Disentangling fine- and large-scale colonization processes in metapopulation dynamics: a case study on a threatened epiphytic bryophyte

Understanding the colonization process of species living in a dynamic fragmented habitat is essential to assess their persistence. In the metapopulation theory, the colonization of a species can be quantified using the turnover of occupancy in habitat patches. However, this approach is often limited by the feasible size of surveyed areas. Because many species are capable of long-distance dispersal, such areas often constitute open systems undergoing colonization of propagules coming from outside, the 'background deposition'. We focus on disentangling background deposition from local colonization among surveyed patches when analyzing turnover. We consider two spatial scales: 1) focal areas where all patches are monitored over time; 2) a larger extent, encompassing the focal areas, over which the distribution of the target species is quantified with a coarse spatial grain. Our key idea is to use the regional connectivity of focal areas within the larger scale as a covariate when analyzing colonization events within focal areas. A positive effect of regional connectivity on the colonization probability of patches may indicate background deposition. We applied this approach to the epiphytic bryophyte Dicranum viride in a managed temperate deciduous forest, considering phorophyte trees as patches, forest stands as focal areas and the whole forest as the larger scale. We combined a fine-grained turnover survey of occupied trees within three forest stands (similar to 3 ha) with a coarse-grained snapshot of D. viride distribution over the forest (similar to 15 000 ha). Regional connectivity came out as the most significant factor, with a strong positive effect on colonization probability within stands. However, it was attributed to sources in the immediate vicinity of focal stands, suggesting a short-ranged colonization process occurring across stands' borders rather than long-distance background deposition. Our results thus call for maintaining a stepping-stone of habitat across the forest through time to improve D. viride persistence.