Does attraction to conspecifics explain the patch‐size effect? An experimental test

Recent theory suggests that attraction to conspecifics during habitat selection can be one potential, yet untested, mechanism for animal sensitivity to habitat fragmentation. The least flycatcher Empidonax minimus , a highly territorial migratory bird, has previously been shown to be attracted to conspecifics and sensitive to patch size by avoiding small patches of riparian forest in Montana, USA. I used a large-scale field experiment in this region to test the conspecific attraction hypothesis for explaining sensitivity to patch size, and I supplemented this experiment by estimating whether vegetation structure, nest predation, or nest parasitism rates could better explain patterns of sensitivity to patch size. Vegetation structure did not vary consistently with patch size, based on a random sample of patches across 150  km of the Madison and Missouri Rivers, Montana. Nest predation and parasitism rates by brown-headed cowbirds Molothrus ater also did not vary with patch size during the experiment. However, when conspecific cues were simulated across a gradient of patch sizes, flycatchers settled in all patches – and their sensitivity to patch size vanished – providing strong support for the conspecific attraction hypothesis. These results provide the first experimental evidence that attraction to conspecifics can indeed help explain area sensitivity in nature and highlight how understanding the role of animal behavior in heterogeneous landscapes can aid in interpreting pressing conservation issues.     

The role of biotic and abiotic cues in stimulating aggregation by larval cane toads (Rhinella marina)

Tadpoles of the cane toad (Rhinella marina) form dense aggregations in the field, but the proximate cues eliciting this behavior are not well understood. We sampled water‐bodies in the Northern Territory of Australia, finding that the density of cane toad tadpoles increased with increasing temperature. Furthermore, we conducted laboratory experiments to explore the roles of biotic factors (attraction to conspecifics; chemical cues from an injured conspecific; food) and spatially heterogeneous abiotic factors (light levels, water depth, physical structure) to identify the cues that induce tadpole aggregation. Annulus and binary choice trials demonstrated weak but significant attraction between conspecifics. Tadpoles decreased swimming speeds, but did not increase grouping in response to cues from an injured conspecific. Larvae aggregated in response to abiotic cues (high levels of illumination and proximity to physical structures) and were strongly attracted to feeding conspecifics. Overall, aggregation by cane toad tadpoles is likely driven by weak social attraction coupled with a shared preference for specific abiotic features, creating loose aggregations that are then reinforced by movement toward feeding conspecifics.     

Spatial patterns of the frog Oophaga pumilio in a plantation system are consistent with conspecific attraction

The conspecific attraction hypothesis predicts that individuals are attracted to conspecifics because conspecifics may be cues to quality habitat and/or colonists may benefit from living in aggregations. Poison frogs (Dendrobatidae) are aposematic, territorial, and visually oriented—three characteristics which make dendrobatids an appropriate model to test for conspecific attraction. In this study, we tested this hypothesis using an extensive mark‐recapture dataset of the strawberry poison frog (Oophaga pumilio) from La Selva Biological Station, Costa Rica. Data were collected from replicate populations in a relatively homogenous Theobroma cacao plantation, which provided a unique opportunity to test how conspecifics influence the spatial ecology of migrants in a controlled habitat with homogenous structure. We predicted that (1) individuals entering a population would aggregate with resident adults, (2) migrants would share sites with residents at a greater frequency than expected by chance, and (3) migrant home ranges would have shorter nearest‐neighbor distances (NND) to residents than expected by chance. The results were consistent with these three predictions: Relative to random simulations, we observed significant aggregation, home‐range overlap, and NND distribution functions in four, five, and six, respectively, of the six migrant–resident groups analyzed. Conspecific attraction may benefit migrant O. pumilio by providing cues to suitable home sites and/or increasing the potential for social interactions with conspecifics; if true, these benefits should outweigh the negative effects of other factors associated with aggregation. The observed aggregation between migrant and resident O. pumilio is consistent with conspecific attraction in dendrobatid frogs, and our study provides rare support from a field setting that conspecific attraction may be a relevant mechanism for models of anuran spatial ecology.     

Alternative models of conspecific attraction in flies and crabs

Animals are often attracted to one another when selecting habitats,but little is known about the rules governing conspecific attraction.We use Akaike Information Criterion to evaluate alternativemodels of the effects of conspecifics on individual choice inthe context of habitat selection. One set of models was testedusing data collected on virgin female flies, Drosophila melanogaster,selecting habitats in the laboratory; a second set of modelswas tested using data collected on crabs, Ocypode rotundata,selecting foraging patches in the field. Patterns of space usein the flies were most consistent with models indicating thatindividuals were attracted to other females that selected trapsduring the same hour, as well as to cues left by females thathad entered traps during the previous hour. Results for thecrabs were most consistent with a model which assumes that individualspreferred to join the patch with the most crabs but that theirability to assess the number of crabs in alternative patcheswas constrained by basic psychophysical principles (Weber'slaw). These results provide support for hypotheses about thefunctional significance of conspecific attraction in the fliesand the crabs and illustrate the richness of information aboutconspecific attraction that can be obtained when the same dataset is confronted with alternative models of the ways that animalsrespond to one another when selecting habitats.     

Female Attraction to Conspecific Chemical Cues in the Palmate Newt Triturus helveticus

Although chemosignals are largely used in sexual communication in urodeles, olfactometer studies in newts provided contrasting results about the sex specificity of female behavioural responses. Because long‐range sexual advertisement is believed to be costly, some species might restrain this activity to close interactions with conspecifics. We tested chemical‐mediated sexual attraction in female palmate newt (Triturus helveticus) by measuring the attraction to male and female odours in a linear water olfactometer. Unexpectedly, females were attracted towards conspecifics regardless of sex. They did not show attraction towards Limnaea stagnalis, a common sympatric aquatic gastropod. These results do not support the use of long‐range male sexual signalling in the palmate newt. Instead, conspecific attraction is likely to promote aggregation of males and females in breeding ponds. Observations in the field and in the laboratory tend to support the aggregative behaviour of this species. We discuss the possible function of conspecific attraction in this context. Heading towards any conspecific would increase the probability of finding potential mates. Chemical cues do not need to be sex‐specific at that stage so that long‐range sexual advertisement might be unnecessary. This work emphasizes the need for studies investigating the evolutionary relationships between sexual signalling systems and population‐distribution patterns.     

Testing the conspecific attraction hypothesis with Dupont's Larks,a resident species of songbird in central Spain

Some birds use social cues, such as the presence of conspecifics, when selecting breeding habitat. This phenomenon, known as conspecific attraction, has been well‐documented in migratory species, but has not been assessed for resident species of birds. We used Dupont's Larks (Chersophilus duponti) as a model species to determine if conspecific attraction plays a role in habitat selection by resident species of birds. At our study site in Soria province in central Spain, we monitored two potential habitat patches and one managed site where management actions had provided apparently suitable habitat. At each site, we broadcast recordings of the songs and calls of male Dupont's Larks, and monitored their presence during the breeding season and dispersal period in 2018 using automated recorders and field surveys. No birds were attracted to our study sites. Our results suggest that management of patches of suitable habitat should occur close to areas (within 1 km) already occupied by Dupont's Larks to encourage natural colonization because, based on our results, playback of conspecific vocalizations may not attract the species to new breeding areas. However, additional studies are needed before drawing conclusions about the effectiveness of conspecific attraction for this and other resident species of birds.     

Sex‐specific responses of Asian citrus psyllid to volatiles of conspecific and host‐plant origin

Diaphorina citri Kuwayama (Hemiptera: Psyllidae) is the primary vector of Candidatus Liberibacter spp. bacteria that cause citrus greening, a disease of worldwide importance. Olfactometry was employed to test responses of D. citri to odours from intact citrus plants (Mexican lime, Citrus aurantifolia, sour orange, Citrus aurantium, Marsh grapefruit, Citrus paradisi and Valencia orange, Citrus sinensis), citrus plants previously infested with D. citri, and odours of conspecifics including nymphs, adult insects of same and opposite sex, and their products (honeydew), both alone and in combination. In contrast to other studies, psyllids of both sexes were attracted to volatiles of undamaged Mexican lime leaves, whereas undamaged grapefruit attracted only females, and leaves of Valencia and sour orange did not attract either sex. All four plant species attracted female psyllids when previously infested, but only Mexican lime and sour orange‐attracted males. Thus, Citrus species appear to vary in the production of both constituitive and induced volatiles that attract adult psyllids. Volatiles emitted by nymphs did not attract either sex, but psyllid honeydew was attractive to males, likely due to female pheromone residues. Males oriented to the odour of females, whereas the reverse was not true, and neither males nor females oriented to same‐sex volatiles. The addition of conspecific cues (adults, nymphs or honeydew) did not increase female attraction to previously infested leaves, but male response was increased by the presence of adults and honeydew, regardless of plant species. Thus, female psyllids appear to orient more strongly to volatiles of plant origin, whereas males respond more strongly to cues emanating from females and conspecific excretions. These results suggest that female psyllids drive the initial colonization of host plants, whereas males orient to females and infested plants. Identification of the specific volatiles involved may permit their use in monitoring and management of this pest.     

Conspecific attraction in a grassland bird, the Baird's Sparrow

ABSTRACT.   Territorial songbirds generally use song to defend territories and attract mates, but conspecific song may also serve as a cue to attract other male songbirds to a breeding site. Although known to occur in some colonial and forest-associated species, only recently have investigators examined conspecific attraction in grassland species. We used a playback experiment to examine the possible role of conspecific attraction for males searching for potentially suitable breeding habitat in a grassland specialist, the Baird's Sparrow ( Ammodramus bairdii ). Experimental playback plots and control plots with similar landscape and vegetation characteristics were established at two sites in North Dakota. Baird's Sparrows colonized three of six experimental plots and none of six control plots. Males on experimental plots established territories adjacent to the playback stations and were sometimes observed counter-singing with the playback of conspecific songs. Vegetation characteristics were similar on all study plots, and did not explain differences in bird density on our treatment plots. Although we found that playback of conspecific songs attracted male Baird's Sparrows to previously unoccupied, potentially suitable habitat, further experiments are needed to examine the importance of conspecific attraction relative to other cues that birds may use, such as vegetation features. The conservation and management implications of conspecific attraction are not completely understood, but the presence of conspecifics should be considered as a potential cue in habitat selection by all species of birds.     

Space use and social association in a gregarious ungulate: Testing the conspecific attraction and resource dispersion hypotheses

Animals use a variety of proximate cues to assess habitat quality when resources vary spatiotemporally. Two nonmutually exclusive strategies to assess habitat quality involve either direct assessment of landscape features or observation of social cues from conspecifics as a form of information transfer about forage resources. The conspecific attraction hypothesis proposes that individual space use is dependent on the distribution of conspecifics rather than the location of resource patches, whereas the resource dispersion hypothesis proposes that individual space use and social association are driven by the abundance and distribution of resources. We tested the conspecific attraction and the resource dispersion hypotheses as two nonmutually exclusive hypotheses explaining social association and of adult female caribou (Rangifer tarandus). We used location data from GPS collars to estimate interannual site fidelity and networks representing home range overlap and social associations among individual caribou. We found that home range overlap and social associations were correlated with resource distribution in summer and conspecific attraction in winter. In summer, when resources were distributed relatively homogeneously, interannual site fidelity was high and home range overlap and social associations were low. Conversely, in winter when resources were distributed relatively heterogeneously, interannual site fidelity was low and home range overlap and social associations were high. As access to resources changes across seasons, caribou appear to alter social behavior and space use. In summer, caribou may use cues associated with the distribution of forage, and in winter caribou may use cues from conspecifics to access forage. Our results have broad implications for our understanding of caribou socioecology, suggesting that caribou use season‐specific strategies to locate forage. Caribou populations continue to decline globally, and our finding that conspecific attraction is likely related to access to forage suggests that further fragmentation of caribou habitat could limit social association among caribou, particularly in winter when access to resources may be limited.     

Evidence for a male-produced aggregation pheromone in Scyphophorus acupunctatus Gyllenhal (Coleoptera: Curculionidae)

Olfactory response of male and female Scyphophorus acupunctatus Gyllenhal (Coleoptera: Curculionidae) to volatiles released from the same or opposite conspecifics alone, or combined with host plant volatiles, was evaluated in the laboratory and field. We also evaluated the response to synthetic Rhynchophorinae pheromones in the laboratory. In laboratory tests, attraction of males and females in Y-tube olfactometer to conspecific males was greater than to females and clean air. Males and females preferred the combination males + agave over agave alone. Both sexes were significantly attracted to 2-methyl-4-heptanol and 2-methyl-4-octanol compared with hexane control. In field trials, weevils were successfully caught in the traps baited with conspecifics and plant material. These field results support those of the laboratory bioassays, showing that males attracted conspecific males and females and addition of plant material enhanced the attraction. These results further suggest that S. acupunctatus produces an aggregation pheromone.     

Plant volatiles and density-dependent conspecific female odors are used by Asian citrus psyllid to evaluate host suitability on a spatial scale

We investigated how chemical cues derived from female Asian citrus psyllid, Diaphorina citri Kuwayama (Hemiptera: Liviidae) and their host plants affect host acceptance choices by conspecifics. In four-choice cage and two-choice olfactometer assays, female psyllids avoided conspecific female cues in a density-dependent manner. However, odors from citrus plants actively damaged by psyllid feeding were attractive to conspecific females. When odors from feeding-damaged plants were presented simultaneously with odors from female D. citri, attraction of female conspecifics was no longer observed as compared with a clean air control in olfactometer assays. In subsequent experiments, D. citri females were released within arenas that contained actively feeding-damaged or non-damaged (control) citrus plants, each with previously psyllid-infested and uninfested young leaves. D. citri development is linked to the presence of these newly emerging leaves which is the only site of nymphal development. Female D. citri were initially attracted by the actively damaged plants as compared with non-damaged controls. After acceptance of plants that were actively damaged by feeding, D. citri females preferentially chose and settled on uninfested young leaves as compared with previously infested young leaves. A herbivore-induced plant volatile attractant and a female-specific odor repellent appear to be complementary foraging cues providing psyllids with information at two spatial scales: (1) the whole plant level for choosing a plant potentially harboring male conspecifics for mating, and (2) the within plant level to reduce intra-conspecific competition by identifying previously exploited resources.     

Finding a place to live: conspecific attraction affects habitat selection in juvenile green and golden bell frogs

Conspecific attraction plays an important role in habitat selection of several taxa and can affect and determine distribution patterns of populations. The behaviour is largely studied and widespread among birds, but in amphibians, its occurrence seems limited to breeding habitats of adults and gregarious tadpoles. The Australian green and golden bell frogs (Litoria aurea) have suffered considerable shrinking of their original distribution in south-eastern Australia since the 1970s. Currently, with only about 40 populations remaining, the species is considered nationally threatened. In natural conditions, these frogs are aggregated in the landscape and do not seem to occupy all suitable ponds within the occurrence area. To date, studies focusing on the frogs’ habitat have failed in finding a general habitat feature that explains current or past occupancy. This led us to the hypothesis that social cues may play a key role in habitat selection in this species. Using two choice experiments, we tested the preference of juvenile green and golden bell frogs for habitats containing cues of conspecifics of similar size versus habitats without conspecific cues. Tested frogs did not show a preference for habitats containing only scent from conspecifics but did prefer habitats where conspecifics were present. Our results show that conspecific attraction is a determining factor in juvenile green and golden bell frog habitat selection. To our knowledge, this is the first time the behaviour is shown to occur in juvenile frogs in the habitat selection context. From a conservation management point of view, the behaviour may help to explain the failure of reintroductions to areas where the frogs have been extinct, and the non-occupation of suitable created habitats in areas where they still inhabit and develop appropriated management strategies.     

Species interactions and population density mediate the use of social cues for habitat selection

1. The perspective that populations and communities are structured by antagonistic interactions among individuals has dominated much of ecology. Yet how animals use social information to guide decisions, such as habitat selection, may be influenced by both positive and negative interactions among individuals. Recent theory also suggests that the way animals use social information may be substantially influenced by population density, which alters the potential costs and benefits of such behaviours. 2. I manipulated cues of two competitors, the dominant least flycatcher Empidonax minimus (Baird & Baird) and the subordinate American redstart Setophaga ruticilla (Linnaeus), to assess the use of conspecific and heterospecific cues during habitat selection, and if population density influences these strategies. The experiment consisted of surveying birds during a pre-treatment year, which allows for the control and testing the effect of baseline densities, and a treatment year, in which treatments were applied just prior to settlement. Treatments included broadcasting songs of flycatchers and redstarts, and were compared with controls. 3. When controlling for pre-treatment densities, bird densities, and to a lesser extent arrival dates, during the treatment year suggested that flycatchers were attracted to both conspecific and heterospecific cues during settlement. Furthermore, attraction was strongest for flycatchers in plots with moderate pre-treatment densities. American redstarts were rare in the study area but showed apparent attraction to conspecifics and avoidance of heterospecifics. 4. These results provide experimental evidence for the use of multiple social cues in habitat selection and suggest that heterospecific attraction may operate under broader contexts than originally envisioned. In such instances, nontarget effects can potentially occur when manipulating social cues to elicit settlement in conservation strategies. The impact of population density on the use of social cues shown here can also influence our understanding of metapopulation dynamics by causing complex threshold effects on the likelihood of rescue, which may influence metapopulation stability and the likelihood of local extinction.     

Foraging and spatial dispersion in protein-scavenging workers ofVespula germanica andV. maculifrons (Hymenoptera: Vespidae)

A series of experiments on the discovery of meat bait by scavenging workers ofV. germanica andV. maculifrons showed that workers oriented to meats by visual cues and olfactory cues. Scavening workers ofV. germanica andV. maculifrons responded positively to the sight of conspecifics at meat baits; i.e., meat with (confined) wasps received more visits than meat without wasps. This attraction to conspecific workers on meat baits provides evidence of local enhancement, in that the presence of conspecifics at meat bait directs other wasps to a particular spatial locality. Local enhancement by yellowjackets foraging for meat may contribute to the clumped distribution of worker populations noted in studies of workers captured in meat-baited traps. We hypothesize that local enhancement of foraging is a ubiquitous behavior in species of theV. vulgaris species group and may be one attribute explaining their ecological success.     

Spatial Nest‐Settlement Decisions in Digger Wasps: Conspecifics Matter more than Heterospecifics and Previous Experience

Research into the driving forces behind spatial arrangement of wasp nests has considered abiotic environmental factors, but seldom investigated attraction or repulsion towards conspecifics or heterospecifics. Solitary female digger wasps (Hymenoptera) often nest in dense aggregations, making these insects good models to study this topic. Here, we analysed the nesting patterns in an area shared by three species of the genus Bembix, in a novel study to discover whether female wasps are attracted to or repulsed by conspecific nests, heterospecific nests or their own previously established nests when choosing nest‐digging locations. Early in the season, each species showed a clumping pattern of nests, but later in the season, a random distribution of nests was more common, suggesting an early conspecific attraction. Such behaviour was confirmed by the fact that females started building their nests more frequently where other females of their species were simultaneously digging. The distances between subsequent nests dug by individual females were shorter than those obtained by random simulations. However, this pattern seemed to depend on the tendency to dig close to conspecifics rather than remain in the vicinity of previous nests, suggesting that females' experience matters to future decisions only on a large scale. Nesting patches within nest aggregations largely overlapped between species, but the nests of each species were generally not closer to heterospecific nests than expected by chance, suggesting that females are neither repulsed by, nor attracted to, congenerics within nest aggregations. A role of the spatial distribution of natural enemies on the observed nesting patterns seemed unlikely. Bembix digger wasp nest aggregations seem thus to be primarily the result of female–female attraction during nest‐settlement decisions, in accordance with the ‘copying’ mechanisms suggested for nesting vertebrates.     

Does Conspecific Attraction Affect Nest Choice in Budgerigars (Melopsittacus undulatus: Psittacidae: Aves)?

In their native Australia the budgerigar's breeding season is very short and rapid nest establishment is crucial. To minimize the time required to locate and establish nesting sites, female budgerigars were predicted to use cues on nest site locations provided by other females in their colony. Experimental results from captive birds indicated that, while females can be attracted to nests that are occupied by other females, the social and breeding experience of individuals affects this use of conspecific cues. Females that have had altercations with other females that resulted in physical injuries showed an aversion to apparently occupied nestboxes. Females that had no such social experience prior to testing were attracted to apparently occupied nestboxes. Breeding experience, as well as social experience, may have an effect on the use of conspecifics as cues. Experienced breeding females were less likely to be attracted to apparently occupied nestboxes than were females that had never bred before.     

Cues for cannibalism: cane toad tadpoles use chemical signals to locate and consume conspecific eggs

Although animals of many species kill and consume conspecifics, most such cases probably involve serendipitous encounters between the individuals concerned. In some taxa, however, cannibalism is an active process, with predatory individuals searching out and consuming specific types of conspecific prey items. Although anuran tadpoles often have been reported to consume conspecific eggs, this behaviour has been interpreted as a by‐product of usual foraging behaviours rather than a result of targeted searching. Our field and laboratory studies in tropical Australia show that the tadpoles of invasive cane toads Bufo marinus are strongly attracted to chemical cues from conspecific eggs; the effective cues are released late in embryonic development, as the jelly coat breaks down. Tadpoles of native Australian frog species were attracted to the eggs of toads only rarely. If deployed as bait in traps, chemical cues from toad eggs could provide a way to selectively remove toad larvae from waterbodies.     

A stochastic locomotor control model for the nurse shark,ginglymostoma cirratum

Summary The locomotor behavior of the nurse shark (Ginglymostoma cirratum) is characterized by 17 variables (frequency and ratios of left, right, and total turns; their radians; straight paths (steps); distance travelled; and velocity) Within each of these variables there is an internal time dependency the structure of which was elaborated together with an improved statistical model predicting their behavior within 90% confidence limits. The model allows for the sensitive detection of subtle locomotor response to sensory stimulation as values of variables may exceed the established confidence limits within minutes after onset of the stimulus. The locomotor activity is well described by an autoregression time series model and can be predicted by only seven variables. Six of these form two independently operating clusters. The first one consists of: the number of right turns, the distance travelled, and the mean velocity; the second one of: the mean size of right turns, of left turns, and of all turns. The same clustering is obtained independently by a cluster analysis of cross-sections of the seven time series. It is apparent that, among a total of 17 locomotor variables, seven behave as individually independent agents, presumably controlled by seven separate and independent centers. The output of each center can only be predicted by its own behavior. In spite of the individual behavior of the seven variables, their internal structure is similar in important aspects which may result from control by a common command center. The shark locomotor model differs in important aspects from that previously constructed for the goldfish. The interdependence of the locomotor variables in both species may be related to the control mechanisms postulated by von Holst for the coordination of rhythmic fin movements in fishes. A locomotor control model for the nurse shark is proposed.     

Species recognition in mormyrid weakly electric fish

We investigated group cohesion in four species of African weakly electric fish (Brienomyrus niger, Gnathonemus petersii, Marcusenius cyprinoides and Pollimyrus isidori). The attraction among members of the same and different species served as the criterion for species recognition. To identify possible mechanisms underlying this ability, conspecifics were allowed to interact through a wide-meshed plastic screen with either a pair of confined conspecifics or a pair from a related species. Members of each of the four test species were optimally attracted to their own kind, but also responded selectively to the presence of the other species. These interspecific interactions ranged from attraction to avoidance. The difference in the fish's preference to aggregate in inter- and intraspecific interactions pointed to species-specific social cues that facilitate group cohesion in mormyrid fish. Since all four species respond to each other's electric organ discharge with changes in their own discharge rate, species recognition cannot be merely a function of electroreceptor characteristics but must involve the integration of electroreceptive and other sensory cues.     

Responses of larval dragonflies to conspecific and heterospecific predator cues

Abstract.  1. In cannibalistic populations, smaller individuals are subject to predation by larger conspecifics, and small individuals commonly alter their behaviour in response to cannibals. Little is known, however, about the underlying cues that trigger such responses and how the behavioural responses to conspecific cannibals differ from heterospecific predators.
2. This study tests which cues are used for the detection of conspecific predators in the larva of the dragonfly Plathemis lydia and how the behavioural response to cannibals differed from the response to heterospecific predators.
3. Individuals were exposed to chemical cues, visual cues, and a combination of both cues from conspecifics as well as no predator and heterospecific predator controls during which their activity and feeding rates were observed.
4. Individuals increased their activity, spatial movement and feeding behaviour in response to either visual or chemical cues from conspecific predators, which was opposite to responses displayed with cues from heterospecific predators. Interestingly, the responses to visual and chemical cues from conspecifics combined were weaker than to either cue in isolation and similar to the no cue control.
5. The results clearly indicate that individuals are able to use chemical and visual cues to detect even very subtle differences in phenotype of conspecific predators.
6. The opposite response in behaviour when exposed to conspecific cannibals vs. heterospecific predators suggests that the presence of cannibals will increase the mortality risk of small individuals due to heterospecific predation. This risk-enhancement is likely to have important consequences for the dynamics of predator–prey interactions.