Wall lizards combine chemical and visual cues of ambush snake predators to avoid overestimating risk inside refuges

The threat sensitivity hypothesis assumes that multiple cues from a predator should contribute in an additive way to determine the degree of risk-sensitive behaviour. The ability to use multiple cues in assessing the current level of predation risk should be especially important to prey exposed to multiple predators. Wall lizards, Podarcis muralis, respond to predatory attacks from birds or mammals by hiding inside rock crevices, where they may encounter another predator, the smooth snake, Coronella austriaca. We investigated in the laboratory whether chemical cues may be important to wall lizards for detection of snakes. The greater tongue-flick rate and shorter latency to first tongue-flick in response to predator scents indicated that lizards were able to detect the snakes' chemical cues. We also investigated the use of different predatory cues by lizards when detecting the presence of snakes within refuges. We simulated successive predator attacks and compared the propensity of lizards to enter the refuge and time spent within it for predator-free refuges, refuges containing either only visual or chemical cues of a snake, or a combination of these. The antipredatory response of lizards was greater when they were exposed to both visual and chemical cues than when only one cue was presented, supporting the threat sensitivity hypothesis. This ability may improve the accuracy of assessments of the current level of predation risk inside the refuge. It could be especially important in allowing lizards to cope with threats posed by two types of predators requiring conflicting prey defences.     

To run or hide? Age-dependent escape behaviour in the common flat lizard (Platysaurus intermedius wilhelmi)

Flat lizards Platysaurus intermedius wilhelmi occur on small discrete rock outcrops in Mpumalanga Province, South Africa. These rock outcrops are structurally simple and this, combined with the lizard's behaviour (ambush foraging in the open), make them ideal for field studies of anti-predatory behaviour. Lizards were approached in the field and how escape behaviour was influenced by habitat and age-sex class was recorded. Juveniles ( c. 4 months of age) responded quite differently to an approaching human 'predator' compared with adult males and females (which responded similarly). Compared to adults, juveniles allowed a closer approach by the investigator; took longer to find a refuge and therefore fled further; were more likely to remain visible in the open and maintain visual contact with the investigator; and more likely to flee into vegetation when given the opportunity to take refuge in a crevice. We suggest that because a greater suite of predators (including arthropods living in rock crevices) feed on small juvenile lizards, this may affect their choice of refuge and result in the avoidance of crevices when chased. Finally, because juveniles were frequently found on small rock outcrops, the influence of rock outcrop area on anti-predatory behaviour was tested. Escape behaviour (time to refuge) was independent of rock outcrop area.     

Belly up: Reduced crevice accessibility as a cost of reproduction caused by increased girth in a rock‐using lizard

Costs of reproduction are any aspect of current reproduction that has the potential to reduce survivorship or reproductive output, and may include physiological costs or increased risks. Females of many species experience increased body mass, and increased girth, when gravid. Increased body mass reduces running speed and increases the cost of locomotion during pregnancy, but few studies have examined the cost of increased girth. If increased girth of gravid females reduces access to shelter from predators or the elements, increased girth could constitute a cost of reproduction. In the laboratory, we experimentally tested whether access to crevices was limited in viviparous, saxicolous female lizards (Eulamprus brachysoma), which use crevices for shelter, by measuring access to artificial crevices of known widths, and body height during and after pregnancy. Gravid E. brachysoma had significantly greater body height (11.2% on average), and as a result were forced to use significantly wider crevices (18.4% wider on average) than post‐parturition. Females with larger clutch sizes had wider mid‐bodies and required larger crevices. Control females, which were not gravid at either time of testing, showed no significant change in the size of crevice they could enter over time. If access to narrow crevices provides advantages such as protection from predators, or is important for thermoregulation, then gravid females may suffer a cost of reproduction because their access to narrower crevices is limited.     

Fleeing towards death – leech‐induced behavioural defences increase freshwater snail susceptibility to predatory fish

Prey species are often exposed to multiple predators, which presents several difficulties to prey species. This is especially true when the response to one predator influences the prey’s susceptibility to other predators. Predator‐induced defences have evolved in a wide range of prey species, and experiments involving predators with different hunting strategies allow researchers to evaluate how prey respond to multiple threats. Freshwater snails are known to respond to a variety of predators with both morphological and behavioural defences. Here we studied how freshwater snails Radix balthica responded behaviourally to fish and leech predators, both separately and together. Our aim was to explore whether conflicting predator‐induced responses existed and, if so, what effect they had on snail survival when both predatory fish and leeches were present. We found that although R. balthica increased refuge use when exposed to predatory fish, they decreased refuge use when exposed to predatory leeches. When both predators were present, snails showed a stronger response towards leech than fish and responded by leaving the refuge. This response made the snails more susceptible to fish predation, which increased snail mortality when exposed to both fish and leech compared to fish only. We show that predators that have a relatively low predation rate can substantially increase mortality rates by indirect effects. By forcing snails out of refuges such as rock and macrophyte habitats, leeches can indirectly increase predation from molluscivorous fish and may thus affect snail densities.     

Long-range visibility of greater sage grouse leks: a GIS-based analysis

We investigated whether male greater sage grouse, Centrocercus urophasianus, select lek locations on the basis of topographic features that affect their visibility to both conspecific females and a major avian predator, the golden eagle, Aquila chrysaetos. We mapped locations of displaying males at all leks in a local population and used a Geographic Information System (GIS) and digital elevation model (DEM) to generate ‘viewsheds’ around male locations within a boundary set by the estimated maximal visual acuity of the viewer. Areas visible around leks were compared to those visible around random sites with the same conformation of displaying males. Male sage grouse displayed at sites where surrounding topography both diminished long-range visibility (>1000 m) and enhanced short-range visibility (<500 m) to ground observers. These characteristics could (1) force eagles to monitor lek activity from the air, where they may be more visible to their prey, (2) make displaying males more visible to females and (3) allow males to monitor predators approaching the lek more easily. These results suggest that, in open habitats, visually signalling animals may exploit local topography to control both their visibility to receivers and the visibility of their immediate surroundings.     

Antipredator defence of the hermit crab Pagurus filholi induced by predatory crabs

Hermit crabs have two antipredator tactics: taking refuge in its shell and fleeing. We examined the following two hypotheses using the hermit crab Pagurus filholi : (1) hermit crabs change their preference for shell types that they take refuge in and/or change the timing of fleeing (i.e. the duration of refuge in the shell) when they perceive a predator threat; (2) the type of shell that a hermit crab occupies affects the fleeing tactic of the individual. Under the stimulus of a crushed conspecific, hermit crabs changed neither their preference for shell species nor their refuge duration. On the other hand, under the stimulus of the predatory crab Gaetice depressus , hermit crabs increased their preference for Batillaria cumingi shells, which provide superior protection against predators, and shortened their refuge duration in the shells even when they occupied those effective against predation. Refuge duration was longer in B. cumingi shells than in the more vulnerable shells of Homalopoma sangarense . These results suggest that both antipredator defences (changing shell and timing of fleeing) are induced by the stimulus of a predator, and the timing of fleeing is affected by the shell type occupied.     

Can Wall Lizards Combine Chemical and Visual Cues to Discriminate Predatory from Non‐Predatory Snakes Inside Refuges?

The ability to use multiple cues in assessing predation risk is especially important to prey animals exposed to multiple predators. Wall lizards, Podarcis muralis, respond to predatory attacks from birds in the open by hiding inside rock crevices, where they may encounter saurophagous ambush smooth snakes. Lizards should avoid refuges with these snakes, but in refuges lizards can also find non‐saurophagous viperine snakes, which lizards do not need to avoid. We investigated in the laboratory whether wall lizards used different predator cues to detect and discriminate between snake species within refuges. We simulated predatory attacks in the open to lizards, and compared their refuge use, and the variation in the responses after a repeated attack, between predator‐free refuges and refuges containing visual, chemical, or visual and chemical cues of saurophagous or non‐saurophagous snakes. Time to enter a refuge was not influenced by potential risk inside the refuge. In contrast, in a successive second attack, lizards sought cover faster and tended to increase time spent hidden in the refuge. This suggests a case of predator facilitation because persistent predators in the open may force lizards to hide faster and for longer in hazardous refuges. However, after hiding, lizards spent less time in refuges with both chemical and visual cues of snakes, or with chemical cues alone, than in predator‐free refuges or in refuges with snake visual cues alone, but there were no differences in response to the two snake species. Therefore, lizards could be overestimating predation risk inside refuges. We discuss which selection pressures might explain this lack of discrimination of predatory from similar non‐predatory snakes.     

The Effect of Refuge on Dermestes ater (Coleoptera: Dermestidae) Predation on Musca domestica (Diptera: Muscidae): Refuge for Prey or the Predator?

It is believed that habitat heterogeneity can change the extent of predator-prey interactions. Therefore, in this study we examined the effect of habitat heterogeneity (characterized here as an addition of refuge) on D. ater predation on M. domestica. Predation of D. ater on M. domestica larvae was carried out in experimental habitats with and without refuge, and examined at different prey densities. The number of prey eaten by beetles over 24 h of predator-prey interaction was recorded, and we investigated the strength of interaction between prey and predator in both experimental habitats by determining predator functional response. The mean number of prey eaten by beetles in the presence of refuge was significantly higher than in the absence of refuge. Females had greater weight gains than males. Logistic regression analyses revealed the type II functional response for both experimental habitats, even though data did not fit well into the random predator model. Results suggest that the addition of refuge in fact enhanced predation, as prey consumption increased in the presence of refuge. Predators kept in the presence of refuge also consumed more prey at high prey densities. Thus, we concluded that the addition of refuge was an important component mediating D. ater-M. domestica population interactions. Refuge actually acted as a refuge for predators from prey, since prey behaviors detrimental to predators were reduced in this case.     

The influence of predator presence on utilization of artificial seagrass habitats by juvenile walleye pollock,Theragra chalcogramma

Synopsis Behavioral preference for a structured habitat (artificial seagrass) by juvenile walleye pollock,Theragra chalcogramma, was tested in controlled laboratory experiments. We monitored position of fish in 2000 1 tanks with and without artificial seagrass present in one half of the tank. In addition, we exposed walleye pollock to a predator model, assessing their response when a grass plot was available or unavailable as a potential refuge. In the absence of predators, the fish avoided the artificial seagrass, displaying a preference for the open water side of the experimental tanks. In the presence of a predator model, however, juvenile walleye pollock readily entered the artificial seagrass plots. In addition, they often remained in the grass canopy in proximity to the predator instead of moving out of the grass to avoid the predator (when no grass was present they consistently moved to the opposite side of the tank from the predator). The behavioral choices exhibited in this study suggest that juvenile walleye pollock modify habitat selection in response to perceived predation risk, and recognize the structure provided by artificial seagrass as a potential refuge.     

Costs and benefits of defences induced by predators differing in dangerousness

While theoretical studies predict that inducible defences should be fine-tuned according to the qualities of the predator, very few studies have investigated how dangerousness of predators, i.e. the rate at which predators kill prey individuals, affects the strength of phenotypic responses and resulting benefits and costs of induced defences. We performed a comprehensive study on fitness consequences of predator-induced responses by involving four predators (leech, water scorpion, dragonfly larva and newt), evaluating costs and benefits of responses, testing differences in dangerousness between predators and measuring responses in several life history traits of prey. We raised Rana dalmatina tadpoles in the presence of free-ranging predators, in the presence of caged predators, and exposed naive and experienced tadpoles to free-ranging predators. Tadpoles adjusted the intensities of their behavioural and morphological defences to predator dangerousness. Survival was lower in the nonlethal presence of the most dangerous predator, while we could not detect costs of induced defences at or after metamorphosis. When exposed to free-ranging predators, small, but not large, tadpoles benefited from exhibiting an induced phenotype in terms of elevated survival when compared to naive tadpoles, but we did not observe higher survival either in tadpoles exhibiting more extreme phenotypes or in tadpoles exposed to the type of predator they were raised with. These results indicate that while predator-induced defences can mirror dangerousness of predators, costs and benefits do not necessarily scale to the magnitude of plastic responses.     

Predation Risk and Opportunity Cost of Fleeing While Foraging on Plants Influence Escape Decisions of an Insular Lizard

Cost‐benefit models of escape behaviour predict how close a prey allows a predator to approach [flight initiation distance (FID)] based on cost of not fleeing (predation risk) and cost of fleeing (loss of opportunities). Models for FID have been used with some success to predict distance fled (DF). We studied effects of foraging opportunity cost of fleeing and examined differences between age‐sex groups in the omnivorous Balearic Lizard, Podarcis lilfordi. Balearic lizards forage on the ground for invertebrate prey and climb the thistle Carlina corymbosa to forage on its inflorescences. We studied escape behaviour in three experimental groups, with human beings as simulated predators: lizard foraging above ground on C. corymbosa, foraging on the ground away from thistles and on the ground with cut inflorescences. Flight initiation distance was shorter for lizards with cut inflorescences than for (1) lizards above ground due to the greater risk above ground due to conspicuousness of black lizards on yellow flowers; and (2) lizards on ground away from flowers due to the cost of leaving while feeding. The only age‐sex difference was slightly greater FID for adult males than subadults, presumably because larger adult males are more likely to be attacked by predators. Other potential factors affecting this difference are discussed. Experimental group and age‐sex group did not interact for FID or DF. Because lizards foraging on inflorescences above ground fled to the base of the plants to refuge provided by spiny thistle leaves, their DF was shorter than in the other groups, which fled across the ground, usually without entering refuge. DF did not differ between groups on the ground or among age‐sex groups. The predicted shorter DF for lizards with cut inflorescences than on ground without inflorescences did not occur. We hypothesize that the opportunity cost was small due to the abundance of blooming thistles and that DF may be less sensitive to opportunity cost than FID.     

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     

Development of predator defences in fishes

Summary A variety of development characteristics, morphological, behavioural, and experiential, contribute to the extreme vulnerability of young fishes to predation. The influence of these characteristics is complicated by the fact that the larval period is one of substantial and rapid change. Yet survival is the ultimate goal;-it is only by reaching maturity that individual fish have the opportunity to reproduce. With such high stakes it is not surprising that predator defences are of major importance during all phases of life. Developmental constraints may limit the defensive options for young fishes. Avoidance behaviours, which reduce the likelihood of encountering a predator or of being attacked by it, are particulaly evident in the youngest stages. Here size, coloration and dispersal are used to help elude the predator's attention. As fishes grow and acquire greater morphological and behavioural sophistication, there is more scope for predator evasion when avoidance fails. Older fishes are increasingly able to respond to external stimuli and can detect and react to predators or join conspecifics in common defence (schooling). Behavioural development is not simply a consequence of growth and the concomitant physical alterations of the body; it is also mediated by experience that comes through interaction with the physical and biotic environment. Predispositions to respond to experience may be a product of evolutionary history. Although mortality rates decline markedly with development and maturity, changes in size or behaviour can render fishes vulnerable to new suites of predators. Effective predator avoidance can compromise other activities, such as foraging, and individuals may be forced to reconcile conflicting demands. Developmental niche shifts that occur, for example, when certain size classes take refuge in less profitable feeding habitats, represent one such trade-off. Niche shifts may also be mediated by the influence of the programme for morphological development on sensory or behavioural capabilities. In addition to all of these developmental consderations, natural variations in environmental conditions — such as temperature, photoperiod, predator density and variety, and presence of alternative prey — represent additional challenges to predator defences during the rite of passage from birth to reproduction.     

Crevice occupation by musk turtles: taxonomic distribution and crevice attributes

Musk turtles, Sternotherus, of all taxa preferentially used crevices as resting sites in laboratory pools. Crevice use was greater during the day. Larger, darker crevices that were low enough to touch the carapace were preferred. Positional preferences within crevices were for full coverage of the shell despite the head being exposed, for the low half of crevices, and for axial orientation that maximized contact between the carapace and crevice ceiling. Light but touchable portions of crevices were preferred over dark but untouchable portions, so the possibility of establishing carapacial contact with the ceiling is the definitive attribute of a satisfactory crevice. The behavioural tendency to use the lowest possible crevice may have an anti-predator function, and it is one necessary condition for existence of a selective regime in favour of depressed shells.     

Interactions between the information content of different chemical cues affect induced defences in tadpoles

Animals often alter their behaviour, morphology and physiology in the presence of predators. These induced defences can be fine‐tuned by a variety of environmental factors such as predator species, acute predation risk or food availability. It has, however, remained unclear what cues influence the extent and quality of induced defences and how the information content of these cues interact to determine the development of antipredator defences. We performed an experiment to study the significance of direct chemical cues, originating from the predators themselves, and indirect cues, released by attacked or consumed prey, for phenotypic responses in Rana dalmatina tadpoles. We reared tadpoles in the presence of caged predators (Triturus vulgaris, Aeshna cyanea) fed either one or three tadpoles every other day outside the tadpole‐rearing tanks. Fifteen hours after food provisioning, predators were put back into the tanks containing focal tadpoles either after washing (direct + digestion‐released cues) or with the water containing remnants of the prey (direct + all types of indirect cues). Our results suggest that direct cues together with digestion‐released cues can be sufficient to induce strong antipredator responses. Induced defences depended on both direct cues, affecting predator‐specific responses, and the quantity of indirect cues, resulting in graded responses to differences in predation threat. Moreover, direct and indirect cues interacted in behaviour, resulting in predator‐specific graded responses. We also observed a decrease in the extent of predator‐induced responses in large tadpoles as compared to small ones. Our results, thus, suggest that prey integrate multiple cues about predators to optimize induced defences and that this process changes during ontogeny.     

Do anuran larvae respond behaviourally to chemical cues from an invasive crayfish predator? A community-wide study

Antipredator behaviour is an important fitness component in most animals. A co-evolutionary history between predator and prey is important for prey to respond adaptively to predation threats. When non-native predator species invade new areas, native prey may not recognise them or may lack effective antipredator defences. However, responses to novel predators can be facilitated by chemical cues from the predators’ diet. The red swamp crayfish Procambarus clarkii is a widespread invasive predator in the Southwest of the Iberian Peninsula, where it preys upon native anuran tadpoles. In a laboratory experiment we studied behavioural antipredator defences (alterations in activity level and spatial avoidance of predator) of nine anurans in response to P. clarkii chemical cues, and compared them with the defences towards a native predator, the larval dragonfly Aeshna sp. To investigate how chemical cues from consumed conspecifics shape the responses, we raised tadpoles with either a tadpole-fed or starved crayfish, or dragonfly larva, or in the absence of a predator. Five species significantly altered their behaviour in the presence of crayfish, and this was largely mediated by chemical cues from consumed conspecifics. In the presence of dragonflies, most species exhibited behavioural defences and often these did not require the presence of cues from predation events. Responding to cues from consumed conspecifics seems to be a critical factor in facilitating certain behavioural responses to novel exotic predators. This finding can be useful for predicting antipredator responses to invasive predators and help directing conservation efforts to the species at highest risk.     

Optimal escape theory predicts escape behaviors beyond flight initiation distance： risk assessment and escape by striped plateau lizards Sceloporus virgatus

Escape theory predicts that flight initiation distance (FID=distance between predator and prey when escape begins) is longer when risk is greater and shorter when escape is more costly. A few tests suggest that escape theory applies to distance fled. Escape models have not addressed stochastic variables, such as probability of fleeing and of entering refuge, but their economic logic might be applicable. Experiments on several risk factors in the lizard Sceloporus virgatus confirmed all predictions for the above escape variables. FID was greater when approach was faster and more direct, for lizards on ground than on trees, for lizards rarely exposed to humans, for the second of two approaches, and when the predator turned toward lizards rather than away. Lizards fled further during rapid and second consecutive approaches. They were more likely to flee when approached directly, when a predator turned toward them, and during second approaches. They were more likely to enter refuge when approached rapidly. A novel finding is that perch height in trees was unrelated to FID because lizards escaped by moving out of sight, then moving up or down unpredictably. These findings add to a growing body of evidence supporting predictions of escape theory for FID and distance fled. They show that two probabilistic aspects of escape are predictable based on relative predation risk levels. Because individuals differ in boldness, the assessed optimal FID and threshold risks for fleeing and entering refuge are exceeded for an increasing proportion of individuals as risk increases[Current Zoology 55(2):123-131,2009].     

Freezing behaviour facilitates bioelectric crypsis in cuttlefish faced with predation risk

Cephalopods, and in particular the cuttlefish Sepia officinalis, are common models for studies of camouflage and predator avoidance behaviour. Preventing detection by predators is especially important to this group of animals, most of which are soft-bodied, lack physical defences, and are subject to both visually and non-visually mediated detection. Here, we report a novel cryptic mechanism in S. officinalis in which bioelectric cues are reduced via a behavioural freeze response to a predator stimulus. The reduction of bioelectric fields created by the freeze-simulating stimulus resulted in a possible decrease in shark predation risk by reducing detectability. The freeze response may also facilitate other non-visual cryptic mechanisms to lower predation risk from a wide range of predator types.     

The effect of natural variation in substrate architecture on the survival of juvenile bicolor damselfish

Coral reefs offer settling fish larvae a spatial mosaic of microhabitats that differ not only in structural complexity but also in the abundance and diversity of predators. This paper provides evidence that interactions between predators and prey are causally linked to variation in relevant architectural characteristics of natural substrates. Juvenile bicolor damselfish, Stegastes partitus, experienced greater mortality on Montastrea annularis boulder coral than on piles of Porites porites coral rubble. This pattern was consistent on both back and fore reef habitats. Architectural differences, variable encounter rates with predators, and access to different food sources all contributed to higher mortality. Spatial and structural differences among refuges provided by the two substrates were most important in affecting survival. Porites rubble contained almost three times the number of crevices (39.9 vs. 14.3 crevices m^–2AM²), had a much smaller mean crevice size (15.9 vs. 48.5 cm²), and had a more complex internal structure than Montastrea coral. Survival of juvenile fish living on Montastrea coral was positively correlated with crevice density (r² = 0.41) whereas survival of fish living on Porites rubble was negatively correlated with crevice size (r² = 0.43). These patterns were evident on fore reef habitats only, whereas on back reef habitats no clear patterns emerged. The effect of these natural differences in architecture on mortality rates of juvenile S. partitus was experimentally tested in the field using a combination of natural and artificial substrates. As the number of large shelters was increased I found that the density of potential predators increased but the survival rates of juvenile bicolor damselfish declined. These results highlight the importance of structural architecture among common reef substrates in affecting predator-prey relationships and in determining the survivorship and small-scale distribution patterns of juvenile reef fish.     

Anti-Predator Signals as Advertisements: Evidence in White-Throated Magpie-Jays

Calls and displays elicited by predators usually function as alarms or to inform predators of their detection. However, predator encounters may afford some individuals the opportunity to demonstrate quality or signal their availability. Here, I report on a class of vocal signals produced in predator-elicited displays that share many characteristics with sexually selected song. White-throated magpie-jays ( Calocitta formosa ) display at low-threat predators while producing 'loud display calls' (LDCs). I use this term because the calls occur primarily in two display contexts (see below) though occasionally in other contexts as well. Such calls and displays are primarily produced by males, and also occur in one other context, at dawn. Playback experiments showed that despite being elicited by predators, males were more likely than females to respond to LDCs, and more likely to respond when their mate was fertile. Over 134 different call types were produced in over 200 displays by 34 males; the largest minimum repertoire size was 67. Presentations of taxidermic raptor mounts elicited some LDCs, but fewer calls and lower diversity than at dawn or in predator approach displays. The male bias and high diversity suggest that LDCs are an outcome of intersexual selection, while their elicitation by predators suggests an alarm function. I propose that male magpie-jays use predator encounters as opportunities to advertise their presence and availability as mates; they use LDCs as songs. Such a communication system seems to have been favored by the unusual social system of magpie-jays, in which female groups defend territories and males have little opportunity to defend resources for mate attraction, forcing them to advertise when females are paying the most attention, during predator encounters.