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1.
Escape theory predicts that flight initiation distance (predator–prey distance when escape begins) increases as predation risk increases and decreases as cost of fleeing increases. Scant information is available about the effects of some putative predation risk factors and about interaction between simultaneously operating risk and cost of fleeing factors on flight initiation distance and distance fled. By simulating an approaching predator, I studied the effects of body temperature (BT), distance to nearest refuge, and eye contact with a predator, as well as simultaneous effects of predator approach speed and female presence/absence on escape behavior by a small ectothermic vertebrate, the lizard Sceloporus virgatus. Flight initiation distance decreased as BT increased, presumably because running speed increases as BT increases, facilitating escape. Distance to nearest refuge was unrelated to BT or flight initiation distance. Substrate temperature was only marginally related, and air temperature was not related to flight initiation distance. Eye contact did not affect flight initiation during indirect approaches that bypassed lizards by a minimum of 1 m, but an effect of eye contact found in other studies during direct approach might occur. Predator approach speed and presence of a female interactively affected flight initiation distance, which increased as speed increased and decreased when a female was present. In the presence of a female, flight initiation distance was far shorter than when no female was present. The high cost of forgoing a mating opportunity accounts for the interaction because the difference between female presence and absence is greater when risk is greater.  相似文献   

2.
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].  相似文献   

3.
Theoretical models of anti-predator escape behaviour suggest that prey may adjust their escape response such that the optimal flight distance is the point at which the costs of staying exceed the costs of fleeing. Anti-predatory decisions should be made based also on consequences for long-term expected fitness, such as the costs of refuge use. For example, in lizards, the maintenance of an optimal body temperature is essential to maximize physiological processes. However, if unfavourable thermal conditions of refuges can decrease the body temperature of lizards, their escape decision should be influenced by refuge conditions. Analyses of the variation in flight distances and emergence latency from a refuge for the lizard Lacerta monticola under two different predation risk levels, and their relationship with the thermal environment, supported these predictions. When risk increased, lizards had longer emergence latencies, and thus costs of refuge use increased (a greater loss of time and body temperature). In the low-risk situation, lizards that were farther from the refuge had longer flight distances, whereas thermal conditions were less important. When risk increased, lizards had longer flight distances when refuges were farther off, but also when the external heating rate and the refuge cooling rate were lower. The results suggest that, in addition to the risk of predation, expected long-term fitness costs of refuges can also affect escape decisions.  相似文献   

4.
The amount of risk animals perceive in a given circumstance (i.e. their degree of 'fear') is a difficult motivational state to study. While many studies have used flight initiation distance as a proxy for fearfulness and examined the factors influencing the decision to flee, there is no general understanding of the relative importance of these factors. By identifying factors with large effect sizes, we can determine whether anti-predator strategies reduce fear, and we gain a unique perspective on the coevolution of predator and anti-predator behaviour. Based on an extensive review and formal meta-analysis, we found that predator traits that were associated with greater risk (speed, size, directness of approach), increased prey distance to refuge and experience with predators consistently amplified the perception of risk (in terms of flight initiation distance). While fish tolerated closer approach when in larger schools, other taxa had greater flight initiation distances when in larger groups. The presence of armoured and cryptic morphologies decreased perception of risk, but body temperature in lizards had no robust effect on flight initiation distance. We find that selection generally acts on prey to be sensitive to predator behaviour, as well as on prey to modify their behaviour and morphology.  相似文献   

5.
Animals often evaluate the degree of risk posed by a predator and respond accordingly. Since many predators orient their eyes towards prey while attacking, predator gaze and directness of approach could serve as conspicuous indicators of risk to prey. The ability to perceive these cues and discriminate between high and low predation risk should benefit prey species through both higher survival and decreased energy expenditure. We experimentally examined whether Indian rock lizards (Psammophilus dorsalis) can perceive these two indicators of predation risk by measuring the variation in their fleeing behaviour in response to type of gaze and approach by a human predator. Overall, we found that the gaze and approach of the predator influenced flight initiation distance, which also varied with attributes of the prey (i.e. size/sex and tail-raise behaviour). Flight initiation distance (FID) was 43% longer during direct approaches with direct gaze compared with tangential approaches with averted gaze. In further, exploratory, analyses, we found that FID was 23% shorter for adult male lizards than for female or young male (FYM) lizards. In addition, FYM lizards that showed a tail-raise display during approach had a 71% longer FID than those that did not. Our results suggest that multiple factors influence the decision to flee in animals. Further studies are needed to test the generality of these factors and to investigate the proximate mechanisms underlying flight decisions.  相似文献   

6.
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.  相似文献   

7.
During encounters with predators, prey must balance the degreeof risk against the loss of fitness-enhancing benefits suchas feeding and social activities. Most studies of tradeoffsbetween risk and cost of escaping have measured flight initiationdistance and time to emerge from refuge, for which theory providesrobustly supported predictions. Tradeoffs involving other aspectsof encounters, including distance fled and time between escapeand return to a food source, have received little theoreticalor empirical attention. By adapting models of flight initiationdistance and time between entry into refuge and emergence, wepredict effects of predation risk and cost on distance fledand time to return to a source of benefit after fleeing. Actingas simulated predators that approached at a fixed speed, weconducted an experimental field study to test the hypothesesthat flight initiation distance, distance fled, and time toreturn to food by Balearic lizards (Podarcis lilfordi) decreasewith the presence and amount of insect food. Predictions ofthe models were strongly supported, including those for distancefled and return time, but predictions for other cost factorsand predation risk factors remain to be tested.  相似文献   

8.
Some aspects of escape predicted by theoretical models are intended to apply universally. For example, flight initiation distance (distance between an approaching predator and prey when escape begins) is predicted from predation risk and the costs of escaping. Escape tactics and refuge selection are not currently predicted by theoretical models, but are expected to vary with structural features of the habitat. One way of studying such variation is to compare aspects of antipredatory behavior among sympatric species that differ in habitat or microhabitat use. In an assemblage of lizards in northwestern Namibia, we conducted experiments to test predictions of escape theory for three risk factors in representatives of three families and observed escape tactics in additional species. As predicted by escape theory, flight initiation distance increased with directness of a predator's approach and predator speed in Agama planiceps, Mabuya acutilabris, and Rhotropus boultoni, and with distance from refuge in M. acutilabris. As predicted by theory, the probability of entering refuge increased with risk in R. boultoni. All available data indicate that flight initiation distance and refuge entry by lizards conform to theoretical predictions. Escape tactics varied greatly as a function of habitat type: (1) arboreal species fled up and around trees and sometimes entered tree holes; (2) saxicolous species used rock crevices as refuges, but differed in tactics prior to entering refuges; and (3) terrestrial species fled into bushes or other vegetation, often to the far sides of them. Some M. acutilabris entered small animal burrows or buried themselves in sand beneath bushes. Escape tactics varied even among congeners in Mabuya, highlighting the important effect of habitat structure on them. Although habitat partitioning has traditionally been viewed as favoring species coexistence, an interesting by‐product appears to be structuring of escape tactics in lizard communities.  相似文献   

9.
Flight initiation distance describes the distance at which an animal flees during the approach of a predator. This distance presumably reflects the tradeoff between the benefits of fleeing versus the benefits of remaining stationary. Throughout ontogeny, the costs and benefits of flight may change substantially due to growth-related changes in sprint speed; thus ontogenetic variation in flight initiation distance may be substantial. If escape velocity is essential for surviving predator encounters, then juveniles should either tolerate short flight initiation distances and rely on crypsis, or should have high flight initiation distances to remain far away from their predators. We examined this hypothesis in a small, short-lived lizard (Sceloporus woodi). Flight initiation distance and escape velocity were recorded on an ontogenetic series of lizards in the field. Maximal running velocity was also quantified in a laboratory raceway to establish if escape velocities in the field compared with maximal velocities as measured in the lab. Finally a subset of individuals was used to quantify how muscle and limb size scale with body size throughout ontogeny. Flight initiation distance increased with body size; larger animals had higher flight initiation distances. Small lizards had short flight initiation distances and remained immobile longer, thus relying on crypsis for concealment. Escape velocity in the field did not vary with body size, yet maximum velocity in the lab did increase with size. Hind limb morphology scaled isometrically with body size. Isometric scaling of the hind limb elements and its musculature, coupled with similarities in sprint and escape velocity across ontogeny, demonstrate that smaller S. woodi must rely on crypsis to avoid predator encounters, whereas adults alter their behavior via larger flight initiation distance and lower (presumably less expensive) escape velocities.  相似文献   

10.
Organisms generally have many defenses against predation, yet may lack effective defenses if from populations without predators. Evolutionary theory predicts that “costly” antipredator behaviors will be selected against when predation risk diminishes. We examined antipredator behaviors in Aegean wall lizards, Podarcis erhardii, across an archipelago of land-bridge islands that vary in predator diversity and period of isolation. We examined two defenses, flight initiation distance and tail autotomy. Flight initiation distance generally decreased with declining predator diversity. All predator types had distinctive effects on flight initiation distance with mammals and birds having the largest estimated effects. Rates of autotomy observed in the field were highest on predator-free islands, yet laboratory-induced autotomy increased linearly with overall predator diversity. Against expectation from previous work, tail autotomy was not explained solely by the presence of vipers. Analyses of populations directly isolated from rich predator communities revealed that flight initiation distance decreased with increased duration of isolation in addition to the effects of current predator diversity, whereas tail autotomy could be explained simply by current predator diversity. Although selection against costly defenses should depend on time with reduced threats, different defenses may diminish along different trajectories even within the same predator–prey system.  相似文献   

11.
There are many anti‐predatory escape strategies in animals. A well‐established method to assess escape behavior is the flight initiation distance (FID), which is the distance between prey and predator at which an animal flees. Previous studies in various species throughout the animal kingdom have shown that group size, urbanization, and distance to refuge and body mass affect FID. In most species, FID increases if body mass, group size or distance to refuge decreases. However, how age and sexual dimorphism affect FID is rather unknown. Here, we assess the escape behavior and FID of the black redstart (Phoenicurus ochruros), a small turdid passerine. When approached by a human, males initiated flights later, that is allowing a closer approach than females. Males of this species are more conspicuous, and therefore, may exhibit aposematism to deter potential predators or are less fearful than females. Additionally, juveniles fled at shorter distances and fled to lower heights than adults. Lastly, concerning escape strategy, black redstarts, unless other passerine birds, fled less often into cover, but rather onto open or elevated spots. Black redstarts are especially prone to predation by ambushing predators that might hide in cover. Hence, this species most likely has a higher chance of escaping by fleeing to an open spot rather than to a potentially risky cover.  相似文献   

12.
In predator-prey encounters, many factors influence risk perceptionby prey and their decision to flee. Previous studies indicatethat prey take flight at longer distances when they detect predatorsat longer distances and when the predator's behavior indicatesthe increased likelihood of attack. We examined the flight decisionsof Columbian black-tailed deer (Odocoileus hemionus columbianus)using an approaching human whose speed, directness of approach,directness of gaze, and simulated gun carrying varied. Deerfled at greater distances when approached more quickly and directly,and there was a concave-down quadratic trend in the relationshipbetween the distances at which the predator began its approachand at which the deer became alert (alert distance [AD]), indicatingthat deer have a zone of awareness beyond which there is a delayin detecting an approaching predator. Time spent assessing theapproacher (assessment time) was shorter during faster approachesand was positively related with AD. Deer fled at longer distancesand had shorter assessment times when they were already alertto the predator at the initiation of approach. Males fled atshorter distances than females when approached during the gun-holdingcondition, and males had shorter assessment times than femaleswhen the approacher averted his gaze. Such sex differences inrisk assessment might reflect male motivation during the matingseason as well as exposure to human hunting. We suggest thatrisk assessment is affected the by the predator's behavior,the state of awareness of the prey, and the distance at whichthey detect the predator.  相似文献   

13.
Exotic predators have driven the extinction of many island species. We examined impacts of feral cats on the abundance and anti-predator behaviours of Aegean wall lizards in the Cyclades (Greece), where cats were introduced thousands of years ago. We compared populations with high and low cat density on Naxos Island and populations on surrounding islets with no cats. Cats reduced wall lizard populations by half. Lizards facing greater risk from cats stayed closer to refuges, were more likely to shed their tails in a standardized assay, and fled at greater distances when approached by either a person in the field or a mounted cat decoy in the laboratory. All populations showed phenotypic plasticity in flight initiation distance, suggesting that this feature is ancient and could have helped wall lizards survive the initial introduction of cats to the region. Lizards from islets sought shelter less frequently and often initially approached the cat decoy. These differences reflect changes since islet isolation and could render islet lizards strongly susceptible to cat predation.  相似文献   

14.
Males of many species show conspicuous breeding colours that are important for status signalling, but that may decrease crypsis and increase predation risk. However, prey may adjust their escape response, such that the optimal distance at which an animal starts to flee (approach distance) would be the point where the costs of staying exceed the cost of fleeing. We examined in the field the escape response of Psammodromus algirus lizards, to test the hypothesis that more conspicuous old males, showing orange nuptial coloration on most of the head, which presumably have a higher probability of being detected (i.e. increased costs of staying), have longer approach distances, independent of other environmental variables. As predicted, old brightly coloured males had significantly longer approach and flight distances than young dull males, and young males had longer ones than females. In contrast, neither the distance to the nearest refuge nor the air temperature when the lizards escaped were significantly different. In addition, although male and female lizards differed in their use of microhabitats, old and young males did not differ. We also found that old males guarding a female (i.e. increased costs of fleeing) had shorter approach distances than old males that were found alone.  相似文献   

15.
Most ecomorphological studies use a comparative approach to examine adaptation by studying variation among species. A question of considerable interest is whether ecomorphological patterns observed among species also exist at the population level. We studied variation in morphology, performance, and behaviour in four populations of Leiocephalus personatus and two populations of Leiocephalus barahonensis in the Dominican Republic. We combined these data with measurements of predation intensity and habitat structure to test for convergence at the population level. We predicted that predation intensity would be higher in open habitats and that lizards in these habitats would have traits conferring higher predator evasion capacity (increased wariness, faster sprint speeds, and longer limbs). Principal components analysis suggests that sites tend to differ with respect to the abundance and spacing of low-lying vegetation (i.e. percentage of shrub cover and distance to nearest vegetation), but we did not detect any striking differences among sites in tail-break frequencies or attacks on clay lizard models. Consistent with predictions we find that in open habitats, lizards tend to have longer limbs, faster sprint speeds (relative to body size), and longer approach distances. These patterns corroborate findings in other ground-dwelling lizard species and indicate that they have evolved at least twice among populations of Leiocephalus lizards. The results of this study also suggest that these traits have evolved rapidly despite recent or ongoing gene flow.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 445–456.  相似文献   

16.
Island tameness (reduced escape behaviour on islands where prey have experienced prolonged relaxation of predation pressure) is known in several taxa, although the relationships between recent predation pressure and escape on islands are poorly known. We investigated escape by numerous populations exposed to differing predation pressure of two sister species of Podarcis lizards in the Balearic Islands. Our main findings are that flight initiation distance was greater in Podarcis pityusensis than Podarcis lilfordi and increased as predation pressure increased in P. pityusensis. Island tameness led to extinction of P. lilfordi on Menorca and Mallorca following anthropogenic introduction of predators; this species is extant only on nearby islets. The lack of relationship between recent predation pressure and flight initiation distance in P. lilfordi indicates that the historically acquired deficit in the ability to adjust escape behaviour to predation pressure still exists. Podarcis pityusensis, which was exposed to greater natural predation pressure before human introduction of predators, survives on Ibiza and Formentera, as well as on islets. Retention of the ability to respond to predation pressure is consistent with our finding that flight initiation distance increases as predation pressure increases among current populations. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ?? , ??–??.  相似文献   

17.
Most studies of predator avoidance behaviours have focussed on single‐predator systems, despite the fact that prey often are confronted with predator rich environments. In the presence of more than one predator, prey may have to choose between avoiding one predator over another. How prey cope with exposure to several enemies simultaneously remains largely untested. In this study I set out to investigate if skinks showed preferential avoidance of snake odours based on the relative predation risk posed by different snake species. This relative predation risk was estimated using information on density, diet specificity and foraging habit of each snake species. I tested retreat‐site selection in two‐choice tests, where lizards chose between different combinations of control and snake treated retreat‐sites as well as two retreat‐sites treated with different snake species odours. Lizards preferred control–treated retreat‐sites to those treated with snake odours and showed a differential avoidance response to refuges treated with odours from different snake species. There was strong evidence to suggest that lizards preferentially avoided refuges with the odours of the snake that posed the greatest predation risk, the white‐lipped snake (Drysdalia coronoides). Naïve juvenile lizards were also tested and their response was similar to the adults demonstrating that the behaviour is innate and not the result of higher encounter rates of more common snake odours. To my knowledge this is one of the first studies to demonstrate that prey can prioritize avoidance to a single most dangerous predator in the face of several predators and conflicting avoidance responses.  相似文献   

18.
Some prey may signal to deter pursuit by predators. Because deterrence is not needed when risk is low or useful when capture is imminent, most signaling should occur at intermediate risk. Probability of fleeing increases with risk for various risk factors. At low–intermediate risk, more frequent signaling should occur as assessed risk associated with risk factors increases. I examined the effects of three risk factors related to immobility and movement by a predator: standing distance (distance from prey to immobile predator), directions of walking, and turning by the predator. Risk is greater when the predator stands nearer, walks toward prey vs. retreating, and turns toward prey vs. away. In the lizard Callisaurus draconoides, which signals by elevating and waving its tail, signaling was more frequent before fleeing when I stood immobile at the shorter of two distances. All the lizards fled when I walked toward them, regardless of standing distance. Fewer fled when I moved away and only at the shorter standing distance. At the shorter standing distance, signal probability was high and did not differ between movement directions. At the longer standing distance, fewer lizards signaled and only when I moved toward them. Patterns of response of signaling and escape to combinations of standing distance and turn direction were qualitatively identical. When I turned away from lizards, none displayed or fled at the longer standing distance. At the shorter standing distance, probabilities of displaying and fleeing were higher when I turned toward than away from lizards. Standing distance affected signaling interactively with directions of movement and turning in manners readily interpretable from risk. Signaling was affected by risk associated with all factors, being absent or infrequent at both high‐ and low‐risk levels but frequent at intermediate risk, strengthening evidence for pursuit‐deterrent signaling.  相似文献   

19.
The ability to assess and respond to predation risk is a strong selective force. Detection of predators is carried out by one or more sensory modalities, but the use of chemoreception has significant advantages. This study examines the chemosensorial assessment of snake predation risk and corresponding behaviours in different species and populations of Liolaemus lizards naturally exposed to different levels of snake predation pressure. The species studied were sympatric (Liolaemus lemniscatus), parapatric (L. nigroviridis) and allopatric (L. fitzgeraldi) to the saurophagous snake, Philodryas chamissonis. Additionally, two populations of L. lemniscatus from areas differing in snake densities were compared. Chemo‐assessment of predation risk was determined by comparing the number of tongue‐flicks (TF) and the behavioural responses of lizards submitted to three treatments (with semiochemicals of snake, conspecifics, and without semiochemicals – control). The results suggest that Liolaemus lizards can chemo‐assess snake predation risk, but this was modulated by the predation pressure experienced by lizards in their natural habitats. When exposed to snake semiochemicals, the sympatric prey showed less chemical exploratory behaviour (i.e. lower number of TF), a higher frequency of antipredator behaviours that would reduce its detection by a predator, and did not show the behaviour triggered by conspecific semiochemicals. The parapatric prey showed similar number of TF across different treatments, suggesting an absence of recognition of snake semiochemicals; however, it did show antipredatory behaviours when confronted with snake semiochemicals. The allopatric prey showed similar behaviour in all treatments. Both populations of the sympatric species, L. lemniscatus, showed a similar ability to detect predation risk when confronted with snake semiochemicals (i.e. similar number of TF), but antipredatory behaviours were diminished, and marking behaviours were present in the population subject to lower predation pressure. Relaxed predation pressure from a predator that releases and detects semiochemicals had similar consequences at species and population levels.  相似文献   

20.
The evolution of striking phenotypes on islands is a well‐known phenomenon, and there has been a long‐standing debate on the patterns of body size evolution on islands. The ecological causes driving divergence in insular populations are, however, poorly understood. Reduced predator fauna is expected to lower escape propensity, increase body size and relax selection for crypsis in small‐bodied, insular prey species. Here, we investigated whether escape behaviour, body size and dorsal coloration have diverged as predicted under predation release in spatially replicated islet and mainland populations of the lizard species Podarcis gaigeae. We show that islet lizards escape approaching observers at shorter distances and are larger than mainland lizards. Additionally, we found evidence for larger between‐population variation in body size among the islet populations than mainland populations. Moreover, islet populations are significantly more divergent in dorsal coloration and match their respective habitats poorer than mainland lizards. These results strongly suggest that predation release on islets has driven population divergence in phenotypic and behavioural traits and that selective release has affected both trait means and variances. Relaxed predation pressure is therefore likely to be one of the major ecological factors driving body size divergence on these islands.  相似文献   

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