The importance of including habitat-specific behaviour in models of butterfly movement

Dispersal is a key process affecting population persistence and major factors affecting dispersal rates are the amounts, connectedness and properties of habitats in landscapes. We present new data on the butterfly Maniola jurtina in flower-rich and flower-poor habitats that demonstrates how movement and behaviour differ between sexes and habitat types, and how this effects consequent dispersal rates. Females had higher flight speeds than males, but their total time in flight was four times less. The effect of habitat type was strong for both sexes, flight speeds were similar to 2.5 x and similar to 1.7 x faster on resource-poor habitats for males and females, respectively, and flights were approximately 50% longer. With few exceptions females oviposited in the mown grass habitat, likely because growing grass offers better food for emerging caterpillars, but they foraged in the resource-rich habitat. It seems that females faced a trade-off between ovipositing without foraging in the mown grass or foraging without ovipositing where flowers were abundant. We show that taking account of habitat-dependent differences in activity, here categorised as flight or non-flight, is crucial to obtaining good fits of an individual-based model to observed movement. An important implication of this finding is that incorporating habitat-specific activity budgets is likely necessary for predicting longer-term dispersal in heterogeneous habitats, as habitat-specific behaviour substantially influences the mean (> 30% difference) and kurtosis (1.4 x difference) of dispersal kernels. The presented IBMs provide a simple method to explicitly incorporate known activity and movement rates when predicting dispersal in changing and heterogeneous landscapes.