Diel vertical migration (DVM) by zooplankton to deep, poorly illuminated habitats during the day is thought to reduce the probability of attack by visual predators at the cost of decreased net energy gain. Herein, I develop a trade-off model used to predict the timing of vertical migration of zooplankton. The model is based on ''Gilliam's rule''-the notion that animals will select habitats that minimize the ratio of mortality risk/energy gain-and incorporates light-and size-dependent vulnerability to visual predators. The model predicts that smaller, less vulnerable prey should ascend into food-rich surface waters earlier and descend later than larger, more conspicuous organisms over a wide range of conditions. The model has been parameterized for a population of Euphausia pacifica in Saanich Inlet, British Columbia, and predicts that 7-mm (larval) animals will ascend/descend ϳ35 min earlier/later than more conspicuous 24-mm adults. These predictions are consistent with acoustic observations of the timing of DVM in this population. The proposed mechanism for size-dependent timing of DVM is based on the potential for increased energy gain in surface waters and diel changes in vulnerability to size-selective predators. These conditions prevail in both freshwater and marine pelagic environments, and size-dependent DVM is likely to be widespread in zooplankton.Diel vertical migration (DVM), in which pelagic organisms migrate away from food-rich surface waters during the day, is a common behavior that occurs in a wide range of taxa and environments. The adaptive value of DVM is thought to be a trade-off concerning predator avoidance (Ohman 1990;Lampert 1993), because organisms forgo feeding opportunities in the food-rich surface waters to avoid spatiotemporal overlap with visual predators under high light intensities during the day. Visual predators (planktivorous fish) are highly selective and feed preferentially on large-bodied and pigmented zooplankton prey, which are more optically conspicuous (Brooks and Dodson 1965;Batty et al. 1990;Brodeur 1998). Although vertical migration behavior in zooplankton has been the subject of substantial study in the past, the focus of previous work has been on the amplitude and 1 Present address: Northwest Fisheries Science Center, Hatfield Marine Science Center, Newport, Oregon 97365 (Alex.DeRobertis@ noaa.gov).
AcknowledgmentsI am indebted to M. D. Ohman for guidance and criticism in all stages of this work. The comments of D.