Nonconsumptive effects (NCEs) have been shown to occur in numerous systems and are regarded as important mechanisms by which predation structures natural communities. Sensory ecology-that is, the processes governing the production, propagation, and masking of cues by ambient noise-provides insights into the strength of NCEs as functions of the environment and modes of information transfer. We discuss how properties of predators are used by prey to encode threat, how the environment affects cue propagation, and the role of single sensory processes versus multimodal sensory processes. We discuss why the present body of literature documents the potential for strong NCEs but does not allow us to easily determine how this potential is expressed in nature or what factors or environments produce strong versus weak NCEs. Many of these difficulties stem from a body of literature in which certain sensory environments and modalities may be disproportionately represented and in which experimental methodologies are designed to show the existence of NCEs. We present a general framework for examining NCEs to identify the factors controlling the number of prey that respond to predator cues and discuss how the properties of predators, prey, and the environment may determine prey perceptive range and the duration and frequency of cue production. We suggest how understanding these relationships provides a schema for determining where, when, why, and how NCEs are important in producing direct and cascading effects in natural communities.

The lethal and nonlethal impacts of predators in marine systems are often mediated via reciprocal detection of waterborne chemical signals between consumers and prey. Local flow environments can enhance or impair the chemoreception ability of consumers, but the effect of hydrodynamics on detection of predation risk by prey has not been investigated. Using clams as our model organism, we investigated two specific questions: (1) Can clams decrease their mortality by responding to predators? (2) Do fluid forces affect the ability of clams to detect approaching predators? Previous research has documented a decrease in clam feeding (pumping) in response to a neighboring predator. We determined the benefits of this behavior to survivorship by placing clams in the field with knobbed whelk or blue crab predators caged nearby and compared mortality between these clams and clams near a cage-only control. Significantly more clams survived in areas containing a caged predator, suggesting that predator-induced alterations in feeding reduce clam mortality in the field. We ascertained the effect of fluid forces on clam perception of predators in a laboratory flume by comparing the feeding (pumping) behavior of clams in response to crabs and whelks in flows of 3 and 11 cm/s. Clams pumped significantly less in the presence of predators, but their reaction to blue crabs diminished in the higher velocity flow, while their response to whelks remained constant in both flows. Thus, clam reactive distance to blue crabs was affected by fluid forces, but hydrodynamic effects on clam perceptive distance was predator specific. After predators were removed, clams exposed to whelks took significantly longer to resume feeding than those exposed to blue crabs. Our results suggest that prey perception of predators can be altered by physical forces. Prey detection of predators is the underlying mechanism for trait-mediated indirect interactions (TMIIs), and recent research has documented the importance of TMIIs to community structure. Since physical forces can influence prey perception, the prevalence of TMIIs in communities may, in part, be related to the sensory ability of prey, physical forces in the environment that impact sensory performance, and the type of predator detected.

Abstract-Hard clams, Mercenaria mercenaria, are sessile, filter-feeding organisms that are heavily preyed upon by blue crabs, which find their clam prey using chemical cues. Clams may evade blue crabs by reducing their pumping (feeding) behavior when a threat is perceived. The purpose of this study was to determine the type of signals that clams use to detect consumers. Clams decreased their pumping time in response to blue crabs and blue crab effluent, but not to crab shells, indicating that chemical signals and not mechanical cues mediated the response of clams to distant predators. Because predator diet can influence prey evaluation of predatory threats, we compared clam responses to blue crabs fed a steady diet of fish, clams, or that were starved prior to the experiment. In addition, we used injured clams as a stimulus because many organisms detect predators by sensing the odor of injured con-or heterospecifics. Clams reduced feeding in response to injured conspecifics and to blue crabs that had recently fed. Clams reacted similarly to fed crabs, regardless of their diet, but did not respond to starved blue crabs. Because blue crabs are generalist predators and the threat posed by these consumers is unrelated to the crab's diet, we should expect clam reactions to blue crabs to be independent of the crab's diet. The failure of clams to react to starved blue crabs likely increases their vulnerability to these consumers, but clam responses to injured conspecifics may constitute a strategy that allows animals to detect an imminent threat when signals emanating from blue crabs are not detectable.

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