Behaviour and choice of refuge by voles under predation risk

Animals often show a strong antipredatory response when they are exposed to their most deadly predator. In northern vole populations, the least weasel, Mustela nivalis nivalis, is probably the most important predator of voles. Because of its small size and slender body shape, the least weasel is capable of hunting voles in their burrows. However, small voles can potentially escape weasel predation by selecting holes smaller than those weasels can enter. We studied the choice of nest holes and refuges by two species of voles under simulated predation risk. In a laboratory experiment, voles were provided with four nest boxes as refuges, with individually adjusted entrance sizes. When exposed to the odour of a weasel, voles did not choose the smallest opening; they rather seemed to trade full protection for easy and immediate access by choosing the nest box with an intermediate entrance size. When outside the nest at the time when a weasel entered the arena, voles avoided the refuges with the smallest holes. In addition to using refuges on ground level, voles climbed on top of the boxes as an escape reaction, as well as exhibiting a variety of behavioural responses, such as fast running, freezing and sneaking.Communicated by P.A. Bednekoff     

Shifted balance of risk and cost after autotomy affects use of cover,escape, activity,and foraging in the keeled earless lizard (<Emphasis Type="Italic">Holbrookia propinqua</Emphasis>)

Autotomy of expendable body parts is often a successful last-chance antipredatory defense for a variety of invertebrates and vertebrates, especially lizards. However, loss of a body part imposes costs and risks, some of which may be reduced by modifying subsequent behavior. I studied effects of experimentally induced autotomy of the tail in the keeled earless lizard (Holbrookia propinqua) on use of cover, escape behavior, activity, and feeding. Autotomized lizards stayed closer to plant cover than sham-autotomized lizards (having intact tails) when approached by an investigator simulating a predator. Autotomized males, but not females, fled further than sham-autotomized lizards. Autotomy did not affect the distance from an approaching predator when the lizards began to flee. After autotomy, the number of days on which lizards were active differed between sexes and lizards made fewer attempts to catch prey per unit time than lizards with intact tails. Tail loss did not affect the percentage of time spent moving. Staying closer to refuge, fleeing further after autotomy, and decreasing surface activity may compensate for decreased maximum escape speed or agility, and may reflect use of a greater margin of safety when the tail is unavailable for autotomy. Decrease in surface activity may be comparable to use of safer microhabitats reported for other lizards and damselflies subsequent to autotomy. Several possible reasons for the lower rate of feeding attempts by autotomized lizards are discussed.     

Do mountain log skinks (<Emphasis Type="Italic">Pseudemoia entrecasteauxii</Emphasis>) modify their behaviour in the presence of two predators?

Prey often adopt antipredator strategies to reduce the likelihood of predation. In the presence of predators, prey may use antipredator strategies that are effective against a single predator (specific) or that are effective against several predators (nonspecific). Most studies have been confined to single predator environments although prey are often faced with multiple predators. When more than one predator is present, specific antipredator behaviours can conflict and avoidance of one predator may increase vulnerability to another. To test how prey cope with this dilemma, I recorded the behaviours of lizards responding to the nonlethal cues of a bird and snake presented singly and simultaneously. Lizards use specific and conflicting antipredator tactics when confronted with each predator, as evidenced by refuge use. However, when both predators were present, lizards refuge use was the same as in the predator-free environment, indicating that they abandoned refuge use as a primary mechanism for predator avoidance. In the presence of both predators, they reduced their overall movement and time spent thermoregulating. This shift in behaviour may represent a compromise to minimize overall risk, following a change in predator exposure. This provides evidence of plasticity in lizard antipredator behaviour and shows that prey responses to two predators cannot be accurately predicted from what is observed when only one predator is present.Communicated by W. Cooper     

Assessing the Distribution and Protection Status of two Types of Cool Environment to Facilitate Their Conservation under Climate Change

Strategies to mitigate climate change can protect different types of cool environments. Two are receiving much attention: protection of ephemeral refuges (i.e., places with low maximum temperatures) and of stable refugia (i.e., places that are cool, have a stable environment, and are isolated). Problematically, they are often treated as equivalents. Careful delineation of their qualities is needed to prevent misdirected conservation initiatives; yet, no one has determined whether protecting one protects the other. We mapped both types of cool environments across a large (～3.4M ha) mixed‐use landscape with a geographic information system and conducted a patch analysis to compare their spatial distributions; examine relations between land use and their size and shape; and assess their current protection status. With a modest, but arbitrary, threshold for demarcating both types of cool environments (i.e., values below the 0.025 quantile) there were 146,523 ha of ephemeral refuge (62,208 ha) and stable refugia (62,319 ha). Ephemeral refuges were generally aggregated at high elevation, and more refuge area occurred in protected areas (55,184 ha) than in unprotected areas (7,024 ha). In contrast, stable refugia were scattered across the landscape, and more stable‐refugium area occurred on unprotected (40,135 ha) than on protected land (22,184 ha). Although sensitivity analysis showed that varying the thresholds that define cool environments affected outcomes, it also exposed the challenge of choosing a threshold for strategies to address climate change; there is no single value that is appropriate for all of biodiversity. The degree of overlap between ephemeral refuges and stable refugia revealed that targeting only the former for protection on currently unprotected land would capture ～17% of stable refugia. Targeting only stable refugia would capture ～54% of ephemeral refuges. Thus, targeting one type of cool environment did not fully protect the other. Evaluación de la Distribución y Estado de Protección de Dos Tipos de Ambientes Fríos para Facilitar su Conservación bajo el Cambio Climático     

Evidence of habitat structure-generated bottleneck in the recruitment process of the SW Atlantic crab<Emphasis Type="Italic"> Cyrtograpsus angulatus</Emphasis>

Shelters generated by the introduced reef-building polychaete Ficopomatus enigmaticus (Serpulidae) significantly enhance settlement of the crab Cyrtograpsus angulatus (Grapsidae) in a SW Atlantic coastal lagoon (Mar Chiquita, 37°32S; 57°26W; Argentina). However, their ultimate role in recruitment does not appear to be very important, suggesting a habitat-generated bottleneck that occurs between settlement and recruitment. Laboratory and field experiments show that newly settled crabs actively select refuges similar in size. Inside the reefs, decreases in crab density of each newly settled cohort mirror the ratio of decreases in the number of reef refuges of similar sizes, suggesting that habitat fractal structure determines the mortality rate after settlement. Field experiments using artificial shelters show that as crabs increase in size, they move outside reefs. Field tethering of juvenile- and adult-sized crabs without access to refuges showed that juvenile crabs suffer a much higher risk of mortality than adults. Our results show that the availability of small refuges provided by polychaete reefs enhances crab settlement, but, then, due to a decrease in the number of refuges as size increases, produces a demographic bottleneck during recruitment. Thus, independent of settlement intensity, enhanced survivorship in the smallest size classes due to refuge is not transmitted to larger size classes. This is likely one reason why stock-recruitment relationships may not hold in species that use shelters during both recently settled and juvenile life stages. These results provide a good example of why habitat enhancement programs need to incorporate a comprehensive evaluation of the species ontogenic ecology to avoid class size-specific bottlenecks.Communicated by P.W. Sammarco, Chauvin     

Boldness and body size of male Spanish terrapins affect their responses to chemical cues of familiar and unfamiliar males

Recognition and avoidance of conspecifics based on chemical cues could reduce the risk of aggressive interactions between males. Success in agonistics encounters with unfamiliar males should be lower than with previously known familiar males. Then, males should avoid the chemicals from unfamiliar males with respect to those from familiar males. However, boldness and size could affect the outcome of encounters between males and, consequently, the response to chemical cues of conspecific males. We compared the time spent by male turtles Mauremys leprosa in water pools with chemical stimuli from unfamiliar or familiar males or with their own chemical stimuli. We also performed a behavioral test to estimate boldness of turtles in an antipredatory situation. Turtles avoided the chemicals from unfamiliar males respect to familiar ones and their own odors, but their responses depended on boldness and size of the tested turtle. Bold turtles avoided water with chemicals of unfamiliar males, but not with chemicals of familiar males, whereas shy turtles avoided chemicals of both familiar and unfamiliar males. On the other hand, large males avoided the odor from unfamiliar males, but small males did not avoid water with the odor from other males. Results suggest that male M. leprosa can detect chemicals released to water from conspecific males and discriminate between familiar and unfamiliar males. However, responses to these chemicals depended on boldness and body size of the responding turtle because these factors may affect intrasexual competition.     

Distance to neighbours influences the trade-off between hiding after disturbance and defending food patches in convict cichlids (<Emphasis Type="Italic">Archocentrus nigrofasciatus</Emphasis>)

Territories are often aggregated. Because of this, distance to neighbours should influence how territory-holders balance safety from predators with the use and defence of resources. I examined the influence of distance to a neighbour on refuge use by pairs of convict cichlids (Archocentrus nigrofasciatus) faced with a conflict between hiding and defending food patches. Neighbours could reduce the rate of intrusions by strangers as a by-product of their own resource defence. This should allow fish with near neighbours to spend more time in the refuge. Neighbours could also steal from patches that are left undefended. This should lead to a reduction in use of the refuge. When one fish was confined to its refuge (so that its patch was undefended), theft by the other increased as inter-patch distance decreased. Distance between patches did not influence the rate of intrusion by non-territorial fish. When both fish defended patches, body mass influenced the effect of inter-patch distance on refuge use. Large fish rarely used the refuge, but small territory-holders spent more time in the refuge when patches were close together, as predicted. However, when one fish was dominant at both patches, distance between patches did not influence refuge use. These results suggest that, despite increased opportunity for theft, there is no realised foraging or defensive benefit to settling near neighbours that are of similar competitive ability.Communicated by J. Krause     

Refuge sharing network predicts ectoparasite load in a lizard

Living in social groups facilitates cross-infection by parasites. However, empirical studies on indirect transmission within wildlife populations are scarce. We investigated whether asynchronous overnight refuge sharing among neighboring sleepy lizards, Tiliqua rugosa, facilitates indirect transmission of its ectoparasitic tick, Amblyomma limbatum. We fitted 18 neighboring lizards with GPS recorders, observed their overnight refuge use each night over 3 months, and counted their ticks every fortnight. We constructed a transmission network to estimate the cross-infection risk based on asynchronous refuge sharing frequencies among all lizards and the life history traits of the tick. Although self-infection was possible, the network provided a powerful predictor of measured tick loads. Highly connected lizards that frequently used their neighbors’ refuges were characterized by higher tick loads. Thus, indirect contact had a major influence on transmission pathways and parasite loads. Furthermore, lizards that used many different refuges had lower cross- and self-infection risks and lower tick loads than individuals that used relatively fewer refuges. Increasing the number of refuges used by a lizard may be an important defense mechanism against ectoparasite transmission in this species. Our study provides important empirical data to further understand how indirectly transmitted parasites move through host populations and influence individual parasite loads.     

Intraspecific brood parasitism as a conditional reproductive tactic in the treehopper<Emphasis Type="Italic"> Publilia concava</Emphasis>

In species exhibiting egg guarding as well as communal egg laying, females may adopt the strategy of laying eggs in the nests of conspecifics and leaving without providing care (termed intraspecific brood parasitism). This study is the first to describe such a behavior in the insect Publilia concava (Hemiptera: Membracidae) through field studies that followed 849 marked females across 1,828 brood associations. While brood parasitism increased the total number of eggs in a host brood, it did not reduce the overall hatching success of host broods. Solitary females exhibited a range of guarding durations while parasitic females rarely remained to guard eggs. Females exhibiting the parasitic tactic increased their lifetime number of clutches without decreasing the number of solitary clutches that they were able to initiate. Estimates of egg number for these individual broods revealed that females adopting the parasitic tactic (in addition to solitary breeding) had higher lifetime fecundity relative to females that did not parasitize. In females that exhibited both tactics (solitary and parasitic), the parasitic tactic yielded a higher rate of oviposition. The major component of oviposition rate was the time to find hosts and this time decreased with increasing host availability across 222 replicate groups. Females exhibited a shift toward the parasitic tactic when host broods were more abundant (i.e. in larger groups and later in the season). However, the time to find hosts increased as the frequency of the parasitic tactic increased, suggesting that this tactic may be maintained through negative frequency dependence. The results of this study suggest that brood parasitism may be the preferred tactic, as part of a conditional strategy, when hosts are readily available with solitary breeding being the preferred tactic when hosts are in short supply.     

Vigilance and foraging substrate: anti-predatory considerations in a non-standard environment

Summary The commonly studied standard anti-predatory environment presents animals with spatially-distinct feeding sites and refuges from attack, neither of which necessarily obstructs predator detection. In contrast, tree-trunks provide animals with a markedly non-standard environment in which the foraging substrate itself may be a refuge from attack that unavoidably obstructs predator detection. Thus anti-predatory behavior in this environment should be influenced not only by a perceived risk of attack, but also by the nature of the refuge/foraging substrate itself. Downy woodpeckers (Picoides pubescens) are a common tree-trunk foraging animal, and an experimental analysis of their behavior suggests that they respond appropriately to their non-standard anti-predatory environment. In particular, anti-predatory vigilance varies strongly with changes in tree trunk diameter. Two modes of vigilance were apparent. In stationary vigilance, woodpeckers maintained the position of their feet while rotating their bodies side-to-side to peer around the trunk; mobile vigilance involved movement around the trunk itself. Both the frequency and angle of rotation of stationary vigilance increased with trunk diameter, as did the frequency of mobile vigilance. The woodpeckers also held their heads farther away from the trunk surface as diameter increased. All of these measures of vigilance increased under a greater perceived risk of predation. As might be expected given these results, downy woodpeckers avoided thick trunks; they did not, however, prefer the thinnest (least obstructive) available trunks. These preferences may reflect the influence of trunk diameter on thermo-ecological and/or anti-predator considerations not related to vigilance. Overall, this arboreal environment provides an unusual perspective on anti-predator decision-making with several implications for tree-trunk foraging animals in general.     

Shark-inflicted injury frequencies, escape ability, and habitat use of green and loggerhead turtles

Interactions between large marine predators and their prey are difficult to observe and little is known about the risk of predation faced by sea turtles. The frequency of predator-inflicted injuries, however, has afforded insights into the predation risk faced by many taxa. We measured the frequency of shark-inflicted injuries on green (Chelonia mydas) and loggerhead (Caretta caretta) sea turtles in Shark Bay, Western Australia with a view to determining differences between species and sex-classes in the risk of predation from tiger sharks (Galeocerdo cuvier). Furthermore, we investigated how escape ability and habitat use might influence the probability of turtles being injured by sharks. Shark-inflicted injuries were more frequent on loggerhead than on green turtles, and most frequent on adult male loggerhead turtles. Species effects could not be attributed to differences in habitat use, since green turtles were found in habitats favored by tiger sharks more often than were loggerhead turtles. Green turtles, however, were faster and maneuvered better than loggerhead turtles, suggesting that escape ability is a factor in interspecific differences in injury frequency. The sex-class difference in injury frequency of loggerhead turtles suggests that males face greater predation risk than females and may take more risks. For green turtles, the lack of a sex difference in injury frequency might be due to greater escape ability lowering overall predation risk or to no differences between sexes in the benefits of risk-taking.     

Localized defecation by pike: a response to labelling by cyprinid alarm pheromone?

Fathead minnows (Pimephales promelas) that have never encountered a predatory pike (Esox lucius), are able to detect conspecific alarm pheromone in a pike's diet if the pike has recently consumed minnows. It remains unclear how this minnow alarm pheromone is secreted by pike and if a pike is able to avoid being labelled as a potential predator by localizing these cues away from its foraging range. The first experiment determined that minnow alarm pheromone is present in pike feces when pike are fed minnows. Individual fathead minnows exhibited a fright response to a stimulus of pike feces if the pike had been fed minnows, but not if the pike had been fed swordtails, which lack alarm pheromone. Individual minnows also exhibited a fright reaction to alarm pheromone in the water (which contained no feces) housing pike which had been fed minnows, suggesting that alarm pheromone is also released in urine, mucous secretions and/or via respiration. The second experiment determined that test pike spent a significantly greater proportion of time in the home area of the test tanks (i.e. where they were fed) but the majority of feces were deposited in the opposite end of the test tank. By localizing their defecation away from the home or foraging area, pike may be able to counter the effects of being labelled as a predator by the alarm pheromone of the prey species.     

Species-specific antipredator capacities and prey refuges: interactions between piscivorous perch (Perca fluviatilis) and juvenile perch and roach (Rutilus rutilus)

The outcome of predator-prey interactions depends on the characteristics of predators and prey as well as the structure of the environment. In a replicated field enclosure experiment, we tested the effects of quantity and quality of different prey refuges (no structure, structure forming a partial refuge, and structure forming a complete refuge) on the interaction between piscivorous perch (Perca fluviatilis) and juvenile perch and roach (Rutilus rutilus). We quantified the behaviour of the predators and the prey and predator-induced prey mortality. The piscivores stayed in or close to the prey refuge and were more dispersed in the presence than in the absence of prey refuges. Survival of juvenile perch and roach decreased in the presence of predators and was higher for juvenile roach than for juvenile perch. In addition, juvenile perch survival increased with refuge efficiency Roach formed schools which were denser in the presence of predators, had a higher swimming speed (both in the open water and in the refuge) and used a larger area than juvenile perch. Both prey species decreased their distance to the prey refuge and increased the proportion of their time spent in the refuge in the presence of predators. The number of switches between the open-water habitat and the prey refuge was higher for juvenile roach than for juvenile perch. Juvenile perch used different parts of the prey refuge in a flexible way depending both on presence of predators and refuge type whereas juvenile roach used the different parts of the prey refuge in fixed proportions over all refuge treatments. Our results suggest that juvenile roach had a overall higher capacity to avoid predation than juvenile perch. However, in the presence of qualitatively different prey refuges juvenile perch responded to predators with more flexible refuge use than juvenile roach. The differences in antipredator capacities of juvenile perch and roach when subjected to piscivorous perch predation may depend on differences in life history patterns of the two species.     

Pursuit deterrence,predation risk,and escape in the lizard <Emphasis Type="Italic">Callisaurus draconoides</Emphasis>

The frequency of pursuit-deterrent signaling should vary with predation risk for factors affecting decisions to flee and hide. Distance to refuge, temperature, and microhabitat types affect risk, and their effects may differ among defenses. Because risk is greater farther from refuge and at lower temperatures that impair escape ability in ectotherms, I predicted that when farther from refuge ectothermic prey flee sooner, enter refuge more frequently, and signal more frequently. At low temperatures, prey should flee sooner, enter refuge, and signal less frequently. Because signaling sometimes deters attack, lowered risk might allow prey to permit closer approach when signaling. In the zebra-tailed lizard Callisaurus draconoides, which signals by waving its tail, signaling and the other behaviors were affected by the same risk factors. Probability of signaling before fleeing increased with distance to refuge and temperature, but variation in temperature accounted for the effect of distance to refuge. Lizards signaled not at all at low temperatures, after starting to flee at intermediate temperatures, and before, during, and after fleeing at higher temperatures. This pattern is consistent with honest signaling of escape ability. Refuge entry was more frequent nearer refuge (lower risk) and at lower temperature (higher risk). Display was less frequent on rock than ground, possibly due to conspicuousness or delay in attaining high speed. Flight initiation distance was shorter after signaling. This novel finding suggests that signaling reduces risk and prey alter escape decisions based on lower risk as a consequence of their own signaling behavior.     

Discrimination of predatory versus nonpredatory mammals by box turtles, <Emphasis Type="Italic">Terrapene carolina</Emphasis>

Summary. Although terrestrial turtles have served as a model for studies of olfactory neurophysiology, little is known about how they use chemical information in an ecological sense. We tested whether box turtles (Terrapene carolina) use chemical information to distinguish between predatory and nonpredatory mammals. Box turtles in our study exhibited more escape behavior when exposed to urine from a predator (coyote, Canis latrans) than when exposed to urine from a nonpredator (white-tailed deer, Odocoileus virginianus) or a blank control. Escape behavior is consistent with an antipredator response. In addition, the turtles decreased their handling time for food when in the presence of urine from either species of mammal in comparison to the blank, indicating that chemical cues from mammals in general may result in increased vigilance by terrestrial turtles. Examination of a variety of response variables may be important for adequate assessment of the ecological role of chemosensory behavior.     

Paternity assurance by “helpful” males: adaptations to sperm competition in burying beetles

Summary Male burying beetles invest parentally by participating in the burial of a carcass and in provisioning and guarding the larvae that come to the carcass. Since most of the females arriving at a carcass have stored fertile sperm within their spermathecae, sperm transferred by such helpful males must compete with other males' ejaculates for the fertilization of the female's eggs. We showed that these males are able to achieve a high level of paternity (mean=92%). The mechanism they employ is a repeat-mating tactic, i.e., the female is mated very frequently shortly before and during oviposition. Repeated matings are essential for a high reliability of paternity, since single copulations result in the fertilization of only a very small proportion of the female's eggs.     

The Evolution of the U.S. National Wildlife Refuge System and the Doctrine of Compatibility

This paper reviews the history and evolution of the United States National Wildlife Refuge System, perhaps the largest and oldest wildlife conservation system in the world. The refuge system is guided by the doctrine of compatibility, which permits only those uses on a refuge which are "compatible with the purposes for which the refuge was established." Memoranda and documents spanning the last 100 years were examined, and key refuge personnel interviewed, to understand the historical basis for current refuge policies. Topics of particular importance were the origins of the compatibility doctrine, the origin and meaning of the sanctuary concept, and the means by which refuges were established.
Through time, the refuge system has shifted from a policy of strict protection of wildlife to a policy of integrating public uses of refuges. The doctrine of compatibility has evolved to facilitate public use of wildlife refuges. Increasingly, the needs for recreation and resource use have interfered with the purposes far which refuges were established. A legislative solution may be needed to ensure that these areas are protected and remain viable reserves for wildlife and natural communities.
The sustained conservation of reserve areas often represents a delicate balance between political constraints and biological realities. The examination of the National Wildlife Refuge System not only has ramifications for North American conservation, but illustrates many of the problems and pitfalls facing conservation and land management worldwide.     

Maintenance of variable responses for coping with wetland drying in freshwater turtles

Aquatic animals inhabiting temporary wetlands must respond to habitat drying either by estivating or moving to other wetlands. Using radiotelemetry and capture mark recapture, we examined factors influencing the decisions made by individuals in a population of freshwater turtles (Chelodina longicollis) in response to wetland drying in southeastern Australia. Turtles exhibited both behaviors, either remaining quiescent in terrestrial habitats for variable lengths of time (terrestrial estivation) or moving to other wetlands. Both the proportion of individuals that estivated terrestrially and the time individuals spent in terrestrial habitats increased with decreasing wetland hydroperiod and increasing distance to the nearest permanent wetland, suggesting behavioral decisions are conditional or state dependent (i.e., plastic) and influenced by local and landscape factors. Variation in the strategy or tactic chosen also increased with increasing isolation from other wetlands, suggesting that individuals differentially weigh the costs and benefits of residing terrestrially vs. those of long-distance movement; movement to other wetlands was the near universal strategy chosen when only a short distance must be traveled to permanent wetlands. The quality of temporary wetlands relative to permanent wetlands at our study site varies considerably and unpredictably with annual rainfall and with it the cost-benefit ratio of each strategy or tactic. Residency in or near temporary wetlands is more successful during wet periods due to production benefits, but movement to permanent wetlands is more successful, or least costly, during dry periods due to survival and body condition benefits. This shifting balance may maintain diversity in response of turtles to the spatial and temporal pattern in wetland quality if their response is in part genetically determined.     

Role of Genetic Refuges in the Restoration of Native Gene Pools of Brown Trout

Abstract:  Captive-bred animals derived from native, alien, or hybrid stocks are often released in large numbers in natural settings with the intention of augmenting harvests. In brown trout ( Salmo trutta ), stocking with hatchery-reared non-native fish has been the main management strategy used to maintain or improve depleted wild brown trout populations in Iberian and other Mediterranean regions. This measure has become a serious threat to the conservation of native genetic diversity, mainly due to introgressive hybridization. Aware of this risk, the agency responsible for management of brown trout in the eastern Pyrenees (Spain) created "brown trout genetic refuges" to preserve the integrity of brown trout gene pools in this region. Within refuge areas, the prerefuge status with respect to fishing activities has been maintained, but hatchery releases have been banned completely. We evaluated this management strategy through a comparison of the stocking impact on native populations that accounted for stocking histories before and after refuge designations and fishing activities. In particular we examined the relevant scientific, cultural, and political challenges encountered. Despite agency willingness to change fishery policies to balance exploitation and conservation, acceptance of these new policies by anglers and genetic monitoring of refuge populations should also be considered. To improve management supported by genetic refuges, we suggest focusing on areas where the public is more receptive, considering the situation of local native diversity, and monitoring of adjacent introgressed populations. We recommend the use of directional supportive breeding only when a population really needs to be enhanced. In any case, management strategies should be developed to allow for protection within the context of human use.     

Health-dependent vulnerability to predation affects escape responses of unguarded chinstrap penguin chicks

Predators may select more often to attack the more vulnerable prey or those with an inferior health status. Thus, prey should be able to assess their own vulnerability to predation and modify their antipredatory behavior accordingly. When approached by predator skuas, unguarded penguin chicks flee short distances, and usually aggregate in dense packs, but there is a clear interindividual variability in their responses under similar conditions. We hypothesized that this variability in escape responses might be related to the perceived vulnerability to predation of each individual chick. We simulated predator attacks to chinstrap penguin chicks and analyzed the sources of variation in their escape response, such as the presence of adults or the density of other chicks, and the sex, age, body condition, and health status of responding chicks. Chicks allowed shorter approach distances when they had a better health condition (i.e., a greater T-cell-mediated immunity, CMI), when they were younger, and when the density of adults around was higher. Sex and density of other chicks were not important. Similarly, chicks fled from the experimenter to longer distances when they had a lower CMI and when the density of adults was lower. Therefore, escape characteristics of chicks depended on the presence of adults that can deter predators and on the health-dependent vulnerability of chicks.