Diel vertical migration in zooplankton: rapid individual response to predators

While diel vertical migration in zooplankton has been shownrecently to be a predator avoidance behavior, the mechanismby which predators induce and maintain such behavior has beendebated. We report results of an in situ predator manipulationexperiment during which enclosed populations of the marine planktomccopepod Acaraa hudsonica rapidly changed their vertical distributionand diel migration behavior depending on presence or absenceof the planktivorous fish Casterosteus aculeatus These resultspoint unambiguously to phenotypic behavioral plasticity of individualplanktonic prey, not, as previously hypothesized, population-geneticlevel behavioral changes caused by selective fish predation,as the mechanism underlying changes in diel vertical migrationin this copepod.     

Predator-induced diet vertical migration in a planktonic copepod

Predator evasion is the most commonly hypothesized reason fordiel vertical migrations undertaken by a wide variety of planktonicorganisms in lakes and seas, yet direct evidence remains elusive.We tested the predation hypothesis by exposing enclosed populationsof a marine copepod Acartia hudsonica to caged or free-rangingindividuals of their natural predator, the planktivorous fishGasterosteus aculeatus. After little more than a week, adultcopepods changed their vertical distribution and migration behaviordepending on the presence or absence of predation. Only free-rangingfish induced vertical migration in the copepod population. Cagedfish had no effect, indicating that vertical migration was nota simple chemically mediated response of copepods to the predator.Rather, copepods seemed to react to the presence of predatorsby other means, perhaps visual or mechanical stimuli, and toexhibit a downward escape response which, because encounterswith visually orienting fish occur chiefly in the daytime, effectivelylimited the copepods' occurrence in the upper water column tothe night-time hours. Alternatively, because fish imposed heavymortality on copepods, it is possible that selective predationaltered the proportions of individuals with fixed, geneticallydetermined migration behaviors. We suggest experiments to distinguishthese alternatives.     

Escape behavior of Acartia hudsonica copepods during interactions with scyphomedusae

Calanoid copepods possess remarkable abilities to detect andescape from hydrodynamic disturbances, such as those createdby approaching predators. At the same time, a number of studiesin coastal ecosystems have suggested that gelatinous predators,including medusae in the Class Scyphozoa, exert top-down controlon copepod populations. Although prey escape behavior playsa critical role in predation models, we have relatively littleempirical data on how copepods respond to encounters with scyphomedusae.In this study, I used video to quantify encounter rates andescape behaviors of the copepod Acartia hudsonica during interactionswith two scyphomedusae, Aurelia aurita and Cyanea sp., in twoflow regimes. Escapes were complex, variable and effective.Fewer than 1% of encounters resulted in ingestion. Typically,A.hudsonica avoided contact by responding when predators remainedseveral body lengths (4 to 10 mm) distant and stringing togethermany escape jumps at submaximum velocities (33 to 59 mm s^–1).In addition, copepodite stages behaved passively—or failedto respond—following encounters with medusae more oftenthan did adults. Because escape behavior exhibited by A.hudsonicawas so variable, it is unlikely that medusae capture copepodsusing a single, quantifiable mechanism. A range of responseswithin populations and individuals may be the best strategyfor zooplankton faced with strong predation pressure from avariety of predators.     

Photophobic responses of calanoid copepods: possible adaptive value

A genual pattern of photophobic responses has been observedwhich differs for calanoid copepods from freshwater, estuarineand oceanic environments. Using a video-computer system formotion analysis, the photophobic responses of light and darkadapted calanoid copepods were compared. Dark-adapted copepodswere exposed to 600 ms flashes of dim blue light at 5 s intervalswhich simulated the flashes of biolumines-cent marine zooplankton.Light-adapted copepods were exposed to 600 ms intervals of darknessat 5 s intervals to simulate the shadows of organisms passingoverhead. Four species of coastal marine copepods (Acartia hudsonica,Centropages hamatus, Pseudocalanus minutus and Temora longicornis)all showed photophobic responses to both flashes and shadows.These responses may have adaptive value to the copepods sincethey live in an environment with predators that are bioluminescentat night and cast shadows on their prey during the day (e.g.ctenophores and cnidarian medusae). Two species of oceanic copepods(Euchaeta marina, Pleuromamma abdominalis) showed strong photophobicresponses to flashes but no response to shadows. This may correspondto the abundance of bioluminescent predators on copepods inthe oceanic environment (fish, ctenophores, siphonophores, etc.)and their lack of exposure to the shadows of predators, sinceboth these species are rarely found in the euphoric zone duringthe day. Two species of freshwater copepods (Diaptomus sanguineus,Epishwa massachusettsensis) showed no similar photophobic responseto flashes of light. This lack of startle response may relateto the lack of bioluminescence in the freshwater environment.Freshwater copepods showed a weak photophobic response to shadows.The adaptive value of this behavior is unclear, however, sincethe responses seem to be too weak to function for escape, andthe dominant predators large enough to cast shadows (fish) tendto approach their prey laterally. ¹Present address: Marine Science Institute, University of Texasat Austin, Port Aransas, TX 78373–1267, USA     

Salinity tolerance of the gemmules of Eunapius fragilis (Leidy) and the inhibition of germination by various salts

We report results of a field test of the predator avoidance hypothesis as an explanation of the adaptive significance of diel vertical migration in zooplankton. We determined the vertical distribution and diel migration of the planktonic copepod Acartia hudsonica, concurrently with the abundance of pelagic fish, transparency and thermal stratification of the water column, on six cruises over a one year period in a temperate marine lagoon (Jakles Lagoon, San Juan Island, Washington, USA). Striking seasonal variability was observed in all biological and environmental variables. Linear regressions of the strength of diel vertical migration in A. hudsonica on these environmental variables resulted in only one statistically significant relationship, that between copepod diel vertical migration and predator abundance. These results, together with those of previous studies, point to diel vertical migration as a widespread behavioral response of planktonic prey to the presence of their predators.     

The role of amino acids in the chemosensory swarming and feeding of marine copepods

Behavioural observations were made on two copepods, Eurytemoraherdmani and Acartia hudsonica, presented with 18 dissolvedL-amino acids the concentrations of which ranged from 10^–8M to 10^–2 M. The onset and duration of the swarming behaviourwere determined by the structure and concentration of a restrictednumber of molecules, which differed depending on the copepodspecies. Dicarboxylic amino acids were the most stimulatoryagent for E. herdmani, whereas A. hudsonica responded preferentiallyto aliphatic amino acids. Both the feeding and swarming of A.hudsonica could be induced by the same kind of molecules. Thethreshold sensitivity of the copepods appeared to be compatiblewith the natural concentrations of the dissolved free aminoacids measured at sea. These findings are discussed with referenceto the chemically mediated interactions between phytoplanktonand zooplankton.     

Phenotypic plasticity in oysters (Crassostrea virginica) mediated by chemical signals from predators and injured prey

Prey organisms reduce predation risk by altering their behavior, morphology, or life history. Avoiding or deterring predators often incurs costs, such as reductions in growth or fecundity. Prey minimize costs by limiting predator avoidance or deterrence to situations that pose significant risk of injury or death, requiring them to gather information regarding the relative threat potential predators pose. Chemical cues are often used for risk evaluation, and we investigated morphological responses of oysters (Crassostrea virginica) to chemical cues from injured conspecifics, from heterospecifics, and from predatory blue crabs (Callinectes sapidus) reared on different diets. Previous studies found newly settled oysters reacted to crab predators by growing heavier, stronger shells, but that adult oysters did not. We exposed oysters at two size classes (newly settled oyster spat and juveniles ~2.0 cm) to predation risk cue treatments including predator or injured prey exudates and to seawater controls. Since both of the size classes tested can be eaten by blue crabs, we hypothesized that both would react to crab exudates by producing heavier, stronger shells. Oyster spat grew heavier shells that required significantly more force to break, an effective measure against predatory crabs, when exposed to chemical exudates from blue crabs as compared to controls. When exposed to chemical cues from injured conspecifics or from injured clams (Mercenaria mercenaria), a sympatric bivalve, shell mass and force were intermediate between predator treatments and controls, indicating that oysters react to injured prey cues but not as strongly as to cues released by predators. Juvenile oysters of ~ 2.0 cm did not significantly alter their shell morphology in any of the treatments. Thus, newly settled oysters can differentiate between predatory threats and adjust their responses accordingly, with the strongest responses being to exudates released by predators, but oysters of 2.0 cm and larger do not react morphologically to predatory threats.     

Direct and indirect cues of predation risk influence behavior and reproduction of prey: a case for acarine interactions

Little is known about the flow of chemical information fromhigher to lower levels within the animal food chain. However,this information may determine the behavior and distributionof many animals (e.g., that of potential prey) when exposedto direct and indirect cues of predation risk. We used herbivorousspider mites, Tetranychus urticae Koch (Tetranychidae) as amodel to examine the foraging and oviposition decisions thatprey make when exposed to these cues. We conducted laboratorytests to determine if the previous presence of predators (directcues) on leaf discs or the presence of injured conspecifics(indirect cues) alters the distribution of adults and eggs ofT. urticae. When given a choice, after 24 h, fewer adults and eggswere found on leaf discs that had previously contained specialistspider mite predators, Phytoseiulus persimilis Athias-Henriot (Phytoseiidae),than on discs unexposed to predators. Also, more T. urticaeemigrated from predator-exposed discs than from unexposed discs orfrom those that had previously contained nonpredatory mites(Tyrophagus putrescentiae, Acaridae). Finally, fewer T. urticaeforaged and laid eggs on predator-exposed discs or on thosewith artificially damaged conspecifics (eggs or dead adults)than on discs with intact conspecifics. Tetranychus urticaeprobably recognizes infochemicals (kairomones) from its predatorsor cues from injured spider mites and consequently avoids feedingor ovipositing in areas exposed to these cues. Recognition and avoidanceof kairomones from specialist predators by this prey are likelyto be hereditary, but avoidance of injured conspecifics maybe an adaptation to avoid predators that are not inherentlyrecognized. We discuss the behavioral and ecological implicationsof our findings.     

Ocean acidification and responses to predators: can sensory redundancy reduce the apparent impacts of elevated CO2 on fish?

Carbon dioxide (CO₂) levels in the atmosphere and surface ocean are rising at an unprecedented rate due to sustained and accelerating anthropogenic CO₂ emissions. Previous studies have documented that exposure to elevated CO₂ causes impaired antipredator behavior by coral reef fish in response to chemical cues associated with predation. However, whether ocean acidification will impair visual recognition of common predators is currently unknown. This study examined whether sensory compensation in the presence of multiple sensory cues could reduce the impacts of ocean acidification on antipredator responses. When exposed to seawater enriched with levels of CO₂ predicted for the end of this century (880 μatm CO₂), prey fish completely lost their response to conspecific alarm cues. While the visual response to a predator was also affected by high CO_2, it was not entirely lost. Fish exposed to elevated CO_2, spent less time in shelter than current‐day controls and did not exhibit antipredator signaling behavior (bobbing) when multiple predator cues were present. They did, however, reduce feeding rate and activity levels to the same level as controls. The results suggest that the response of fish to visual cues may partially compensate for the lack of response to chemical cues. Fish subjected to elevated CO₂ levels_, and exposed to chemical and visual predation cues simultaneously, responded with the same intensity as controls exposed to visual cues alone. However, these responses were still less than control fish simultaneously exposed to chemical and visual predation cues. Consequently, visual cues improve antipredator behavior of CO₂ exposed fish, but do not fully compensate for the loss of response to chemical cues. The reduced ability to correctly respond to a predator will have ramifications for survival in encounters with predators in the field, which could have repercussions for population replenishment in acidified oceans.     

Effect of prey size on vulnerability of copepods to predation by the scyphomedusae Aurelia aurita and Cyanea sp.

A growing number of studies correlate changes in zooplanktonpopulations with abundance of medusae, but we cannot yet explainor predict the specific factors driving these interactions.This study demonstrates that the size of copepods has a significantinfluence on their vulnerability to predation by scyphomedusae.This finding is important because prey size, independent ofbehavior, has been neglected in theoretical models of predationby medusae. In experiments in a planktonkreisel, we used liveand heat-killed prey (Acartia hudsonica adults and copepodites)to separate the effects of copepod size and behavior on feedingrates by two medusae (Aurelia aurita and Cyanea sp.). Resultsrevealed that: differences in copepod size had a significantimpact on feeding rates, and thus small size can provide a refugefrom predation; behavior of adults diminished the liabilityassociated with larger size; and medusae with different morphologiesingested A.hudsonica at similar rates. Other experiments demonstratedthat medusae digested copepods at different rates based on preysize and predator species, findings that have implications forall future laboratory and field studies that assess feedingby scyphomedusae. Finally, this study illustrates how laboratorystudies serve as critical supplements to field observations.The effect of prey size on feeding rates can be confounded bydifferences in prey behavior, yet explains why small copepodswere typically ingested at relatively low rates by medusae.Size was clearly a dominant factor influencing copepod vulnerabilityacross scyphomedusan species, even those with very differentmorphologies. Future work should focus on the mechanisms ofsize selection, or the factors influencing contact and retentionrates.     

The prevalence and implications of copepod behavioral responses to oceanographic gradients and biological patchiness

Several species and developmental stages of calanoid copepodswere tested for responses to environmental cues in a laboratoryapparatus that mimicked conditions commonly associated withpatches of food in the ocean. All species responded to the presenceof phytoplankton by feeding. All species responded by increasingproportional residence time in one, but not both, of the treatmentsdefined by gradients of velocity or density. Most species increasedswimming speed and frequency of turning in response to the presenceof chemical exudates or gradients of velocity. Only one species,Eurytemora affinis, increased proportional time of residencein response to gradients in density of the water. Responsesof E. affinis to combined cues did not definitively demonstratea hierarchical use of different cues as previously observedfor Temora longicornis and Acartia tonsa. A simple foragingsimulation was developed to assess the applicability in thefield of the behavioral results observed in the laboratory.These simulations suggest that observed fine-scale behaviorscould lead to copepod aggregations observed in situ. The presentstudy demonstrates that behavioral response to cues associatedwith fine-scale oceanographic gradients and biological patchinessis functionally important and prevalent among copepods and likelyhas significant impacts on larger-scale distributional patterns.     

Species recognition by male swordtails via chemical cues

Species recognition can often play a key role in female matingpreferences. Far less is known about conspecific mate recognitionfrom the male perspective. In many closely related taxa, femalesexhibit few obvious visual differences and males might haveto attend to chemical cues in mate recognition, a possibilitythat has rarely been explored in vertebrates. Here, we examinemale species recognition via odor cues in the swordtail fish,Xiphophorus birchmanni. In dichotomous choice experiments wefirst tested whether males respond to female odor cues. We foundthat males were attracted to conspecific female odor and thoseof a related allopatric congener, Xiphophorus malinche, overa water control. Males did not, however, respond to the femaleodor of the more distantly related sympatric platyfish, Xiphophorusvariatus. We then gave male X. birchmanni the choice betweenconspecific and heterospecific female stimuli. Males, in thisscenario, significantly preferred the conspecific odor whenthe alternative was platyfish. However, when offered odor cuesof X. malinche, male X. birchmanni actually preferred the heterospecificfemale cue. The complex array of preferences reported here,previously documented only in females, underscores the needto consider the behavior of both sexes in dictating actual matingoutcomes.     

Implications of the feeding current structure of Euchaeta rimana, a carnivorous pelagic copepod, on the spatial orientation of their prey

Many marine planktonic organisms create water currents to entrainand capture food items. Rheotactic prey entrained within thesefeeding currents often exibit escape reactions. If the directionof escape is away from the feeding current, the prey may successfullydeter predation. If the escape is towards the center of thefeeding current, the prey will be re-entrained towards its predatorand remain at risk of predation. The direction of escape isdependent on (i) the ability of the prey to escape in a directiondifferent than its pre-escape orientation and (ii) the orientationcaused by the interaction of the prey's body with the movingfluid. In this study, the change in orientation of Acartia hudsonicanauplii as a result of entrainment within the feeding currentof Euchaeta rimana, a planktonic predatory copepod, was examined,When escaping in still water, A.hudsonica nauplii were ableto vary their pre-escape direction by only 10. This allowsonly a limited ability to escape in a direction different thantheir pre-escape orientation. Analyses of the feeding currentof E.rimana show the flow speed to be most rapid in the centralregion with an exponential decrease in speed distally. In contrast,flow vorticity is minimal in the center of the feeding currentand maximal at 1.75 mm along the antennae. As a result, thedegree of rotation of the prey towards the center of the feedingcurrent shows a strong dependency on the prey's location withinthe feeding current. The feeding current of E.rimana rotatedthe prey 14 when near the center of the flow field and up to160 when located more distal in the feeding current Since theprey's escape abilities cannot compensate for the rotation dueto the flow, this mechanism will maintain the escaping preywithin the feeding current of their predator. Therefore, thefeeding current facilitates predatory copepods in capturingprey by (i) increasing the amount of water which passes overtheir sensors and through their feeding appendages and (ii)controlling the spatial orientation of their prey prior to escape.     

Prey perception of predation risk: volatile chemical cues mediate non-consumptive effects of a predator on a herbivorous insect

Predators can affect prey in two ways—by reducing their density (consumptive effects) or by changing their behavior, physiology or other phenotypic traits (non-consumptive effects). Understanding the cues and sensory modalities prey use to detect predators is critical for predicting the strength of non-consumptive effects and the outcome of predator–prey encounters. While predator-associated cues have been well studied in aquatic systems, less is known about how terrestrial prey, particularly insect larvae, detect their predators. We evaluated how Colorado potato beetle, Leptinotarsa decemlineata, larvae perceive predation risk by isolating cues from its stink bug predator, the spined soldier bug, Podisus maculiventris. When exposed to male “risk” predators that were surgically manipulated so they could hunt but not kill, beetles reduced feeding 29 % compared to controls. Exposure to risk females caused an intermediate response. Beetles ate 24 % less on leaves pre-exposed to predators compared to leaves never exposed to predators, indicating that tactile and visual cues are not required for the prey’s response. Volatile odor cues from predators reduced beetle feeding by 10 % overall, although male predators caused a stronger reduction than females. Finally, visual cues from the predator had a weak effect on beetle feeding. Because multiple cues appear to be involved in prey perception of risk, and because male and female predators have differential effects, beetle larvae likely experience tremendous variation in the information about risk from their local environment.     

Tadpoles of the bronze frog (Rana temporalis) assess predation risk before evoking antipredator defense behavior

Predation threat-associated behavioral response was studied in Rana temporalis tadpoles to discover the importance of predators’ visual and chemical cues (kairomones and diet-derived metabolites of consumed prey) in evoking antipredator behavior. The caged predators (dragonfly larvae) fed on prey tadpoles or insects (Notonecta spp.) and water conditioned with the predators provided the threat stimuli to the tadpole prey. The predators’ visual cues were ineffective in evoking antipredator behaviors in the tadpole prey. However, exposure to caged tadpole-fed predators or water conditioned with tadpole-fed predators elicited predator avoidance behavior in the tadpoles; they stayed away from the predators, significantly reduced swimming activity (swimming time and distance traveled), and increased burst speed. Interestingly, exposure to water conditioned with starved predators did not elicit any antipredator behavior in the prey. Further, the antipredator responses of predator-experienced tadpoles were significantly greater than those exhibited by predator-na?ve tadpoles. The study shows that R. temporalis tadpoles assess predation threat based exclusively on chemical cues emanating from the predators’ dietary metabolites and that the inclusion of conspecific prey items in the diet of the predators is perceived as a threat. The study also shows that antipredator behavior in these tadpoles is innate and is enhanced during subsequent encounters with the predators.     

Rodent foraging is affected by indirect, but not by direct, cues of predation risk

We used foraging trays to determine whether oldfield mice, Peromyscuspolionotus, altered foraging in response to direct cues of predationrisk (urine of native and nonnative predators) and indirectcues of predation risk (foraging microhabitat, precipitation,and moon illumination). The proportion of seeds remaining ineach tray (a measure of the giving-up density [GUD]) was usedto measure risk perceived by mice. Mice did not alter theirGUD when presented with cues of native predators (bobcats, Lynxrufus, and red foxes, Vulpes vulpes), recently introduced predators(coyotes, Canis latrans), nonnative predators (ocelots, Leoparduspardalis), a native herbivore (white-tailed deer, Odocoileusvirginianus), or a water control. Rather, GUD was related tomicrohabitat: rodents removed more seeds from foraging trayssheltered beneath vegetative cover compared with exposed traysoutside of cover. Rodents also removed more seeds during nightswith precipitation and when moon illumination was low. Our resultssuggest that P. polionotus used indirect cues rather than directcues to assess risk of vertebrate predation. Indirect cues maybe more reliable than are direct scent cues for estimating riskfrom multiple vertebrate predators that present the most riskin open environments.     

Latitudinal differentiation in the effects of the toxic dinoflagellate Alexandrium spp. on the feeding and reproduction of populations of the copepod Acartia hudsonica

Blooms of the dinoflagellate Alexandrium spp. increase in their frequency, toxicity and historical presence with increasing latitude from New Jersey (USA) to the Gaspé peninsula (Canada). Biogeographic variation in these blooms results in differential exposure of geographically separate copepod populations to toxic Alexandrium. We hypothesize that the ability of copepods to feed and reproduce on toxic Alexandrium should be higher in copepods from regions that are frequently exposed to toxic Alexandrium blooms. We tested this hypothesis with factorial common environment experiments in which female adults of the copepod Acartia hudsonica from five separate populations ranging from New Jersey to New Brunswick were fed toxic and non-toxic strains of Alexandrium, and the non-toxic flagellate Tetraselmis sp. Consistent with the hypothesis, when fed toxic Alexandrium we observed significantly higher ingestion and egg production rates in the copepods historically exposed to toxic Alexandrium blooms relative to copepods from regions in which Alexandrium is rare or absent. Such differences among copepod populations were not observed when copepods were fed non-toxic Alexandrium or Tetraselmis sp. These results were also supported by assays in which copepods from populations both historically exposed and naïve to toxic Alexandrium blooms were fed mixtures of toxic Alexandrium and Tetraselmis sp. Two-week long experiments demonstrated that when copepods from populations naïve to toxic Alexandrium were fed a toxic strain of Alexandrium they failed to acclimate, such that their ingestion rates remained low throughout the entire two-week period. The differences observed among populations suggest that local adaptation of populations of A. hudsonica from Massachusetts (USA) to New Brunswick (Canada) has occurred, such that some populations are resistant to toxic Alexandrium.     

Inducible reproductive plasticity of the guppy Poecilia reticulata in response to predation cues

Predation has long been described as one of the major driving forces in evolution. Guppies (Poecilia reticulata) from natural populations exposed to different predation pressures, were found to have different life history traits. Reproductive plasticity in response to direct predation cues has mainly been reported for invertebrates. The goals of the present study were to determine whether exposure to predation cues would induce reproductive phenotypic plasticity in female guppies and to determine whether the effective cues are visual, chemical, or a combination of both. In our first experiment, female guppies exposed to predation cues of the african cichlids Aulonocara nyassae increased their reproductive output by almost two fold, having larger brood-sizes and shorter brood-interval at the first spawn. This effect disappeared in the second spawn in the absence of predators. In the second experiment we found that exposure to the predators induced an increase in the brood-size regardless of whether the cue was: only visual, only chemical, visual and chemical or visual, chemical and tactile. The impacts of these cues were equally powerful on the tested variables and they did not have any cumulative effect. Similar to the results of the first experiment, this effect disappeared in the second spawn, in the absence of predation cues. The present study demonstrates a direct immediate and reversible effect of predation cues on guppy reproduction.     

The Scent of Danger: the Impact of Predator Chemical Cues on Emergence from Refuge and Willingness to Autotomize Limbs in the House Cricket (<Emphasis Type="Italic">Acheta domesticus</Emphasis>)

Prey can accurately assess predation risk via the detection of chemical cues and take appropriate measures to survive encounters with predators. Research on the chemical ecology of terrestrial invertebrate predator-prey interactions has repeatedly found that direct chemical cues can alter prey organisms’ antipredator behavior. However, much of this research has focused on the chemical mediation of avoidance and immobility by cues from lycosid spiders neglecting other prominent invertebrate predators and behavior such as autotomy. In our study, house crickets (Acheta domesticus) were exposed to cues from cricket-fed orange-footed centipedes (Cormocephalus aurantiipes), red-back spiders (Latrodectus hasselti), an odorous (cologne) control, and a non-odorous control to determine whether direct chemical cues had any influence on two types of anti-predatory behavior: the willingness (latency) to emerge from a refuge and to autotomize limbs. Exposure to C. aurantiipes cues resulted in a significantly slower emergence from a refuge, but exposure to L. hasselti cues did not. Direct chemical cues had no influence on initial autotomy, but exposure to L. hasselti cues did significantly decrease the latency to autotomize a second limb. That cues from L. hasselti had an influence on a second autotomy, but not initial autotomy may be because crickets that undergo autotomy for a second time may perceive themselves to be already at a higher risk of predation as they were already missing a limb. Variation in responses to cues from different predators demonstrates a need to examine the influence of chemical cues from a wider variety of invertebrate predators on anti-predator behavior.     

Assessment of local predation risk: the role of subthreshold concentrations of chemical alarm cues

The ability to accurately assess local predation risk is criticalto prey individuals, as it allows them to maximize threat-sensitivetrade-offs between predator avoidance and other fitness relatedactivities. A wide range of taxonomically diverse prey (includingmany freshwater fishes) relies on chemical alarm cues (alarmpheromones) as their primary information source for local riskassessment. However, the value of chemical alarm cues has beenquestioned due to the availability of additional sensory inputs(i.e., visual cues) and the lack of an overt antipredator responseunder conditions of low perceived risk. In this paper, we testthe hypothesis that chemical alarm cues at concentrations belowthe point at which they elicit an overt behavioral responsefunction to increase vigilance towards other sensory modalities(i.e., visual alarm cues). Shoals of glowlight tetras (Hemigrammuserythrozonus) exposed to the subthreshold concentration of hypoxanthine-3-N-oxide(the putative Ostariophysan alarm pheromone) did not exhibitan overt antipredator response in the absence of secondary visualcues (not different than the distilled water control). However,when exposed to the sight of a visually alarmed conspecific,they significantly increased the intensity of their antipredatorresponse (not different from shoals exposed to the suprathresholdalarm cue). This study demonstrates that prey may benefit fromresponding to low concentration alarm cues by increasing vigilancetowards secondary cues during local risk assessment, even inthe absence of an overt behavioral response. By increasing vigilancetowards secondary risk assessment cues in the presence of alow risk chemical cue, individuals are likely able to maximizethe threat-sensitive trade-offs between predator avoidance andother fitness related activities.