, managed interior forests were homogenized/constrained in biodiversity, with dominance of shade/cold adapted commercial tree species. Comparisons with recently unmanaged stands (> 40 a) revealed first indications for nascent conversion to oldgrowth conditions, with larger variability in light conditions and subsequent community composition. Reactions to microclimatic conditions, the relationship between associated species traits and the corresponding species pool, as well as facilitative/constraining effects by foresters were discussed as underlying mechanisms. Reactions of herbivore assemblages to forest fragmentation and the subsequent changes in host plant communities were assessed by comprehensive sampling of > 1,000 live herbivores from 134 species in the forest understory. Diversity was – similarly to plant communities - higher in fragmentation affected habitats, particularly in edges of continuous control forests. Furthermore, average trophic specialization showed an identical pattern. Mechanistically, benefits from microclimatic conditions, host availability, as well as pronounced niche differentiation are deemed responsible. While communities were heterogeneous, with no segregation across habitats, (smallforest fragments, edges, and interior of control forests), vegetation diversity, herbivore diversity, as well as trophic specialization were identified to shape community composition. This probably reflected a gradient from generalistic/species poor vs. specialist/species rich herbivore assemblages. Insect studies conducted in forest systems are doomed to incompleteness without considering ‘the last biological frontier’, the tree canopies. To access their biodiversity, relationship to edge effects, and their conservational value, the arboricolous arthropod fauna of 24 beech (Fagus sylvatica) canopies was sampled via insecticidal knockdown (‘fogging’). This resulted in an exhaustive collection of > 46,000 specimens from 24 major taxonomic/functional groups. Abundance distributions were markedly negative exponential, indicating high abundance variability in tree crowns. Individuals of six pertinent orders were identified to species level, returning > 3,100 individuals from 175 species and 52 families. This high diversity did marginally differ across habitats, with slightly higher species richness in edge canopies. However, communities in edge crowns were noticeably more heterogeneous than those in the forest interior, possibly due to higher variability in environmental edge conditions. In total, 49 species with protective value were identified, of which only one showed habitat preferences (for near-natural interior forests). Among them, six species (all beetles, Coleoptera) were classified as ‘priority species’ for conservation efforts. Hence, beech canopies of the Northern Palatinate highlands can be considered strongholds of insect biodiversity, incorporating many species of particular protective value. The intricacy of plant-herbivore interaction networks and their relationship to forest fragmentation is largely unexplored, particularly in Central Europe. Illumination of this matter is all the more important, as ecological networks are highly relevant for ecosystem stability, particularly in the face of additional anthropogenic disturbances, such as climate change. Hence, plant-herbivore interaction networks (PHNs) were constructed from woody plants and their associated herbivores, sampled alive in the understory. Herbivory verification was achieved using no-choice-feeding assays, as well as literature references. In total, networks across small forest fragments, edges, and the forest interior consisted of 696 interactions. Network complexity and trophic niche redundancy were compared across habitats using a rarefaction-like resampling procedure. PHNs in fragmentation affected forest habitats were significantly more complex, as well as more redundant in their realized niches, despite being composed of relatively more specialist species. Furthermore, network robustness to climate change was quantified utilizing four different scenarios for climate change susceptibility of involved plants. In this procedure, remaining herbivores in the network were measured upon successive loss of their host plant species. Consistently, PHNs in edges (and to a smaller degree in small fragments) withstood primary extinction of plant species longer, making them more robust. This was attributed to the high prevalence of heat/drought-adapted species, as well as to beneficial effects of network topography (complexity and redundancy). Consequently, strong correlative relationships were found between realized niche redundancy and climate change robustness of PHNs. This was both the first time that biologically realistic extinctions (instead of e.g.random extinctions) were used to measure network robustness, and that topographical network parameters were identified as potential indicators for network robustness against climate change. In synthesis, in the light of global biotic degradation due to human forest modification, the necessity to differentiate must be claimed. Ecosystems react differently to anthropogenic disturbances, and it seems the particular features present in Central European forests (ancient deforestation, extensive management, and, most importantly, high richness in open-forest plant species) cause partly opposed patterns to other biomes. Lenient microclimates and diverse plant communities facilitate equally diverse herbivore assemblages, and hence complex and robust networks, opposed to the forest interior. Therefore, in the reality of extensively used cultural landscapes, fragmentation affected forest ecosystems, particularly forest edges, can be perceived as reservoir for biodiversity, and ecosystem functionality. Nevertheless, as practically all forest habitats considered in this thesis are under human cultivation, recommendations for ecological enhancement of all forest habitats are discussed.

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Author:Kevin Bähner
Advisor:Burkhard Büdel, Rainer Wirth
Document Type:Doctoral Thesis
Language of publication:English
Publication Date:2016/07/27
Date of Publication:2016/07/27
Publishing Institute:Technische Universität Kaiserslautern
Granting Institute:Technische Universität Kaiserslautern
Acceptance Date of the Thesis:2016/07/21
Date of the Publication (Server):2016/07/27
Tag:anthropogenic effects; biodiversity; climate change; diversity; ecology; forest fragmentation; forest management; interaction networks; plant-herbivore interactions; stability
GND-Keyword:Pflanzenfressende Insekten; Nahrungsnetz
Number of page:206
Faculties / Organisational entities:Fachbereich Biologie
CCS-Classification (computer science):A. General Literature
DDC-Cassification:5 Naturwissenschaften und Mathematik / 570 Biowissenschaften, Biologie
MSC-Classification (mathematics):92-XX BIOLOGY AND OTHER NATURAL SCIENCES / 92-02 Research exposition (monographs, survey articles)
PACS-Classification (physics):00.00.00 GENERAL
Licence (German):Standard gemäß KLUEDO-Leitlinien vom 30.07.2015