The effect of predator and prey density on the induced defence of a ciliate

1. The level of antipredator defence should be proportional to the actual attack probability to minimize the cost of defence and maximize the net benefit.
2. The hypothesis that the induced antipredator morphology of Euplotes octocarinatus is a graded response to the actual risk of predation by Stylonychia mytilus was tested by manipulating the density of both prey and predator populations.
3. The magnitude of the response was graded according to both predator and prey density. A dense prey population may be protective since a prey is more exposed to a predator's attack as a solitary individual.
4. The results suggest that Euplotes is able to 'estimate' the real risk of predation and respond appropriately, without mobilizing more resources than needed.
5. Separation of the prey and predator with a nylon net revealed that the response was not induced by a water-transmitted factor but that direct cell-to-cell contacts were important. This finding departs from those of other studies.     

Threat-sensitive anti-predator defence in precocial wader,the northern lapwing <Emphasis Type="Italic">Vanellus vanellus</Emphasis>

Birds exhibit various forms of anti-predator behaviours to avoid reproductive failure, with mobbing—observation, approach and usually harassment of a predator—being one of the most commonly observed. Here, we investigate patterns of temporal variation in the mobbing response exhibited by a precocial species, the northern lapwing (Vanellus vanellus). We test whether brood age and self-reliance, or the perceived risk posed by various predators, affect mobbing response of lapwings. We quantified aggressive interactions between lapwings and their natural avian predators and used generalized additive models to test how timing and predator species identity are related to the mobbing response of lapwings. Lapwings diversified mobbing response within the breeding season and depending on predator species. Raven Corvus corax, hooded crow Corvus cornix and harriers evoked the strongest response, while common buzzard Buteo buteo, white stork Ciconia ciconia, black-headed gull Chroicocephalus ridibundus and rook Corvus frugilegus were less frequently attacked. Lapwings increased their mobbing response against raven, common buzzard, white stork and rook throughout the breeding season, while defence against hooded crow, harriers and black-headed gull did not exhibit clear temporal patterns. Mobbing behaviour of lapwings apparently constitutes a flexible anti-predator strategy. The anti-predator response depends on predator species, which may suggest that lapwings distinguish between predator types and match mobbing response to the perceived hazard at different stages of the breeding cycle. We conclude that a single species may exhibit various patterns of temporal variation in anti-predator defence, which may correspond with various hypotheses derived from parental investment theory.     

Neural correlates of threat perception: neural equivalence of conspecific and heterospecific mobbing calls is learned

Songbird auditory areas (i.e., CMM and NCM) are preferentially activated to playback of conspecific vocalizations relative to heterospecific and arbitrary noise. Here, we asked if the neural response to auditory stimulation is not simply preferential for conspecific vocalizations but also for the information conveyed by the vocalization. Black-capped chickadees use their chick-a-dee mobbing call to recruit conspecifics and other avian species to mob perched predators. Mobbing calls produced in response to smaller, higher-threat predators contain more "D" notes compared to those produced in response to larger, lower-threat predators and thus convey the degree of threat of predators. We specifically asked whether the neural response varies with the degree of threat conveyed by the mobbing calls of chickadees and whether the neural response is the same for actual predator calls that correspond to the degree of threat of the chickadee mobbing calls. Our results demonstrate that, as degree of threat increases in conspecific chickadee mobbing calls, there is a corresponding increase in immediate early gene (IEG) expression in telencephalic auditory areas. We also demonstrate that as the degree of threat increases for the heterospecific predator, there is a corresponding increase in IEG expression in the auditory areas. Furthermore, there was no significant difference in the amount IEG expression between conspecific mobbing calls or heterospecific predator calls that were the same degree of threat. In a second experiment, using hand-reared chickadees without predator experience, we found more IEG expression in response to mobbing calls than corresponding predator calls, indicating that degree of threat is learned. Our results demonstrate that degree of threat corresponds to neural activity in the auditory areas and that threat can be conveyed by different species signals and that these signals must be learned.     

Prey attack and predators defend: counterattacking prey trigger parental care in predators

That predators attack and prey defend is an oversimplified view. When size changes during development, large prey may be invulnerable to predators, and small juvenile predators vulnerable to attack by prey. This in turn may trigger a defensive response in adult predators to protect their offspring. Indeed, when sizes overlap, one may wonder "who is the predator and who is the prey"! Experiments with "predatory" mites and thrips "prey" showed that young, vulnerable prey counterattack by killing young predators and adult predators respond by protective parental care, killing young prey that attack their offspring. Thus, young individuals form the Achilles' heel of prey and predators alike, creating a cascade of predator attack, prey counterattack and predator defence. Therefore, size structure and relatedness induce multiple ecological role reversals.     

Predator-induced morphological shift in the pea aphid

Aphids exhibit a polymorphism whereby individual aphids are either winged or unwinged. The winged dispersal morph is mainly responsible for the colonization of new plants and, in many species, is produced in response to adverse environmental conditions. Aphids are attacked by a wide range of specialized predators and predation has been shown to strongly influence the growth and persistence of aphid colonies. In two experiments, we reared two clones of pea aphid (Acyrthosiphon pisum) in the presence and absence of predatory ladybirds (Coccinella septempunctata or Adalia bipunctata). In both experiments, the presence of a predator enhanced the proportion of winged morphs among the offspring produced by the aphids. The aphid clones differed in their reaction to the presence of a ladybird, suggesting the presence of genetic variation for this trait. A treatment that simulated disturbance caused by predators did not enhance winged offspring production. The experiments indicate that aphids respond to the presence of a predator by producing the dispersal morph which can escape by flight to colonize other plants. In contrast to previous examples of predator-induced defence this shift in prey morphology does not lead to better protection against predator attack, but enables aphids to leave plants when mortality risks are high.     

Predator diet breadth influences the relative importance of bottom-up and top-down control of prey biomass and diversity

We investigated the effects of predator diet breadth on the relative importance of bottom-up and top-down control of prey assemblages, using microbial food webs containing bacteria, bacterivorous protists and rotifers, and two different top predators. The experiment used a factorial design that independently manipulated productivity and the presence or absence of two top predators with different diet breadths. Predators included a "specialist" predatory ciliate Euplotes aediculatus, which was restricted to feeding on small prey, and a "generalist" predatory ciliate Stentor coeruleus, which could feed on the entire range of prey sizes. Both total prey biomass and prey diversity increased with productivity in the predator-free control and specialist predator treatments, a pattern consistent with bottom-up control, but both remained unchanged by productivity in the generalist predator treatment, a pattern consistent with top-down control. Linear food chain models adequately described responses in the generalist predator treatment, whereas food web models incorporating edible and inedible prey (which can coexist in the absence of predators) adequately described responses in the specialist predator treatment. These results suggest that predator diet breadth can play an important role in modulating the relative strength of bottom-up and top-down forces in ecological communities.     

Divergent phenotypic responses to predators and cyanobacteria in Daphnia lumholtzi

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Non-lethal presence of predators modifies morphology and movement rates in <Emphasis Type="Italic">Euplotes</Emphasis>

Many species are able to modify aspects of their behaviour and morphology in the presence of predators. The aim of this study was to investigate the relationship between the expression of morphological and behavioural defences according to the framework proposed by DeWitt et al (1999). Experiments were carried out using hypotrich ciliates of the genus Euplotes as prey and turbellarians of the genus Stenostomum as predators. The smaller species Euplotes octocarinatus showed a greater proportional increase in width, a reduction in foraging movement rates and an increase in maximum movement rates following exposure to predator cues. The larger Euplotes aediculatus induced lesser changes in width, similar reductions in movement during foraging and no change in maximum speed following predator exposure. These results provide evidence of a cospecialised relationship between morphological and behavioural defences. Despite substantial differences in the absence of predators, movement rates and lateral body width were similar in both species following predator exposure. The observed changes may be considered adaptive, gape limited flatworm predators are unable to ingest large Euplotes and a reduction in movement rates during foraging reduces predator encounter rates, while an increase in maximal movement rates increases chances of predator evasion. Handling editor: S. I. Dodson     

Predator recognition and differential behavioural responses of adult wood warblers <Emphasis Type="Italic">Phylloscopus sibilatrix</Emphasis>

Birds often engage in nest defence against predators to improve breeding success, but defence efficiency requires the capability to assess the threat level posed by potential predators. For species with low breeding-site tenacity, which may encounter varying occurrence and density of predators in different areas, threat recognition could be compromised due to naivety, and so predator recognition may focus on broad key features to diminish the risk of misidentification. We experimentally tested this hypothesis by recording behavioural reactions of the nomadic wood warbler Phylloscopus sibilatrix to objects reflecting various levels of threat: least weasel and Eurasian jay taxidermy mounts, an inanimate object and an empty display mount. To assess actual nest predators, we used remote cameras to record predation events at wood warbler nests. As in other studies in Western Europe, Eurasian jay was found to be the main nest predator, with occasional predation by least weasel. The reaction of adult warblers to the models was generally to remain silent and on nests during the incubation stage presumably due to the need to maintain efficient nest camouflage and concealment. During the nestling stage, behavioural responses of adult warblers, calling and suspended feeding of young, showed the strongest effects from the jay taxidermy mount, moderate to the weasel and weakest to the inanimate object and empty mount. As the reaction of wood warblers reflected the degree of genuine threat posed by the predators depicted by the models, we conclude that predator recognition may be present in this species.     

Non‐pathogenic aquatic bacteria activate the immune system and increase predation risk in damselfly larvae

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THE EVOLUTIONARY ECOLOGY OF AN ANTIPREDATOR REACTION NORM: DAPHNIA PULEX AND CHAOBORUS AMERICANUS

Ponds containing the parthenogenetic zooplankter Daphnia pulex with and without chaoborid predators were sampled over the course of a season. A significant (P < 0.05) Spearman rank correlation was found between predator density and the expression of an antipredator defense (neckteeth) by the Daphnia. The reaction norms (percent induction of a single genotype versus predator density) of clones isolated from predator-free and predator-rich habitats were determined in a laboratory setting. There was a statistically significant different response among the six clones tested (P < 0.05). Clones isolated from chaoborid ponds showed significantly greater sensitivity to the presence of predator than clones from predator-free ponds (P < 0.05). In the laboratory, food levels under which prey were cultured affected induction of the antipredator response. Highest induction was found at the lowest food level used.     

Effects of Sibship and the Presence of Multiple Predators on the Behavior of Green Frog (Rana clamitans) Tadpoles

In nature, prey are exposed to multiple predators simultaneously. We examined the effects of the cues of two potential predators, mosquitofish and odonate larvae, individually and in combination on the behavior of green frog (Rana clamitans) tadpoles. In addition to examining the behavioral response of green frog tadpoles to multiple predators, we examined variation in behavior among tadpoles from different egg masses (i.e. different sibships). Sibships differed in activity level and there was a significant predator cue by sibship interaction. Two sibships were relatively more active in the control and odonate predator cue treatments but showed reduced activity in treatments containing mosquitofish cues, whereas the remaining sibships showed consistently low levels of activity in all predator cue treatments, including the control. The use of the vegetated side of the aquarium did not differ between tadpoles exposed to the different predator cues. Sibship had no effect on tadpoles’ use of the vegetated side of the aquarium, and there was no interaction between sibship and predator cue. Our results suggest that green frogs did not respond to simultaneous exposure to multiple predator cues any differently than they did to exposure to individual predator cues. More importantly, our results suggest variation, possibly genetically based, in behavioral responses of tadpoles to predators, and thus selection on these behaviors is possible. Of particular interest is that there was variation in behavioral responses to a non‐native predator (Gambusia affinis), suggesting an evolutionary response to an invasive predator is possible.     

Predator-mediated plasticity in morphology, life history, and behavior of Daphnia: the uncoupling of responses

We studied the way 12 traits responded to fish kairomones in a set of 16 Daphnia magna clones derived from four different habitats-two where daphnids co-occur with fish and two without fish. These clones differed widely in their response to predator kairomones, with none of the clones showing a significant response in all traits and all clones showing a response for at least one trait. Most of the clones showed a significant response in one to four traits, with no evidence for an association between different traits. Clones from fish habitats were slightly more responsive to the presence of fish kairomones than clones from fishless locations. We conclude that most clones show an induced response to the presence of their predators (fish) but that there is a large genetic variability with respect to the traits for which clones show a response. Our results indicate that the major distinction is not between inducible and noninducible genotypes but rather that the genotypes differ in the combination of traits for which they show inducible responses.     

Increased nest defence of upland‐nesting ducks in response to experimentally reduced risk of nest predation

Parent birds should take greater risks defending nests that have a higher probability of success. Given high rates of mammalian nest predation, therefore, parents should risk more for nests in areas with a lower risk of mammalian predation. We tested this hypothesis using nest defence data from over 1300 nests of six species of dabbling ducks studied in an area where predation risk had been reduced through removal of mammalian predators. When predator removal reduced nest predation, the ducks increased risk taking as predicted. Also as predicted, risk taking varied inversely with body size, an index of annual survival, among species. For ducks to vary nest defence in response to variation in predation risk they must be able to assess the risk of nest predation. Because ducks modified nest defence in the breeding season immediately following predator removal, ducks may be able to assess predator abundance indirectly (e.g. by UV reflection from urine) rather than by seeing or interacting directly with the predators.     

Express yourself: bold individuals induce enhanced morphological defences

Organisms display an impressive array of defence strategies in nature. Inducible defences (changes in morphology and/or behaviour within a prey''s lifetime) allow prey to decrease vulnerability to predators and avoid unnecessary costs of expression. Many studies report considerable interindividual variation in the degree to which inducible defences are expressed, yet what underlies this variation is poorly understood. Here, we show that individuals differing in a key personality trait also differ in the magnitude of morphological defence expression. Crucian carp showing risky behaviours (bold individuals) expressed a significantly greater morphological defence response when exposed to a natural enemy when compared with shy individuals. Furthermore, we show that fish of different personality types differ in their behavioural plasticity, with shy fish exhibiting greater absolute plasticity than bold fish. Our data suggest that individuals with bold personalities may be able to compensate for their risk-prone behavioural type by expressing enhanced morphological defences.     

Induction of defensive morphological changes in ciliates

Freshwater ciliates of the genusEuplotes change their morphology in response to the presence of predators. The morphological transformations limit the ability of the predators to ingest the ciliates. Induction of defensive morphology by the predatorsStenostomum sphagnetorum (Turbellaria),Lembadion bullinum (Ciliata),L. magnum, andAmoeba proteus (Rhizopoda) was studied inEuplotes aediculatus andE. octocarinatus. The results suggest the possibility of natural occurrence of predator-induced defensive morphology inEuplotes. A density of 1 predator ml⁻¹ was sufficient to induce significant changes in the morphology ofEuplotes. L. magnum was found at natural population densities of 5 individuals ml⁻¹. Transformations can take place within 2–4 h. Morphological changes are induced by signal substances released from the predators; direct contact between prey and predators is not necessary. The extent of transformation depends on the concentration of the signal substance. The size frequency distribution of the populations only had one peak that was related to predator density. All individuals of a population ofEuplotes changed their morphology according to the predator abundance. This may reduce energy costs for the prey by prevention of unnecessary morphological changes.Amoeba proteus induces morphological changes inE. octocarinatus, but not inE. aediculatus.     

Disease in group-defending prey can benefit predators

Infectious diseases have the capacity not only to influence the host population but also to interacting species like predators. In particular, they can reduce host densities, which can have knock-on effects on predators. Here, we consider how an infectious disease in the prey affects the predator–prey relationship where the prey exhibit some kind of group defence against the predator (using a Holling type IV functional response). We find that the disease can reduce prey densities to levels where the group defence is weaker. This weakened group defence allows predators to survive in many situations where they could not without the disease.     

Tasting the difference: do multiple defence chemicals interact in Müllerian mimicry?

Müllerian mimicry, where two unpalatable species share a warning pattern, is classically believed to be a form of mutualism, where the species involved share the cost of predator education. The evolutionary dynamics of Müllerian mimicry have recently become a controversial subject, after mathematical models have shown that if minor alterations are made to assumptions about the way in which predators learn and forget about unpalatable prey, this textbook case of mutualism may not be mutualistic at all. An underlying assumption of these models is that Müllerian mimics possess the same defence chemical. However, some Müllerian mimics are known to possess different defence chemicals. Using domestic chicks as predators and coloured crumbs flavoured with either the same or different unpalatable chemicals as prey, we provide evidence that two defence chemicals can interact to enhance predator learning and memory. This indicates that Müllerian mimics that possess different defence chemicals are better protected than those that share a single defence chemical. These data provide insight into how multiple defence chemicals are perceived by birds,and how they influence the way birds learn and remember warningly coloured prey. They highlight the importance of considering how different toxins in mimicry rings can interact in the evolution and maintenance of Müllerian mimicry and could help to explain the remarkable variation in chemical defences found within and between species.     

Effects of predator-specific defence on biodiversity and community complexity in two-trophic-level communities

Summary Antipredator strategies employed by prey may be specific (effective against only one type of predator) or non-specific (effective against all predators). To examine the effects of the specificity of antipredator behaviour on biodiversity and community complexity, we analyse mathematical models including both evolutionary and population dynamics of a system including multiple prey species and multiple predator species. The models assume that all predator species change in their prey choice and all prey species have evolutionary change in their antipredator effort in evolution. The traits of each species change in an adaptive manner, whose rate is proportional to the slope of their fitness function. We calculate community complexity, resource-overlap between predators, an index of biodiversity and other properties of the coevolutionarily stable community for two cases: (1) all prey species have non-specific antipredator behaviour and (2) all prey species have predator-specific defence. Predator-specificity in defence increases community complexity, resource-overlap between predators, the total abundance of predators and the ratio of predator to prey abundance. Specific defence also decreases the number of isolated subwebs within the entire foodweb.     

Ontogenetic change in effectiveness of chemical defence against different predators in Oxycarenus true bugs

Many prey species change their antipredator defence during ontogeny, which may be connected to different potential predators over the life cycle of the prey. To test this hypothesis, we compared reactions of two predator taxa – spiders and birds – to larvae and adults of two invasive true bug species, Oxycarenus hyalinipennis and Oxycarenus lavaterae (Heteroptera: Oxycarenidae) with life-stage-specific chemical defence mechanisms. The reactions to larvae and adults of both true bug species strikingly differed between the two predator taxa. The spiders were deterred by the defences of adult bugs, but the larval defences were ineffective against them. By contrast, birds attacked the larvae considerably less often than the adult bugs. The results indicate a predator-specific ontogenetic change in defence effectiveness of both Oxycarenus species. The change in defence is likely linked to the life-stage-specific composition of secretions in both species: whereas secretions of larvae are dominated by unsaturated aldehydes, secretions of adults are rich in terpenoids, which probably serve dual function of defensive chemicals and pheromones. Our results highlight the variation in defence between different life stages and the importance of testing responses of different types of predators.