Stop-and-go approach by a predator: a novel predation risk factor for the phrynosomatid lizards Sceloporus virgatus and Callisaurus draconoides

2015 ◽  
Vol 36 (4) ◽  
pp. 401-409
Author(s):  
William E. Cooper ◽  
Wade C. Sherbrooke
Keyword(s):  
Field Experiments ◽  
Empirical Studies ◽  
Pause Duration ◽  
Predator Prey ◽  
Escape Theory ◽  
Stop And Go ◽  
Approach Speed ◽  
Sceloporus Virgatus ◽  
Escape Behaviors ◽  
Flight Initiation

As an immobile prey monitors an approaching predator, the predator may move at a constant speed directly toward the prey or on a path that bypasses the prey. These scenarios have been studied extensively. Economic escape theory successfully predicts flight initiation distance (FID = predator-prey distance when escape begins). However, predators often alter their speed and may exhibit stops and starts during approaches. Empirical studies have shown that prey rapidly adjust assessed risk to a predator’s changes in approach speed and direction, but effects of interrupted (stop-start) approach are unknown. Because a prey is likely to assess that a nearby predator that resumes approaching has detected it and is attacking, escape theory predicts that assessed risk is greater at a given predator-prey distance when approach resumes than is continuous. Therefore, we predicted that FID is longer when a predator approaches, stops nearby, and renews its approach than when it approaches continuously. Second, although assessed risk increases as duration of the predator’s stop nearby increases, as indicated by latency to flee, we predicted that pause duration does not affect FID because prey interpret resumed approach as attack. Field experiments with two lizards, Sceloporus virgatus and Callisaurus draconoides, verified the predictions: FID was longer for discontinuous than continuous approaches and pause duration did not affect FID. We also observed distance fled and probability of entering refuge, escape behaviors for which theory is undeveloped. Distance fled was unrelated to continuity of approach in both species, as was refuge entry in S. virgatus.

Compensatory changes in escape and refuge use following autotomy in the lizard Sceloporus virgatus

2007 ◽  
Vol 85 (1) ◽  
pp. 99-107 ◽  
Author(s):  
W.E. Cooper, Jr.
Keyword(s):  
Body Part ◽  
Control Group ◽  
Flight Initiation Distance ◽  
Refuge Use ◽  
Predator Prey ◽  
Escape Theory ◽  
Antipredatory Strategies ◽  
Sceloporus Virgatus ◽  
Flight Initiation ◽  
Predator Effects

Following autotomy of a body part to escape from a predator, prey may alter antipredatory strategies to compensate for the inability to use autotomy and impaired escape ability. Because prey that have been captured may increase their assessment of risk posed by a predator, effects of capture may have been attributed to autotomy. I conducted an experiment using three groups of striped plateau lizards ( Sceloporus virgatus Smith, 1938): control, captured, and autotomized. Captured and autotomized lizards were less active on the day after autotomy than controls. Flight initiation distance and distance fled were greater in both experimental groups than in the control group, but did not differ between experimental groups. Flight initiation distance was greater in autotomized than in captured lizards only in males. No sex difference occurred for distance fled. Autotomized lizards entered refuges more than other groups. Escape strategy changed after autotomy to increased reliance on refuge and, in males, to increased flight initiation distance; behaviours that are appropriate to compensate for impaired escape ability. Decreased activity and increased distance fled might erroneously have been considered effects of autotomy, had effects of capture not been assessed. Predictions of escape theory that flight initiation distance and distance fled increase with predation risk were supported.

Effects of predation risk factors on escape behavior by Balearic lizards (Podarcis lilfordi) in relation to optimal escape theory

2009 ◽  
Vol 30 (1) ◽  
pp. 99-110 ◽  
Author(s):  
Dror Hawlena ◽  
Valentín Pérez-Mellado ◽  
William Cooper
Keyword(s):  
Risk Factors ◽  
Predation Risk ◽  
Escape Behavior ◽  
Weather Conditions ◽  
Predator Prey ◽  
Escape Theory ◽  
Approach Speed ◽  
Podarcis Lilfordi ◽  
Effects Of Temperature ◽  
Flight Initiation

AbstractEscape theory predicts that flight initiation distance (FID = predator-prey distance when escape begins) increases as predation risk increases. We tested effects of variation of approach speed and directness, predator persistence, concealment, and weather conditions on FID in the Balearic lizard (Podarcis lilfordi) by ourselves simulating predators. We examined effects of directness of approach on probability of fleeing and of repeated approach on entering refuge and distance fled. As predicted, FID was greater for faster approach speed, more direct approach, during second than first approaches, and when lizards were exposed than partially concealed. Other effects of directness of approach and repeated approach also were as predicted by greater assessed risk by the lizards. The proportion of individuals that fled was greater for direct than indirect approaches. The proportion of lizards that entered refuges and distance fled were greater during the second of two successive approaches. Effects of weather on FID were complex. FID was shortest in the warmest conditions with no noticeable wind, when lizards were active. Lizards were inactive and basked in the other conditions. FID was longest at 20°C without wind, and intermediate FID occurred at 18°C in windy conditions. We present hypotheses for weather effects. Tests are needed to unravel effects of temperature and wind speed. All predictions of escape theory for simple risk factors, i.e., all except than weather conditions, were confirmed. Escape theory successfully predicts FID for these risks in P. lilfordi, other lacertids, and more broadly, in ecologically and taxonomically diverse lizards.

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

2009 ◽  
Vol 55 (2) ◽  
pp. 123-131 ◽  
Author(s):  
William E. Cooper
Keyword(s):  
Risk Factors ◽  
Risk Assessment ◽  
Predation Risk ◽  
Flight Initiation Distance ◽  
Risk Levels ◽  
Escape Theory ◽  
Growing Body ◽  
Sceloporus Virgatus ◽  
Escape Behaviors ◽  
Flight Initiation

Abstract 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.

Age, sex and escape behaviour in the Striped Plateau Lizard (Sceloporus virgatus) and the Mountain Spiny Lizard (S. jarrovii), with a review of age and sex effects on escape by lizards

Behaviour ◽  
2011 ◽  
Vol 148 (11-13) ◽  
pp. 1215-1238 ◽  
Author(s):  
William E. Cooper Jr.
Keyword(s):  
Sex Differences ◽  
Relevant Literature ◽  
Lizard Species ◽  
Escape Behaviour ◽  
Predator Prey ◽  
Escape Strategy ◽  
Escape Theory ◽  
Multiple Components ◽  
Sceloporus Virgatus ◽  
Age And Sex

AbstractEscape behaviour often differs between sexes, reproductive states and ages. Escape theory predicts that flight initiation distance (FID = predator–prey distance when escape begins) increases as predation risk and fitness increase, and decreases as cost of escaping increases. Similar predictions hold for distance fled and refuge entry, suggesting that age and sex differences in escape behaviour may occur when risk, fitness, and opportunity costs differ. I studied such differences in two lizard species and reviewed relevant literature on escape by lizards. In Sceloporus virgatus no difference occurred between sexes or female reproductive states in FID, distance fled, distance from refuge, or probability of entering refuge. In S. jarrovii juveniles had shorter FID and distance fled than adults; juveniles were closer than females to refuge, but this did not affect FID or distance fled. Juveniles were more likely than adults to be on rocks and use them as refuges. The literature review showed that sexual dimorphism in FID occurs in about 1/5 of species (male FID usually > female FID), but distance fled differed between sexes in only 1 of 21 species. Juveniles had shorter FID than adults in all of five species; the relationship between age and distance fled was highly variable. Reasons for patterns of age/sex differences are discussed. Because age and sex differences in these factors and escape strategy can alter multiple components affecting optimality, sometimes in opposite ways, these factors and escape strategy must be known to predict effects of age, sex and reproductive state on escape.

scholarly journals Empirical studies of escape behavior find mixed support for the race for life model

2021 ◽  
Author(s):  
Kwasi Wrensford ◽  
Jahaziel Gutierrez ◽  
William E Cooper ◽  
Daniel T Blumstein
Keyword(s):  
Escape Behavior ◽  
Empirical Studies ◽  
Relative Orientation ◽  
Similar Amount ◽  
Predator Prey ◽  
Future Studies ◽  
Escape Theory ◽  
Mixed Support ◽  
Life Model ◽  
The Relationship

Abstract Escape theory has been exceptionally successful in conceptualizing and accurately predicting effects of numerous factors that affect predation risk and explaining variation in flight initiation distance (FID, predator-prey distance when escape begins). Less explored is the relative orientation of an approaching predator, prey, and its eventual refuge. The relationship between an approaching threat and its refuge can be expressed as an angle we call the “interpath angle” or “Φ”, which describes the angle between the paths of predator and prey to the prey’s refuge and thus expresses the degree to which prey must run towards an approaching predator. In general, we might expect that prey would escape at greater distances if they must flee toward a predator to reach its burrow. The ‘race for life’ model makes formal predictions about how interpath angle should affect FID. We evaluated the model by studying escape decisions in yellow-bellied marmots Marmota flaviventer, a species which flees to burrows. We found support for some of the model’s predictions, yet the relationship between interpath angle and FID was less clear. Marmots may not assess interpath angle in a continuous fashion; but we found that binning angle into four 45° bins explained a similar amount of variation as models that analyzed angle continuously. Future studies of interpath angle, especially those that focus on how different species perceive relative orientation, will likely enhance our understanding of its importance in flight decisions.

Beyond optimal escape theory: microhabitats as well as predation risk affect escape and refuge use by the phrynosomatid lizard Sceloporus virgatus

Behaviour ◽  
2007 ◽  
Vol 144 (10) ◽  
pp. 1235-1254 ◽  
Author(s):  
Dawn Wilson ◽  
William Cooper
Keyword(s):  
Risk Factors ◽  
Individual Differences ◽  
Predation Risk ◽  
Flight Initiation Distance ◽  
Refuge Use ◽  
Escape Theory ◽  
Sceloporus Virgatus ◽  
Flight Initiation

AbstractEscape studies often focus on one variable, but tactics and refuge use vary with microhabitats, exposure, distance to refuge, and temperature. We studied these effects and effects of microhabitats and risk factors (distance from refuge, temperature) on flight initiation distance (FID, distance between predator and prey when escape begins) and distance fled (DF) in the lizard Sceloporus virgatus. FID increased as distance to refuge increased and temperature decreased. DF increased as FID increased, supporting the hypothesis that individual differences in boldness are consistent among encounter phases. Refuges were rock crevices, trees, logs, and grass clumps. Interhabitat differences in FID and DF matched those in distance to refuge. FID was longer for lizards on rocks and ground than trees due to proximity to and use of refuge. Lizards on trees rarely changed microhabitats, moving to the far side and unpredictably up or down. Lizards on slopes fled long distances up slopes. Most lizards on rocks entered crevices or switched microhabitats. Lizards on ground usually changed microhabitats. Optimal escape theory accurately predicted effects of risk on FID, but initial microhabitats and final microhabitats and refuges affected tactics, FID and DF. DF was affected by risk, being longer when lizards remained visible.

Risk factors and escape strategy in the grasshopper Dissosteira carolina

Behaviour ◽  
2006 ◽  
Vol 143 (10) ◽  
pp. 1201-1218 ◽  
Author(s):  
William E. Cooper
Keyword(s):  
Risk Factors ◽  
Field Experiments ◽  
Escape Behavior ◽  
Sudden Change ◽  
Flight Initiation Distance ◽  
Initial Direction ◽  
Escape Strategy ◽  
Approach Speed ◽  
Lateral Escape ◽  
Flight Initiation

AbstractTo determine aspects of escape strategy by the Carolina grasshopper, Dissosteira carolina, and applicability of models of escape behavior applied primarily to vertebrates, I conducted three field experiments by simulating an approaching predator. Escape theory predicts that flight initiation distance (distance from predator when escape begins) and distance fled increase as predation risk increases. Some aspects of escape are not predicted, and theory does not identify escape strategies including several components. I examined effects of risk factors (predator approach speed, directness of approach, and repeated approach) on flight initiation distance, distance fled, and the initial direction of escape. Flight initiation distance and distance fled were predicted to increase with approach speed. Because predators approaching directly may bypass prey without detecting it, probability of fleeing and flight initiation distance were predicted to increase with directness of approach. Because a persistent predator poses greater threat, flight initiation distance and distance fled were predicted to be greater for the second of two successive approaches. All findings were consistent with predictions of the Ydenberg & Dill (1986) model, suggesting that risk assessment and escape decisions by visually oriented insects may be similar to those of vertebrates. Although escape directly away from the predator might be expected to minimize risk, most grasshoppers escaped by flying at nearly right angles to the approach path. Lateral escape may be part of an escape strategy in which dark wing colour during flight rapidly disappears upon landing. With sudden change in colour and movement, the grasshopper becomes cryptic and difficult to relocate. Lateral escape may increase difficulty of maintaining visual contact with the grasshopper until it lands. It also avoids need for further escape from a predator that continues in its initial direction.

When and how do predator starting distances affect flight initiation distances?

2005 ◽  
Vol 83 (8) ◽  
pp. 1045-1050 ◽  
Author(s):  
W E Cooper, Jr.
Keyword(s):  
Bird Species ◽  
Ecological Factors ◽  
Flight Initiation Distance ◽  
Lizard Species ◽  
Urosaurus Ornatus ◽  
Approach Speed ◽  
Monitoring Costs ◽  
Sceloporus Virgatus ◽  
The Relationship ◽  
Flight Initiation

The distance separating predator and prey when the predator begins to approach, starting distance, was recently shown to affect flight initiation distance in many bird species, raising questions about the effect's generality, variation with ecological factors, and economic basis. I studied the effect in two lizard species that forage by ambush and escape into nearby refuges. Monitoring costs during approach are absent because ambushers remain immobile while scanning for prey and predators. Risks are minimized because of the proximity to refuge. Flight initiation distance increased weakly with starting distance in Sceloporus virgatus Smith, 1938 significantly only at rapid approach speed. It was not significant in Urosaurus ornatus (Baird and Girard, 1852) at slow approach speed. Flight initiation distance is predicted to increase with starting distance, owing to monitoring costs and assessment by prey of greater risk during prolonged approaches. The significant effect in S. virgatus, which lacks monitoring costs, is the first indication that risk affects the relationship between starting distance and flight initiation distance. Conditions in which starting distance is important and its possible effects in earlier studies are discussed, as well as standardizing approaches and possible artifactual effects of starting distance.

FEAR and DREAD: starting distance, escape decisions and time hiding in refuge

Behaviour ◽  
2015 ◽  
Vol 152 (10) ◽  
pp. 1371-1389 ◽  
Author(s):  
William E. Cooper ◽  
Wade C. Sherbrooke
Keyword(s):  
Flight Initiation Distance ◽  
Predator Prey ◽  
Sceloporus Virgatus ◽  
Flight Initiation

Flight initiation distance (FID = predator–prey distance when escape begins) increases as starting distance (predator–prey distance when approach begins) increases. The flush early and avoid the rush (FEAR) hypothesis proposes that this relationship exists because monitoring an approach is costly. Hypothesized causes are increase in assessed risk and decrease in obtainable benefits while monitoring as starting distance increases. We propose the delay risking emergence and avoid dying (DREAD) hypothesis: hiding time in refuge increases as starting distance increases because prey use risk assessed during approach to estimate risk upon emerging. In the lizard Callisaurus draconoides, FID increased as standardized starting distance increased at faster approach speeds, supporting the FEAR hypothesis. In its first test, the DREAD hypothesis was supported: hiding time in the lizard Sceloporus virgatus increased as standardized starting distance increased. No large benefits were attainable, suggesting that dynamic increase in assessed risk accounts for these findings.

Conspicuousness and vestigial escape behaviour by two dendrobatid frogs, Dendrobates auratus and Oophaga pumilio

Behaviour ◽  
2009 ◽  
Vol 146 (3) ◽  
pp. 325-349 ◽  
Author(s):  
Janalee Caldwell ◽  
Laurie Vitt ◽  
William Cooper
Keyword(s):  
Defensive Behavior ◽  
Close Approach ◽  
Flight Initiation Distance ◽  
Escape Behaviour ◽  
Predator Prey ◽  
Poison Frogs ◽  
Oophaga Pumilio ◽  
Approach Speed ◽  
Dendrobates Auratus ◽  
Flight Initiation

AbstractAposematic prey are thought to move slowly and openly near predators, but exhibit reduced escape behaviour. We studied conspicuousness and escape by aposematic poison frogs (Dendrobates auratus and Oophaga pumilio). In circles of leaf litter, observers detected poison frogs quickly. Flight initiation distance (FID, predator-prey distance when escape begins) increases with approach speed in non-cryptic palatable prey, but not for frogs in clearings, which permitted close approach. On trails frogs moved slowly into forest and FID in D. auratus increased with approach speed. Distance from cover and handling exposing predators to distastefulness may account for greater reliance on aposematism in clearings. We observed responses to a simulated predator (stick with painted face) in three conditions: not approached, approached, and touched. Latency to hop and time to exit circles decreased and exit from circles was directed further away from the approach path in the order not approached, approached, touched. Oophaga pumilio changed directions less when approached than not; many exhibited no escape behaviour. Aposematic dendrobatids move slowly near predators, but retain risk-assessment mechanisms due to occasional predation. Differences in escape between dendrobatids and palatable Craugastor frogs suggest that dendrobatid defensive behavior may have been molded to maximize the effectiveness of aposematism.

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