Foraging strategies of dogwhelks, Nucella lapillus (L.): interacting effects of age, diet and chemical cues to the threat of predation.

The effects of diet history, hunger and predation risk on short-term behavioral decisions of dogwhelks were tested in a specially designed test apparatus, termed a linear feeding array (LFA). The LFA consists of a sequential series of prey items mounted in a flume with unidirectional current directed towards a test (predatory) animal, and into which potential olfactory cues regarding predation risk are introduced. For dogwhelks the array was constructed vertically to accomodate intertidal foraging movements and is termed a vertical linear array (VLA). The behaviors exhibited by the dogwhelks were interpreted from distribution patterns in the VLA. Recent experimental studies and advances in optimal foraging theory provided the basis for the hypotheses tested in the VLA, which included: foraging and other behaviors are affected by predation, animals will avoid risk in the presence of predation threat, responses to predation threat will be proportional to the number and kinds of predator cues present, and starved animals will take greater risks than fed animals. We also test the proposition that foraging decisions are further modified by age. Three groups of juvenile and adult animals were maintained on diets of barnacles, mussels or no food (starved). The scent of crabs and damaged conspecifics served as olfactory cues to predation risk. Dogwhelks exhibited a range of behaviors in the VLA including: sheltering, searching, feeding, and aerial climbing. Distribution of animals in the tank assumed a relatively stable pattern after 2-3 h. These patterns were interpreted as the consequence of heirarchial decision making including: (i) a decision to become active, leaving the resting place or water refuge adopted during initial placement, followed by (ii) a decision to move vertically upwards or downwards, and (iii) a decision to attack prey when encountered. Analysis of movement patterns revealed that the initial decision, analogous to leaving a crevice as the tide comes in, was influenced in adults by predator cues and in juveniles by both predator cues and diet history. Perceived risk, as crab and damaged-conspecific odors, made individuals more likely to remain inactive, a risk-avoiding strategy for animals already in a refuge. Starved animals were more likely to descend into the tank and attack prey than fed animals. Our results support the hypotheses that higher-order predators affect the foraging decisions of dogwhelks and that juveniles and satiated animals are more sensitive to predation risk than starved ones. Together, these and earlier studies suggest that dogwhelks assess their environment before foraging, and that they are attuned to reducing the risks of mortality.