Dispersal across habitat boundaries: uncovering the demographic fates of populations in unsuitable habitat

Patchy landscapes are characterized by abrupt transitions among distinct habitat types, forcing species to cross habitat boundaries in order to spread. Since seed dispersal is a probabilistic process, with a kernel that decays with distance, most individuals will fail to reach new, suitable habitat. Although failed dispersers are presumed dead in population models, their demographic fates may not be so simple. If transient survival is possible within unsuitable habitat, then through time, individuals may be able to reach distant, suitable habitat, forming new populations and buffering species from extinction. In a fragmented Californian grassland, we explored the fates of individuals that crossed habitat boundaries, and if those fates differed among specialists dispersing from two habitat types: serpentine habitat patches and the invaded non-serpentine matrix. We surveyed the diversity of seedbank and adult life stages along transects that crossed boundaries between patches and the matrix. First, we considered how patch specialists might transiently survive in the matrix via seed dormancy or stepping-stone populations. Second, we investigated the dispersal of an invasive matrix specialist (Avena fatua) into patches, to assess if sink populations existed across the habitat boundary. We found that dormancy maintained populations of patch specialists deep into the matrix, as abundances of seedbanks and of adult plant communities differed with distance into the matrix. We found evidence that these dormant seeds disperse secondarily with vectors of material flows in the landscape, suggesting that they could eventually reach suitable patches even if they first land in the matrix. We found that A. fatua were largely absent deep in patches, where reproductive outputs plummeted and there was no evidence of a dormant seedbank. Our results not only reveal the demographic fates of individuals that land in unsuitable habitat, but that their ecological consequences differ depending on the direction by which the boundary is crossed (patch [->] matrix = matrix [->] patch). Dormancy is often understood as a mechanism for persisting in face of temporal variability, but it may serve as a means of traversing unsuitable habitat in patchy systems, warranting its consideration in estimates of habitat connectivity.