The authors wish to thank Peter Rhines, David Dusenbery, Daniel Grubaum, and the participants in Friday Harbor Lab’s 1999 Marine Chemosensory Ecology Course for productive discussions; Fritz Star, Christian Sarason Parker, and Christopher Krembs for critical reviews of earlier drafts; and Eric Lin-dahl, Michael O’Donnell, and Rex Johnson for help with the model implementation. This work was conducted with support from the Office of Naval Research and Friday Harbor Labs, University of Washington.