Hunger rapidly overrides the risk of predation in the subtidal scavenger Nassarius siquijorensis (Gastropoda: Nassariidae): an energy budget and a comparison with the intertidal Nassarius festivus in Hong Kong

This study shows that, as with its intertidal counterpart, Nassarius festivus, the rate at which subtidal Nassarius siquijorensis moves towards food bait is similar for starved and well-fed individuals. This study also investigates another facet of nassariid nutrition related to the degree of hunger, i.e. the effect of simulated predation upon a feeding assemblage. Individuals which fed within 7 days, cease feeding and depart palatable food if crushed conspecifics are added. Between 7 and 13 days since its last meal, however, N. siquijorensis will feed when food is available, despite the possibility of predation. For the intertidal N. festivus, the critical time for hunger to override the risk of predation is between 14 and 21 days. The difference between subtidal and intertidal species may be due to a difference, in terms of days, that a meal can provide for their energy expenditure, particularly with regard to respiration. The bigger, subtidal, N. siquijorensis needs to feed more frequently than the smaller, intertidal, N. festivus.     

THE PHYSIOLOGY AND FEEDING BEHAVIOUR OF TWO MARINE SCAVENGING GASTROPODS IN HONG KONG: THE SUBTIDAL BABYLONIA LUTOSA (LAMARCK) AND THE INTERTIDAL NASSARIUS FESTIVUS (POWYS)

Aspects of the physiology and feeding behaviour of the scavengingsnails, Babylonia lutosa and Nassarius festivus have been studiedand compared. N. festivus has a wider physiological tolerancethan B. lutosa; upper lethal temperatures being 31.5°C and28.5°C respectively and lower lethal salinities 15.5 and11.5. N. festivus was also more resistant to desiccation. N.festivus is adapted to an intertidal environment that is morevariable than the subtidal habitat of B. lutosa. The metabolicrate of adult N. festivus was two to three times higher thanthat of adult B. lutosa, the specific oxygen consumption rateof the former being 7.91±0.49x10^–6 mol of O₂.g^–1wet wt.hr^–1. This correlates with estimates of consumptionobtained for the two animals, B. lutosa eating, on average,6.57% of its wet tissue weight per day, N. festivus 15.33%.Standardization of the data for 2 g animals of both species,however, reveals no significant difference in terms of oxygenconsumption and thus the enhanced consumption by N. festivussimply reflects its smaller size. Both species are opportunisticfeeders and can survive long periods of starvation (>100days). The feeding strategies of the two are also different,N. festivus eating faster than B. lutosa, possibly because ofthe more ephemeral nature of carrion in the intertidal. Starvationaffected the feeding behaviour of N. festivus more than B. lutosaand N. festivus was able to perceive carrion from greater relativedistances than B. lutosa. (Received 27 February 1989; accepted 24 July 1989)     

Overriding the oddity effect in mixed-species aggregations: group choice by armored and nonarmored prey

Because "odd" individuals often suffer disproportionately highrates of predation, solitary individuals should join groupswhose members are most similar to themselves in appearance.We examined group-choice decisions by individuals in armoredand nonarmored species and predicted that either (1) the oddityeffect would result in preference for conspecific groups forsolitary individuals of both species, or (2) individuals inthe armored species would prefer to associate with groups containingindividuals of the more vulnerable species. Armored brook sticklebacks(Culaea inconstans) and nonarmored fathead minnows (Pimephalespromelas) have the same predators and often occur together instreams. In mixed-species shoals, yellow perch (Perca flavescens)attacked minnows earlier and more often than sticklebacks. Wetested whether solitary minnows and sticklebacks preferred toassociate with conspecific or heterospecific shoals under conditionsof both low and high predation risk. When predation risk washigh, minnows preferred to associate with conspecifics overheterospecifics, as predicted by the oddity effect. In contrast,sticklebacks preferentially associated with groups of minnowsover groups of conspecifics when predation risk was high. Whenpredation risk was low, solitary individuals of both speciespreferentially associated with conspecific over heterospecificshoals. Stickleback shoal choices under low-risk conditionsmay have been influenced by interspecific competition for food.In feeding experiments, minnows were more efficient foragersthan sticklebacks, so it should benefit sticklebacks to avoidminnows unless predation risk is high. Therefore, for armoredprey, the benefits of associating with more vulnerable preyappear to override the costs of both the oddity effect and foodcompetition when predation risk is high.     

Is It Worth the Risk? Food Deprivation Effects on Tadpole Anti-Predatory Responses

Predation risk affects foraging behavior and, particularly, the amount of time devoted to the search for food. When exposed to predation risk, food deprived animals should be risk prone and relax behavioral defenses to a wider extent than well fed individuals. To test for this prediction, we used a 2?×?2 factorial design experiment, manipulating both the energetic state (fed vs. fasted) and exposure to an attack-released cue (injured vs. uninjured conspecifics) of common water frog tadpoles (Pelophylax kl. esculentus). Contrary to expectations, food deprivation significantly lowered the activity level of predator-exposed tadpoles. As in this experiment no food resource was added to test containers, energy conserving behavior might have both delayed starvation and lowered the probability of encountering the potential predator. To test for the effect of food availability on behavioral responses, we performed a second experiment, using the same protocol and procedures, except for adding food to all test containers. All tadpoles showed similar levels of activity, while fed tadpoles exposed to alarm cues tended to swim farther from the cage containing the stimulus than in the first experiment. As many anuran larvae can feed on dead conspecifics, prey-borne cues may have been interpreted as the potential presence of both a food source and a predator, fed tadpoles possibly being more confident than fasted tadpoles in their ability to escape predation in case an actual attack occurs.     

Predation Risk Perception,Food Density and Conspecific Cues Shape Foraging Decisions in a Tropical Lizard

When foraging, animals can maximize their fitness if they are able to tailor their foraging decisions to current environmental conditions. When making foraging decisions, individuals need to assess the benefits of foraging while accounting for the potential risks of being captured by a predator. However, whether and how different factors interact to shape these decisions is not yet well understood, especially in individual foragers. Here we present a standardized set of manipulative field experiments in the form of foraging assays in the tropical lizard Anolis cristatellus in Puerto Rico. We presented male lizards with foraging opportunities to test how the presence of conspecifics, predation-risk perception, the abundance of food, and interactions among these factors determines the outcome of foraging decisions. In Experiment 1, anoles foraged faster when food was scarce and other conspecifics were present near the feeding tray, while they took longer to feed when food was abundant and when no conspecifics were present. These results suggest that foraging decisions in anoles are the result of a complex process in which individuals assess predation risk by using information from conspecific individuals while taking into account food abundance. In Experiment 2, a simulated increase in predation risk (i.e., distance to the feeding tray) confirmed the relevance of risk perception by showing that the use of available perches is strongly correlated with the latency to feed. We found Puerto Rican crested anoles integrate instantaneous ecological information about food abundance, conspecific activity and predation risk, and adjust their foraging behavior accordingly.     

Modelling the effect of feeding-related mortality on the feeding strategy of tsetse (Diptera: Glossinidae)

Abstract. Free-living haematophagous insects risk death through host grooming responses or through increased susceptibility to predation whenever they take a bloodmeal. In this paper we investigate the effects of these risks on the feeding strategy of tsetse. A model is presented that allows for death of tsetse by starvation if they do not succeed in feeding within a fixed time (set at 6 days in the first instance) and for mortality specifically associated with feeding. In addition there is background mortality that applies to all flies at all times.
The model is used to compute the individual life-time fertility (number of female puparia per female) as a function of the probability of obtaining a meal (indicated by field data to be very high, usually > 0.85 per day) and the day on which flies start to search for a meal. We suggest that the feeding strategy that would be selected for is that which allows the maximum reproductive output. The model shows that this strategy involves making no attempts to feed for 3–4 days after the previous meal and then attempting to feed with the greatest possible probability until a meal is obtained. The predicted feeding interval, obtained independently of any trapping data, agrees closely with all previous estimates from field studies using a variety of methods. Preliminary results from a laboratory experiment reveal an increased risk of predation of recently fed as compared with hungry tsetse. The lower the actual feeding mortality the more frequently will flies be able to feed should conditions so demand. It is adaptive, however, for tsetse to delay attempting to feed for as long as they can, which is made possible by the near certainty of locating and feeding on a host within 1 day, using their sophisticated sensory systems.     

FEEDING A VOIDANCE OF TERRESTRIAL GASTROPODS TO CONSPECIFIC AND NONSPECIFIC MATERIAL

In laboratory tests the terrestrial gastropods Arion fasciatus,Deroceras reticulatum, Arianta arbustorum and Bradybaena fruticumpreferred control leaf discs over discs treated with an aqueoussuspension of crushed conspecifics. The response to suspensionsof different species was weaker. The strength of the responseincreased with the concentration of the suspension. The suspension'seffect in deterring feeding was inactivated within 18 to 32hours. Hunger reduced normal responsiveness to conspecific fluids;the quality of the control food also affected the response.The results also imply that ranking among individuals is labile.Spraying lettucc with the suspension affected the dispersalof the animals only in early summer. The article discusses thebiological significance of the response and the use of substancesobtained from slugs and snails in the control of pest gastropods. (Received 21 March 1991; accepted 5 September 1991)     

Effects of pre- and post-prandial starvation on meal size and evacuation rate of juvenile Atlantic salmon, Salmo salar L.

After a pre-prandial period of starvation or feeding with unlabelled food, 0+ salmon parr (0.8–11.7 g) were fed a test meal of iron particle labelled food and subsequently were again either starved or fed unlabelled food. The quantity of labelled food consumed and the evacuation rate was determined by serial radiographs. In fish of all sizes, pre-prandial starvation causes a larger test meal (as a percentage of body weight) to be consumed when compared to pre-prandially fed fish. In addition, pre-prandial starvation results in relatively larger meals as a percentage of body weight being taken by smaller compared to larger fish. This result was not evident for pre-prandially fed fish. Evacuation rate was unrelated to body size irrespective of feeding history. Post-prandial starvation decreased evacuation rate but this effect was inversely related to the quantity of food consumed. Larger meals were not evacuated differently from smaller meals if feeding occurred post-prandially, irrespective of pre-prandial starvation.     

捕食风险对不同饥饿状态下艾虎取食行为的影响

在室内条件下,将大鵟作为艾虎的天敌动物,通过双通道选择实验确定6 只成体艾虎在3 个捕食风险水平和4 种饥饿状态条件下的取食行为,探讨艾虎在取食过程中对饥饿风险与捕食风险的权衡策略。研究结果表明:在无捕食风险存在时,艾虎被剥夺食物0 d 和1 d 后对食物量不同的两个斑块中的取食量和利用频次均无明显不同(P > 0. 05),但对高食物量斑块的利用时间均明显高于低食物量斑块的(P <0.05),而艾虎被剥夺食物2 d和3 d后对高食物量斑块中的取食量和利用时间均明显高于低食物量斑块中的(P < 0.05),但在利用频次上均无明显差异(P > 0.05)。在面临低风险时,艾虎在4 种饥饿状态下均只利用无天敌动物存在的低食物量斑块,而基本不利用有天敌动物存在的高食物量斑块。在面临高风险时,艾虎不得不利用有天敌动物存在的食物斑块,被剥夺食物0 d 时艾虎对无风险、无食物量斑块的利用时间基本相同于对高风险、有食物量斑块的利用时间(P>0.05),而被剥夺食物1d、2 d 和3 d 后艾虎对高风险、有食物量斑块的利用时间明显高于无风险、无食物量斑块的(P< 0. 05)。在相同风险条件下,随着饥饿程度增加,艾虎在斑块中的取食量均明显增加(P< 0.05),而对斑块的利用时间和利用频次明显降低(P<0.05)。在相同的饥饿状态下,不同风险水平时,艾虎在斑块中的取食量无明显的差异(P>0.05),但在低风险和高风险时对斑块的利用时间和频次均明显低于无风险时的(P <0.05)。以上结果说明艾虎能够根据食物摄取率和自身的能量需求在捕食风险和饥饿风险之间做出权衡,当饥饿风险小于捕食风险时,艾虎趋于躲避捕食风险,当饥饿风险大于捕食风险时,艾虎趋于面对捕食风险,所采用的取食策略是减少活动时间和能量消耗,最大程度地提高单位时间内获得的能量。     

Ecological Risk Aversion and Foraging Behaviors of Juvenile Squirrel Monkeys (Saimiri sciureus)

The ‘ecological risk aversion hypothesis’ [C.H. Janson and C.P. van Schaik, Juvenile Primates, Oxford Univ. Press, New York (1993), pp. 57–74] proposes that the pattern of slow growth characteristic of juvenile primates is a response to ecological risks (predation and starvation) experienced by juveniles. Juveniles are thought to avoid predation risk by positioning themselves near conspecifics, therefore experiencing high levels of feeding competition with older individuals, reduced access to resources and, consequently, high starvation risks during periods of food scarcity. The present study compared the foraging behaviors of juvenile and adult squirrel monkeys, a small neotropical primate characterized by a long juvenile period, to determine how predation and starvation risks affected juvenile behaviors. The study was conducted in Eastern Amazonia, in a seasonal environment. Due to their slow development, small body size and large group sizes, it was expected that juveniles in this species would behave in a manner consistent with the risk aversion hypothesis. However, age differences in foraging efficiency and foraging success were smaller than predicted. There was also no evidence that juveniles sacrificed access to food for predator protection. Adults did not have preferential access to fruit patches and direct competition was rare. Feeding competition for prey, the most common resource in the troop's diet, was negligible. Therefore, the slow growth and long juvenile period of squirrel monkeys do not correspond with evidence of predation or starvation risk, as predicted by the risk aversion hypothesis.     

The decision to feed by a scavenger in relation to the risks of predation and starvation

Recent theoretical models have considered how nutritional status and predation risk affect the decision by an individual to forage. We consider the feeding behaviour of a scavenger under risk of predation. The intertidal gastropod Nassarius dorsatus Röding was used to test the following hypotheses: (1) a damaged conspecific should be a less preferred meal than a damaged predator or another species; (2) a scavenger should be able to discriminate between a damaged conspecific in the presence and one in the absence of its predator, preferring the latter; and (3) the decision by a scavenger to feed should reflect a trade-off between nutritional status and the above preferences. Results from the laboratory and field were consistent with predictions. Two hypotheses are suggested from the work: scavengers may generally be less willing to feed on carcases of conspecifics than on those of other species, and intertidal scavengers may commonly face a relative or absolute shortage of food.     

Predation cues rather than resource availability promote cryptic behaviour in a habitat-forming sea urchin

It is well known that predators often influence the foraging behaviour of prey through the so-called “fear effect”. However, it is also possible that predators could change prey behaviour indirectly by altering the prey’s food supply through a trophic cascade. The predator–sea urchin–kelp trophic cascade is widely assumed to be driven by the removal of sea urchins by predators, but changes in sea urchin behaviour in response to predators or increased food availability could also play an important role. We tested whether increased crevice occupancy by herbivorous sea urchins in the presence of abundant predatory fishes and lobsters is a response to the increased risk of predation, or an indirect response to higher kelp abundances. Inside two New Zealand marine reserves with abundant predators and kelp, individuals of the sea urchin Evechinus chloroticus were rarer and remained cryptic (i.e. found in crevices) to larger sizes than on adjacent fished coasts where predators and kelp are rare. In a mesocosm experiment, cryptic behaviour was induced by simulated predation (the addition of crushed conspecifics), but the addition of food in the form of drift kelp did not induce cryptic behaviour. These findings demonstrate that the ‘fear’ of predators is more important than food availability in promoting sea urchin cryptic behaviour and suggest that both density- and behaviourally mediated interactions are important in the predator–sea urchin–kelp trophic cascade.     

The benefits of stealing from a predator: foraging rates, predation risk, and intraspecific aggression in the kleptoparasitic spider Argyrodes antipodiana

In the trade-off between food and safety, the role of aggressiveintraspecific interactions has not been extensively examined.Here I present information on this system using a kleptoparasiticspider, Argyrodts antipodiana, and its host spider and potentialpredator, Eriophora pustulosa. A. antipodiana can feed eitherat a potentially dangerous site (the hub of its host's web withthe host), or at a relatively safe site (on food bundles aroundthe edge of the host's web). I found that A. antipodiana cangain food very quickly when feeding with the host, apparentlyby exploiting the host's ability to digest the prey. Thus A.antipodiana follows predictions based on foraging models inthat it accepts a higher predation risk at the hub because ofthe higher food payoff. A. antipodiana also aggressively competesfor access to more food. However, aggressive competition increasesthe predation risk from the host, especially at tile hub wherethe host is very close. Consequently, A. antipodiana modifiesits level of intraspecific aggressiveness in accordance withits position on the web: at the hub, where the cost of aggressionis high (due to predation risk), A. antipodiana reduces itsaggressiveness, but it is aggressive away from the hub whencompeting for food bundles. The ability of A. antipodiana tochange interaction intensity as a function of its position onthe web enables it to exploit a rich, but risky, food sourceand provides a new angle for examining food and safety tradeoffs in light of intraspecific competition for food     

Effects of diet and starvation on digestive enzyme activity and feeding behavior of the marine copepod Calanus pacificus

In this paper we look at the response of Calanus pacificus overa period of several weeks to both starvation and diet as reflectedby survival rates, feeding rates and digestive enzyme activities.Phytoplankton species used as food were the diatom Thalassiosiraweissflogii, two dinoflagellates, Gymnodinium simplex and Scrippsiellatrochoidea, and the chlorophyte Dunaliella tertiolecta. In general,C.pacificus appeared to be resistant to starvation for periodsup to 1 week. Survival remained high for at least 6 days ofstarvation, while maximum feeding rates remained constant forseveral days and then declined by 50% after a week of starvation.When initial digestive enzyme activities were low, they didnot change appreciably during the experiments. However, activitylevels dropped rapidly during starvation experiments in whichthey were initially high. Other evidence indicates that starvationmay induce changes in the digestive system before it affectsingestion rates or survival. In general, long-term changes indigestive enzyme activity under different dietary regimes weregradual. An exception to this pattern was S. trochoidea, whosepresence in the diet induced a precipitous, though reversible,decline in digestive enzyme activity, a phenomena possibly relatedto the toxic reaction to this species reported by other researchers.A positive induction of digestive enzymes occurred over a time-scaleof 1–5 days in response to a change in diet. The absenceof laminarin in the diet of C.pacificus for nearly 3 weeks (whenfeeding on G.simplex), during which time laminarinase activitydeclined by {small tilde}50%, did not adversely affect subsequentassimilation of a laminarin-rich food (T. weissflogii).     

Copepod reaction to odor stimuli influenced by cestode infection

The cestode Schistocephalus solidus uses copepods as first andsticklebacks as second intermediate hosts. For transmission,an infected copepod has to be preyed upon by a stickleback.We used copepods of the species Macrocy albidus to test whetherinfected and uninfected copepods differ in their reaction totwo kind of simultaneously presented odors: odors of sticklebacksand odors of sticklebacks and conspecificz. By giving this choice,we attempted to force the copepods to make a trade-off betweenthe benefit of risk dilution and possible predator confusionand the costs of food competition and other disadvantages inducedby conspecifics. Within 1–8 h after last feeding, uninfectedcopepods clearly preferred the odors of conspeciflcs under thechemically simulated threat of predation. This was in contrastto the infected copepods, who tended to avoid the odor of conspecifics.When the time between experiment and last feeding varied, infectedcopepods showed an increas preference for fish water only (oravoided conspecthcs) with increasing hunger level This suggeststhat S. solidus benefits from hunger-induced behavioral changesof its copepod host by influencing its microhabitat selection.The same effect could be found in both sexes; however, it wassignificantly more pronounced in male than in female copepods.We propose several hypotheses that could explain the differencebetween the sexes in their infection-dependent microhabitatselection.     

Subtle cues of predation risk: starlings respond to a predator's direction of eye-gaze

For prey animals to negotiate successfully the fundamental trade-off between predation and starvation, a realistic assessment of predation risk is vital. Prey responses to conspicuous indicators of risk (such as looming predators or fleeing conspecifics) are well documented, but there should also be strong selection for the detection of more subtle cues. A predator's head orientation and eye-gaze direction are good candidates for subtle but useful indicators of risk, since many predators orient their head and eyes towards their prey as they attack. We describe the first explicit demonstration of a bird responding to a live predator's eye-gaze direction. We present wild-caught European starlings (Sturnus vulgaris) with human 'predators' whose frontal appearance and gaze direction are manipulated independently, and show that starlings are sensitive to the predator's orientation, the presence of eyes and the direction of eye-gaze. Starlings respond in a functionally significant manner: when the predator's gaze was averted, starlings resumed feeding earlier, at a higher rate and consumed more food overall. By correctly assessing lower risk and returning to feeding activity earlier (as in this study), the animal gains a competitive advantage over conspecifics that do not respond to the subtle predator cue in this way.     

Public information for solitary foragers: timber rattlesnakes use conspecific chemical cues to select ambush sites

Many animals use public information (PI) gathered from conspecificsto assess the quality of potential foraging locations. To date,research on this phenomenon has focused almost exclusively onsocial foragers that live in groups and monitor nearby individuals.PI is potentially available to solitary foragers as well, inthe form of cues (such as chemical cues) that persist in theenvironment after conspecifics are no longer present. In thisstudy, I examined the response of a solitary sit-and-wait predator,the timber rattlesnake (Crotalus horridus), to chemical cuesfrom conspecifics that had recently fed as opposed to thosethat had been deprived of food. Experiments with a T-maze indicatedthat timber rattlesnakes always follow conspecific chemicaltrails out of the maze, regardless of whether or not the individualleaving the trail had recently fed. However, an enclosure choicetest found that individuals are more likely to select ambushsites in areas with chemical cues from conspecifics that hadrecently fed. These results indicate that snakes may use conspecificchemical cues not only to find mates, shelter sites, and hibernaculabut also profitable food patches. Additionally, this study highlightsthe possibility that other solitary foragers may use PI to guidetheir foraging behavior.     

Behavioral syndrome over the boundaries of life--carryovers from larvae to adult damselfly

Activity is an important behavioral trait that mediates a trade-offbetween obtaining food for growth and avoiding predation. Activeindividuals usually experience a higher encounter rate withfood items and suffer higher predation pressure than less activeindividuals. I investigated how activity of the damselfly Lestescongener is affected by larval state and predator presence andif larval behavioral type (BT) can be used to predict larvalboldness, foraging success, and adult BT. Activity level ofindividual larvae was studied without predator at 2 differentphysiological states (hungry and fed) and in 2 predator treatments:familiar predator cues and unfamiliar predator cues. Larvaedid not adjust their activity depending on state or when subjectedto unfamiliar predator cues, but a general reduction in activitywas seen in the familiar predator treatment. Hence, active individualsremained active compared with their conspecifics, independentof state or predator treatment. Active individuals were alsobolder and more efficient foragers than their less active conspecifics.Furthermore, both adult activity and boldness were correlatedwith larval BT. The results illustrate that BT of a larvae iscarried over many different situations keeping active larvaeactive even in maladaptive situations, demonstrating how a behavioralsyndrome may constrain behavioral plasticity. Furthermore, resultsshowed that behavioral syndromes can carry over from larvaethrough metamorphosis and dictate the BT of the adult.     

Nitrogen excretion in rats on a protein-free diet and during starvation

Nitrogen balances (six days) were determined in male Wistar rats during feeding a diet with sufficient protein or a nearly protein-free diet (n = 2×24), and then during three days of starvation (n = 2×12). The objective was to evaluate the effect of protein withdrawal on minimum nitrogen excretion in urine (UN), corresponding to endogenous UN, during feeding and subsequent starvation periods. The rats fed the protein free-diet had almost the same excretion of urinary N during feeding and starvation (165 and 157 mg/kg W^0.75), while it was 444 mg/kg W^0.75 in rats previously fed with protein, demonstrating a major influence of protein content in a diet on Nexcretion during starvation. Consequently, the impact of former protein supply on Nlosses during starvation ought to be considered when evaluating minimum N requirement necessary to sustain life.     

Predator Avoidance and Alarm-response Behaviour in Kin-discriminating Tadpoles (Rana cascadae)

When captured by predators, the tadpoles of some species of frogs and toads may release an alarm substance that alters the behaviour of conspecifics. Such ‘alarm response’ behaviour has been proposed as a potential mechanism whereby related conspecifics may ‘warn’ relatives of a predator's presence and thus, improve their inclusive fitness. We examined predator avoidance and alarm response behaviour in tadpoles of the Cascades frog (Rana cascadae) and tested whether such behaviour is influenced by kinship factors. Tadpoles reduced activity when in the presence of a predatory newt (Taricha granulosa). Individuals in sibling groups were more active than both solitary tadpoles and individuals in mixed groups of siblings and nonsiblings. However, we found no evidence of an alarm response in R. cascadae. Behaviour of tadpoles in groups exposed only to predators was not different from that of tadpoles in groups exposed to predators plus crushed conspecifics. Tadpoles in groups exposed to crushed tadpoles were as active as tadpoles in groups exposed to water controls, and some test individuals fed upon the dead tadpoles. Thus, while R. cascadae tadpoles reduce activity in response to newt predators, crushed tadpoles appear to initiate a feeding response rather than an alarm response as has been previously proposed.