Nonrandom movement behavior at habitat boundaries in two butterfly species: implications for dispersal

We observed meadow brown (Maniola jurtina) and gatekeeper (Pyronia tithonus) butterflies at habitat boundaries and observed spontaneous movements out of suitable habitat in order to investigate such movements in relation to dispersal. We found that butterflies of both species were aware of the position of a highly permeable habitat boundary without needing to cross it. Nevertheless, a considerable proportion of butterflies close to the boundary left their habitat (25-43%). Butterflies that crossed the boundary, and moved substantial distances into unsuitable habitat (up to 350 m in M. jurtina and 70 m in P. tithonus), usually returned to their original habitat patch (98-100%). Movement trajectories, at least in M. jurtina, were significantly different from, and more directed and systematic than, a correlated random walk. Approximately 70-80% of spontaneous movements into unsuitable habitat in both species were "foray" loops comparable to those described in mammals and birds. We conclude that, since migrants seemed to have considerable control over leaving their patch and over their subsequent movement trajectories, chance encounter rates with habitat boundaries, and indeed habitat leaving rates, might be less crucial in determining dispersal rates than is usually assumed. In addition, random dispersal trajectories should not be taken for granted in population or evolution models.