Dynamics of ecological communities in variable environments : local and spatial processes

Abstract: The ecosystems of the world are currently facing a variety of anthropogenic perturbations, such as climate change, fragmentation and destruction of habitat, overexploitation of natural resources and invasions of alien species. How the ecosystems will be affected is not only dependent on the direct effects of the perturbations on individual species but also on the trophic structure and interaction patterns of the ecological community. Of particular current concern is the response of ecological communities to climate change. Increased global temperature is expected to cause an increased intensity and frequency of weather extremes. A more unpredictable and more variable environment will have important consequences not only for individual species but also for the dynamics of the entire community. If we are to fully understand the joint effects of a changing climate and habitat fragmentation, there is also a need to understand the spatial aspects of community dynamics. In the present work we use dynamic models to theoretically explore the importance of local (Paper I and II) and spatial processes (Paper III-V) for the response of multi-trophic communities to different kinds of perturbations.In paper I we investigate how species richness and correlation in species responses to a highly variable environment affect the risk of extinction cascades. We find that the risk of extinction cascades increases with increasing species richness especially when the correlation among species is low. Initial stochastic extinctions of primary producer species unleash bottomup extinction cascades, where specialist consumers are especially vulnerable. Although the risks of extinction cascades were higher in the species-rich systems, we found that the temporal stability of aggregate abundance of primary producers increased with increasing richness. Thus, species richness had a two-sided effect on community stability. Also during the extinction cascades it is possible that more robust species and interaction patterns will be selected which would further act to stabilize the post-extinction communities. In paper II we explore how the process of disassembly affects the structure of the interaction network and the robustness of the community to additional disturbances. We find that the disassembled communities are structurally different and more resistant to disturbances than equally sized communities that have not gone through a phase of disassembly. The disassembled communities are topologically as well as dynamically more stable than non-disassembled communities.In paper III, IV and V we expand the analysis to incorporate the spatial dimension. In paper III we analyze how metacommunities (a set of local communities coupled by species dispersal) in spatially explicit landscapes respond to environmental variation. We examine how this response is affected by varying 1) species richness in the local communities, 2) the degree of correlation in species response to the environmental variation, between species within patches (species correlation) and among patches (spatial correlation) and 3) dispersal pattern of species. First we can confirm that our previous findings from paper I regarding local species richness and correlation among species within a patch are robust to the inclusion of a spatial dimension. However our results also show that the spatial dynamics are of great importance: first we find that the risk of global extinctions increases with increasing spatial correlation. Second we find that the pattern and rate of dispersal are important; a high migration rate in combination with localized dispersal decrease the risk of global extinctions whereas a global dispersal pattern increases the risk of global extinctions. When dispersal is global the subpopulations of a species become more synchronized which reduces the potential for a patch to become recolonized following extinctions. We also demonstrate the importance of both local and spatial processes when examining the temporal stability of primary production at the scale of metapopulations, local communities and metacommunities.In paper IV we investigate how the spatial structure of the landscape (number of patches) and dispersal pattern of species affect a metacommunities response to increased mortality during dispersal and local loss of species. We find a two-sided effect of dispersal on metacommunity persistence; on the one hand, high migration rate significantly reduces the risk of bottom-up extinction cascades following the removal of a species when dispersal involves no risk. On the other hand, high migration rate increases extinction risks when dispersal imposes a risk to the dispersing individuals, especially when dispersal is global. Species with long generation times at the highest trophic level are particularly vulnerable to extinction when dispersal involves a risk. These results suggest that decreasing the mortality risk of dispersing individuals by constructing habitat corridors or by improving the quality of the habitat matrix might greatly increase the robustness of metacommunities to local loss of species by enhancing recolonisations and rescue effects.In paper V we use network theory to identify keystone patches in the landscape, patches that are of critical importance for the local and global persistence of species in the metacommunity. By deleting patches one at a time and investigating the risk of local and global extinctions we quantified the importance of a patch’s position in the landscape for the persistence of species within the metacommunity. A selection of indices were used including some local indices that measure the connectedness of a patch in the intact network and some indices which measure the decrease in a global index after the deletion of the patch from the network. Global indices are those that give an impression of the connectivity of the entire patch network. We find that deletion of patches contributing strongly to the connectivity of the entire patch network had the most negative effect on species persistence.