Differences in the nestling begging calls of hosts and host-races of the common cuckoo, Cuculus canorus

We compared nestling begging calls of four hosts (reed warbler, Acrocephalus scirpaceus; great reed warbler, A. arundinaceus; dunnock, Prunella modularis; and meadow pipit, Anthus pratensis) and the respective host-races of the common cuckoo. Note structure varied between host species, but not between cuckoo host-races, so cuckoos did not vary their call note structure to match that of their hosts' chicks. Call rate increased with age, but there were marked differences between both host species and cuckoo host-races. Dunnock-cuckoos called more rapidly than reed warbler-cuckoos despite growing at the same rate. We suggest this difference reflects how cuckoos tune into the way these host species respond to begging signals from their own young, because dunnock chicks called much more rapidly than reed warbler chicks. Great reed warbler-cuckoos called at a lower rate than reed warbler-cuckoos when young, but at a greater rate when older than 8 days. This could also result from the cuckoo chicks tuning into differences in the way these hosts respond to begging signals. However, great reed warbler-cuckoos grew at a faster rate than the other cuckoo host-races, so they may also call faster to demand higher provisioning rates from this larger host. To test these hypotheses critically, data are needed on how the different host species integrate visual and vocal begging signals from their own broods. We discuss how differences in cuckoo begging might develop, given that cuckoo host-races are restricted to female cuckoo lineages. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.      

Who is in the neighborhood? Conspecific and heterospecific responses to perceived density for breeding habitat selection

Theoretical models of habitat selection often incorporate negative density dependence. Despite strong negative density‐dependent effects on habitat selection, more recent studies indicate that animals settle near members of their own (conspecific) and other species (heterospecific) when selecting habitat with social cues. Social cue use for habitat selection is particularly common among songbirds, but few studies have investigated if songbirds use social cues to assess conspecific or heterospecific density (as opposed to just presence/absence) when making settlement decisions. We conducted a playback experiment to evaluate if yellow warblers (Setophaga petechia) and willow flycatchers (Empidonax traillii), two potential competitors for breeding habitat, use social cues to assess density (conspecific for warblers and heterospecific for flycatchers) when selecting breeding locations at two spatial scales. We simulated yellow warbler density to be high or low at multiple treatment plots (3.14 ha) with song playback and then evaluated settlement decisions by comparing yellow warbler and willow flycatcher abundances across plots (broad‐scale habitat selection) and individual space use within plots (fine‐scale territory establishment). Yellow warbler density treatments did not affect habitat selection by yellow warblers at the broad scale, but caused individuals to cluster territories at high‐density treatments. Willow flycatchers were most abundant at high‐density treatment plots, but yellow warbler density treatments did not affect territory locations. The results indicate that perceived density affects the habitat selection process for both conspecifics and heterospecifics.     

Reinforcement generates reproductive isolation between neighbouring conspecific populations of spadefoot toads

Reproductive character displacement is the adaptive evolution of traits that minimize deleterious reproductive interactions between species. When arising from selection to avoid hybridization, this process is referred to as reinforcement. Reproductive character displacement generates divergence not only between interacting species, but also between conspecific populations that are sympatric with heterospecifics versus those that are allopatric. Consequently, such conspecific populations can become reproductively isolated. We compared female mate preferences in, and evaluated gene flow between, neighbouring populations of spadefoot toads that did and did not occur with heterospecifics (mixed- and pure-species populations, respectively). We found that in mixed-species populations females significantly preferred conspecifics. Such females also tended to prefer a conspecific call character that was dissimilar from heterospecifics. By contrast, females from pure-species populations did not discriminate conspecific from heterospecific calls. They also preferred a more exaggerated conspecific call character that resembles heterospecific males. Moreover, gene flow was significantly reduced between mixed- and pure-species population types. Thus, character displacement (and, more specifically, reinforcement) may initiate reproductive isolation between conspecific populations that differ in interactions with heterospecifics.     

A host-race of the cuckoo Cuculus canorus with nestlings attuned to the parental alarm calls of the host species

The common cuckoo has several host-specific races, each with a distinctive egg that tends to match its host's eggs. Here, we show that the host-race specializing on reed warblers also has a host-specific nestling adaptation. In playback experiments, the nestling cuckoos responded specifically to the reed warbler's distinctive 'churr' alarm (given when a predator is near the nest), by reducing begging calls (likely to betray their location) and by displaying their orange-red gape (a preparation for defence). When reed warbler-cuckoos were cross-fostered and raised by two other regular cuckoo hosts (robins or dunnocks), they did not respond to the different alarms of these new foster-parents. Instead, they retained a specific response to reed warbler alarms but, remarkably, increased both calling and gaping. This suggests innate pre-tuning to reed warbler alarms, but with exposure necessary for development of the normal silent gaping response. By contrast, cuckoo chicks of another host-race specializing on redstarts showed no response to either redstart or reed warbler alarms. If host-races are restricted to female cuckoo lineages, then chick-tuning in reed warbler-cuckoos must be under maternal control. Alternatively, some host-races might be cryptic species, not revealed by the neutral genetic markers studied so far.     

Ontogeny of conspecific and heterospecific alarm call recognition in wild Verreaux's sifakas (Propithecus verreauxi verreauxi)

The production of vocalizations in nonhuman primates is predominantly innate, whereas learning influences the usage and comprehension of vocalizations. In this study, I examined the development of alarm call recognition in free-ranging infant Verreaux's sifakas. Specifically, I investigated their ability to recognize conspecific alarm calls as well as those of sympatric redfronted lemurs (Eulemur fulvus rufus) in Kirindy forest, western Madagascar. Both species have functionally referential alarm calls for aerial predators and give general alarm calls for both aerial and general predators and also other kinds of threats, such as intergroup encounters with conspecifics. I conducted playback experiments with members of two birth cohorts (nine and ten individuals) to determine the age at which infant Verreaux's sifakas discriminate between conspecific alarm calls, heterospecific alarm calls, and non-alarm vocalizations (parrot song). Most 3-4 months old infants fled toward adults after hearing any playback stimuli, whereas 4-5-month-old infants did so only after presentation of alarm calls. Moreover, all infants of these age classes showed a longer latency to flee after the parrot song indicating their emerging ability to discriminate between alarm calls and non-alarm stimuli. At an age of about 6 months, infants switched from fleeing toward adults to performing adult-like escape responses after presentation of conspecific and heterospecific alarm calls. Thus, the ability to discriminate between alarm from non-alarm stimuli precedes the appearance of adult-like responses. The transition to adult-like escape behavior was coincident with the physical independence of infants from their mothers.     

A host-race difference in begging calls of nestling cuckoos Cuculus canorus develops through experience and increases host provisioning

The structure of common cuckoo nestling begging calls differs between the two host-races parasitizing reed warblers (reed warbler-cuckoos) and dunnocks (dunnock-cuckoos; longer syllable duration, lower peak and maximum frequency, narrower bandwidth). Cross-fostering experiments demonstrated that this difference is not genetically fixed but develops through experience. When newly hatched reed warbler-cuckoos were transferred to dunnock nests, they developed begging calls more like those of dunnock-cuckoos, whereas controls transferred to the nests of robins or left to be raised by reed warblers developed calls more typical of reed warbler-cuckoos. We tested the effectiveness of these different calls in stimulating host provisioning by placing in host nests a single blackbird or song thrush nestling (of similar size to a young cuckoo, but lacking its exuberant begging calls); when it begged we broadcast, from a small loudspeaker on the nest rim, recordings of either dunnock-cuckoo or reed warbler-cuckoo begging calls. Playback of dunnock-cuckoo begging calls induced higher levels of provisioning by dunnocks, whereas playback of reed warbler-cuckoo begging calls did so for both reed warblers and robins. We suggest that the young cuckoo (which ejects the host's eggs/chicks and so is raised alone) learns by experience which calls best stimulate host provisioning.     

Asymmetric reproductive character displacement in male aggregation behaviour

Reproductive character displacement--the evolution of traits that minimize reproductive interactions between species--can promote striking divergence in male signals or female mate preferences between populations that do and do not occur with heterospecifics. However, reproductive character displacement can affect other aspects of mating behaviour. Indeed, avoidance of heterospecific interactions might contribute to spatial (or temporal) aggregation of conspecifics. We examined this possibility in two species of hybridizing spadefoot toad (genus Spea). We found that in Spea bombifrons sympatric males were more likely than allopatric males to associate with calling males. Moreover, contrary to allopatric males, sympatric S. bombifrons males preferentially associated with conspecific male calls. By contrast, Spea multiplicata showed no differences between sympatry and allopatry in likelihood to associate with calling males. Further, sympatric and allopatric males did not differ in preference for conspecifics. However, allopatric S. multiplicata were more variable than sympatric males in their responses. Thus, in S. multiplicata, character displacement may have refined pre-existing aggregation behaviour. Our results suggest that heterospecific interactions can foster aggregative behaviour that might ultimately contribute to clustering of conspecifics. Such clustering can generate spatial or temporal segregation of reproductive activities among species and ultimately promote reproductive isolation.     

Reproductive character displacement generates reproductive isolation among conspecific populations: an artificial neural network study

When interactions with heterospecifics prevent females from identifying conspecific mates, natural selection can promote the evolution of mating behaviours that minimize such interactions. Consequently, mating behaviours may diverge among conspecific populations in sympatry and in allopatry with heterospecifics. This divergence in conspecific mating behaviours-reproductive character displacement-can initiate speciation if mating behaviours become so divergent as to generate reproductive isolation between sympatric and allopatric conspecifics. We tested these ideas by using artificial neural networks to simulate the evolution of conspecific mate recognition in populations sympatric and allopatric with different heterospecifics. We found that advertisement calls diverged among the different conspecific populations. Consequently, networks strongly preferred calls from their own population to those from foreign conspecific populations. Thus, reproductive character displacement may promote reproductive isolation and, ultimately, speciation among conspecific populations.     

Interspecific responses to distress calls in bats (Chiroptera: Vespertilionidae): a function for convergence in call design?

Distress calls were recorded from three sympatric species of pipistrelle bat (Pipistrellus nathusii, P. pipistrellus and P. pygmaeus) in England and Northern Ireland. At foraging sites, we conducted playback experiments, consisting of experimental distress call sequences from each species and control sequences of random noise and sound recorded with no bats present. We measured response by simultaneously recording ultrasound during playbacks and counting the echolocation pulses above a predetermined threshold which were then identified to species. All three species responded to each other's calls. The number of recorded echolocation pulses of all species increased eight-fold, on average, during the playback of distress call sequences compared with the playback of ultrasonic noise, and four-fold compared with the playback of silence. In a separate playback experiment, the number of echolocation pulses of P. pygmaeus increased 14-fold during the playback of distress calls of four endemic species of bat from Madagascar (Emballonura atrata, Myotis goudoti, Miniopterus majori and M. manavi) compared with the playback of silence. This increased response might have been caused by the high calling rates of the Malagasy species. Distress calls of P. nathusii, P. pipistrellus and P. pygmaeus were structurally convergent, consisting of a series of downward-sweeping, frequency-modulated elements of short duration and high intensity with a relatively strong harmonic content. Selection may favour convergence in the structure of distress calls among bat species, if attracting heterospecifics increases the chance of repelling predators by mobbing.     

Differential Response to Interspecific and Intraspecific Signals Amongst Chickadees

Black‐capped chickadees (Poecile atricapillus) and mountain chickadees (P. gambeli) have a similar vocal repertoire and share many other life history traits; yet, black‐capped chickadees are socially dominant to mountain chickadees where populations overlap. Previous research suggested that in contact zones, both species respond weakly to heterospecific songs during the breeding season, and have suggested minimal interspecific competition. However, both black‐capped and mountain chickadees discriminate between conspecific and heterospecific chick‐a‐dee calls, suggesting attention is paid to interspecific signals. We compared the responses of both black‐capped and mountain chickadees to conspecific and heterospecific chick‐a‐dee calls during the winter, when both species compete for the same food resources. We conducted an aviary playback experiment exposing both species to playback composed of heterospecific and conspecific chick‐a‐dee calls, which had been recorded in the context of finding food sources. Responses from the tested birds were measured by recording vocalizations and behaviour. Black‐capped chickadees responded significantly more to conspecific than to heterospecific stimuli, whereas the subordinate mountain chickadees responded to both mountain and black‐capped chickadee calls. Based upon the reactions to playbacks, our results suggest these two closely related species may differ in their perception of the relative threat associated with intra‐ versus interspecific competitors.     

Alarm responses are maintained during captive rearing in chicks of endangered kaki

The maintenance of species-specific behavioural repertoires and traditions is an important but often implicit goal of conservation efforts. When captive rearing is used as a conservation practice, it becomes critical to address its possible implications for the social and behavioural traits of developing individuals. In particular, animals must retain or acquire many of their behavioural abilities to increase the likelihood of survival upon release into the wild. This study investigated the behavioural development of critically endangered kaki (black stilt: Himantopus novaezelandiae ) chicks reared without live adult conspecifics. The captive rearing programme included playbacks of adult kaki alarm calls during cleaning and handling of precocial chicks housed as groups. We used videotaped observations and playback experiments to address the following questions: do kaki chicks respond differentially to (1) familiar versus unfamiliar adult kaki alarm calls and (2) conspecific alarm versus heterospecific control vocalizations. Adult-naïve kaki chicks exhibited a varied behavioural repertoire over their early development. In multivariate analyses, when age was statistically controlled, chicks showed responses to familiar and unfamiliar alarm calls that were similar in magnitude. In contrast, following conspecific alarm calls chicks had longer average latencies to resume pre-playback activities than following heterospecific vocalizations. Although the generality of these conclusions is limited by experimental constraints stemming from working with an endangered species, the findings suggest that current management techniques produce captive-reared kaki for release into the wild that possess many of the behavioural and auditory recognition skills that are required for survival.     

Interspecific Response to Playback of Bird Song

Responses to bird song have usually only been studied at the intraspecific level. I experimentally tested whether playback of the song of the black wheatear Oenanthe leucura in an area in S Spain resulted in responses from conspecifics as well as heterospecific birds by comparing the numbers of individuals singing before and after playback. The number of singing male black wheatears increased considerably, but also the number of singing males of five other passerine species increased significantly. The heterospecific response to playback may be due (1) to interspecific territoriality, (2) to black wheatear song signalling the absence of predators, or (3) to heterospecifics confusing the species-identity of the singer. The second alternative is considered more likely, since an ecologically wide array of species increased their song rate following playback. The conspicuous dawn (and dusk) chorus of bird song may be augmented by social facilitation due to the singing of conspecifics as well as heterospecifics.     

Simulated chorus attracts conspecific and heterospecific Amazonian explosive breeding frogs

In tropical regions, some anuran species breed "explosively", reproducing in massive and highly diverse aggregations during a brief window of time. These aggregations can serve as acoustic beacons, attracting other anurans toward seasonal ponds. We hypothesize that conspecific and heterospecific calls play a role in navigation toward ponds and synchronization of reproduction among species. We simulated a chorus of two species (Trachycephalus coriaceus and Chiasmocleis shudikarensis) with contrasting call characteristics (low‐frequency vs. high‐frequency) and reproductive strategies (strict pond breeder vs. opportunistic) near known explosive breeding sites. We predicted that choruses of T. coriaceus are more attractive to heterospecifics than of C. shudikarensis because the first provides a more reliable indicator of a suitable breeding pond and a better long‐distance signal. We found that both choruses attracted conspecific frogs to the playback outside a natural breeding event. As predicted, heterospecifics were attracted only by low‐frequency calls of T. coriaceus that breeds exclusively in large ponds, but not by higher frequency calls of C. shudikarensis that also breeds in small pools not suitable for other species. Our study presents the first experimental evidence that tropical explosive breeding anurans are attracted to conspecific and heterospecific choruses. The contrasting effect of the playback of the two species on heterospecifics suggests that the attractive effect of a chorus depends on the reproductive strategy of both the sender and the receiver. Given the abundance and diversity of communities in tropical ecosystems, the use of heterospecific acoustic cues may prove widespread and requires further investigation. Abstract in Portuguese is available with online material.     

Early singers attend to conspecific but not heterospecific behavioural cues at dawn

During the dawn chorus songbirds initiate singing activities just prior to sunrise. Environmental factors affect the timing of the dawn chorus, but relatively little is known about how behavioural cues influence chorus timing. We assessed whether early playback of conspecific and heterospecific song influenced chorus start times in two early‐singing temperate species, hermit thrush Catharus guttatus and veery C. fuscescens. We used arrays of GPS time‐synchronized recorders to record the dawn chorus at 30 locations. In each location, thirty minutes prior to natural chorus initiation, we broadcast: 1) an early singing species (hermit thrush or veery), 2) a late singing species (black‐throated green warbler Setophaga virens or Nashville warbler Oreothlypis ruficapilla), 3) a noise control, and 4) also recorded without playback (silent control). Playback treatments were separated by two‐three days at each location. Both hermit thrush and veery sang significantly earlier in response to conspecific playback compared to noise or silent controls. Both species also sang earlier in response to conspecific playback compared to the day before or the day after playback. Neither species sang earlier in response to heterospecific playback (regardless of whether the broadcast species was a naturally early or late song initiator) compared to noise or silent controls. Our results support the theory that the dawn chorus is a communication network where individuals attend primarily to conspecific cues; heterospecific song appears to have minimal impacts on the chorus start times of the two early‐singing species we investigated.     

Nestling cuckoos,Cuculus canorus,exploit hosts with begging calls that mimic a brood

Nestling cuckoos, Cuculus canorus, eject host eggs or young from the nest and are then raised alone by the hosts. Using reed warblers, Acrocephalus scirpaceus, as hosts, we investigated how the single cuckoo chick can command the same provisioning rate as a whole brood of host young. Large size alone is not sufficient to stimulate adequate provisioning because single blackbird, Turdus merula, or song thrush, T. philomelos, chicks of the same mass as a cuckoo were fed at a lower rate. Our experiments show that the key stimulus is the cuckoo chick''s rapid begging call (''si, si, si, si ...''), which sounds remarkably like a whole brood of host chicks, and which it matched in calling rate. When single blackbird or song thrush chicks were accompanied by loudspeakers that broadcast either cuckoo begging calls or calls of a brood of reed warblers, the hosts increased their provisioning rate to that for a cuckoo chick. We suggest that the cuckoo needs vocal trickery to stimulate adequate care to compensate for the fact that it presents a visual stimulus of just one gape.     

Reproductive isolation inProkelisia planthoppers (Homoptera: Delphacidae): Acoustic differentiation and hybridization failure

Males and females of Prokelisia marginata (Van Duzee) and Prokelisia dolusWilson communicate through substrate-transmitted vibrations. The acoustic signals (attraction and courtship calls) of these planthoppers are effective in mate location, attraction, and mate choice. Attraction calls are structurally distinct for both species and differ in pulse type, pulse repetition rate, and pulse duration. Using playback of prerecorded calls, individuals discriminated between conspecific and heterospecific signals. Depending on the sex and species, response calls were produced three to eight times more frequently to conspecifics than to heterospecifics. However, acoustic signals alone did not explain reproductive isolation and hybridization failure in these two congeners. Some heterospecific pairs called, courted, and attempted to join genitalia, but no connections were successful and no progeny were produced. Thus, acoustic behavior is not a guaranteed premating isolating mechanism in no-choice situations. Other courtship behaviors and possibly morphological differences in genitalia also contributed to their isolation. Females displayed a variety of rejection behaviors to conspecific and heterospecific males, suggesting that sexual selection (female choice), in addition to species recognition, may be an important force in the evolution of the acoustic signals of planthoppers. Although signal structure was not dependent on wing form (planthoppers exhibit wing dimorphism), the age when females first began to call was related to wing form. Brachypterous (flightless) females of both species began calling early in adult life (day 2), whereas macropterous (migratory) females began calling later in adult life (day 6). This pattern is consistent with the oogenesis-flight syndrome, in which reproductive maturity is delayed until after migration occurs.     

Conspecific and heterospecific information use in bumblebees

Heterospecific social learning has been understudied in comparison to interactions between members of the same species. However, the learning mechanisms behind such information use can allow animals to be flexible in the cues that are used. This raises the question of whether conspecific cues are inherently more influential than cues provided by heterospecifics, or whether animals can simply use any cue that predicts fitness enhancing conditions, including those provided by heterospecifics. To determine how freely social information travels across species boundaries, we trained bumblebees (Bombus terrestris) to learn to use cues provided by conspecifics and heterospecific honey bees (Apis mellifera) to locate valuable floral resources. We found that heterospecific demonstrators did not differ from conspecifics in the extent to which they guided observers'' choices, whereas various types of inorganic visual cues were consistently less effective than conspecifics. This was also true in a transfer test where bees were confronted with a novel flower type. However, in the transfer test, conspecifics were slightly more effective than heterospecific demonstrators. We then repeated the experiment with entirely naïve bees that had never foraged alongside conspecifics before. In this case, heterospecific demonstrators were equally efficient as conspecifics both in the initial learning task and the transfer test. Our findings demonstrate that social learning is not a unique process limited to conspecifics and that through associative learning, interspecifically sourced information can be just as valuable as that provided by conspecific individuals. Furthermore the results of this study highlight potential implications for understanding competition within natural pollinator communities.     

Breeding success of common cuckoos Cuculus canorus parasitising four sympatric species of Acrocephalus warblers

We investigated the level of parasitism, egg mimicry and breeding success of cuckoos parasitising four sympatric species of Acrocephalus warblers in southern Moravia, Czech Republic. The parasitism rate was highest in the marsh warbler Acrocephalus palustris (44.8%) followed by great reed warbler A. arundinaceus (33.8%), sedge warbler A. schoenobaenus (26.5%) and reed warbler A. scirpaceus (11.6%). Although the cuckoo eggs showed a high level of mimicry the eggs of the marsh warbler this host species rejected 72% of the cuckoo eggs, resulting in a cuckoo breeding success of only 4.3%. Cuckoo eggs laid in great reed warbler and reed warbler nests showed a similar hatching success, but the cuckoo chicks survived better in great reed warbler nests, resulting in a breeding success of 30.4%, as compared to 16.4% in nests of the reed warbler. The relationship between the level of parasitism, host rejection of cuckoo eggs, cuckoo chick survival and breeding success is discussed for the four host species.     

Wild fledgling tits do not mob in response to conspecific or heterospecific mobbing calls

Mobbing, where birds harass a predator through a combination of vocalizations and stereotyped behaviours, is an effective anti-predator behaviour for many species. Mobbing may be particularly important for juveniles, as these individuals are often more vulnerable than adults. Although the component behaviours of mobbing are often considered to be un-learned, there are few confirmatory data, and the developmental trajectory of mobbing is unknown. In this study, we tested whether conspecific or heterospecific mobbing calls initiated mobbing behaviour in juvenile Blue Tits Cyanistes caeruleus. We located wild adult and recently fledged juvenile Blue Tits and presented them with playback recordings of adult conspecific (Blue Tit) and heterospecific (Great Tit Parus major) mobbing alarm calls. Although adult birds readily mob in response to these types of playbacks, juveniles did not exhibit characteristic mobbing behaviour. Some juveniles did, however, exhibit individual components of mobbing behaviour found in mobbing, despite not producing adult-like mobbing behaviour in response to either conspecific or heterospecific playback. These results suggest that, although birds might be capable of mobbing as juveniles, the associations between the non-vocal stereotyped mobbing behaviours and mobbing calls may be learned.     

Survival Benefit to Antipredator Behavior in the Amphipod Gammarus minus (Crustacea: Amphipoda) in Response to Injury-released Chemical Cues from Conspecifics and Heterospecifics

A wide range of aquatic taxa use environmental chemical cues for the assessment of predation risk. We examined whether Gammarus minus (Crustacea: Amphipoda) exhibit antipredator behavior in response to injury-released chemicals from conspecifics or heterospecifics (Crustacea: Isopoda). We then examined whether behavioral responses to these cues conferred survival benefits to the amphipods. In the first part of this study, we tested the behavioral response of G. minus to the following treatments: 1. water containing injury-released cues of conspecifics; 2. water containing injury-released cues of a sympatric isopod crustacean, Lirceus fontinalis; or 3. water containing no cues (control). Relative to the control, Gammarus responded to the conspecific cue by moving to the substratum and decreasing activity. In contrast, Gammarus responded to the heterospecific cue by moving up into the water column and increasing activity. In the second part of this study, we tested if the behavioral response to these cues confers a survival benefit to Gammarus when exposed to a predator. A green sunfish ( Lepomis cyanellus ) was retained behind a partition in the test tanks. Two minutes after the introduction of the chemical cues in the first test, the barrier was lifted and predation events recorded. Relative to the control, the time to the first attack increased for Gammarus exposed to conspecific cues and decreased for those exposed to heterospecific cues. These data indicate that Gammarus distinguish between chemical cues from conspecific and heterospecific crustaceans, and that the antipredator response to conspecific cues confers a fitness benefit (i.e. increased survival due to increased time to the first attack).