Actual Predation Risk Research Articles

The impact of increasing turbidity on the predator–prey interactions of freshwater fishes

Abstract Human activities are exacerbating environmental change globally. Turbidity in freshwater systems is increasing due to extreme weather events and intensification of activities such as deforestation, urbanisation, agriculture and altering water flow with dams and other structures. Prey species may benefit from turbid habitats as turbidity creates a visual barrier to predators; such predators thus face reduced foraging success as water clarity decreases. However, the literature on the effects of water turbidity on predator–prey interactions in fish is often contrasting. Environmentally driven changes in predator and prey behaviour, foraging and survival ultimately have the potential to affect community structure and ecosystem functioning, thus understanding the impact of changes in turbidity on predator–prey interactions should be a priority. Given the contrasting results in the primary literature, the aim of this review is to provide a broad overview of the current knowledge on the impacts of increased turbidity on predator–prey interactions in freshwater fishes, summarise the published literature and identify knowledge gaps on this topic. We collated 281 studies from the peer‐reviewed literature that tested the impact of water turbidity on the foraging behaviour of predatory fish and anti‐predator behaviour of prey fish. We recorded changes in predator and prey behaviours in response to changes in turbidity levels. Furthermore, we reported whether these behavioural changes were considered to have a positive (e.g. enhanced foraging success) or negative (e.g. higher mortality) impact on the focal species, or whether the impact was unspecified (e.g. changes in anti‐predator behaviour without quantifying actual predation risk). The literature has some strong taxonomic, geographical and environmental biases, where most research focused on commercially important, temperate species. A large proportion of studies (47%) reported a negative (versus a positive or unspecified) impact of turbidity on the focal species (either predator or prey freshwater fish). Negative impacts were reported more often for predators (58%) compared to prey (19%) and while changes in behaviour were frequently reported in prey species, if these changes are adaptive or not is unclear, as actual predation risk was not assessed in the majority of relevant studies (55%). Increases in turbidity have complex effects on predator–prey interactions, with negative impacts more often reported for adult piscivore predators, while prey species may gain a relative advantage compared to visual predators. However, whether prey behavioural responses to increased turbidity are adaptive is generally unclear. We suggest that a more holistic and ecologically relevant experimental approach is needed to disentangle the underlying mechanisms and determine the adaptive nature of prey behavioural responses to increased turbidity. Our review serves as a broad and comprehensive summary of the empirical research conducted on impacts of increasing water turbidity on predator–prey interactions in freshwater fishes. It highlights knowledge gaps and provides a reference point to guide future experimentation and research synthesis in this crucial and growing area of research.

Conspicuous cruciform silk decorations deflect avian predator attacks.

Although camouflage as an effective antipredator defense strategy is widespread across animals, highly conspicuous color patterning is not uncommon either. Many orb-web spiders adorn their webs with extra bright white silk. These conspicuous decorations are hypothesized to deter predators by warning the presence of sticky webs, camouflaging spiders, acting as a decoy, or intimidating predators by their apparent size. The decorations may also deflect predator attacks from spiders. However, empirical evidence for this deflection function remains limited. Here, we tested this hypothesis using the X-shaped silk cruciform decorations built by females of Argiope minuta. We employed visual modeling to quantify the conspicuousness of spiders and decorations from a perspective of avian predators. Then, we determined actual predation risk on spiders using naïve chicks as predators. Spider bodies and decorations were conspicuous against natural backgrounds to the avian visual systems. Chicks attacked the spider main bodies significantly less frequently on the decorated webs than on the undecorated webs, thus reducing predation risk. When both spiders and decorations were present, chicks also attacked the spider main bodies and their legs or decorations, and not randomly: they attacked the legs or decorations sooner and more frequently than they attacked the main bodies, independent of the ratio of the surface area between the decoration and spider size. Despite the increase in detectability, incorporating a conspicuous cruciform decoration to the web effectively defends the spider by diverting the attack toward the decoration or leg, but not by camouflaging or intimidating, thus, supporting the deflection hypothesis.

Contextual behavioural plasticity in Italian agile frog (Rana latastei) tadpoles exposed to native and alien predator cues.

Predation is a strong driver for the evolution of prey behaviour. To properly assess the actual risk of predation, anuran tadpoles mostly rely on water-borne chemical cues, and their ability to evaluate environmental information is even more crucial when potential predators consist of unknown alien species. Behavioural plasticity - that is, the capacity to express changes in behaviour in response to different environmental stimuli - is crucial to cope with predation risk. We explored the defensive behaviour of Italian agile frog (Rana latastei) tadpoles when exposed to the chemical cues of two predator species, one native (dragonfly larvae) and one alien (red swamp crayfish). Firstly, we observed whether a plastic life history trait (i.e. hatching time) might be affected by native predatory cues. Secondly, we recorded a suite of behavioural responses (activity level, lateralization and sinuosity) to each cue. For assessing lateralization and sinuosity, we developed a C++ code for the automatic analysis of digitally recorded tadpole tracks. Hatching time seemed not to be affected by the potential risk of predation, while both predator species and diet affected tadpoles' defensive behaviour. Tadpoles responded to a predator threat by two main defensive strategies: freezing and 'zig-zagging'. While the first behaviour had previously been reported, the analysis of individual trajectories indicated that tadpoles can also increase path complexity, probably to prevent predators from anticipating their location. We also recorded a decrease in lateralization intensity, which suggests that under predation risk, tadpoles tend to scrutinize the surrounding environment equally on both sides.

The role of hunting on Sapajus xanthosternos' landscape of fear in the Atlantic Forest, Brazil.

Habitats with spatial variation in food availability, predation risk, and hunting pressure allow us to study how animals resolve the trade-off between food searching and predator avoidance. We investigated the influence of food availability, predation risk, and the perceived predation risk on habitat use by a primate living under high hunting pressure, the yellow-breasted capuchin monkeys, Sapajus xanthosternos, at Una Biological Reserve (ReBio Una). We hypothesized that the hunting pressure occurring in the capuchins' home range would favor predator avoidance to the detriment of searching for food. We characterized a set of covariates related to resource availability (fruit and invertebrate biomasses, feeding on dispersed and clumped food items, sleeping sites), perceived predation risk (alarm calls given to terrestrial and aerial predators, silent group movement, and vigilance behavior), and actual predation risk (evidence of hunting) and estimated their effects on how one group of capuchin monkeys uses its habitat. The group divides its time among three major forest types within their home range: agroforest, mature, and secondary. Our results suggest that the actual and perceived risk of hunting by humans, as well as the perceived predation risk by both terrestrial and aerial predators, were significant determinants of capuchin monkeys' space use. Yellow-breasted capuchin monkeys' space use was negatively related to the risk of hunting by humans (actual evidence and silent behavior), the perceived risk of predation by both aerial and terrestrial predators, and the presence of sleeping sites. Capuchin monkeys' use of space was not related to the biomass of fruits in the habitat, and the biomass of invertebrates had a very low positive effect. We confirmed our prediction that in a habitat with high hunting pressure, the risk of predation, both perceived and actual, had a more significant impact on how yellow-breasted capuchins used space than did food availability.

Habitat structure and association with ungulates drive habitat selection and grouping behaviour of lesser rhea (Rhea pennatasubsp.pennata)

AbstractSocial prey species respond to predation risk by modifying habitat selection and grouping behaviour. These responses may depend on both actual predation risk (predator probability of occurrence) and/or on perceived predation risk associated with habitat structure. Other factors like food availability and co‐occurrence with other species may also affect habitat selection and group formation. We analyse habitat selection and grouping behaviour (group size and cohesion) of lesser rhea (Rhea pennatasubsp.pennata), a ratite endemic of South America inhabiting steppe shrublands and grasslands, in relation to actual (puma probability of occurrence) and perceived (habitat structure: openness, visibility) predation risk, co‐occurrence with other herbivore species and forage availability in the Chilean Patagonia. We used data from 9 sampling seasons in 5 years. Results show that habitat selection, group size and cohesion in lesser rhea were mainly driven by variables associated with perceived predation risk and by co‐occurrence with other herbivores both during breeding and non–breeding season. As expected, lesser rhea preferred open habitats (vegas and grasslands) that allow a behaviour of ‘watch and run’ to avoid predation and formed larger groups in them. Moreover, lesser rhea positively selected year‐round habitats where livestock occur, forming large groups during non–breeding season there. Group size and co‐occurrence with other herbivores significantly decreased group cohesion, suggesting a reduction of perceived predation risk. Therefore, lesser rhea seems to take advantage of forming mixed interspecific groups to reduce predation risk. These results suggest that lesser rhea habitat selection and grouping behaviour are preferentially driven by factors related to perceived predation risk than by actual predator occurrence or food availability.

Nest-site selection and breeding success of passerines in the world's southernmost forests.

BackgroundBirds can maximize their reproductive success through careful selection of nest-sites. The ‘total-foliage’ hypothesis predicts that nests concealed in vegetation should have higher survival. We propose an additional hypothesis, the ‘predator proximity’ hypothesis, which states that nests placed farther from predators would have higher survival. We examined these hypotheses in the world’s southernmost forests of Navarino Island, in the Cape Horn Biosphere reserve, Chile (55°S). This island has been free of mammalian ground predators until recently, and forest passerines have been subject to depredation only by diurnal and nocturnal raptors.MethodsDuring three breeding seasons (2014–2017), we monitored 104 nests for the five most abundant open-cup forest-dwelling passerines (Elaenia albiceps, Zonotrichia capensis, Phrygilus patagonicus, Turdus falcklandii, and Anairetes parulus). We identified nest predators using camera traps and assessed whether habitat characteristics affected nest-site selection and survival.ResultsNest predation was the main cause of nest failure (71% of failed nests). Milvago chimango was the most common predator, depredating 13 (87%) of the 15 nests where we could identify a predator. By contrast, the recently introduced mammal Neovison vison, the only ground predator, depredated one nest (7%). Species selected nest-sites with more understory cover and taller understory, which according to the total-foliage hypothesis would provide more concealment against both avian and mammal predators. However, these variables negatively influenced nest survival. The apparent disconnect between selecting nest-sites to avoid predation and the actual risk of predation could be due to recent changes in the predator assemblage driven by an increased abundance of native M. chimango associated with urban development, and/or the introduction of exotic mammalian ground predators to this island. These predator assemblage changes could have resulted in an ecological trap. Further research will be needed to assess hypotheses that could explain this mismatch between nest-site selection and nest survival.

Risk assessment in the plateau pika (Ochotona curzoniae): intensity of behavioral response differs with predator species

BackgroundThe ability of a prey species to assess the risk that a predator poses can have important fitness advantages for the prey species. To better understand predator–prey interactions, more species need to be observed to determine how prey behavioral responses differ in intensity when approached by different types of predators. The plateau pika (Ochotona curzoniae) is preyed upon by all predators occurring in its distribution area. Therefore, it is an ideal species to study anti-predator behavior. In this study, we investigated the intensity of anti-predator behavior of pikas in response to visual cues by using four predator species models in Maqu County on the eastern Qinghai-Tibetan Plateau.ResultsThe behavioral response metrics, such as Flight Initiation Distance (FID), the hiding time and the percentage of vigilance were significantly different when exposed to a Tibetan fox, a wolf, a Saker falcon and a large-billed crow, respectively. Pikas showed a stronger response to Saker falcons compared to any of the other predators.ConclusionsOur results showed that pikas alter their behavioral (such as FID, the hiding time and the vigilance) response intensity to optimally balance the benefits when exposed to different taxidermy predator species models. We conclude that pikas are able to assess their actual risk of predation and show a threat-sensitive behavioral response.

Risk of social colours in an agamid lizard: implications for the evolution of dynamic signals.

The forces of sexual and natural selection are typically invoked to explain variation in colour patterns of animals. Although the benefits of conspicuous colours for social signalling are well documented, evidence for their ecological cost, especially for dynamic colours, remains limited. We examined the riskiness of colour patterns of Psammophilus dorsalis, a species in which males express distinct colour combinations during social interactions. We first measured the conspicuousness of these colour patterns on different substrates based on the visual systems of conspecifics and predators (bird, snake, canid) and then quantified actual predation risk on these patterns using wax/polymer lizard models in the wild. The black and red male state exhibited during courtship was the most conspicuous to all visual systems, while the yellow and orange male aggression state and the brown female colour were least conspicuous. Models bearing the courtship colour pattern experienced the highest predator attacks, irrespective of the substrate they were placed on. Thus, social colours of males are not only conspicuous but also risky. Using physiological colours to shift in and out of conspicuous states may be an effective evolutionary solution to balance social signalling benefits with predation costs.

Ultrasound avoidance by flying antlions (Myrmeleontidae).

The acoustic arms race between insectivorous bats and their invertebrate prey has led to the convergent evolution of ultrasound hearing in seven orders of nocturnal insects. Upon hearing the echolocation calls of an approaching bat, such insects take defensive action. Here, we document a previously unknown sense of ultrasound hearing and phonotactic flight behaviour in the neuropteran family Myrmeleontidae (antlions). The antlion Myrmeleon hyalinus was presented with sound pulses at ultrasonic frequencies used by echolocating bats and its response thresholds in tethered flight determined. Behaviours included abdominal twitches, wing flicks, brief pauses in flight and flight cessation. Such behaviours create erratic evasive flight manoeuvres in other eared insects, particularly mantids and lacewings. Antlions responded best to ultrasound between 60 and 80 kHz (75 dB peSPL at 80 kHz), showing response thresholds similar to those of the related lacewings (Neuroptera, Chrysopidae). Yet, at lower ultrasonic frequencies (20-50 kHz), antlions were far less sensitive than lacewings. Based on calculated response distances, we conclude that antlions respond only after having been detected by bats rather than using early evasive flights. We argue that the high response threshold for low-frequency ultrasound is adaptive for an insect that is mainly active close to and within vegetation, because a behavioural response to the lower ultrasonic frequencies used by high-flying bats would result in evasive action in the absence of actual predation risk.

Testing the link between perceived and actual risk of predation: mosquito oviposition site selection and egg predation by native and introduced fish

SummaryAccording to the threat‐sensitivity hypothesis, prey avoidance behaviour should reflect the magnitude of predation risk. Since predation can strongly affect reproduction success, ovipositing females are expected to adaptively adjust their predator avoidance response, or local breeding patch selectivity, in accordance with the perceived level of threat posed for their progeny by specific predators. However, association between avoidance and predation can be disrupted when the prey and the predator lack spatiotemporal opportunities to co‐evolve, such as in cases of non‐native predator introductions.We examined the interactions between mosquitoes (from the genusCulex) and three species of sympatric predaceous freshwater fish, a native cyprinidBarbus paludinosus, a cichlidPseudocrenilabrus philanderand an introduced poeciliid, the western mosquitofishGambusia affinis.In an outdoor mesocosm experiment, we quantified patterns ofCulexoviposition site selection across fish species using free‐roaming, caged and fish‐free treatments. In a complementary laboratory experiment, we tested the effectiveness of each fish species as predators of mosquito eggs and larvae.Synthesis and applications. We found evidence for: (i) mosquito egg raft predation by free‐roaming fish; (ii) fish‐specific avoidance by ovipositingCulex; and (iii) a positive association between fish‐specific oviposition avoidance and fish‐specific efficiency as an egg predator. These results contribute towards a better understanding of predator–prey co‐evolution, predator‐borne cue recognition, and suggest local native fish, the southern mouthbrooderPseudocrenilabrus philander, as an alternative toGambusiafor the biocontrol ofCulexmosquitoes.

The effect of predation risk on spermatophore deposition rate of the eriophyoid mite, Aculops allotrichus.

Eriophyoids are minute herbivores in which males deposit spermatophores on a substrate while females, independent of the presence of males, pick up sperm (sex dissociation). Their most dangerous enemies are phytoseiid mites. Eriophyoids can successfully avoid the predation by, e.g., forming galls in which they live, by inhabiting narrow spaces on plants, or by climbing up leaf trichomes for the time of quiescence. All these behaviours, however, are fixed and independent of the actual risk of predation. The aim of this study was to examine whether eriophyoids can respond to the cues of predation risk and how this could affect their spermatophore deposition rate. Aculops allotrichus is a vagrant eriophyoid which inhabits leaves of the black locust tree, Robinia pseudoacacia. On leaf arenas with injured conspecifics (pierced with a fine needle which simulated the attack of phytoseiids), single males of Ac. allotrichus deposited a similar number of spermatophores as on control, ‘clean’ leaves. They did not respond to the cues left by the non-enemy, yeast-fed acarid mite Tyrophagus putrescentiae either. However, they deposited significantly fewer spermatophores on leaf arenas previously exposed to the presence of the eriophyoid-fed phytoseiid mite Amblyseius swirskii. This is a first report indicating that eriophyoids can respond to the cues left by predators and change their reproductive activity accordingly. The ultimate and proximate factors that may influence the behaviour of Ac. allotrichus males are discussed.

On plasticity of aggression: influence of past and present predation risk, social environment and sex

Behaviours are phenotypically very plastic traits, allowing fast changes and fine-tuning of trait expression in response to situational demands. Aggression is a behaviour for which the right decisions can have immediate fitness consequences. Hence, the right behavioural decision is expected to be influenced by both past experience and the current environment. Using a factorial common garden experiment, we tested how past and presently perceived predation risk, as well as the social environment during development, and sex affected two components of intraspecific aggression (viz. hesitation to attack and attack intensity) in a social fish species, the three-spined stickleback (Gasterosteus aculeatus). Exposure to predator cues during development affected how fish responded to actual predation risk: exposed fish hesitated more, while predator-naive fish attacked more in the presence of predators. Sociality also had a strong effect: fish grown in pairs hesitated less than fish grown in solitude or in shoals, whereas solitary-grown fish attacked more than conspecific-experienced (i.e. pair or shoal reared) fish. Female aggression was even across the different developmental environments and lower than that of males. Predator- and conspecific-naive males were more aggressive than other males, suggesting that males have high innate aggression levels that are mediated by environmental risk. The results demonstrate complex underpinnings to plasticity-driven behavioural variation and draw attention to the fact that interpretations of presence/absence of an expected behavioural response may be difficult if it is influenced by both past and present environmental conditions.

Conspecific density affects predator‐induced prey phenotypic plasticity

The risk‐assessment hypothesis (R‐AH) states that prey must consider conspecific density and not simply the concentration of predation cues to evaluate actual predation risk. However, little is known about whether the R‐AH might serve to predict predator‐inducible plastic responses involving different prey phenotypes. We approached this question through an experiment in outdoor mesocosms, manipulating predation risk (with caged predators) and prey conspecific density to test the importance of R‐AH for the expression of predator‐induced morphological and behavioral phenotypes. We found behavioral (swimming activity) and morphological (tail width and tail muscle depth) responses of bullfrog tadpoles (Lithobates catesbeianus) to be affected by chemical predation cues. However, only the morphological phenotype responded to the predation‐risk:conspecific‐density interaction. The width of the tail and the muscle depth of tadpoles were significantly greater when individuals were exposed to predators. However, this effect was not significant in treatments with high prey density in an experimental setup that minimized intraspecific competition. We interpret the differences found among the responses of the phenotypes in terms of the effects of three factors: the potential costs related to each phenotype expression, how these phenotypes are affected by environmental conditions and the immediate response of the phenotypes to actual and perceived prey risk. Our results shed light on the fact that prey individuals use information about population density to estimate actual predation risk from chemical cues. However, different traits are differentially affected, suggesting that trade‐off mechanisms associated with the costs of the expression in terms of anti‐predator defenses may interfere with the predictions of R‐AH for multiple prey phenotypes.

Resident but not transient Eurasian Siskins reduce body mass in response to increasing predation risk: a natural experiment

The body mass of birds is the result of a trade-off between predation and starvation risks. According to the mass-dependent predation hypothesis, birds reduce their body mass when predation risk increases. Many studies have supported this hypothesis, but it has rarely been possible to investigate the mass-dependent predation risk response by comparing individuals of the same species foraging simultaneously at the same location, in the field and facing a real (not simulated) risk of predation. Wintering Siskins (Carduelis spinus) are divided in two subpopulations: residents, which stay in a foraging area for several weeks or months, and transients, which only remain a few hours or days. In this paper, we present a natural experiment by comparing the body mass variation of resident and transient Siskins between a period of the wintering season without avian predators and another period with a Sparrowhawk (Accipiter nisus) that was hunting regularly. We found that resident Siskins reduced their body mass with the presence of the hawk, while transients did not. Since in a previous study with no avian predators at the study area it was found that the difference in body condition between residents and transients did not vary throughout the wintering season, we can confidently say that the reduction in body mass observed in this study was directly linked to the presence of the hawk. We additionally found that this reduction was not associated with the dominance of residents over transients. We discuss the advantages of residence related to the knowledge of actual predation risk. Our results strongly support the mass-dependent predation hypothesis.

Plasticity in incubation behaviour under experimentally prolonged vulnerability to nest predation

Nest predation is the main cause of nest failures in many bird species. To counter this, birds have evolved different behavioural strategies to decrease the visibility of their nests, thus reducing the probability of nest detection. We manipulated the long-term perception of nest predation risk in pied flycatchers (Ficedula hypoleuca) by experimentally increasing the nest vulnerability to predators. We placed treatment and control nest-boxes for breeding pied flycatchers that appeared identical during the initial phase of breeding. But after the removal of a front panel, treatment boxes had an enlarged entrance hole, almost twice the initial diameter. This treatment increases actual predation risk and presumably parental perception of risk. Control boxes presented instead an entrance hole of the same size both before and after the manipulation. When breeding in enlarged entrance holes, females doubled the vigilance at the nest while males reduced the time spent at the nest, compared to pied flycatchers breeding in control boxes. Increased vulnerability of the nest site to predation risk, thus, induced pied flycatcher parents to increase nest vigilance while reducing their activity at the nest. These results highlight the existence of plasticity in incubation behaviours under long-term experimentally increased nest predation risk.

Compensation for predator-induced reduction in nestling feeding rate in the Great Spotted Woodpecker

Parent birds make efforts to prevent the immediate costs of predation through plastic behavioural responses to the actual predation risk, but this may incur future costs for offspring due to reduced parental care. However, the temporary nature of predator encounters suggests that nestling feeding reduced during the risky periods may be later compensated for by an increased feeding effort (the predation risk allocation hypothesis). We tested this prediction in the Great Spotted Woodpecker (Dendrocopos major) confronted with its major nest predator/competitor, the European Starling (Sturnus vulgaris). A brief encounter with a live starling was followed by a reduction in the nestling feeding rate, but the lost feedings were subsequently compensated for by an increased feeding rate. This compensatory effect was higher in older nestlings that are highly demanding in terms of energy requirements and fitness value from the parents’ perspective. Thus, birds are potentially able to respond not only to the immediate risk of nest predation by nest defence but also to compensate for the potential costs of nest defence in terms of unfulfilled nestling demands. However, data on the amount of delivered food are necessary to distinguish whether parents truly compensate for lost feeding or whether the increased feeding frequency represents a nest guarding strategy.

Does Reed Bunting (Emberiza schoeniclus) predict the risk of nest predation when choosing a breeding territory? An experimental study

Increasing evidence suggests that birds minimize the risk of nest predation by preferentially settling in territories with low predator encounter rate. However, little is known about whether they are also able to assess the actual risk of nest predation at the time of habitat occupancy and choose their breeding territories accordingly. Here, we tested this prediction experimentally, using artificial nests placed in 29 Reed Bunting territories and 29 in randomly-selected non-territories located in the same habitat. Simultaneously, we estimated the spatial distribution of the main Reed Bunting nest predator in the study area, viz. the Marsh Harrier. Nests located in Reed Bunting territories were predated less frequently (14%) than were nests placed in non-territories (41%), and this was also associated with the relative abundance of Marsh Harriers in territories (average 3.7) and in non-territories (average 5.8). These findings support the hypothesis that Reed Buntings may be capable of choosing their territories according to the actual predation risk.

Effect of Direct and Indirect Cues of Predation Risk on the Foraging Behavior of the White-Footed Mouse (Peromyscus leucopus)

Understanding predator-prey dynamics requires an understanding of how prey assess predation risk. This study tested the effect of microhabitat, moon stages, and mammalian predator urines (Vulpes vulpes [Red Fox], Mustela vison [Mink], and Procyon lotor [Raccoon]) on the degree of predation risk perceived by Peromyscus leucopus (White-footed Mouse). Giving-up densities from artificial food patches were used to quantify perceived predation risk. White-footed Mice exhibited a strong preference for cover microhabitat and for the new moon stage. However, the mice did not significantly alter their foraging behavior in response to the predator urines compared to a water control. Additionally, mice foraged less on colder nights. The results suggest that mammalian predator urines may not provide reliable information on actual predation risk for the White-footed Mice and that the mice extensively use indirect cues to assess predation risk.

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.

Testing the threat-sensitive predator avoidance hypothesis: physiological responses and predator pressure in wild rabbits

Predation is a strong selective force with both direct and indirect effects on an animal's fitness. In order to increase the chances of survival, animals have developed different antipredator strategies. However, these strategies have associated costs, so animals should assess their actual risk of predation and shape their antipredator effort accordingly. Under a stressful situation, such as the presence of predators, animals display a physiological stress response that might be proportional to the risk perceived. We tested this hypothesis in wild European rabbits (Oryctolagus cuniculus), subjected to different predator pressures, in Doñana National Park (Spain). We measured the concentrations of fecal corticosterone metabolites (FCM) in 20 rabbit populations. By means of track censuses we obtained indexes of mammalian predator presence for each rabbit population. Other factors that could modify the physiological stress response, such as breeding status, food availability and rabbit density, were also considered. Model selection based on information theory showed that predator pressure was the main factor triggering the glucocorticoid release and that the physiological stress response was positively correlated with the indexes of the presence of mammalian carnivore predators. Other factors, such as food availability and density of rabbits, were considerably less important. We conclude that rabbits are able to assess their actual risk of predation and show a threat-sensitive physiological response.