Repeated predatory attacks and multiple decisions to come out from a refuge in an alpine lizard

Prey often respond to predator presence by increasing theiruse of refuges. However, because the use of refuges may entailseveral costs, the decision of when to come out from a refugeshould be optimized. In some circumstances, if predators remainwaiting outside the refuge and try new attacks or if predator density increases, the prey may suffer successive repeated attacksin a short time. Successive attacks may represent an increasein the risk of predation, but the costs of refuge use alsomay increase with time spent in the refuge. Thus, prey shouldmake multiple related decisions on when to emerge from the refuge after each new attack. We simulated in the field repeatedpredatory attacks to the same individuals of the lizard Lacertamonticola and specifically examined the variation in successivetimes to emergence from a refuge under different thermal conditions(i.e., different costs of refuge use). The results showed thatrisk of predation but also thermal costs of refuge use affectedthe emergence decisions. Lizards increased progressively theduration of time spent in the refuge between successive emergencetimes when the costs of refuge use were lower, but tended tomaintain or to decrease the duration of time spent in the refugebetween successive emergence times when cost of refuge useincreased. Additionally, lizards that entered the refuge withhigher body temperatures had overall emergence times of longer duration. Optimization of refuge use and flexibility in theantipredator responses might help lizards to cope with increasedpredation risk without incurring excessive costs of refugeuse.     

Costs of Refuge Use Affect Escape Decisions of Iberian Rock Lizards Lacerta monticola

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.     

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.     

Pregnant female lizards Iberolacerta cyreni adjust refuge use to decrease thermal costs for their body condition and cell-mediated immune response

Lizards often respond to increased predation risk by increasing refuge use, but this strategy may entail a loss of thermoregulatory opportunities, which may lead to a loss of body condition. This may be especially important for pregnant oviparous female lizards, because they need to maintain optimal body temperatures as long as possible to maximize developmental embryos rate until laying. However, little is known about how increased time spent at low temperatures in refuges affects body condition and health state of pregnant female lizards. Furthermore, it is not clear how initial body condition affects refuge use. Female Iberian rock lizards forced to increase time spent at low temperatures showed lower body condition and tended to show lower cell-mediated immune responses than control females. Therefore, the loss of thermoregulatory opportunities seems to be an important cost for pregnant females. Nevertheless, thereafter, when we simulated two repeated predatory attacks, females modified refuge use in relation to their body condition, with females with worse condition decreasing time hidden after attacks. In conclusion, female lizards seemed able to compensate increased predation risk with flexible antipredatory strategies, thus minimizing costs for body condition and health state.     

Wall Lizards Modulate Refuge Use through Continuous Assessment of Predation Risk Level

Prey species might use several possible ways to assess predation risk when encountering a predator. Animals may consider the risk level estimated in a first encounter to remain unchanged across subsequent encounters (fixed risk response), or they may update and change their responses across encounters in accordance with short‐term changes in risk levels (flexible risk response). We examined in the field how wall lizards assess risk level by analyzing time spent in refuges after simulated predator attacks. We first examined how risk was assessed when multiple consecutive sources of risk were present simultaneously. The results suggest that wall lizards assess risk based on multiple cues, such as approach speed, directness, and persistence (measured as the distance of the predator to their refuge after an attack). When risk was high lizards remained longer in their refuges. The first decision to appear partly from the refuge depended on both approach speed and persistence, whereas the decision to emerge completely depended only on persistence and not on approach speed. This suggests that wall lizards update information on predator threat and adjusted their emergence accordingly. In a second experiment, we analyzed how short‐term changes in risk level of successive attacks affected refuge use. Successive emergence times varied as a function of current risk level of each repeated attack, independently of the risk level of previous attacks. This indicated that lizards could track short‐term changes in risk level through time and modify their initial responses when required. Fine adjustments of refuge use may help lizards to minimize costs of refuge use in unfavorable and variable environments where antipredatory responses are costly.     

Optimal time to emerge from refuge

Factors affecting emergence by prey that enter refuges when approached by predators have been studied intensively, but only two theoretical models predict how long prey should remain in a refuge before emerging. We argue that prey can make better decisions than allowed by one model; the other model describes cases in which predators wait for prey to emerge. We present optimality models that permit prey to select a time to emerge that maximizes fitness. When in a refuge, a prey cannot obtain benefits outside; emerging too soon can be catastrophic, but delaying emergence entails loss of fitness. If predators resume foraging quickly rather than engaging in strategic waiting games, current theory suggests that prey emerge when the costs of remaining in a refuge and of emerging are equal. However, prey often can do better by emerging at the time maximizing fitness rather than when benefits equal costs (i.e. when prey break even). Optimal emergence time depends on initial fitness, benefits lost by remaining in refuge, and the decay rate of predation risk. Benefits lost if a prey is killed are modelled separately from benefits that contribute to lifetime fitness, even if the prey is killed (individual reproduction, altruism). Fitness of prey emerging at the optimal emergence time may be greater than, equal to or less than initial fitness. Break-even and optimality models base predictions on the opposing effects of risk and loss of benefits. Thus, many empirically verified predictions are identical at the ordinal level although differing quantitatively. Optimality models provide novel testable predictions for the effects of initial fitness, benefits, and, for ectotherms, the rate of cooling in refuge. They predict earlier emergence for equal retainable benefits than for those lost upon death.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 375–382.     

Foraging and refuge use by a pond snail: Effects of physiological state,predators, and resources

The costs and benefits of anti-predator behavioral responses should be functions of the actual risk of predation, the availability of the prey's resources, and the physiological state of the prey. For example, a food-stressed individual risks starvation when hiding from predators, while a well-fed organism can better afford to hide (and pay the cost of not foraging). Similarly, the benefits of resource acquisition are probably highest for the prey in the poorest state, while there may be diminishing returns for prey nearing satiation. Empirical studies of state-dependent behavior are only beginning, however, and few studies have investigated interactions between all three potentially important factors. Here I present the results of a laboratory experiment where I manipulated the physiological state of pond snails (Physa gyrina), the abundance of algal resources, and predation cues (Belostoma flumineum waterbugs consuming snails) in a full factorial design to assess their direct effects on snail behavior and indirect effects on algal biomass. On average, snails foraged more when resources were abundant, and when predators were absent. Snails also foraged more when previously exposed to physiological stress. Snails spent more time at the water's surface (a refuging behavior) in the presence of predation cues on average, but predation, resource levels, and prey state had interactive effects on refuge use. There was a consistent positive trait-mediated indirect effect of predators on algal biomass, across all resource levels and prey states.     

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.     

Effect of Risk on Aspects of Escape Behavior by a Lizard, Holbrookia propinqua, in Relation to Optimal Escape Theory

Prey must balance gains from activities such as foraging and social behavior with predation risk. Optimal escape theory has been successful in predicting escape behavior of prey under a range of risk and cost factors. The optimal approach distance, the distance from the predator at which prey should begin to flee, occurs when risk equals cost. Optimal escape theory predicts that for a fixed cost, the approach distance increases as risk increases. It makes no predictions about approach distance for prey in refuges that provide only partial protection or about escape variables other than approach distance, such as the likelihood of stopping before entering refuge and escape speed. By experimentally simulating a predator approaching keeled earless lizards, Holbrookia propinqua, the predictions of optimal escape theory for two risk factors, predator approach speed and directness of approach were tested. In addition, predictions that the likelihood of fleeing into refuge without stopping and the speed of escape runs increase with risk, in this case predator approach speed, and that lizards in incompletely protective refuges permit closer approach than lizards not in refuges were also tested. Approach distance increased with predator approach speed and directness of approach, confirming predictions of optimal escape theory. Lizards were more likely to enter refuge and ran faster when approached rapidly, verifying that predation risk affects escape decisions by the lizards for escape variables not included in optimal escape theory. They allowed closer approach when in incompletely protective refuges than when in the open, confirming the prediction that risk affects escape decisions while in refuge. Optimal escape theory has been highly successful, but testing it has led to relative neglect of important aspects of escape other than approach distance.     

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.     

When to Come Out from a Refuge: Balancing Predation Risk and Foraging Opportunities in an Alpine Lizard

Prey often respond to predator presence by increasing their use of refuges, but because this strategy may be costly, the decision regarding when to come out from a refuge should be optimized. The loss of foraging opportunities may be one of the main costs when safer microhabitats (i.e. refuges) are also the poorest in terms of their foraging profitability. We present the results of an experimental field study to test whether emergence times from a refuge of the Iberian rock lizard, Lacerta monticola, vary as a function of expected foraging opportunities and level of satiation of the lizard. As predicted, short‐term fluctuations in availability of food influenced emergence times; when a lizard had just detected some food in the recent past, emergence times decreased greatly, because the loss of opportunities for foraging increased costs of refuge use. Furthermore, the characteristics and success of the encounter with food, nutritional state of lizards, and the added possibility of capturing new food items influenced the duration of hiding times. Therefore, foraging requirements and avoidance of predators may be conflicting demands that L. monticola lizards balance by modifying the duration of time spent in refuges.     

Size-dependent predation by snakes: selective foraging or differential prey vulnerability?

I staged replicate encounters between unrestrained lizards andsnakes in outdoor enclosures to examine size-dependent predationwithin the common garden skink (Lampropholis guichenoti). Yellow-facedwhip snakes (Demansia psammophis) forage widely for activeprey and most often consumed large skinks, whereas death adders(Acanthophis antarcticus) ambush active prey and most oftenconsumed small skinks. Small-eyed snakes (Rhinoplocephalusnigrescens) forage widely for inactive prey and consumed bothsmall and large skinks equally often. Differential predationmay reflect active choice by the predator, differential preyvulnerability, or both. To test for active choice, I presentedforaging snakes with an inert small lizard versus an inertlarge lizard. They did not actively select lizards of a particularbody size. To test for differential prey vulnerability, I quantifiedvariation between small and large lizards in behavior thatis important for determining the outcome of predator—prey interactions. Snakes did not differentiate between integumentarychemicals from small and large lizards. Large lizards tendto flee from approaching predators, thereby eliciting attackby the visually oriented whip snakes. Small lizards were moremobile than large lizards and therefore more likely to passby sedentary death adders. Additionally, small skinks were more effectively lured by this sit-and-wait species and less likelyto avoid its first capture attempt. In contrast, overnightretreat site selection (not body size) determined a lizard'schances of being detected by small-eyed snakes. Patterns ofsize-dependent predation by elapid snakes may arise not becauseof active choice but as a function of species-specific predatortactics and prey behavior.     

Anti-predator behaviour changes following an aggressive encounter in the lizard Tropidurus hispidus

Avoiding predators may conflict with territorial defence because a hiding territorial resident is unable to monitor its territory or defend it from conspecific intrusions. With persistent intruders, the presence of an intruder in the near past can indicate an increased probability of future intrusions. Therefore, following a conspecific-intrusion, territorial residents should minimize costs from future intrusions at the cost of higher predation risks. I conducted experiments with males of the territorial lizard Tropidurus hispidus recording approach distance (distance between predator and prey when the prey escapes) and time to re-emergence from a refuge after hiding. Past aggressive interactions affected anti-predator behaviour: lizards re-emerged sooner (compared to a control) when the predator attacked 5 min after an aggressive encounter. If the predator attacked while an aggressive encounter was ongoing, there was also a reduction in approach distance. The results are consistent with an economic hypothesis which predicts that T. hispidus incur greater predation risks to minimize future territorial intrusion; additionally they show that the effects of past and ongoing aggressive interactions are different, consistent with the minimization of present intrusion costs. These results are relevant for studies of the changes in aggressive behaviour due to changes in the social environment and for studies of the costs and (co) evolution of aggressive and anti-predator strategies.     

Difference in response by two cyprinid species to predatory threat from the nocturnal catfish Silurus asotus

Response to predators may not be identical between different prey species with different life histories and body sizes, particularly when the threat of predation is not great. To clarify this hypothesis, we introduced two prey species (10 Japanese dace, Tribolodon hakonensis, and 10 pale chub, Zacco platypus) into each experimental pond (in total, 8 ponds×4 trials) in which benthic algae had been allowed to grow. The presence or absence of Far Eastern catfish, Silurus asotus, and a refuge for prey fish was used to produce four treatments. The presence of catfish and/or a refuge did not affect either the feeding behavior or growth rate of Japanese dace. In contrast, when catfish were present and no refuge was available, the incidence of bottom feeding for pale chub greatly decreased. Pale chub growth rate was low when catfish were present and a refuge was available, indicating that pale chub spent more of their time in the refuge and lost opportunities of acquiring food. Japanese dace can reach a threshold size at which the prey are safe from predation, but pale chub cannot, and this may explain the differences in response to predators of the two species.     

Loss of body mass under predation risk: cost of antipredatory behaviour or adaptive fit-for-escape?

Predation risk may compromise the ability of animals to acquire and maintain body reserves by hindering foraging efficiency and increasing physiological stress. Locomotor performance may depend on body mass, so losing mass under predation risk could be an adaptive response of prey to improve escape ability. We studied individual variation in antipredatory behaviour, feeding rate, body mass and escape performance in the lacertid lizard Psammodromus algirus. Individuals were experimentally exposed to different levels of food availability (limited or abundant) and predation risk, represented by reduced refuge availability and simulated predator attacks. Predation risk induced lizards to reduce conspicuousness behaviourally and to avoid feeding in the presence of predators. If food was abundant, alarmed lizards reduced feeding rate, losing mass. Lizards supplied with limited food fed at near-maximum rates independently of predation risk but lost more mass when alarmed; thus, mass losses experienced under predation risk were higher than those expected from feeding interruption alone. Although body mass of lizards varied between treatments, no component of escape performance measured during predator attacks (endurance, speed, escape strategy) was affected by treatments or by variations in body mass. Thus, the body mass changes were consistent with a trade-off between gaining resources and avoiding predators, mediated by hampered foraging efficiency and physiological stress. However, improved escape efficiency is not required to explain mass reduction upon predator encounters beyond that expected from feeding interruption or predation-related stress. Therefore, the idea that animals may regulate body reserves in relation to performance demands should be reconsidered.     

Climate dependent heating efficiency in the common lizard

Regulation of body temperature is crucial for optimizing physiological performance in ectotherms but imposes constraints in time and energy. Time and energy spent thermoregulating can be reduced through behavioral (e.g., basking adjustments) or biophysical (e.g., heating rate physiology) means. In a heterogeneous environment, we expect thermoregulation costs to vary according to local, climatic conditions and therefore to drive the evolution of both behavioral and biophysical thermoregulation. To date, there are limited data showing that thermal physiological adjustments have a direct relationship to climatic conditions. In this study, we explored the effect of environmental conditions on heating rates in the common lizard (Zootoca vivipara). We sampled lizards from 10 populations in the Massif Central Mountain range of France and measured whether differences in heating rates of individuals correlated with phenotypic traits (i.e., body condition and dorsal darkness) or abiotic factors (temperature and rainfall). Our results show that heat gain is faster for lizards with a higher body condition, but also for individuals from habitats with higher amount of precipitation. Altogether, they demonstrate that environmentally induced constraints can shape biophysical aspects of thermoregulation.     

Plesiomorphic Escape Decisions in Cryptic Horned Lizards (Phrynosoma) Having Highly Derived Antipredatory Defenses

Escape theory predicts that the probability of fleeing and flight initiation distance (predator–prey distance when escape begins) increase as predation risk increases and decrease as escape cost increases. These factors may apply even to highly cryptic species that sometimes must flee. Horned lizards (Phrynosoma) rely on crypsis because of coloration, flattened body form, and lateral fringe scales that reduce detectability. At close range they sometimes squirt blood‐containing noxious substances and defend themselves with cranial spines. These antipredatory traits are highly derived, but little is known about the escape behavior of horned lizards. Of particular interest is whether their escape decisions bear the same relationships to predation risk and opportunity costs of escaping as in typical prey lacking such derived defenses. We investigated the effects of repeated attack and direction of predator turning on P. cornutum and of opportunity cost of fleeing during a social encounter in P. modestum. Flight initiation distance was greater for the second of two successive approaches and probability of fleeing decreased as distance between the turning predator and prey increased, but was greater when the predator turned toward than away from a lizard. Flight initiation distance was shorter during social encounters than when lizards were solitary. For all variables studied, risk assessment by horned lizards conforms to the predictions of escape theory and is similar to that in other prey despite their specialized defenses. Our findings show that these specialized, derived defenses coexist with a taxonomically widespread, plesiomorphic method of making escape decisions. They suggest that escape theory based on costs and benefits, as intended, applies very generally, even to highly cryptic prey that have specialized defense mechanisms.     

Predicted fitness consequences of threat-sensitive hiding behavior

In studies of refuge use as a form of antipredator behavior,where prey hide in response to a predator's approach, factorssuch as foraging costs and the perceived risk in a predator'sapproach have been shown to influence the hiding behavior ofprey. Because few studies of waiting games have focused on mammals,we studied the hiding behavior of the yellow-bellied marmot(Marmota flaviventris), a ground-dwelling rodent. We testedthe prediction that marmots vary hiding time as a function ofpredator approach speed and presence and absence of food outsidetheir refuge and that marmots hide differently depending ontheir relative condition. We conducted "fast approaches" and"slow approaches" in the presence and absence of extra foodand evaluated hiding times. Multiple regression analyses demonstratedthat the interaction between the approach speed and the presenceand absence of food influenced hiding behavior; body conditionhad a smaller, but nonsignificant effect. We then developeda state-dependent dynamic model to explore potential fitnessconsequences of these decisions. The model suggested that theoverall survival of a population is substantially reduced whenindividuals make suboptimal decisions. Our research builds onprevious studies, indicating that animals integrate both costsand benefits of hiding when determining their hiding times.     

Ontogenetic differences of herbivory on woody and herbaceous plants: a meta-analysis demonstrating unique effects of herbivory on the young and the old, the slow and the fast

Defensive modifications in prey traits that reduce predation risk can also have negative effects on prey fitness. Such nonconsumptive effects (NCEs) of predators are common, often quite strong, and can even dominate the net effect of predators. We develop an intuitive graphical model to identify and explore the conditions promoting strong NCEs. The model illustrates two conditions necessary and sufficient for large NCEs: (1) trait change has a large cost, and (2) the benefit of reduced predation outweighs the costs, such as reduced growth rate. A corollary condition is that potential predation in the absence of trait change must be large. In fact, the sum total of the consumptive effects (CEs) and NCEs may be any value bounded by the magnitude of the predation rate in the absence of the trait change. The model further illustrates how, depending on the effect of increased trait change on resulting costs and benefits, any combination of strong and weak NCEs and CEs is possible. The model can also be used to examine how changes in environmental factors (e.g., refuge safety) or variation among predator–prey systems (e.g., different benefits of a prey trait change) affect NCEs. Results indicate that simple rules of thumb may not apply; factors that increase the cost of trait change or that increase the degree to which an animal changes a trait, can actually cause smaller (rather than larger) NCEs. We provide examples of how this graphical model can provide important insights for empirical studies from two natural systems. Implementation of this approach will improve our understanding of how and when NCEs are expected to dominate the total effect of predators. Further, application of the models will likely promote a better linkage between experimental and theoretical studies of NCEs, and foster synthesis across systems.     

Behavioural interactions between the lizard <Emphasis Type="Italic">Takydromus tachydromoides</Emphasis> and the praying mantis <Emphasis Type="Italic">Tenodera aridifolia</Emphasis> suggest reciprocal predation between them

The Japanese lacertid lizard Takydromus tachydromoides and the praying mantis Tenodera aridifolia are sympatric generalist predators feeding on similar prey. To confirm reciprocal predation between them, we observed the behavioural interactions between the lizards and the mantises of different sizes in a laboratory condition. The lizards caught small mantises (from first to fifth instars), but sometimes escaped from large mantises (from sixth instar to adult). Large mantises occasionally showed catch responses to the lizards. The lizards sometimes caught the mantis without a tongue-flick response (sampling of chemical cues), and they sometimes did not catch the small mantises showing immobile or cryptic responses that prevent visual detection. These results suggested the primary role of vision on recognition of the mantis as a prey. The lizards spent a longer time to approach larger mantises. The time from orienting to catch was longer when the lizards showed tongue-flick responses. The lizard also spent a longer time before deciding to escape from the mantis than to catch it. Biological significance of these differences in timing was discussed.