Stepping stones are crucial for species' long-distance dispersal and range expansion through habitat networks.
Abstract
Climate and land-use changes will require species to move large distances following shifts in their suitable habitats, which will frequently involve traversing intensively human-modified landscapes. Practitioners will therefore need to evaluate and act to enhance the degree to which habitat patches scattered throughout the landscape may function as stepping stones facilitating dispersal among otherwise isolated habitat areas. We formulate a new generalized network model of habitat connectivity that accounts for the number of dispersing individuals and for long-distance dispersal processes across generations. By doing so, we bridge the gap between complex dynamic population models, which are generally too data demanding and hence difficult to apply in practical wide-scale decision-making, and simpler static connectivity models that only consider the amount of habitat that can be reached by a single average disperser during its life span. We find that the loss of intermediate and sufficiently large stepping-stone habitat patches can cause a sharp decline in the distance that can be traversed by species (critical spatial thresholds) that cannot be effectively compensated by other factors previously regarded as crucial for long-distance dispersal (fat-tailed dispersal kernels, source population size). We corroborate our findings by showing that our model largely outperforms previous connectivity models in explaining the large-scale range expansion of a forest bird species, the Black Woodpecker Dryocopus martius, over a 20-year period. The capacity of species to exploit the opportunities created by networks of stepping-stone patches largely depends on species-specific life-history traits, suggesting that species assemblages traversing fragmented landscapes may be exposed to a spatial filtering process driving long-term changes in community composition. Synthesis and applications. Previous static connectivity models seriously underestimate the importance of stepping-stone patches in sustaining rare but crucial dispersal events. We provide a conceptually broader model that shows that stepping stones (i) must be of sufficient size to be of conservation value, (ii) are particularly crucial for the spread of species (either native or invasive) or genotypes over long distances and (iii) can effectively reduce the isolation of the largest habitat blocks in reserves, therefore largely contributing to species persistence across wide spatial and temporal scales. Previous static connectivity models seriously underestimate the importance of stepping-stone patches in sustaining rare but crucial dispersal events. We provide a conceptually broader model that shows that stepping stones (i) must be of sufficient size to be of conservation value, (ii) are particularly crucial for the spread of species (either native or invasive) or genotypes over long distances and (iii) can effectively reduce the isolation of the largest habitat blocks in reserves, therefore largely contributing to species persistence across wide spatial and temporal scales.