Spatial occurrence and colonisations in patch-tracking metapopulations:: local conditions versus dispersal

We studied the relative importance of local variables and dispersal for the occurrence and colonisation of the epiphytic bryophytes Orthotricum speciosum (spore dispersed), and O. obtusifolium (spore and asexual gemmae) on aspen trees ('patches') in two forest landscapes (one old-growth and one fragmented) using multiple logistic regression. The relative importance of dispersal was quantified as the reduction of residual deviance for a connectivity variable. In modelling dispersal, we assumed that trees with low local abundance were recent colonisations, and that trees with high local abundance were diaspore sources for colonisation. The occurrence of O. speciosum in the fragmented landscape was most affected by shading, but also by connectivity, aspen diameter and vitality. In the old-growth landscape, connectivity was the single most important variable for recent colonisations, but its effect was lower than the sum of the effects of all local environmental variables. The occurrence of O. obtusifolium in the fragmented landscape was related to similar variables but the relative importance of these variables was different, and connectivity did not affect the probability of a recent colonisation in this species. We describe the epiphyte-tree system in the patch-tracking metapopulation model. In this model colonisations are distance dependent, but in contrast to the classical metapopulation model local extinctions are caused by deterministic patch destruction - once the epiphyte has colonised the tree it remains until the tree dies.