When is general wariness favored in avoiding multiple predator types?

Free access to article and electronic appendices via DOI.Adaptive responses to predation are generally studied assuming only one predator type exists, but most prey species are depredated by multiple types. When multiple types occur, the optimal antipredator response level may be determined solely by the probability of attack by the relevant predator: "specific responsiveness." Conversely, an increase in the probability of attack by one predator type might increase responsiveness to an alternative predator type: "general wariness." We formulate a mathematical model in which a prey animal perceives a cue providing information on the probability of two predator types being present. It can perform one of two evasive behaviors that vary in their suitability as a response to the "wrong" predator type. We show that general wariness is optimal when incorrect behavioral decisions have differential fitness costs. Counterintuitively, difficulty in discriminating between predator types does not favor general wariness. We predict that where responses to predator types are mutually exclusive (e.g., referential alarm-calling), specific responsiveness will occur; we suggest that prey generalize their defensive responses based on cue similarity due to an assumption of response utility; and we predict, with relevance to conservation, that habituation to human disturbance should generalize only to predators that elicit the same antipredator response as humans

Proximate factors underpinning receiver responses to deceptive false alarm calls in wild tufted capuchin monkeys: is it counterdeception?

Previous research demonstrates that tufted capuchin monkeys use terrestrial predator alarm calls in a functionally deceptive manner to distract conspecifics when feeding on contestable resources, although the success of this tactic is limited because listeners frequently ignore these calls when given in such situations. While this decreased response rate is suggestive of a counterstrategy to deception by receivers, the proximate factors underpinning the behavior are unclear. The current study aims to test if the decreased response rate to alarm calls in competitive contexts is better explained by the perception of subtle acoustic differences between predator-elicited and deceptive false alarms, or by receivers varying their responses based on the context in which the signal is received. This was tested by first examining the acoustic structure of predator-elicited and deceptive false alarms for any potentially perceptible acoustic differences, and second by comparing the responses of capuchins to playbacks of each of predator-elicited and false alarms, played back in noncompetitive contexts. The results indicate that deceptive false alarms and predator-elicited alarms show, at best, minimal acoustic differences based on the structural features measured. Likewise, playbacks of deceptive false alarms elicited antipredator reactions at the same rate as did predator-elicited alarms, although there was a nonsignificant tendency for false alarms to be more likely to elicit escape reactions. The lack of robust acoustic differences together with the high response rate to false alarms in noncompetitive contexts suggests that the context in which the signal is received best explains receiver responses. It remains unclear, however, if listeners ascribe different meanings to the calls based on context, or if they generally ignore all signals in competitive contexts. Whether or not the decreased response rate of receivers directly stems from the deceptive use of the calls cannot be determined until these latter possibilities are rigorously tested

Cognitive dimensions of predator responses to imperfect mimicry?

Many palatable insects, for example hoverflies, deter predators by mimicking well-defended insects such as wasps. However, for human observers, these flies often seem to be little better than caricatures of wasps – their visual appearance and behaviour are easily distinguishable. This imperfect mimicry baffles evolutionary biologists, because one might expect natural selection to do a more thorough job. Here we discuss two types of cognitive processes that might explain why mimics distinguishable mimics might enjoy increased protection from predation. Speed accuracy tradeoffs in predator decision making might give imperfect mimics sufficient time to escape, and predators under time constraint might avoid time-consuming discriminations between well-defended models and inaccurate edible mimics, and instead adopt a “safety first” policy of avoiding insects with similar appearance. Categorization of prey types by predators could mean that wholly dissimilar mimics may be protected, provided they share some common property with noxious prey

Fear of predation alters clone-specific performance in phloem-feeding prey

Fear of predation has been shown to affect prey fitness and behaviour, however, to date little is known about the underlying genetics of responses to predator-associated risk. In an effort to fill this gap we exposed four naïve clones of green peach aphid (Myzus persicae), maintained on the model crop Brassica oleracea, to different types of cues from aphid lion (Chrysoperla carnea). The respective predation risks, we termed Fear Factors, were either lethal (consumption by predator), or non-lethal (non-consumptive predator- associated cues: plant-tethered predator cadavers and homogenised shoot- sprayed or soil-infused blends of predator remains). Our results show that the non-lethal risk cues differentially impeded prey reproductive success that varied by clone, suggesting genotype-specific response to fear of predation. Furthermore, whether plants were perceived as being safe or risky influenced prey responses as avoidance behaviour in prey depended on clone type. Our findings highlight that intra-specific genetic variation underlies prey responses to consumptive and non-consumptive effects of predation. This allows selection to act on anti-predator responses to fear of predation that may ramify and influence higher trophic levels in model agroecosystems

Analysis of reinforcement learning strategies for predation in a mimic-model prey environment

In this paper we propose a mathematical learning model for a stochastic automaton simulating the behaviour of a predator operating in a random environment occupied by two types of prey: palatable mimics and unpalatable models. Specifically, a well known linear reinforcement learning algorithm is used to update the probabilities of the two actions, eat prey or ignore prey, at every random encounter. Each action elicits a probabilistic response from the environment that can be either favorable or unfavourable. We analyse both fixed and varying stochastic responses for the system. The basic approach of mimicry is defined and a short review of relevant previous approaches in the literature is given. Finally, the conditions for continuous predator performance improvement are explicitly formulated and precise definitions of predatory efficiency and mimicry efficiency are also provided

Indirect effects of primary prey population dynamics on alternative prey

We develop a theory of generalist predation showing how alternative prey species are affected by changes in both mean abundance and variability (coefficient of variation) of their predator's primary prey. The theory is motivated by the indirect effects of cyclic rodent populations on ground-breeding birds, and developed through progressive analytic simplifications of an empirically-based model. It applies nonetheless to many other systems where primary prey have fast life-histories and can become locally superabundant, which facilitates impact on alternative prey species. In contrast to classic apparent competition theory based on symmetric interactions, our results suggest that predator effects on alternative prey should generally decrease with mean primary prey abundance, and increase with primary prey variability (low to high CV) - unless predators have strong aggregative responses, in which case these results can be reversed. Approximations of models including predator dynamics (general numerical response with possible delays) confirm these results but further suggest that negative temporal correlation between predator and primary prey is harmful to alternative prey. We find in general that predator numerical responses are crucial to predict the response of ecosystems to changes in key prey species exhibiting outbreaks, and extend the apparent competition/mutualism theory to asymmetric interactions

THE ART OF WAR: PATTERNS AND MECHANISMS UNDERLYING PREDATOR-INDUCED PLASTICITY OF AMPHIBIANS

Organisms often employ phenotypic plasticity as a strategy to cope with variable environments. This is particularly true of predation threats, wherein prey induce defenses to reduce detection or capture by predators. In order to produce appropriate defenses, prey must be able to discern useful information from environmental cues. Despite the pervasive production of inducible defenses, we understand very little of how much useful information is conveyed to organisms in cues, or how the subsequent plastic responses vary within groups of organisms. To address the need for comparative studies of phenotypic plasticity, we sought to examine morphological and behavioral defenses of five species of Ambystoma salamander larvae in response to larval dragonfly (Anax junius) chemical cues in a common garden environment. Dragonfly cues induced relatively few morphological changes across species. Likewise, salamanders did not vary in their refuge use during the experiment, though several species reduced their activity in the presence of predators early in development. Our results suggest that behavioral and morphological defenses in Ambystoma are highly variable among species and the genus appears to be less plastic than tadpoles and other salamander species. To understand what types of information prey are capable of responding to in their environment, we raised grey treefrog tadpoles (Hyla verisciolor) in the presence of cues isolated from different stages of an attack sequence by larval dragonflies (A. junius) or larval dragonflies THE ART OF WAR: PATTERNS AND MECHANISMS UNDERLYING PREDATOR-INDUCED PLASTICITY OF AMPHIBIANS Heather Michelle Shaffery, M.S. University of Pittsburgh, 2013 iv consuming different combinations of grey treefrog tadpoles and snails (Helisoma trivolvis) across different temporal sequences. When exposed to a predator consuming grey treefrogs, tadpoles reduced their activity, increased their hiding behavior, and induced deeper tails. As we exposed prey to more types of cues from an attack sequence, they also increased tail depth and hiding behavior but did not change their activity. Additionally, treefrog tadpoles generally increased their defense as the biomass of treefrogs consumed increased, regardless of whether heterospecifics were being consumed. Our results suggest that treefrogs can gain cue information from all portions of an attack sequence, and that both temporal patterns of feeding and diet content of predators influence the type and magnitude of induced prey defenses