Process-based approach reveals directional effects of environmental factors on movement between habitats.

1. Understanding the effects of environmental factors on animal distributions is a central issue in ecology. However, movement rules inferred from distribution patterns do not reveal the processes through which animal distribution is realized. 2. We investigated individual movement rules using a process-based approach. In experiments, coastal fish larvae (red drum, Sciaenops ocellatus) were matched with an intraspecific competitor of different sizes, and time series of habitat transition of individuals were fitted with a continuous-time Markov chain model to evaluate the effects of the presence of a competitor, behavioural interactions and habitat quality on the likelihoods of habitat transition. 3. The process-based approach revealed that these factors did not simply act as a 'slope' between habitats that makes it easier to go in one direction and more difficult to return. Rather, these factors modify the movement rules differently depending on the directions of the movement. 4. Individuals were less likely to enter a better habitat in the presence of a larger conspecific, more likely to shift to a poorer habitat when they received aggressive behaviour and more likely to stay in a better habitat in the presence of food. However, no effect was found on the transition intensity for moving in the opposite direction. 5. The process-based approach to evaluating movement rules of animals allowed us to see the contrasting directional effects of different factors on the underlying movement rules used by animals, as opposed to pattern-based fitting of observed distributions. Consideration of these rules would improve the existing habitat-choice models.