Risk factors and escape strategy in the grasshopper Dissosteira carolina

Behaviour ◽  
2006 ◽  
Vol 143 (10) ◽  
pp. 1201-1218 ◽  
Author(s):  
William E. Cooper

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.

2009 ◽  
Vol 30 (1) ◽  
pp. 99-110 ◽  
Author(s):  
Dror Hawlena ◽  
Valentín Pérez-Mellado ◽  
William Cooper

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.


2010 ◽  
Vol 31 (3) ◽  
pp. 363-373 ◽  
Author(s):  
Arián Avalos ◽  
William Cooper, Jr.

AbstractEconomic theories of antipredatory behavior take into account expected fitness losses and gains to predict escape decisions. Prey at greater risk are predicted to have longer flight initiation distance (= distance from predator when escape begins), flee farther, be more likely to enter refuge, and have longer hiding time (= time between entering and exiting refuge). We simulated predators to study effects of risk factors in the lizard Sceloporus jarrovii. Flight initiation distance, distance fled and probability of entering refuge increased with approach speed. Flight initiation distance increased additively with increases in speed and directness of approach. Lizards habituated to human presence had shorter flight initiation distances than unhabituated lizards. As predicted from greater threat posed by a persistent predator, flight initiation distance and hiding time were longer after the second of two approaches. Fleeing was more likely when an investigator stood nearby and turned toward them than when farther way or turned away. These findings verify theoretical predictions of escape and refuge use for several risk factors and most are consistent with those for other lizards. However, speed and directness of approach had additive effects in S. jarrovii, but interacted in other species. Another novel finding was a small interaction between individual investigator and directness of approach. Although outcomes of tests of escape and refuge use theories consistently confirm qualitative predictions, quantitative comparisons between studies by different investigators may have limited value unless information is available about relative responses by prey to each investigator.


2015 ◽  
Vol 36 (4) ◽  
pp. 401-409
Author(s):  
William E. Cooper ◽  
Wade C. Sherbrooke

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.


1999 ◽  
Vol 77 (4) ◽  
pp. 671-674 ◽  
Author(s):  
William E Cooper, Jr.

Current models of optimal antipredation behavior do not apply to prey blocked by a predator from access to the primary refuge because the predator is closer than the optimal approach distance and flight toward the refuge would increase risk. If other alternative refuges are available, the prey should flee toward the best alternative one. I studied the effect of an approaching human simulated predator interposed between prey and refuge on the use of alternative refuges and on flight-initiation distance in the keeled earless lizard, Holbrookia propinqua. When the predator approached on a line between a lizard and its closest refuge, the lizard invariably fled to or toward an alternative refuge. Lizards were significantly more likely to use alternative refuges than lizards approached on a line connecting the closest refuge, prey, and predator, but with the lizard between the predator and the refuge. Flight-initiation distance was significantly greater for lizards having free access to the closest refuge than for those blocked from it, perhaps because of the time required to assess the new risk posed by blockage of the closest refuge, to select the best alternative refuge, or to wait for the predator to commit to a closing pattern before choosing the best flight option.


Behaviour ◽  
2007 ◽  
Vol 144 (10) ◽  
pp. 1235-1254 ◽  
Author(s):  
Dawn Wilson ◽  
William Cooper

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.


2006 ◽  
Vol 84 (4) ◽  
pp. 495-504 ◽  
Author(s):  
W.E. Cooper

Approach distance (flight initiation distance) and escape methods depend on predation risk. I studied escape methods and effects of risk factors (temperature, perch height and orientation, conspicuousness) on approach distance in seven Puerto Rican anoles. Approach distance increased as temperature decreased in Anolis gundlachi Peters, 1876, presumably because of decreasing running speed, but not in other species (probably because of narrow temperature range). Perch height and approach distance varied inversely in four arboreal species that escape upward, positively in two grass–bush species that are more conspicuous when higher and flee downward, and were unrelated in cryptic Anolis stratulus Cope, 1861. Approach distance was shortest in cryptic A. stratulus and shorter intraspecifically in three species for partially concealed lizards and at sites providing more cover in two species. Approach distance was shorter for A. gundlachi on vertical than nonvertical perches, suggesting that ease of escape upward affects assessed risk. Escape behaviours have been proposed to vary among anole ecomorphs. Grass–bush species fled downward as proposed, or horizontally near ground level. Contrary to predictions of escape downward by trunk–ground and upward by trunk–crown anoles, all arboreal species escaped upward. Only trunk anoles were proposed to use squirreling, but species from four ecomorphs did as well.


Ethology ◽  
2019 ◽  
Vol 125 (7) ◽  
pp. 430-438 ◽  
Author(s):  
Nadine Kalb ◽  
Fabian Anger ◽  
Christoph Randler

2005 ◽  
Vol 83 (8) ◽  
pp. 1045-1050 ◽  
Author(s):  
W E Cooper, Jr.

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.


2009 ◽  
Vol 55 (2) ◽  
pp. 123-131 ◽  
Author(s):  
William E. Cooper

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.


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