Footdrumming and other anti-predator responses in the bannertail kangaroo rat (Dipodomys spectabilis)

Summary Anti-predator behaviors of the bannertail kangaroo rat (Dipodomys spectabilis) toward snake predators were investigated. We induced responses by presenting a live (tethered) snake and an inflatable snake decoy to rats of known ages and sex in their territories. Comparisons of behaviors during spontaneous activity (baseline), naturally occurring snake-rat interactions, and experimental procedures further clarified anti-predator behaviors. The tethered snake immediately induced high rates of footdrumming (the repeated striking of the hind feet on the ground) in all rats (n=24) (Table 2), and individuals of both sexes and all ages footdrummed significantly more at the tethered snake than at the decoy. Other antipredator behaviors included alert postures, jumpback, kicking sand and avoidance. Juvenile rats exhibited more avoidance behavior and alert postures than adults and footdrummed at the decoy only if they experienced the snake first. We conclude that footdrumming functions as an individual alarm signal against predation by snakes in D. spectabilis. The drumming probably informs a snake that it has been detected and may cause it to leave. Because rats did not footdrum in response to aerial predators (owls), we suggest that kangaroo rats use different anti-predator strategies that depend on the kind of threat and the type of predator.     

Responses of pikas (Ochotona princeps,Lagomorpha) to naturally occurring terrestrial predators

Summary The responses of individually marked pikas (Ochotona princeps) to terrestrial predators were investigated in 1980 and 1981 in the Rocky Mountains of Colorado. Pikas uttered short call vocalizations in a variety of contexts: preceding or following an individual's movement, and in response to conspecifics, other nonpredaceous mammals and predators. Adult pikas apparently discriminated contexts in which predators were present by short calling more frequently and for longer duration compared with calling in nonpredator contexts. Short calls uttered by juveniles were similar in all contexts.Adults responded differently to two types of terrestrial predators: weasels and pine martens. Pikas called less frequently in response to weasels than to martens and avoided weasels more often than martens. They delayed the initiation of calling following the first sighting of a weasel more often than to martens. Weasels were determined to be more effective predators of pikas than martens, and these asymmetries in behavior and alarm vocalizations may indicate that responses reduce an individual's risk of predaton by weasels.Both male and female pikas called in response to predators, and residents called more often than nonresidents. The possible function of predator-related vocalization in pikas is discussed. It is suggested that calls to predators may function to warn local residents, which in pikas are usually closely related.     

Different antipredator behaviour in two anuran tadpoles: effects of predator diet

Recent investigations have indicated that animals are able to use chemical cues of predators to assess the magnitude of predation risk. One possible source of such cues is predator diet. Chemical cues may also be important in the development of antipredator behaviour, especially in animals that possess chemical alarm substances. Tadpoles of the common toad (Bufo bufo) are unpalatable to most vertebrate predators and have an alarm substance. Tadpoles of the common frog (Rana temporaria) lack both these characters. We experimentally studied how predator diet, previous experience of predators and body size affect antipredator behaviour in these two tadpole species. Late-instar larvae of the dragonfly Aeshna juncea were used as predators. The dragonfly larvae were fed a diet exclusively of insects, R. temporaria tadpoles or B. bufo tadpoles. R. temporaria tadpoles modified their behaviour according to the perceived predation risk. Depending on predator diet, the tadpoles responded with weak antipredatory behaviour (triggered by insect-fed predators) or strong behaviour (triggered by tadpole-fed predators) with distinct spatial avoidance and lowered activity level. The behaviour of B. bufo in predator diet treatments was indistinguishable from that in the control treatment. This lack of antipredator behaviour is probably related to the effective post-encounter defenses and more intense competitive regime experienced by B. bufo. The behaviour of both tadpole species was dependent on body size, but this was not related to predator treatments. Our results also indicate that antipredator behaviour is largely innate in tadpoles of both species and is not modified by a brief exposure to predators. Received: 22 August 1996 / Accepted after revision: 31 January 1997     

A review of predator diet effects on prey defensive responses

Numerous studies have examined how predator diets influence prey responses to predation risk, but the role predator diet plays in modulating prey responses remains equivocal. We reviewed 405 predator–prey studies in 109 published articles that investigated changes in prey responses when predators consumed different prey items. In 54 % of reviewed studies, prey responses were influenced by predator diet. The value of responding based on a predator’s recent diet increased when predators specialized more strongly on particular prey species, which may create patterns in diet cue use among prey depending upon whether they are preyed upon by generalist or specialist predators. Further, prey can alleviate costs or accrue greater benefits using diet cues as secondary sources of information to fine tune responses to predators and to learn novel risk cues from exotic predators or alarm cues from sympatric prey species. However, the ability to draw broad conclusions regarding use of predator diet cues by prey was limited by a lack of research identifying molecular structures of the chemicals that mediate these interactions. Conclusions are also limited by a narrow research focus. Seventy percent of reviewed studies were performed in freshwater systems, with a limited range of model predator–prey systems, and 98 % of reviewed studies were performed in laboratory settings. Besides identifying the molecules prey use to detect predators, future studies should strive to manipulate different aspects of prey responses to predator diet across a broader range of predator–prey species, particularly in marine and terrestrial systems, and to expand studies into the field.     

The effects of nesting stage,sex, and type of predator on parental defense by killdeer (Charadrius vociferous): testing models of avian parental defense

Summary Two models predicting the temporal patterns of parental investment in offspring defense over the nesting cycle were tested. The first is based on offspring age, the other on the vulnerability of offspring to predation. Both models make very similar predictions for altricial species after eggs have hatched, i.e., increases in intensity of parental defense until fledging. For precocial species, however, the post-hatching predictions of each model are different: the offspring age model predicts a continued increase in defense intensity, while the vulnerability model predicts a decline. I examined the temporal patterns of parental defense of a precocial shorebird, the killdeer (Charadrius vociferus), and determined which model was supported. Killdeer responses to human and natural predators were observed. Killdeer were less willing to leave the nest, responded most intensely, and displayed closest to a potential predator around hatching. Defense intensity increased from early to late incubation as predicted by the offspring age model. However, after hatching killdeer parental defense declined for both males and females, thus supporting the vulnerability model for this stage. Males and females responded significantly differently to all types of predators. Males took greater risks, remained on the nest longer, defended offspring more intensely, and displayed closer to the predator than females at the approach of a potential predator. Responses to natural predators depended on the type of predator and the approach made by the predator; a greater range of defense behavior was used for predators approaching on the ground compared to aerial predators. In general, killdeer responses to humans were more intense and less variable than their responses to natural predators. This was most likely because the human intruder approached nests and chicks more directly and closer than natural predators.     

Colonial breeding and nest defence in Montagu's harrier (Circus pygargus)

We assessed whether colonial breeding allows individuals to decrease their investment in predator defence, by presenting decoys of owls, foxes and crows to Montagu's harrier, Circus pygargus. Decoy detection increased with colony size, as did the number of individuals mobbing the decoy. The number of mobbers was greater for predators potentially risky for the adults (owl or fox) than for non-dangerous predators (crow). Recruits (breeding neighbours, fledglings and non-breeders) were present a lower percentage of the time, and attacked and alarm called less frequently than tested individuals. Nevertheless, the overall attack rate on the predator increased with the number of mobbers. When the size of the mobbing group increased, individuals were more likely to attack predators that represented a risk for adults, but did so less intensively and with a lower frequency of close dives. Thus, coloniality decreased the individual costs of defence in terms of risk taken, whilst enhancing defence efficacy. Birds alarm called more intensively when presented with dangerous predators than with the crow. The number of recruits significantly increased with increasing alarm rate of the tested individuals, even when taking colony size into account. Furthermore, the alarm rate of the tested birds also had a significant effect on the proportion of recruits that engaged in attacks against dangerous predators but not against the crow. The higher recruitment and attack rates for dangerous predators were thus apparently modulated through alarm calling. We discuss whether tested birds may manipulate recruits' behaviour to lessen their own risk.     

Anti-predator behavior of group-living Malagasy primates: mixed evidence for a referential alarm call system

Many mammals warn conspecifics with alarm calls about detected predators. These alarm calls are either functionally referential, urgency based, or they can have multiple functions, including predator deterrence. The taxonomic distribution of these alarm call systems is uneven, with primates providing the best-known examples for a functionally referential system and rodents most examples of an urgency-based system. Reports of different alarm call systems in lemurid primates prompted us to examine the anti-predator behavior of two additional lemur species. In an experimental field study we exposed adult redfronted lemurs (Eulemur fulvus rufus) and white sifakas (Propithecus verreauxi verreauxi) to playbacks of vocalizations of their main aerial and terrestrial predators, as well as to their own alarm calls given in response to the presentation of these predators. We scored the subjects' immediate behavioral responses, including alarm calls, from video recordings made during the first minute following a playback. We found that both species gave specific alarm calls only in response to raptor playbacks and the corresponding alarm calls, whereas calls given in response to carnivores and the corresponding alarm calls were also observed in other situations characterized by high arousal. Other behavioral responses, such as gaze and escape directions, corresponded to the hunting strategies of the two predator classes, suggesting that the corresponding vocalizations were categorized correctly. These two lemur species, which represent different families, have therefore independently evolved a mixed alarm call system, characterized by functionally referential calls for diurnal raptors, but not for carnivores. Electronic supplementary material to this paper can be obtained by using the Springer LINK server located at http://dx.doi.org/10.1007/s00265-001-0436-0 Electronic Publication     

Variation in behavioral and hormonal responses of adult male gray-cheeked mangabeys (<Emphasis Type="Italic">Lophocebus albigena</Emphasis>) to crowned eagles (<Emphasis Type="Italic">Stephanoaetus coronatus</Emphasis>) in Kibale National Park,Uganda

Intensive study of arboreal forest-dwelling primates and their predators in Africa is increasingly revealing that crowned eagles (Stephanoaetus coronatus) are major predators of primates. Gray-cheeked mangabeys (Lophocebus albigena) are overrepresented in the diets of crowned eagles in Kibale National Park, Uganda, and adult male mangabeys are represented more than females. We focused on the behavior of adult male gray-cheeked mangabeys living in social groups in Kibale National Park (1) to clarify the interactions between mangabeys and eagles that might put adult males at greater risk and (2) to better understand individual variation in behavioral responses to predators. Adult male mangabeys in five groups responded to observer-confirmed presence of crowned eagles 88 times over a 13-month period. While all males gave alarm calls, only the highest-ranking male in each of four groups chased eagles. These males had elevated levels of fecal cortisol metabolites in the days immediately after they engaged in active defense, suggesting that they perceived such behavior as risky. In the one group where male ranks were unstable and there were no infants, no male was observed to chase eagles. We suggest that males pursue the dangerous tactic of chasing eagles only when they are likely to have offspring in the group. Males in larger groups also spent less time alarm calling to crowned eagles (from first to last call in a group), and our observations confirmed that the duration of their alarm calls was related to eagle presence. Thus, eagles spent less time around larger mangabey groups. Alarm calling by adult male mangabeys may signal to this ambush predator that it has been detected and should move on.     

Tidal regime dictates the cascading consumptive and nonconsumptive effects of multiple predators on a marsh plant

Prey perception of predators can dictate how prey behaviorally balance the need to avoid being eaten with the need to consume resources, and this perception and consequent behavior can be strongly influenced by physical processes. Physical factors, however, can also alter the density and diversity of predators that pursue prey. Thus, it remains uncertain to what extent variable risk perception and antipredator behavior vs. variation in predator consumption of prey underlie prey-resource dynamics and give rise to large-scale patterns in natural systems. In an experimental food web where tidal inundation of marsh controls which predators access prey, crab and conch (predators) influenced the survivorship and antipredator behavior of snails (prey) irrespective of whether tidal inundation occurred on a diurnal or mixed semidiurnal schedule. Specifically, cues of either predator caused snails to ascend marsh leaves; snail survivorship was reduced more by unrestrained crabs than by unrestrained conchs; and snail survivorship was lowest with multiple predators than with any single predator despite interference. In contrast to these tidally consistent direct consumptive and nonconsumptive effects, indirect predator effects differed with tidal regime: snail grazing of marsh leaves in the presence of predators increased in the diurnal tide but decreased in the mixed semidiurnal tidal schedule, overwhelming the differences in snail density that resulted from direct predation. In addition, results suggest that snails may increase their foraging to compensate for stress-induced metabolic demand in the presence of predator cues. Patterns from natural marshes spanning a tidal inundation gradient (from diurnal to mixed semidiurnal tides) across 400 km of coastline were consistent with experimental results: despite minimal spatial variation in densities of predators, snails, abiotic stressors, and marsh productivity, snail grazing on marsh plants increased and plant biomass decreased on shorelines exposed to a diurnal tide. Because both the field and experimental results can be explained by tidal-induced variation in risk perception and snail behavior rather than by changes in snail density, this study reinforces the importance of nonconsumptive predator effects in complex natural systems and at large spatial scales.     

The influence of intraguild predation on prey suppression and prey release: a meta-analysis

Intraguild predation (IGP) occurs when one predator species consumes another predator species with whom it also competes for shared prey. One question of interest to ecologists is whether multiple predator species suppress prey populations more than a single predator species, and whether this result varies with the presence of IGP. We conducted a meta-analysis to examine this question, and others, regarding the effects of IGP on prey suppression. When predators can potentially consume one another (mutual IGP), prey suppression is greater in the presence of one predator species than in the presence of multiple predator species; however, this result was not found for assemblages with unidirectional or no IGP. With unidirectional IGP, intermediate predators were generally more effective than the top predator at suppressing the shared prey, in agreement with IGP theory. Adding a top predator to an assemblage generally caused prey to be released from predation, while adding an intermediate predator caused prey populations to be suppressed. However, the effects of adding a top or intermediate predator depended on the effectiveness of these predators when they were alone. Effects of IGP varied across different ecosystems (e.g., lentic, lotic, marine, terrestrial invertebrate, and terrestrial vertebrate), with the strongest patterns being driven by terrestrial invertebrates. Finally, although IGP theory is based on equilibrium conditions, data from short-term experiments can inform us about systems that are dominated by transient dynamics. Moreover, short-term experiments may be connected in some way to equilibrium models if the predator and prey densities used in experiments approximate the equilibrium densities in nature.     

A forest monkey’s alarm call series to predator models

Some non-human primates produce acoustically distinct alarm calls to different predators, such as eagles or leopards. Recipients respond to these calls as if they have seen the actual predator, which has led to the notion of functionally referential alarm calls. However, in a previous study with free-ranging putty-nosed monkeys (Cercopithecus nictitans martini), we demonstrated that callers produced two acoustically distinct alarm calls to eagle shrieks and leopard growls, but both alarm calls were given to both predators. We can think of two basic explanations for this surprising result, a methodological and theoretical one. Firstly, acoustic predator models may not always be suitable to test alarm call behaviour in primates, sometimes causing uncharacteristic behaviour. Secondly, referential alarm calling may not be a universal feature of primate alarm call systems. Considering the methodological and theoretical importance of these possibilities, we conducted a follow-up study using life-sized leopard, eagle, and human models on the same population and compared the resulting vocal responses to those given to acoustic predator models. We compared the alarm call series given to each of these predator model types and found a considerable degree of consistency suggesting that the mode of presentation did not affect anti-predator calling strategies. However, evidence for audience effects on calling behaviour was inconclusive. While it appears that predator class is reliably encoded by different call series types irrespective of the mode of presentation, observations of these same call series given in non-predatory contexts indicate that predator class is unlikely to be the relevant organising principle underlying the alarm-calling behaviour in this species. We conclude by offering an alternative, non-referential, account of the alarm-calling system exhibited by this species. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Risk level of chemical cues determines retention of recognition of new predators in Iberian green frog tadpoles

In aquatic environments, many prey rely on chemosensory information from injured (alarm cues) or stressed conspecifics (disturbance cues) to assess predation risk. Alarm cues are considered as a sign of higher risk than disturbance cues. These cues could be used by prey to learn potential new predators. In this study, we tested whether Iberian green frog tadpoles (Pelophylax perezi) exhibited antipredator responses to alarm and disturbance cues of conspecifics and whether tadpoles could associate new predators with alarm or disturbance cues. Tadpoles reduced their activity in the presence of disturbance cues, but only weakly when compared with their response to alarm cues. Also, tadpoles learned to recognize new predators from association with alarm or disturbance cues. However, the period of retention of the learned association was shorter for disturbance than alarm cues. Our results indicate that tadpoles are able to modify their antipredatory behavior according to (1) the degree of risk implied by the experimental cues (2) their previous experience of chemical cues of the predator.     

Threat-sensitive learning of predators by larval mosquitoes <Emphasis Type="Italic">Culex restuans</Emphasis>

A prerequisite for prey to show adaptive behavioural responses to predators is that the prey has the ability to recognise predators as threats. While predator recognition can be innate in many situations, learning is often essential. For many aquatic species, one common way to learn about predators is through the pairing of a novel predator odour with alarm cues released from injured conspecifics. One study with fish demonstrated that this mode of learning not only allows the prey to recognise the predatory cues as a threat, but also mediates the level of threat associated with the predator cues (i.e. threat-sensitive learning). When the prey is exposed to the novel predator with a high concentration of alarm cues, they subsequently show a high intensity of antipredator response to the predator cues alone. When exposed to the predator with a low concentration of alarm cues, they subsequently show a low-intensity response to the predator cues. Here, we investigated whether larval mosquitoes Culex restuans have the ability to learn to recognise salamanders as a threat through a single pairing of alarm cues and salamander odour and also whether they would learn to respond to salamander cues in a threat-sensitive manner. We conditioned individual mosquitoes with water or a low, medium or high concentration of crushed conspecific cues (alarm cues) paired with salamander odour. Mosquitoes exposed to salamander odour paired with alarm cues and subsequently exposed to salamander odour alone responded to the salamander as a threat. Moreover, the intensity of antipredator response displayed during the conditioning phase matched the response intensity during the testing phase. This is the first demonstration of threat-sensitive learning in an aquatic invertebrate.     

Ontogenetic differences in chemical alarm cue production determine antipredator responses and learned predator recognition

How individuals assess, respond and subsequently learn from alarm cues is crucial to their survival and future fitness. Yet this information is not constant through time; many individuals are exposed to different predators throughout their life as they outgrow some predators or move to habitats containing different predators. To maximise overall fitness, individuals should discriminate between different cues and respond and learn from only those that are relevant to their current ontogenetic stage. We tested whether juvenile spiny chromis, Acanthochromis polyacanthus, could distinguish between chemical alarm cues from conspecific donors of different ontogenetic stages and whether the cue ontogenetic stage of the cue donor affected the efficacy of learning about predators. Juveniles displayed a significant antipredator response when conditioned with juvenile chemical alarm cues paired with predator odour but failed to respond when conditioned with predator odour paired with either adult alarm cues or with saltwater. Subsequently, individuals only recognised the predator odour alone as a threat when conditioned with juvenile alarm cues. This demonstrates that prey may be highly specific in how they use information from conspecific alarm cues, selectively responding to and learning from only those cues that are relevant to their developmental stage.     

Behavioral resource depression and decaying perceived risk of predation in two species of coexisting gerbils

Summary Behavioral resource depression occurs when the behavior of prey individuals changes in response to the presence of a predator, resulting in a reduction of the encounter rate of the predator with its prey. Here I present experimental evidence on the response of two species of gerbils (Gerbillus allenbyi and G. pyramidum) to the presence of barn owls. I conducted the experiments in a large aviary. Both gerbils responded to the presence of barn owl predators by foraging in fewer resource patches (seed trays) and by quitting foraged resource patches at a higher resource harvest rate (giving-up density of resource; GUD). This reduced the amount of time gerbils were exposed to owl predation, and hence the encounter rate of owls with gerbils, i.e., behavioral resource depression. Thus, the presence of owls imposes a foraging cost on gerbils due to risk of predation, and also on the owls themselves due to resource depression. I then examined how resource depression relaxed over time following exposure to owls. In the days following an encounter with the predator, the reduction in foraging activity for both gerbil species eased. Increasing numbers of trays were foraged each day, and GUDs in seed trays declined. The two gerbils differed in their rate of recovery, with G. pyramidum returning to prepredator levels of foraging after 1 or 2 nights and G. allenbyi taking 5 nights or longer. Interspecific differences in recovery rates may be based on differences between the species in vulnerability to predation and/or ability to detect the presence of predators. The differences in recovery rates may be due to optimal memory windows or decay rates, where differences between species are based on risk of predation or on how perceived risk changes with time since a predator was last encountered. Finally, differences between or among competitors in recovery from resource depression may provide foraging opportunities in time for the species which recover most quickly and may have implications for species coexistence.     

Recognising one’s enemies: a functional approach to risk assessment by prey

Little has been done to compare the relative importance of various mechanisms through which prey assess the potential risk from natural enemies. We used predator-naive spider mites (Tetranychus urticae, Tetranychidae) to (1) compare the responses of prey to chemical cues from enemy and non-enemy species and (2) investigate the source of these cues. In the laboratory, we observed the distribution of T. urticae in response to cues from nine mite species, including (1) predators of spider mites, (2) predators/parasites of other animals, and (3) fungivores/pollen-feeders. When given a choice over 24 h, spider mites foraged and oviposited in fewer numbers on leaf discs that were previously exposed to predatory or parasitic mites (including species incapable of attacking spider mites) than on clean leaf discs (unexposed to mites). Interestingly, previous exposure of leaf arenas to fungivores and pollen-feeders had no significant effect on spider mite distribution. We then observed the response of T. urticae to cues from two species of predator that had been reared on a diet of either spider mites or pollen. T. urticae showed stronger avoidance of leaf discs that were previously exposed to spider-mite-fed predators than of discs exposed to pollen-fed predators. Nevertheless, for one predator species (Amblyseius andersoni), T. urticae still preferred to forage and oviposit on clean (unexposed) discs than on discs exposed to pollen-fed predators. Protein-derived metabolic wastes of predatory or parasitic mites may provide a general cue about potential predation risk for T. urticae. However, T. urticae also avoided areas exposed to pollen-fed predators, suggesting there may be other sources of enemy recognition by the spider mites. We discuss the ecological and evolutionary mechanisms that may influence the scope of information through which animals assess predation risk. Received: 11 January 1999 / Received in revised form: 25 October 1999 / Accepted: 20 November 1999     

Communal burrowing in the hystricognath rodent, Octodon degus: a benefit of sociality?

We examined the hypothesis that a main benefit of group-living in the semifossorial rodent, Octodon degus (Rodentia: Octodontidae), is to decrease individual cost of burrow construction. We contrasted the digging behavior of groups of three same-sex, adult-sized individuals with that of solitary degus. The behavior of singles and trios was recorded inside a large terrarium partially filled with natural soil and under controlled conditions of food, light, and temperature. The observation that degus in groups do not decrease their burrowing time or frequency of digging compared with solitary diggers does not support the hypothesis that communal burrowing is a primary cause of degu sociality. On the other hand, the observation that degus in groups removed significantly more soil per capita than solitary digging degus, and that grouped individuals coordinated their digging – group members burrowed mostly in the same sites and formed digging chains –, suggests that social burrowing may potentially reduce the cost of burrow construction in the long term. We suggest that such long-term benefits will be a consequence rather than a cause of degu group-living. Received: 9 December 1999 / Received in revised form: 2 January 2000 / Accepted: 8 February 2000     

Risk taking during parental care: a test of three hypotheses applied to the pied flycatcher

According to life-history theory, there will often be a conflict between investment in current versus future reproduction. If a predator appears during breeding, parents must make a compromise between ensuring the growth and survival of offspring (nest defence, feeding and brooding of young), and reducing the risk of predation to ensure their own survival. We model three hypotheses for the outcome of this conflict which are particularly relevant for altricial birds. They are not mutually exclusive, but focus on different costs and benefits. (1) Parental investment is determined by the parents’ own risk of predation. This hypothesis predicts that a lone parent should take smaller risks than a parent that has a mate. (2) Parental investment is related to the reproductive value of the offspring: Parents are predicted to take greater risks for larger broods, larger-sized or older offspring. (3) Finally, we present the new hypothesis that parental investment is related to the harm that offspring would suffer during a period of no parental care (incubation, brooding, feeding). This hypothesis predicts that parents should take greater risks for younger offspring, or for offspring in poorer condition, because the marginal benefit of parental care is largest in such cases. Hence, one may also expect that lone parents should take greater risks than two parents because their offspring are more in need of care. We tested these hypotheses on the pied flycatcher (Ficedula hypoleuca) by presenting a stuffed predator of the parents (a sparrowhawk, Accipiter nisus) close to the nest when parents were feeding the young. Risk taking was measured as the time that elapsed until the first visit to the nest. Most support was found for the ‘‘harm to offspring’’ hypothesis. Previous studies have usually measured the intensity of nest defence against typical nest predators, and have found evidence for the ‘‘reproductive value of offspring’’ hypothesis. However, our model predicts that the importance of the reproductive value of the offspring should decrease relative to the harm that offspring would suffer if they were not cared for when the predator type changes from a nest predator to a predator of adults, and when conditions for breeding turn from good to bad. Received: 13 April 1995/Accepted after revision: 11 March 1996     

Predator-induced alarm responses in the common periwinkle, Littorina littorea: dependence on season, light conditions, and chemical labelling of predators

Chemically mediated alarm reactions of the common periwinkle, Littorina littorea (L.), were studied in laboratory experiments during two consecutive summers, and one intermediate autumn season. Responses to chemical stimuli were detected as crawl-out responses, i.e. movements of snails out of the water. Snails were exposed to extracts of injured conspecifics, extracts of the mussel Modiolus modiolus (L.), and water conditioned by the predatory crab Carcinus maenas (L.), which had been maintained on different diets. In experiments carried out during the summer, a significantly larger number of snails moved out of the water when exposed to chemical stimuli from injured conspecifics, compared to chemical stimuli from injured mussels or filtered seawater. These results suggest that chemical alarm substances are present in L. littorea. Water conditioned by crabs that had been fed L. littorea released significantly more crawl-out responses compared to water conditioned by crabs that had been kept on a fish diet. When tested in autumn, no significant differences were found in responses to the above-mentioned water samples. Crawl-out responses under different light regimes were also investigated. All series of experiments carried out in the dark evoked a higher number of responses compared to series that took place in light. These findings may indicate an adaptation of snails to night-active predators. In total, the current results suggest that a L. littorea diet may chemically “label” the predator crab with snail alarm substances, and that predator-induced responses of L. littorea are actually responses to conspecific alarm substances released from crabs that have been maintained on a L. littorea diet. The response to the alarm signal, however, appears to be dependent on season and light conditions; some ecological implications of these findings are also discussed. Received: 8 January 1999 / Accepted: 29 March 1999     

The paradox of risk assessment: comparing responses of fathead minnows to capture-released and diet-released alarm cues from two different predators

Summary. Many aquatic prey are known to use chemical alarm cues to assess their risk of predation. In fishes, such alarm cues can be released either through damage of the epidermis during a predatory attack (capture-released) or through release from the predator feces (diet-released). In our study, we compared the importance of capture- versus diet-released alarm cues in risk assessment by fathead minnows (Pimephales promelas) that were na?ve to fish predators. We utilized two different fish predators: a specialized piscivore, the northern pike (Esox lucius) and a generalist predator, the brook trout (Salvelinus fontinalis). Handling time of pike consuming minnows was much shorter than for trout consuming minnows, likely resulting in less epidermal damage to the minnows during attacks by pike. In accordance with this, minnows showed a less intense antipredator response to capture-released cues from pike than capture-released cues from trout. This represents a paradox in risk assessment for the minnows as they respond to the specialized piscivore, the more dangerous predator, with a less intense antipredator response. In contrast, the minnows showed a stronger antipredator response to the specialized piscivore than to the generalist when given diet cues. This work highlights the need for researchers to carefully consider the nature of the information available to prey in risk assessment.