Density-dependent population dynamics and dispersal in heterogeneous metapopulations.

1. Metapopulation microcosms were constructed to test the effect of four different types of habitat heterogeneity on the dynamics and dispersal in spatially extended systems; homogeneity, spatial heterogeneity, temporal heterogeneity and spatio-temporal heterogeneity. Resources were distributed across discrete habitat patches in bruchid beetle (Callosobruchus maculatus) metapopulations, and long-term time series were recorded. 2. Mathematical models were fitted to the long-term time series from the experimental systems using a maximum likelihood approach. Models were composed of separate birth, death, emigration and immigration terms all of which incorporated stochasticity drawn from different probability distributions. Models with density-dependent and density-independent birth, death and emigration terms were investigated and, in each case, the model that best described the empirical data was identified. 3. At the local scale, population sizes differed between patches depending on the type of heterogeneity. Larger populations were associated with higher resource availabilities. As a result of this, the variation between local population sizes was greatest when there was spatial heterogeneity in which mean resource abundance varied from patch to patch. Variation in population sizes within patches was largest when there was temporal heterogeneity. 4. Density-dependent processes leading to the regulation of local population dynamics in our experimental systems were strongest in homogeneity or temporal heterogeneity treatments. Associated with this, we found that these systems were best described using mathematical models with density dependence acting on mortality. In contrast, spatial and spatio-temporal time series were adequately described using density-independent population processes. 5. Experimental metapopulations showed varying degrees of density-dependent dispersal. Local net dispersal each week was primarily driven by the local population size and secondarily affected by neighbourhood population density. Mathematical population models illustrated the importance of explicit description of density-dependent dispersal in all systems except the homogeneous metapopulations