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Foraging by consumers acts as a biotic filtering mechanism for biodiversity at the trophic level of resources. Variation in foraging behaviour has cascading effects on abundance, diversity, and functional trait composition of the community of resource species. Here we propose diversity at giving-up density (DivGUD), i.e. when foragers quit exploiting a patch, as a novel concept and simple measure quantifying cascading effects at multiple spatial scales. In experimental landscapes with an assemblage of plant seeds, patch residency of wild rodents decreased local alpha-DivGUD (via elevated mortality of species with large seeds) and regional gamma-DivGUD, while dissimilarity among patches in a landscape (beta-DivGUD) increased. By linking theories of adaptive foraging behaviour with community ecology, DivGUD allows to investigate cascading indirect predation effects, e.g. the ecology-of-fear framework, feedbacks between functional trait composition of resource species and consumer communities, and effects of inter-individual differences among foragers on the biodiversity of resource communities.
Biodiversity and abundance of wildlife has dramatically declined in agricultural landscapes. Sown, short-lived wildflower (WF) strips along the margins of crop fields are a widespread and often subsidised in agri-environmental schemes, intended to enhance biodiversity, provide refuges for wild plant and arthropod populations and to provide ecosystem services to crops. Meanwhile, WF elements are also criticised, since their functionality decreases with plant succession, the removal of aged WF strip poses an ecological trap for the attracted arthropod populations and only common and mobile species benefit. Further, insects in WF strips are impacted by pesticides from agricultural fields due to shared boundaries with crop fields and by edge effects. The performance of the measure could be improved by combining several WF strips of different successional stages, each harbouring a unique community of plants and arthropods, into persistent, composite WF block, where successional stages exist in parallel. Monitoring data on many taxa in the literature shows, that a third of species are temporarily present in an ageing WF stip, thus offering composite WF blocks should increase cumulative species richness by 28%-39% compared to annual richness in WF strips. Persistence of composite WF blocks would offer reliable refuge for animal and plant populations, also supporting their predators and herbivores. Further, WF blocks have less boundaries to crops compared to WF strips of the same area, and are less impacted by edge effects and pesticides. Policy implications. Here I suggest a change of conservation practice changing from successional WF strips to composite WF blocks. By regular removal and replacement of aged WF strips either within the block (rotational) or at its margins (rolling), the habitat heterogeneity in composite WF block could be perpetuated. Rolling composite WF blocks change locations over years, and the original location can be reconverted to arable land while a nearby WF block is still available to wildlife. A change in agricultural schemes would be necessary, since in some European countries clustered WF strips are explicitly not subsidised.
In this thesis, a collection of studies is presented that advance research on complex food webs in several directions. Food webs, as the networks of predator-prey interactions in ecosystems, are responsible for distributing the resources every organism needs to stay alive. They are thus central to our understanding of the mechanisms that support biodiversity, which in the face of increasing severity of anthropogenic global change and accelerated species loss is of highest importance, not least for our own well-being.
The studies in the first part of the thesis are concerned with general mechanisms that determine the structure and stability of food webs. It is shown how the allometric scaling of metabolic rates with the species' body masses supports their persistence in size-structured food webs (where predators are larger than their prey), and how this interacts with the adaptive adjustment of foraging efforts by consumer species to create stable food webs with a large number of coexisting species. The importance of the master trait body mass for structuring communities is further exemplified by demonstrating that the specific way the body masses of species engaging in empirically documented predator-prey interactions affect the predator's feeding rate dampens population oscillations, thereby helping both species to survive. In the first part of the thesis it is also shown that in order to understand certain phenomena of population dynamics, it may be necessary to not only take the interactions of a focal species with other species into account, but to also consider the internal structure of the population. This can refer for example to different abundances of age cohorts or developmental stages, or the way individuals of different age or stage interact with other species.
Building on these general insights, the second part of the thesis is devoted to exploring the consequences of anthropogenic global change on the persistence of species. It is first shown that warming decreases diversity in size-structured food webs. This is due to starvation of large predators on higher trophic levels, which suffer from a mismatch between their respiration and ingestion rates when temperature increases. In host-parasitoid networks, which are not size-structured, warming does not have these negative effects, but eutrophication destabilises the systems by inducing detrimental population oscillations. In further studies, the effect of habitat change is addressed. On the level of individual patches, increasing isolation of habitat patches has a similar effect as warming, as it leads to decreasing diversity due to the extinction of predators on higher trophic levels. In this case it is caused by dispersal mortality of smaller and therefore less mobile species on lower trophic levels, meaning that an increasing fraction of their biomass production is lost to the inhospitable matrix surrounding the habitat patches as they become more isolated. It is further shown that increasing habitat isolation desynchronises population oscillations between the patches, which in itself helps species to persist by dampening fluctuations on the landscape level. However, this is counteracted by an increasing strength of local population oscillations fuelled by an indirect effect of dispersal mortality on the feeding interactions. Last, a study is presented that introduces a novel mechanism for supporting diversity in metacommunities. It builds on the self-organised formation of spatial biomass patterns in the landscape, which leads to the emergence of spatio-temporally varying selection pressures that keep local communities permanently out of equilibrium and force them to continuously adapt. Because this mechanism relies on the spatial extension of the metacommunity, it is also sensitive to habitat change.
In the third part of the thesis, the consequences of biodiversity for the functioning of ecosystems are explored. The studies focus on standing stock biomass, biomass production, and trophic transfer efficiency as ecosystem functions. It is first shown that increasing the diversity of animal communities increases the total rate of intra-guild predation. However, the total biomass stock of the animal communities increases nevertheless, which also increases their exploitative pressure on the underlying plant communities. Despite this, the plant communities can maintain their standing stock biomass due to a shift of the body size spectra of both animal and plant communities towards larger species with a lower specific respiration rate. In another study it is further demonstrated that the generally positive relationship between diversity and the above mentioned ecosystem functions becomes steeper when not only the feeding interactions but also the numerous non-trophic interactions (like predator interference or competition for space) between the species of an ecosystem are taken into account. Finally, two studies are presented that demonstrate the power of functional diversity as explanatory variable. It is interpreted as the range spanned by functional traits of the species that determine their interactions. This approach allows to mechanistically understand how the ecosystem functioning of food webs with multiple trophic levels is affected by all parts of the food web and why a high functional diversity is required for efficient transportation of energy from primary producers to the top predators.
The general discussion draws some synthesising conclusions, e.g. on the predictive power of ecosystem functioning to explain diversity, and provides an outlook on future research directions.