The influence of habitat on travel speed, intermittent locomotion, and vigilance in a diurnal rodent

We studied effects of habitat structure on routine travel velocities, intermittent locomotion, and vigilance by the degu (Octodondegus), a diurnal rodent of central Chile. We predicted thattravel speed, pauses during locomotion, and vigilance wouldbe greater in open (riskier) than in shrub (safer) habitats.Video recordings of marked individuals in the wild were used to measure speed and other variables of spontaneous locomotionnot triggered by predatory attack or any other noticeable stimulusduring nonforaging periods. Time spent vigilant while foragingwas also measured. Because degus use bare-ground runways fordistant movements (e.g., between burrow openings and/or foodpatches), data on locomotion decisions were not confounded by effects of obstructive vegetation cover and/or resource abundance.When moving across the habitat between different feeding places,degus showed an intermittent pattern of locomotion, interruptingrunning events with short pauses. As predicted, travel speedand the duration of pauses between locomotion bursts were significantlygreater in open habitats. Further, the duration of locomotionbursts between feeding sites or between feeding sites and burrowswas significantly longer in open habitats. Our assumption that pauses and velocities are independent decisions was supportedby the lack of correlation between pauses and speeds duringlocomotion events. During foraging, degus devoted more timeto vigilance in open than in shrub habitats. The static positionadopted by degus during pauses, the speeds attained during movements, and the concordance between pausing behavior andvigilance across habitats suggest that pausing has an antipredatoryrole and is not limited to orientation and/or physiologicalrecovery. Our results support the view that degus perceivehigher predation risk in open areas and that flexible movement behavior reflects an adaptive antipredator response.     

Predation and the evolution of prey behavior: an experiment with tree hole mosquitoes

We tested for facultative changes in behavior of an aquaticinsect in response to cues from predation and for evolutionof prey behavior in response to experimental predation regimes.Larvae of the tree hole mosquito Aedes triseriatus reducedfiltering, browsing, and time below the surface in responseto water that had held a feeding larva of the predator Toxorhynchitesrutilus. We subjected experimental A. triseriatus populationsto culling of 50% of the larval population, either by T. rutiluspredation or by random removal. After two generations of laboratoryculling, behavior of the two treatment groups diverged. Aedestriseriatus in control-culled lines retained their facultativeshift from filtering to resting, but tended to lose the response of reduced browsing below the surface in water that had helda feeding predator. Predator-culled lines lost their facultativeresponse of reduced filtering in water that had held a feedingpredator and evolved toward more time resting and less timefiltering in both water that had held a feeding predator andwater that had held only A. triseriatus. Predator-culled linesretained their facultative response of reduced browsing belowthe surface in water that had held a feeding predator. Twofield populations and their reciprocal hybrids responded similarlyto cues from predation and did not differ in their evolutionaryresponse to experimental culling. We conclude that consistentpresence or absence of predation can select rapidly for divergencein prey behavior, including facultative behavioral responsesto predators.     

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.     

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.     

Phylogenetic analysis of coadaptation in behavior, diet, and body size in the African antelope

Several authors have suggested that African antelope (familyBovidae) exemplify coadaptation of ecological, behavioral,and morphological traits. We tested four hypotheses relatedto the ecology and behavior of 75 species of African antelopeusing both conventional statistical techniques and techniques that account for the nonindependence of species by consideringtheir phylogenetic relationships. Specifically, we tested thehypotheses that (1) dietary selectivity is correlated negativelywith body mass, (2) dietary selectivity is correlated negativelywith group size, (3) gregarious species either flee or counterattackwhen approached by predators, but solitary and pair-livingspecies seek cover to hide, and (4) body mass and group sizeare correlated positively. Each of these hypotheses was examinedfor the global data set (family Bovidae) and, when possible,within the two antelope subfamilies (Antilopinae and Bovinae)and within 7 of the 10 antelope tribes. The results of ourconventional and phylogenetically corrected analyses supportedthe hypotheses that group and body size vary predictably with feeding style and that antipredator behavior varies with groupsize. The hypothesis that body mass and group size are correlatedpositively was supported by conventional statistics, but thesetwo traits were only weakly related using a phylogeneticallycorrected analysis. Moreover, qualitative and quantitativecomparisons within each of the eight major African antelope tribes generally gave little support for the four hypothesestested. Thus, although our analyses at the subfamily levelprovided results that were consistent with prior hypotheses,our analyses at the level of tribes were equivocal. We discussseveral possible explanations for these differences.     

Predator-induced plasticity in nest visitation rates in the Siberian jay (Perisoreus infaustus)

Bird nestlings may be at risk not only from starvation but alsofrom predators attracted to the nest by parental feeding visits.Hence, parents could trade reduced visitation rates for a lowerpredation risk. Here, through field data and an experiment,we show plasticity in daily patterns of nest visitation in theSiberian jay, Perisoreus infaustus, in response to predatoractivity. In high-risk territories, jay parents avoided goingto the nest at certain times of the day and compensated by allocatingmore feeding effort to periods when predators were less active.Such modifications in provisioning routines allowed parentsin high-risk habitat to significantly lower the risk of providingvisitation cues to visually oriented corvid nest predators.These results indicate that some birds modify their daily nestvisitation patterns as a fourth mechanism to reduce predator-attractingnest visits in addition to the clutch size reduction, maximizationof food load-sizes, and prevention of allofeeding suggestedby Skutch.     

Asset protection in juvenile salmon: how adding biological realism changes a dynamic foraging model

The "asset-protection principle" created by Clark is based ona dynamic programming model and states that individuals should(1) become more averse to predation risk as they accumulatefitness assets but (2) generally be more willing to acceptpredation risk later in the foraging season. To test whetherthese predictions hold under biologically meaningful foraging parameters, I constructed a dynamic model of the optimal trade-offbetween foraging and predator avoidance in juvenile salmon.The model incorporates temperature and body-size dependentbio-energetic constraints typical for juvenile fish, whichgrow by orders of magnitude over a season. In its simplestform using seasonally constant growth potential and a linear over-winter survival function, my results equal those of Clark'smodel. Adding a fitness function and environmental data fromfield studies accentuates the asset-protection effect and fundamentallychanges the seasonal pattern of optimal effort. Simulationof typical poor feeding conditions in mid-summer yields theprediction of increased foraging in the spring in anticipationof worsening conditions. Increasing overall predation riskresults in smaller fish at the end of the season with a trade-offbetween summer and winter survival. The model generates testablepredictions for juvenile salmon and provides insights for otherorganisms (particularly poikilotherms) that are subject tosize-dependent or seasonally changing foraging dynamics.     

Net costs of group living in a solitary forager, the Eurasian badger (Meles meles)

Group living is expected to evolve under two extreme conditions:when there are net benefits to the individual of living in agroup and when there are net costs to living in a group togetherwith strong ecological constraints on dispersal and independentbreeding. A third condition may facilitate group formation:when the territorial defense of resources that are dispersedwidely or renewed at high rates allows groups to form at nocost to group members. Eurasian badgers are solitary over alarge part of their geographic distribution and forage aloneeven where diey live in territorial groups. To determine whichof the three conditions best explains group living in badgers,we analyzed dispersal and breeding patterns. Using these datawe describe the mechanism of group formation and the net costsor benefits of group living in 16 badger groups in one population.Groups form primarily by the retention of young on their natalterritory. Typically, groups contain one or more sexually maturebut nonbreeding females in addition to one or more breedingfemales. Females do not benefit from group living as indicatedby the lack of relationships, or perhaps negative relationships,between the proportion of adult females that bred in a groupand mean reproductive success, on the one hand, and group sizeand components of group size, on the other. A high reproductivefailure rate among females in this and other populations ofsocial badgers, as compared with a population of solitary badgers,suggests that there is a cost of group living to females. Wepropose that if badgers defend resources whose spatial distributionor high renewal rates allow groups to form on a territory atlitde or no cost to group members, weak ecological constraintson dispersal and independent breeding are sufficient to explainthe formation of groups. Badgers appear to have not evolvedcooperative breeding in response to these ecological conditions,perhaps because badger groups represent an early stage in theevolution of carnivore sociality.     

The Effects of Human Age,Group Composition,and Behavior on the Likelihood of Being Injured by Attacking Pumas

ABSTRACT

Documentation from the years 1890 to 2000 of 185 instances of pumas (Puma concolor) attacking humans in the United States and Canada has provided statistical evidence that pumas are less likely to kill or injure humans in certain circumstances. We identified incidents of fatal attacks, severe injuries, light injuries, and no injuries as a function of human age class, group size, body posture, and conspicuous action, such as noise making, running, or shooting. Ordinal multinomial regression revealed that age class (< 13 years old vs. older) was not a statistically reliable predictor of attack severity. This statistical method also revealed that there was no reliable association between the number of individuals present during the attack and attack severity. Nevertheless, examination of specific attack outcomes indicated that the likelihood of escaping injury increased when two or more people were present. The speed that individuals moved during the attack did not predict attack severity, but it was apparent that the lowest likelihood of escaping injury (26%) and greatest frequency of severe injuries (43%) occurred when individuals remained stationary. In contrast, half of the individuals who ran when they were attacked escaped injury, whereas running was associated with only a small increase in the frequency of fatal attacks (28%), compared with remaining stationary (23%). Evidence that half of the individuals who ran escaped injury suggests that pumas are assessing immobility in humans as they might with other prey, using it as an index of prey inattention or disablement and hence greater vulnerability.     

The effect of group size manipulations on the foraging behavior of black-tailed prairie dogs

Individuals foraging in large groups are thought to benefitbecause they are better able to detect and avoid predators.As a consequence, individuals in groups can adopt more risky,but rewarding, foraging behaviors without exposing themselvesto excessive danger. I experimentally manipulated the size ofblack-tailed prairie dog (Cynomys ludovicianus) groups to determineif individuals in large groups do forage in a more risky manner.I found that prairie dogs foraged more alertly and in less riskylocations (nearer to burrows, nearer to the center of the group,and in shorter vegetation) when group size was reduced. Effectsof group-size reductions were reversed when removed individualswere replaced, and persisted for at least three weeks in experimentswhere group size was permanently reduced. My results provideevidence that the relationships between group size and bothalertness and risk-place foraging are causal.     

Mortality by moonlight: predation risk and the snowshoe hare

Optimal behavior theory suggests that prey animals will reduceactivity during intermittent periods when elevated predationrisk outweighs the fitness benefits of activity. Specifically,the predation risk allocation hypothesis predicts that preyactivity should decrease dramatically at times of high predationrisk if there is high temporal variation in predation risk butshould remain relatively uniform when temporal variation inpredation risk is low. To test these predictions we examinedthe seasonably variable response of snowshoe hares to moonlightand predation risk. Unlike studies finding uniform avoidanceof moonlight in small mammals, we find that moonlight avoidanceis seasonal and corresponds to seasonal variation in moonlightintensity. We radio-collared 177 wild snowshoe hares to estimatepredation rates as a measure of risk and used movement distancesfrom a sample of those animals as a measure of activity. Inthe snowy season, 5-day periods around full moons had 2.5 timesmore predation than around new moons, but that ratio of theincreased predation rate was only 1.8 in the snow-free season.There was no significant increase in use of habitats with morehiding cover during full moons. Snowshoe hares' nightly movementdistances decreased during high-risk full-moon periods in thesnowy season but did not change according to moon phase in thesnow-free season. These results are consistent with the predationrisk allocation hypothesis.     

The influence of distance to burrow on flight initiation distance in the woodchuck, Marmota monax

We used woodchucks (Marmota monax) to test predictions of acost-benefit model of antipredator behavior that flight initiationdistance would increase with distance to refuge and with predatorapproach velocity. We also examined the effects of distanceto refuge and predator approach velocity on escape velocityand on both temporal and spatial margin of safety (expectedtime and distance between predator and burrow at the time ofthe woodchuck's arrival). The observer, assumed to be perceivedas a potential predator, approached juvenile woodchucks fromthe direction opposite to the burrow at a slow (1.24 m/s) orfast (1.79 m/s) walking pace. When the woodchuck started toflee, the observer recorded the woodchuck's distance from theobserver and from its burrow, the time spent running, and whetherthe woodchuck stopped before reaching its burrow. Flight initiationdistance increased consistendy with distance to the burrow overthe entire observed range (0–25 m) but was not significantlyaffected by observer approach velocity. Escape velocity wasnot significantly influenced by the observer approach velocityand was approximately constant over the range of 2–25m, but was slower for woodchucks less than 2 m from their burrows.Both temporal and spatial margins of safety increased with distancefrom the burrow. The temporal margin of safety increased withdistance from the burrow more rapidly for slow than for fastobserver approach velocity. Woodchucks fleeing from greaterthan 2 m usually stopped near the burrow before entering, butthose from closer distances usually entered directly. Theseresults support the assumption that antipredator behavior issensitive to the costs and benefits of alternative escape decisions.     

Molecular genetic analysis of Daphnia in the English Lake District: species identity, hybridisation and resting egg banks

• 1 Taxonomic issues in many Daphnia species complexes are often confused by a high degree of phenotypic plasticity and by interspecific hybridisation. Here, we employ molecular genetic tools to confirm the species composition and incidence of hybridisation in extant and resting egg populations of Daphnia from Windermere and Esthwaite Water in the English Lake District.
• 2 A combination of species‐diagnostic allozymes and mtDNA, confirms that contemporary populations are dominated by a single species, D. galeata.
• 3 We present the first account of genetic characterisation of dated ‘resting’ eggs using microsatellites and mtDNA, employing PCR‐based DNA recovery, thus providing a temporal dimension to taxonomic patterns. Thirty years ago, two species were present in Esthwaite, D. galeata and D. hyalina, but Windermere populations were dominated by D. galeata only.
• 4 The use of PCR‐based mtDNA RFLP analysis as a species‐diagnostic tool, and microsatellites to monitor clonal diversity, provide a valuable approach for long‐term studies, especially in populations free from the complicating effect of frequent hybridisation. The detailed limnological records available for many large lakes, and associated changes in land‐use, pollutants and climate, combined with long‐term ephippial molecular genetic data, provide opportunities for exploring natural and anthropogenic impacts on genetic and community structure.

     

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.     

Vigilance and its complex synchrony in the red-necked pademelon, Thylogale thetis

Several adaptive functions, including gaining information fromother group members and detecting predators, are generally ascribedto vigilance in groups of animals subject to predation. Moststudies of the effects of neighbors on vigilance have focusedon individual vigilance. We investigated the effects of neighborson vigilance in wild red-necked pademelons Thylogale thetisforaging at night in nonpersistent aggregations in a clearingin rain forest. Neither the total number of pademelons in theclearing nor the numbers at various distances around focal individualsaffected the individual vigilance of focal animals. However,focal animals’ individual vigilance did change with thedistance to their nearest neighbor and also with distance tocover. Pairs of individuals closer than 10 m apart tended tosynchronize their bouts of individual vigilance and foraging.The degree of synchrony within pairs increased with both distanceto cover and the total number of pademelons foraging in thearea and decreased with increasing distance to the pair's nearestneighbor but did not vary with the distance separating the membersof the pair. Thus, despite their individual vigilance beingunaffected by the number of other pademelons in the feedingaggregation, pademelons were nonetheless sensitive to the presenceof conspecifics and adjusted their behavior in relation to theirseparation from neighbors. Thus, some vigilance benefits maybe obtained from the presence of conspecifics even in speciesthat aggregate only temporarily on food patches without formingmore permanent social groups.     

Asymmetry in size, shape, and color impairs the protective value of conspicuous color patterns

The received view of protective coloration in animals is thatconspicuous colors and patterns have evolved because they elicitavoidance behavior in potential predators. In the present study,we examine the spontaneous response of naive predators (Gallusgallus domesticus) to artificial prey to test the hypothesisthat deviations from bilateral symmetry of signaling patternelements may negatively influence the avoidance-inducing effectof conspicuous color patterns. Chicks displayed stronger aversionsto artificial "butterfly" prey items possessing symmetric colorpattern elements than to those possessing asymmetric signalswith pattern elements of different color or shape. Althoughthey attacked signals with a size asymmetry of 5% at the samerate as symmetric signals, signals with a size asymmetry of7.5% or more were attacked more often than were symmetric signals.These results suggest that the protective value of conspicuouscolor patterns is impaired by asymmetry in color, shape, andsize of color pattern elements. Our findings also argue againstthe notion that animals have inherent preferences for symmetricover asymmetric objects, and demonstrate the existence of athreshold for asymmetry detection, beyond which further incrementsin asymmetry have no influence on signal efficacy.