Magnitude of food reward affects escape behavior and acceptable risk in Balearic lizards, Podarcis lilfordi

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.     

Effects of risk assessment, predator behavior, and habitat on escape behavior in Columbian black-tailed deer

The relationship between preflight risk assessment by prey andthe escape behaviors they perform while fleeing from predatorsis relatively unexplored. To examine this relationship, a humanobserver approached groups of Columbian black-tailed deer (Odocoileushemionus columbianus), varying his behavior to simulate moreor less threatening behavior. We measured the focal deer's angleof escape, distance moved during flight, duration of trottingand stotting behavior, and change in elevation during flight.Analyses revealed positive relationships between the distancemoved during flight and the distance at which they fled. Whenflight was initiated when the approacher was close, deer fledrelatively shorter distances and took flight paths at more acuteangles, a property that would force a real predator to changedirection suddenly. Our results indicate that deer do not compensatefor allowing the observer to approach more closely by fleeinggreater distances. Rather, distance moved and flight initiationdistance are linked by level of reactivity and habituation:more reactive or less habituated deer both flee at a greaterdistance and move away to a greater distance during flight.More threatening behavior by the approacher led to longer durationsof rapid flight behavior (e.g., trotting and stotting), anddeer tended to flee uphill and into taller vegetation, usingthese landscape features as refuge from danger. Finally, weprovide the first evidence for Pitcher's untested "antiambush"hypothesis for the function of stotting and discuss its significance.In general, both preflight predator behavior and habitat featuresinfluence both duration and direction of escape.     

Effects of risk, cost, and their interaction on optimal escape by nonrefuging Bonaire whiptail lizards, Cnemidophorus murinus

Optimal escape theory seeks to explain variation in the distanceto an approaching predator at which the prey initiates escape(flight initiation distance). Flight initiation distance increaseswhen predators pose a greater threat and decreases when escapecosts increase. Although optimal escape theory has been highlysuccessful, its predictions have been tested primarily for speciesthat escape to discrete refuges, and most studies have focusedon single risk or cost factors. We present data from two experimentsin which two risks or a risk and a cost varied in Bonaire whiptaillizards (Cnemidophorus murinus) that escaped without enteringrefuges. Our data verify several predictions about optimal escapefor nonrefuging lizard prey. Two risk factors, speed and directnessof approach by the predator, interacted. Directly approachedlizards had greater flight initiation distances than did indirectlyapproached lizards when approached rapidly, but shorter flightinitiation distances when approached slowly. Flight initiationdistance was shorter in the presence of food and during slowversus rapid approaches, but contrary to expectation, food presenceand approach speed did not interact. This would be explainedif cost curves are nonlinear or if they are parallel ratherthan intersecting when the predator reaches the prey. More empiricalwork is needed to determine which risk and cost factors actadditively and which act synergistically. The absence of interactionbetween the risk and cost factors suggests that cost curveswere nonlinear.     

Island tameness: living on islands reduces flight initiation distance

One of Darwin''s most widely known conjectures is that prey are tame on remote islands, where mammalian predators are absent. Many species appear to permit close approach on such islands, but no comparative studies have demonstrated reduced wariness quantified as flight initiation distance (FID; i.e. predator–prey distance when the prey begins to flee) in comparison with mainland relatives. We used the phylogenetic comparative method to assess influence of distance from the mainland and island area on FID of 66 lizard species. Because body size and predator approach speed affect predation risk, we included these as independent variables. Multiple regression showed that FID decreases as distance from mainland increases and is shorter in island than mainland populations. Although FID increased as area increased in some models, collinearity made it difficult to separate effects of area from distance and island occupancy. FID increases as SVL increases and approach speed increases; these effects are statistically independent of effects of distance to mainland and island occupancy. Ordinary least-squares models fit the data better than phylogenetic regressions, indicating little or no phylogenetic signal in residual FID after accounting for the independent variables. Our results demonstrate that island tameness is a real phenomenon in lizards.     

Sex and social costs of escaping in the striped plateau lizard Sceloporus virgatus

As a predator approaches, prey base decisions about when toflee on a balance between degree of predation risk and costsof escaping. Lost opportunities to perform activities that mayincrease fitness are major escape costs. Paramount among theseare chances to increase fitness by courting and mating and bydriving away sexual rivals. Because sexual selection imposesdifferent social demands on the sexes, social opportunitiescan have different consequences for males and females, but effectsof sex differences in social opportunity costs on escape behaviorare unknown. We conducted a field experiment showing that malestriped plateau lizards (Sceloporus virgatus) given the opportunityto court or perform aggressive behavior permit closer approachbefore fleeing, but females do not. Males allowed a simulatedpredator to approach closer before initiating escape if a tetheredmale or female rather than a control stimulus was introducedto them, but females initiated escape at similar distances inall conditions. For males, a trade-off between the greater predationrisk accepted before fleeing due to the likelihood of enhancingfitness by sexual or aggressive behavior accounts for closerapproach allowed in the presence of conspecifics. Mating opportunitiesare not limiting for females in most species and females oftenhave little to gain by interacting aggressively with other females.Therefore, presence of a conspecific male or female may notjustify taking greater risk. Results confirm the predictionof optimal escape theory that flight initiation decreases ascost of escaping increases. The sex difference in effect ofpresence of conspecifics on flight initiation distance is aconsequence of the sex difference in costs of escaping.     

Does Locomotor Ability Influence Flight Initiation Distance in Yellow‐Bellied Marmots?

Flight initiation distance (FID) is the distance between a potential threat and the point at which a potential prey flees. Animals may modify their FID to compensate for increased risk generated by external/extrinsic factors such as habitat type, visibility, group size, time of year, predator‐approach velocity, and distance to burrow, as well as internal/intrinsic factors such as physical condition, body temperature, crypsis, and morphological antipredator defenses. The intrinsic speed at which an animal can escape a predator is a factor that should influence FID. We studied the relationship between an individual's intrinsic escape speed and FID in yellow‐bellied marmots (Marmota flaviventris) to determine whether marmots compensated for slower escape speeds by fleeing at greater distances. We found no evidence of risk compensation. Rather, we found that slower marmots tolerated closer approaches. This behavioral syndrome may be explained by a coevolution of FID and escape speed in determining an individual's antipredator behavior, an idea upon which we expand.     

Do Yellow‐Bellied Marmots Perceive Enhanced Predation Risk When they are Farther from safety? An experimental study

Whereas there are many studies of the time allocated to antipredator vigilance while animals forage, the vast majority of these studies remain correlative. This is potentially problematic because a variety of factors other than variation in perceived risk might influence putative antipredator behaviors such as time allocated to vigilance and foraging. We conducted an experimental study of yellow‐bellied marmot (Marmota flaviventris) antipredator behavior while marmots foraged at a replicated set of feeding stations established 1, 5, 10, and 20 m from their main burrows. Marmots appeared to perceive a reduced risk of predation when they foraged in the presence of other marmots; they allocated more time to foraging and decreased the time allocated to vigilance. When they foraged farther from their burrows, marmots initiated foraging after a substantially greater amount of time, tended to increase the frequency of their bouts of vigilance, and decreased the duration of each bout. Yearling marmots took less time to begin foraging than adults. Marmot flight initiation distance at our feeding trays was independent of the distance they foraged away from the burrow. Taken together, these experimental results demonstrate that marmots' perceptions of risk increased with distance to the burrow and decreased when other individuals were within 10 m of them while they foraged.     

Effects of predator behavior and proximity on risk assessment by Columbian black-tailed deer

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.     

Evolution of antipredator behavior in an island lizard species,Podarcis erhardii (Reptilia: Lacertidae): The sum of all fears?

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.