Woodland caribou habitat selection patterns in relation to predation risk and forage abundance depend on reproductive state

The ideal free distribution assumes that animals select habitats that are beneficial to their fitness. When the needs of dependent offspring differ from those of the parent, ideal habitat selection patterns could vary with the presence or absence of offspring. We test whether habitat selection depends on reproductive state due to top‐down or bottom‐up influences on the fitness of woodland caribou (Rangifer tarandus caribou), a threatened, wide‐ranging herbivore. We combined established methods of fitting resource and step selection functions derived from locations of collared animals in Ontario with newer techniques, including identifying calf status from video collar footage and seasonal habitat selection analysis through latent selection difference functions. We found that females with calves avoided predation risk and proximity to roads more strongly than females without calves within their seasonal ranges. At the local scale, females with calves avoided predation more strongly than females without calves. Females with calves increased predation avoidance but not selection for food availability upon calving, whereas females without calves increased selection for food availability across the same season. These behavioral responses suggest that habitat selection by woodland caribou is influenced by reproductive state, such that females with calves at heel use habitat selection to offset the increased vulnerability of their offspring to predation risk.     

Alternative forms of competition and predation dramatically affect habitat selection under foraging--predation-risk trade-offs

Habitat selection under foraging—predation-risk trade-offshas been a frequent topic of interest to theoretical behavioraland evolutionary ecologists. However, most habitat selectionmodels assume that individuals compete exploitatively for resourcesand that predation is either density independent or dilutedcompletely by competitor number, despite empirical evidencethat other forms of competition and predation also occur innature. I developed an individual-based model for studyingthe effects of alternative forms of competition and predationon the process of habitat selection under foraging—predation-risktrade-offs. To make the model more relevant to natural populations,I assumed that individuals vary continuously in traits relatedto competitive ability and vulnerability to predation and allowed resources and predators to be distributed across more than twohabitats. The results of my investigation demonstrate thatthe predicted pattern of habitat selection can be affecteddramatically by the form predation is assumed to take. Whenpredation is density dependent or frequency dependent, individuals will tend to be distributed across habitats according to theirabsolute vulnerability to predation. In contrast, when predationis density dependent or vulnerability dependent, individualswill tend to segregate by competitive ability. Whether oneassumes that individuals compete for resources via exploitationor interference also influences the predicted pattern of habitat selection. In general, interference competition results in amore even distribution of competitors across habitats.     

Effects of multiple vertebrate predators on grasshopper habitat selection: trade-offs due to predation risk, foraging, and thermoregulation

Predation risk can influence habitat use and activity of potential prey. I explored how the risk of predation by vertebrates influenced the behavior of grasshoppers. I monitored the height in vegetation and the frequency of resting, moving, and feeding behaviors of both tethered and free-ranging grasshoppers under exposure to various predators. Grasshoppers protected from birds remained high in the vegetation, while those protected only from small mammals and lizards remained low in the vegetation. Grasshoppers exposed to all predators occupied an intermediate height. Lower positions in the vegetation were associated with cooler thermal conditions, lower feeding rates, and lower food availability. My results are consistent with the hypothesis that grasshoppers utilize different microhabitats to balance the trade-off between reducing mortality from predators and experiencing greater food availability, and warmer conditions. This revised version was published online in July 2006 with corrections to the Cover Date.     

Predator behaviour and predation risk in the heterogeneous Arctic environment

1. Habitat heterogeneity and predator behaviour can strongly affect predator-prey interactions but these factors are rarely considered simultaneously, especially when systems encompass multiple predators and prey. 2. In the Arctic, greater snow geese Anser caerulescens atlanticus L. nest in two structurally different habitats: wetlands that form intricate networks of water channels, and mesic tundra where such obstacles are absent. In this heterogeneous environment, goose eggs are exposed to two types of predators: the arctic fox Vulpes lagopus L. and a diversity of avian predators. We hypothesized that, contrary to birds, the hunting ability of foxes would be impaired by the structurally complex wetland habitat, resulting in a lower predation risk for goose eggs. 3. In addition, lemmings, the main prey of foxes, show strong population cycles. We thus further examined how their fluctuations influenced the interaction between habitat heterogeneity and fox predation on goose eggs. 4. An experimental approach with artificial nests suggested that foxes were faster than avian predators to find unattended goose nests in mesic tundra whereas the reverse was true in wetlands. Foxes spent 3.5 times more time between consecutive attacks on real goose nests in wetlands than in mesic tundra. Their attacks on goose nests were also half as successful in wetlands than in mesic tundra whereas no difference was found for avian predators. 5. Nesting success in wetlands (65%) was higher than in mesic tundra (56%) but the difference between habitats increased during lemming crashes (15%) compared to other phases of the cycle (5%). Nests located at the edge of wetland patches were also less successful than central ones, suggesting a gradient in accessibility of goose nests in wetlands for foxes. 6. Our study shows that the structural complexity of wetlands decreases predation risk from foxes but not avian predators in arctic-nesting birds. Our results also demonstrate that cyclic lemming populations indirectly alter the spatial distribution of productive nests due to a complex interaction between habitat structure, prey-switching and foraging success of foxes.     

Hazardous duty pay and the foraging cost of predation

We review the concepts and research associated with measuring fear and its consequences for foraging. When foraging, animals should and do demand hazardous duty pay. They assess a foraging cost of predation to compensate for the risk of predation or the risk of catastrophic injury. Similarly, in weighing foraging options, animals tradeoff food and safety. The foraging cost of predation can be modelled, and it can be quantitatively and qualitatively measured using risk titrations. Giving‐up densities (GUDs) in depletable food patches and the distribution of foragers across safe and risky feeding opportunities are two frequent experimental tools for titrating food and safety. A growing body of literature shows that: (i) the cost of predation can be big and comprise the forager's largest foraging cost, (ii) seemingly small changes in habitat or microhabitat characteristics can lead to large changes in the cost of predation, and (iii) a forager's cost of predation rises with risk of mortality, the forager's energy state and a decrease in its marginal value of energy. In titrating for the cost of predation, researchers have investigated spatial and temporal variation in risk, scale‐dependent variation in risk, and the role of predation risk in a forager's ecology. A risk titration from a feeding animal often provides a more accurate behavioural indicator of predation risk than direct observations of predator‐inflicted mortality. Titrating for fear responses in foragers has some well‐established applications and holds promise for novel methodologies, concepts and applications. Future directions for expanding conceptual and empirical tools include: what are the consequences of foraging costs arising from interference behaviours and other sources of catastrophic loss? Are there alternative routes by which organisms can respond to tradeoffs of food and safety? What does an animal's landscape of fear look like as a spatially explicit map, and how do various environmental factors affect it? Behavioural titrations will help to illuminate these issues and more.     

Microhabitat variation and sexual selection can maintain male color polymorphisms

Male color polymorphism may be an important precursor to sympatric speciation by sexual selection, but the processes maintaining such polymorphisms are not well understood. Here, we develop a formal model of the hypothesis that male color polymorphisms may be maintained by variation in the sensory environment resulting in microhabitat-specific selection pressures. We analyze the evolution of two male color morphs when color perception (by females and predators) is dependent on the microhabitat in which natural and sexual selection occur. We find that an environment of heterogeneous microhabitats can lead to the maintenance of color polymorphism despite asymmetries in the strengths of natural and sexual selection and in microhabitat proportions. We show that sexual selection alone is sufficient for polymorphism maintenance over a wide range of parameter space, even when female preferences are weak. Polymorphisms can also be maintained by natural selection acting alone, but the conditions for polymorphism maintenance by natural selection will usually be unrealistic for the case of microhabitat variation. Microhabitat variation and sexual selection for conspicuous males may thus provide a situation particularly favorable to the maintenance of male color polymorphisms. These results are important both because of the general insight they provide into a little appreciated mechanism for the maintenance of variation in natural populations and because such variation is an important prerequisite for sympatric speciation.     

Adult sex-ratio in ostracods and its implications for sexual selection

ABSTRACT

The Ostracoda – ubiquitous aquatic micro-crustaceans – show an exceptionally high incidence of female-biased adult sex ratio. Intraspecific sex ratio is known to vary in extant species and yet in the fossil record a species’ adult sex ratio can be highly stable across time. Sex ratio conditions the intensity of sexual selection and influences which sex undergoes stronger selective pressure. However, the impact of variation in spatial and temporal intraspecific sex ratio on the evolution of sexual selection remains an open question, calling for further investigations on the factors controlling adaptive sex ratio. This mini-review aims to introduce the system, and explores some of the key literature addressing factors influencing intraspecific variation in adult sex ratio (ASR) and its implication in the intensity of sexual selection and evolution of mating systems.     

Effect of predation risk on the presence and persistence of yellow-bellied marmot (Marmota flaviventris) colonies

Habitat selection may have population level consequences and ultimately may influence a population's local persistence or extinction. We capitalized on a long-term study (1962–2004) of yellow-bellied marmots Marmota flaviventris in and around the Rocky Mountain Biological Laboratory, Gothic, Colorado, USA, and compared habitat characteristics associated with food availability and predation risk to explain variation in persistence of marmots at 27 sites, and their absence at 22 additional, randomly selected sites. We classified sites as persistent, intermittent or null based on whether there was a history of extinction; intermittent sites periodically went extinct and null sites never had marmots. Logistic regression analyses revealed that environmental variables associated with visibility and safety, rather than food, correctly classified sites as persistent or non-persistent as well as persistent or intermittent. Discriminant function analysis that included the null sites revealed that the same visibility-related characteristics predicted where marmots were found. These results highlight the importance of variation in safety among sites in predicting long-term population persistence, as well as a species' distribution.     

Experimental evidence for predation risk sensitive oviposition by a mosquito, Culiseta longiareolata

1. Females should choose to oviposit in habitats where risk of predation and competition are low. The ovipositional responses of a mosquito, Culiseta longiareolata, to a predator and to species sharing the same trophic level as this mosquito (controphic species) were assessed experimentally in outdoor artificial pools. 2. The predator, larval Anax imperator, which strongly reduced larval C. longiareolata survival, resulted in a 52% reduction of C. longiareolata egg rafts. The controphic species (primarily Daphnia magna), which had a small but statistically significant negative effect on the survival of C. longiareolata larvae, did not have a statistically significant influence on the number of egg rafts. 3. Laboratory trials indicated that only a small fraction of the reduced number of egg rafts seen in predator pools may be due to consumption of the egg rafts by A. imperator. 4. The experimental evidence indicates that the reduced number of C. longiareolata egg rafts found in the presence of A. imperator is due largely to oviposition habitat selection, i.e. C. longiareolata females choose pools with low risk of predation for their offspring.     

Predator related oviposition site selection of aquatic beetles (Hydroporus spp.) and effects on offspring life-history

1. Theory predicts that natural selection should favour females that are able to correctly assess the risk of predation and then use that information to avoid high‐risk oviposition sites to reduce the risk of offspring predation. Despite the potential significance of such behaviour on individual fitness, population dynamics and community structure, relatively few studies of oviposition behaviour connected to the risk of predation have been carried out. 2. However, some recent studies suggest that oviposition site selection in response to risk of predation may be a common phenomenon, at least among amphibians and mosquitoes. A vast majority of previous studies have, however, neglected to investigate how the offspring are affected, in terms of fitness related parameters, by the maternal oviposition site choice. 3. In an outdoor artificial pond experiment we tested the oviposition site selection of female aquatic beetles (Hydroporus spp.) in relation to the presence or absence of a predatory fish (Perca fluviatilis). In addition, we monitored how the oviposition site selection affected the behaviour, growth and food resource of the progeny. 4. We show that free‐flying females of the aquatic beetles Hydroporus incognitus and H. nigrita prefer to oviposit in waters without fish compared with waters with fish. Larval activity of Hydroporus spp. was unaffected by fish presence. Our results indicate that beetle larvae from females that do lay eggs in waters with fish show increased growth compared with larvae in waters without fish. We explain this difference in growth by a higher per‐capita food supply in the presence of a fish predator. This finding may have important implications for our understanding of how the variance of oviposition site selection in a population is sustained.     

Temperature modulates the effects of predation and competition on mosquito larvae

1. Predation risk affects interspecific competition by decreasing foraging activity and relative competitive ability. Predation risk is determined by predators' prey choice and prey responses, both of which can be influenced by temperature. Temperature is especially important for larval prey and can result in a trade‐off between predator‐induced decreases in foraging activity and growth. Interspecific competition must also be examined in relation to intraspecific density‐dependent competition; weaker interspecific competition leads to coexistence of competitors. 2. This study explored how temperature (15 and 25 °C) could affect a focal species, larvae of the mosquito Culex quinquefasciatus, by examining prey choice in a shared predator (mosquitofish; Gambusia holbrooki) and the effects of predation risk on interspecific competition with Limnodynastes peronii tadpoles. Intraspecific density‐dependent competition in C. quinquefasciatus at these temperatures was also examined. 3. At 25 °C, G. holbrooki consumption of both C. quinquefasciatus and L. peronii increased; however, the effects of interspecific competition on mosquito survival did not decrease with L. peronii exposure to predation risk. The relationship between intraspecific density‐dependent competition and interspecific competition was temperature‐dependent, with competitive dominance of L. peronii at 25 °C. Male and female mosquitoes had different temperature‐dependent responses, indicating sex‐specific intrinsic responses to starvation and differential selection pressures. At 25 °C, females were susceptible to interspecific competition by L. peronii, while males were susceptible to intraspecific competition. 4. The use of competitors as biological controls has implications for mosquito disease transmission, and these results suggest that control effectiveness may be modified by climate change.     

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 density on frequency-dependent food selection: a comparison of four experiments with wild birds

We compare the results of four experiments, conducted at different times and with different protocols, that explored the relationship between frequency-dependent selection and prey density in wild birds feeding on artificial populations of coloured baits. One (experiment 4) used pastry baits that differed only in the presence or absence of a red stripe, and this experiment provided no evidence for any kind of selective behaviour. The other three experiments used green and brown baits, and they all provided evidence for a trend towards increasing anti-apostatic selection with high densities (>100 baits m^–2). However, one of these (experiment 3) provided no evidence for frequency-dependent selection at low densities (0.5–20 baits m^–2), while the other two experiments concurred in suggesting a trend towards increasing apostatic selection with low densities (down to 2 baits m^–2). Together, these experiments both support and qualify the published findings of experiment 1 that frequency- dependent selection by wild birds on bait populations is modified by density. Experiment 4 indicates that frequency-dependent selection may break down entirely if bait types are too similar, while experiment 3 indicates that some details of this trend with density will depend either on the protocol used or on exogenous changes in the birds’ feeding behaviour. Received: 1 September 1999 / Accepted: 22 March 2000     

Stage-dependent predation on competitors: consequences for the outcome of a mosquito invasion

1. Predator-mediated coexistence occurs when predation allows competitors to coexist, due to preferential consumption of a superior competitor relative to an inferior competitor. Differences between the native treehole mosquito ( Aedes triseriatus ) and the co-occurring Asian tiger mosquito ( Aedes albopictus ) in anti-predatory larval behaviours account, in part, for the greater vulnerability of this invasive species to native predatory midge ( Corethrella appendiculata ). We test the hypothesis that stage-dependent differences in the sizes of A. albopictus and A. triseriatus larvae, relative to the size-limited C. appendiculata , contribute to differential consumption and the likelihood of predator-mediated coexistence of these competitors.
2. In all instars, larvae of A. triseriatus were larger than A. albopictus of the same stage. Third and fourth instar C. appendiculata selectively consumed late-stage A. albopictus in preference to same-stage A. triseriatus . Small, early-stage prey larvae did not differ in vulnerability to predation, but large, late-stage larvae differed significantly in vulnerability to predation, probably owing to size-limited predation by fourth instar C. appendiculata. This effect was less pronounced for third instar C. appendiculata .
3. Prey size, in conjunction with anti-predatory behavioural responses, alters the probability of predator-mediated coexistence. A stage-structured predation model showed that equally vulnerable early stages reduce the range of environmental conditions (productivities) in which predator-mediated coexistence is possible, increasing the likelihood of both competitive exclusion of the resident species or failure of the invasive to establish. These results underscore the importance of stage-dependent interspecific differences in predator–prey interactions for determining how predators may affect community composition.     

Perch selection by singing chaffinches: a better view of surroundings and the risk of predation

Predation risk varies among perches, and the vulnerability ofsinging chaffinches (Fringilla coelebs) might differ dependingon where they perch in a tree. To find out how singing of thechaffinch is associated with antipredatory behavior, I studiedperch selection in mature pine forest and in pine saplings,two habitats differing in the amount of cover for protection from predators. My results show that male chaffinches preferto sing below the canopy of mature pines and in the uppermostparts of sapling pines. Although these are the canopy partsmost exposed to sparrowhawk (Accipiter nisus) attacks, stayingin open sites may be a beneficial strategy for singing chaffinchesbecause it makes it possible to improve antipredatory vigilance.This assumption was supported by the hawk experiments. Afterbeing exposed to the sparrowhawk model, all of the singingbirds selected the conspicuous perch below the canopy of maturepines. The males that perched higher in the canopy before theexperiment moved to the lower canopy, whereas those males thatsang below the canopy did not change their singing perch.     

Beak marks on the wings of butterflies and predation pressure in the field

Beak marks on butterfly wings have been used as an indicator of predation pressure. The relationship between butterflies and their predators in the field was examined to evaluate the beak mark rate as an indicator of predation pressure. Transect censuses were conducted to measure the beak mark rate on butterflies from May to November, 2010, in Shiga Prefecture, Japan. A total of 1216 butterflies of 42 species from five families were observed during the study period. The beak mark rate in the Papilionidae was significantly higher than those of all other families. Analysis of the monthly fluctuations in the beak mark rate and relative abundance of predators revealed that the higher beak mark rates observed for two butterfly families (Papilionidae and Nymphalidae) were significantly and positively related to predation pressure, while no significant relationship was found for other families (Lycaenidae, Pieridae and Hesperiidae). Beak marks in larger butterflies (Papilionidae and Nymphalidae) can be used as an indicator to evaluate the relative intensity of predation pressure in the field.