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.     

The worm re-turns: hiding behavior of a tube-dwelling marine polychaete, Serpula vermicularis

The polychaete worm Serpula vermicularis (Serpulidae) filterfeeds at the mouth of its calcareous tube, but retreats intothe tube when startled by mechanical stimuli likely to be associatedwith predators. While in its tube, a worm is safe but cannotfeed. Thus, hiding has a lost-opportunity cost. We show thatthis cost can be substantial, given that food in the naturalhabitat appears in pulses, and good feeding conditions may notlast long or recur frequently. We expect that a worm's hidingtime will be sensitive to the lost-opportunity cost, and wepresent data from a series of experiments that support thisprediction. The worms seem able to track relatively short-termchanges in food availability, and some evidence suggests thatthey assess food availability on a relative basis, comparingcurrent feeding conditions to those recently experienced. Hidingand other types of cryptic behavior are common antipredatortactics, and animals may commonly adjust the durations of suchbehaviors to current benefits and costs (including lost opportunity),as they perceive them     

Inter‐individual Variation in Antipredator Hiding Behavior of Spanish Terrapins Depends on Sex,Size, and Coloration

Behavioral responses to predation risk are critical for survival but as antipredator behavior is costly, prey animals should flexibly modulate their optimum defensive responses by considering both costs and benefits, which are partly influenced by the individual characteristics of the prey. Turtles have the shell as a morphological structure that may provide partial protection against predators, but hiding into the shell may entail some high costs, and turtles should decide when to switch to an active escape strategy to safe refuges. Here, we examined how gender, body size, and sexual coloration influence inter‐individual variability of antipredatory hiding behavior into the shell of Spanish terrapins (Mauremys leprosa). We simulated predatory attacks under different conditions and measured the time that the turtles spent hidden entirely inside the shell (i.e., appearance times) and from then until the turtle started to flee actively (i.e., waiting times). Our results showed that when risk increased, appearance times increased but waiting times decreased. When turtles were in a prone position, their hiding behavior was related with their body weight with heavier turtles having longer appearance times. Also, the conspicuousness of limb coloration was important for the appearance times of males, but not for females. Thus, males with brighter coloration of the limb stripes had longer appearance times than duller ones. In addition, when turtles were overturned, males appeared out of the shell earlier than females and heavier turtles started to right sooner, but only when risk was low. However, when turtles were overturned and risk was high, they should assume that they have already been detected, making inter‐individual differences in size and coloration apparently unimportant for deciding hiding behavior.     

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.     

A framework for characterizing overlap of communication and computation in parallel applications

Effective overlap of computation and communication is a well understood technique for latency hiding and can yield significant performance gains for applications on high-end computers. In this paper, we propose an instrumentation framework for message-passing systems to characterize the degree of overlap of communication with computation in the execution of parallel applications. The inability to obtain precise time-stamps for pertinent communication events is a significant problem, and is addressed by generation of minimum and maximum bounds on achieved overlap. The overlap measures can aid application developers and system designers in investigating scalability issues. The approach has been used to instrument two MPI implementations as well as the ARMCI system. The implementation resides entirely within the communication library and thus integrates well with existing approaches that operate outside the library. The utility of the framework is demonstrated by analyzing communication-computation overlap for micro-benchmarks and the NAS benchmarks, and the insights obtained are used to modify the NAS SP benchmark, resulting in improved overlap.

Behavioural Consistency in Female Resistance to Male Harassment in a Water Strider Species

Sexual conflict over mating rate often implies that males persist at frequently harassing females to gain matings while females resist mating attempts. In water striders, females can resist by engaging in vigorous pre‐copulatory struggles to dislodge males, but alternative means of resistance have seldom been investigated. Contrary to males, female resistance has not been investigated as a repeatable behaviour. We used Gerris buenoi to investigate the capacity to abbreviate struggles and the tendency to hide off the water as two potential female resistance traits. Specifically, we asked whether these behaviours are repeatable and whether they vary according to sexual conflict intensity and past mating experience. Also, we studied the possible connections between these behaviours and traits linked to fitness, namely endured harassment and mating activity. The capacity to abbreviate struggles was poorly repeatable and decreased with sexual conflict intensity and endured harassment. It seems to be mainly determined by the social environment and by recent events related to sexual conflict. The tendency to hide off the water was significantly repeatable across sexual conflict intensities and can be considered as a repeatable behaviour. Hiding frequently off the water allowed females to decrease the harassment endured by females and may enhance female fitness. In nature, hiding is more readily and more frequently observed than pre‐copulatory struggles. Directly associating hiding off water with female fitness would confirm that this consistent phenotype contributes to sexually antagonistic female resistance.     

Climbing of Leaf Trichomes by Eriophyid Mites Impedes Their Location by Predators

Eriophyid mites are plant parasites that are well adapted to hide away from predators. Tiny and wormlike, they can invade very narrow spaces in plants or form galls that, apart from other functions, serve as a shelter from predation. Previous observations showed that some free-living eriophyids as well as tetranychid mites spend their quiescence on the top of leaf trichomes. Here, I investigated climbing leaf trichomes by the eriophyid, Rhinophytoptus concinnus, and tested whether it enables the herbivores to avoid phytoseiid mites. Climbing behavior took place just prior to the quiescent period of juveniles. Larvae and nymphs raised the hind part of their stiffening bodies and walked, turning around on their axis. Having found a hair, juveniles attached their anal suckers to its tip, and, pushing back from a leaf surface or the base of the hair, they lifted their bodies up to become motionless. As revealed by the playback experiments with the phytoseiid mite, Typhloctonus tiliarum, predatory females needed much more time to find quiescent nymphs perching on hairs than those placed on a leaf surface. The time of nymph handling was similar in both situations. Also, a similar number of predators gave up feeding on nymphs in both locations. I conclude that climbing leaf trichomes enables the herbivorous mite to hide from predators. After detection, however, placement on trichomes does not give the quiescent juveniles any advantage over those placed on a leaf blade.     

The waiting game: a "battle of waits" between predator and prey

Many prey respond to the presence of a predator by retreatinginto a shell or burrow, or by taking refuge in some other waythat guarantees their safety but restricts further informationfrom being obtained about the predator's continued presence.When this occurs, the individual predator and prey involvedbecome opponents in a "waiting game." The prey must decide howlong to wait for the predator to depart before re-emerging andpotentially exposing itself to attack. The predator must decidehow long to wait for the prey to re-emerge before departingin search of other foraging opportunities. I use a numericalapproach to determine the evolutionarily stable waiting strategyof both players and examine the effects of various parameterson the ESS. The model predicts that each player's waiting distribution—thedistribution of waiting times one would expect to observe forindividuals in that role—will have a characteristic shape:the predator's distribution should resemble a negative exponentialfunction, whereas the waiting time of the prey is predictedto be more variable and follow a positively skewed distribution.The model also predicts that very little overlap will occurbetween the players' waiting distributions, and that the predatorwill rarely outwait the prey. Empirical studies relating tothe model and comparisons between the waiting game and the asymmetricwar of attrition are discussed.     

When to come out from a refuge: risk-sensitive and state-dependent decisions in an alpine lizard

Prey often respond to predator presence by increasing theiruse of refuges. However, unfavorable thermal conditions in refugesmight entail physiological costs for an ectothermic prey. Thus,the decision of when to come out from a refuge should be optimizedby considering the expected fitness effects of diminution ofpredation risk with time, but also by considering the cost of theloss of time spent at optimal body temperature maximizing physiological functions.The model of Ydenberg and Dill describes the trade-off betweenrisk and cost for a prey fleeing to a refuge. We present a specialcase of this model to predict how emergence time from the refugein lizards or other ectotherms should vary as a function ofrisk of predation and thermal costs of refuge use. The analysesof the variation in emergence time from a refuge of Lacertamonticola lizards in the field under two different predation risklevels supported the predictions of the model. As predicted,time spent in the refuge was longer when the threat of the initialattack had been higher, and therefore the subsequent diminutionof risk was slower, but only when lizards emerged at the sameplace where they hid. When initial body temperature was high,some lizards decreased emergence time by emerging from a differentplace. In addition, the effects of thermal costs were more relevant inthe high-risk situation. Time spent in the refuge under highrisk increased when thermal conditions of the refuge were moresimilar to thermal conditions outside (i.e., physiological costsof refuge use were lower). We conclude that optimization ofrefuge-use strategies might help lizards cope with changes in predationrisk without incurring excessive physiological costs.     

The significance of shelter for young cutworms (Agrotis segetum)

The importance of shelter for young Agrotis segetum Schiff. (Lep. Noctuidae) larvae was investigated using caged carrots (Daucus carota L.) with the soil surface covered only by paraffin wax, or with a layer of vermiculite resembling dry soil on top of a layer of paraffin wax. In the vermiculite treatment, most cutworms were found in the vermiculite with only a small proportion on the plant foliage, whereas in the paraffin wax treatment, the majority was found on the foliage. Larval growth rate was higher and mortality lower among the cutworms which could shelter in the vermiculite than amongst those with no shelter. These results agree with those from previous investigations which showed that moist soil prevented the larvae from sheltering in it. The importance of shelter in dry soil cannot be fully explained, but it may provide a more favourable temperature for larval development.

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Résumé L'importance des abris pour les jeunes chenilles d'A. segetum a été examinée en utilisant des carottes (Daucus carota) dans des cages dont le sol était recouvert d'une couche de paraffine ou d'une couche de vermiculite mimant du sol sec au-dessus de la couche de paraffine. La plupart des vers gris ont été trouvés dans la couche de vermiculite contre une faible proportion dans le feuillage, tandis qu'en absence de vermiculite la majorité a été retrouvée dans le feuillage. La croissance larvaire était meilleure et la mortalité plus faible avec la vermiculite. Ces résultats sont conformes à des observations antérieures qui avaient montré qu'un sol humide empechait les chenilles de s'y abriter. L'importance de l'abri en sol sec n'est pas entièrement expliquée, mais celui-ci pourrait assurer une température plus favorable au développement larvaire.