Paternal Genetic Effects on Foraging Decision-Making Under the Risk of Predation

Foraging behavior under the risk of predation has interested biologists for decades. Here, we examine paternal genetic effects on foraging decisions of bluegill (Lepomis macrochirus) larvae sired by males adopting alternative life histories. We use split in vitro fertilization to generate maternal half‐siblings sired by either a ‘parental’ male or a ‘cuckolder’ male. Immediately, upon the switch to exogenous feeding, we fed the larvae ad libitum for 2 d. We then starved the larvae for between 12 and 17 h, following which we subjected them to a dichotomous choice foraging trial, where one side of a test tank posed a risky foraging habitat and the other posed a safe foraging habitat. Equal amounts of food were simultaneously introduced to both sides of the tank and the proportion of individuals on either side was recorded. There were significantly fewer cuckolder offspring foraging on the risky side of the tank when compared with their parental half‐siblings indicating that cuckolder offspring took fewer risks than parental offspring. These results demonstrate a paternal genetic effect (sire life history) on foraging behavior. We ruled out energetic state as a possible explanation for this difference because the half‐siblings did not differ in body length or mass. Instead, previous research suggests that cuckolder offspring have higher conversion efficiency (efficiency of converting food into soma) than parental offspring and therefore the differences in foraging behavior observed here may, in part, be attributed to genetic differences in conversion efficiency.     

Sex‐Specific Response of Pardosa milvina (Araneae: Lycosidae) to Experience with a Chemotactile Predation Cue

Predators frequently leave behind chemical information (i.e., semiochemicals such as pheromones or kairomones) that can be detected by their prey and used to avoid areas where predators are likely present. Prey that have interacted indirectly with predators via chemical information thus may gain insight into their risk of being consumed that naïve individuals lack. Pardosa milvina (Araneae: Lycosidae) is a chemosensitive wolf spider that shows adaptive responses to chemotactile cues deposited by the larger wolf spider Tigrosa helluo. We raised offspring from P. milvina to examine the effect of experience with a predation cue on activity, foraging, and antipredator behavior. Spiders differed in activity and foraging behavior across ontogeny and between sexes, but there was no effect of experience with a predation cue. However, a sex‐specific effect of experience was found in antipredator behavior. Male spiders, but not females, used experience with a predator cue to increase their survival in the presence of a live predator. Specifically, naïve males were attacked sooner than experienced males, indicating that prior exposure to predator cues can modify Pardosa antipredator behavior. Intersexual differences in how spiders respond to experience with a predation cue likely reflect the risk of predation faced by males and females in nature.     

动物觅食行为对捕食风险的反应

动物进行任何活动时均面临被捕食的风险 ,分析捕食风险与猎物觅食行为的关系 ,有助于揭示捕食者与猎物的协同进化机制。捕食风险具有限制或调节猎物种群数量的功能。在进化时间内 ,对猎物形态和行为特征的进化是潜在的选择压力之一 ,可利用环境因子作为信息源估测食物可利用性和捕食风险大小的动物 ,具有更大的适合度。信息源可分为包括视觉的、听觉的和化学的。动物进行觅食活动时 ,依据信息源的变化确定环境中捕食风险的大小 ,并根据自身的质量在捕食风险的大小之间做出权衡 ,通过食物选择、活动格局和栖息地利用等行为的变化降低捕食风险     

Behavioural Response of Bats to Perceived Predation Risk While Foraging

The ability to detect and respond to predation risk while foraging may have important fitness consequences for prey organisms. Anti‐predator behaviours may reduce the probability of mortality because of predation, but they may also be associated with reduced foraging efficiency. Several behaviours of bats have been suggested to serve as anti‐predator responses, and there is evidence that predation, particularly by avian predators such as owls, may be an important cause of bat mortality. Previous studies have attempted to determine whether predator presence affects bat behaviour when emerging from roost sites, but few have examined effects of predator presence on bat behaviour while foraging. In this study, we investigated whether foraging bats respond to predator cues by presenting bats with an acoustic cue simulating the presence of an owl. Within matched trials, which were conducted at different locations each of 18 nights, significantly fewer bat detections were recorded at owl playback stations than at control stations (no auditory cue), suggesting an avoidance response by bats. An acoustic control (i.e. station playing woodpecker calls), however, did not have significantly more detections than the stations playing the owl calls, suggesting that bats may simply be avoiding noise and more detailed investigation is warranted. Although evidence for owl predation on bats is minimal in North America, results of this study may indicate that the perceived presence of owls may represent a factor influencing the behaviour of bats while foraging.     

Effects of Structural Refuge and Density on Foraging Behaviour and Mortality of Hungry Tadpoles Subject to Predation Risk

Theoretical models of prey behaviour predict that food‐limited prey engage in risk‐prone foraging and thereby succumb to increased mortality from predation. However, predation risk also may be influenced by factors including prey density and structural cover, such that the presumed role of prey hunger on predation risk may be obfuscated in many complex predator–prey systems. Using a tadpole (prey) – dragonfly larva (predator) system, we determined relative risk posed to hungry vs. sated prey when both density and structural cover were varied experimentally. Overall, prey response to perceived predation risk was primarily restricted to increased cover use, and hungry prey did not exhibit risk‐prone foraging. Surprisingly, hungry prey showed lower activity than sated prey when exposed to predation risk, perhaps indicating increased effort in search of refuge or spatial avoidance of predator cues among sated animals. An interaction between hunger level and predation risk treatments indicated that prey state affected sensitivity to perceived risk. We also examined the lethal implications of prey hunger by allowing predators to select directly between hungry and sated prey. Although predators qualitatively favoured hungry prey when density was elevated and structural cover was sparse, the overall low observed variation in mortality risk between hunger treatments suggests that preferential selection of hungry prey was weak. This implies that hunger effects on prey mortality risk may not be readily observed in complex landscapes with additional factors influencing risk. Thus, current starvation‐predation trade‐off theory may need to be broadened to account for other mechanisms through which undernourished prey may cope with predation risk.     

The Importance of Predation Risk and Missed Opportunity Costs for Context-Dependent Foraging Patterns

Correct assessment of risks and costs of foraging is vital for the fitness of foragers. Foragers should avoid predation risk and balance missed opportunities. In risk-heterogeneous landscapes animals prefer safer locations over riskier, constituting a landscape of fear. Risk-uniform landscapes do not offer this choice, all locations are equally risky. Here we investigate the effects of predation risk in patches, travelling risk between patches, and missed social opportunities on foraging decisions in risk-uniform and risk-heterogeous landscapes. We investigated patch leaving decisions of 20 common voles (M. arvalis) in three experimental landscapes: safe risk-uniform, risky risk-uniform and risk-heterogeneous. We varied both the predation risk level and the predation risk distribution between two patches experimentally and in steps, assuming that our manipulation consequently yield different distributions and levels of risk while foraging, risk while travelling, and costs of missed, social opportunities (MSOCs). We measured mean GUDs (giving-up density of food left in the patch) for both patches as a measure of foraging gain, and delta GUD, the differences among patches, as a measure of the spatial distribution of foraging effort over a period of six hours. Distribution of foraging effort was most even in the safe risk-uniform landscapes and least even in the risk-heterogeneous landscape, with risky risk-uniform landscapes in between. Foraging gain was higher in the safe than in the two riskier landscapes (both uniform and heterogeneous). Results supported predictions for the effects of risk in foraging patches and while travelling between patches, however predictions for the effects of missed social opportunities were not met in this short term experiment. Thus, both travelling and foraging risk contribute to distinct patterns observable high risk, risk-uniform landscapes.     

Effects of Predation Risk on Habitat Selection by Water Column Fish, Benthic Fish and Crayfish in Stream Pools

Predation risk can affect habitat selection by water column stream fish and crayfish, but little is known regarding effects of predation risk on habitat selection by benthic fish or assemblages of fish and crayfish. I used comparative studies and manipulative field experiments to determine whether, (1) habitat selection by stream fish and crayfish is affected by predation risk, and (2) benthic fish, water column fish, and crayfish differ in their habitat selection and response to predation risk. Snorkeling was used to observe fish and crayfish in, (1) unmanipulated stream pools with and without large smallmouth bass predators (Micropterus dolomieui >200 mm total length, TL) and (2) manipulated stream pools before and after addition of a single large smallmouth bass, to determine if prey size and presence of large fish predators affected habitat selection. Observations of microhabitat use were compared with microhabitat availability to determine microhabitat selection. Small fish (60–100 mm TL, except darters that were 30–100 mm TL) and crayfish (40–100 mm rostrum to telson length; TL) had significantly reduced densities in pools with large bass, whereas densities of large fish and crayfish (> 100 mm TL) did not differ significantly between pools with and without large bass. Small orangethroat darters (Etheostoma spectabile), northern crayfish (Orconectes virilis), and creek chubs (Semotilus atromaculatus) showed significantly greater densities in pools without large bass. The presence of large smallmouth bass did not significantly affect depths selected by fish and crayfish, except minnows, which were found significantly more often at medium depths when bass were present. Small minnows and large and small crayfish showed the greatest response to additions of bass to stream pools by moving away from bass locations and into shallow water. Small darters and sunfish showed an intermediate response, whereas large minnows showed no significant response to bass additions. Response to predation risk was dependent on prey size and species, with preferred prey, crayfish and small minnows, showing the greatest response. Small benthic fish, such as darters, are intermediate between small water column fish and crayfish and large water column fish in their risk of predation from large smallmouth bass.     

相关风险因子对高原鼠兔摄食行为的影响

研究了捕食风险环境中集和洞口距离对高原鼠兔摄食行为的影响。结果表明,集群数量的增加不仅降低了警觉行为,同时也减少了摄食行为,在高风险环境中,集群为1时的取食行为强度最大,低风险环境中,为0时最大,警觉行为主要出现在距洞口2m的范围内,其行为强度与洞口踪影职责负相关,当洞口距离大于3m时,风险处理区的高原鼠兔几乎无警觉行为出现,且该处理区的取食区域几乎压缩的洞口旁,研究结果表明,在捕食风险环境中,高原鼠兔摄食行为与集群和洞口距离之间具有复杂的关系,其行为决策反映了降低风险与摄取食物间的权衡,行为目标是在降低捕食风险的同时尽可能地取食食物。     

Group Dynamics of Zebra and Wildebeest in a Woodland Savanna: Effects of Predation Risk and Habitat Density

Background

Group dynamics of gregarious ungulates in the grasslands of the African savanna have been well studied, but the trade-offs that affect grouping of these ungulates in woodland habitats or dense vegetation are less well understood. We examined the landscape-level distribution of groups of blue wildebeest, Connochaetes taurinus, and Burchell''s zebra, Equus burchelli, in a predominantly woodland area (Karongwe Game Reserve, South Africa; KGR) to test the hypothesis that group dynamics are a function of minimizing predation risk from their primary predator, lion, Panthera leo.

Methodology/Principal Findings

Using generalized linear models, we examined the relative importance of habitat type (differing in vegetation density), probability of encountering lion (based on utilization distribution of all individual lions in the reserve), and season in predicting group size and composition. We found that only in open scrub habitat, group size for both ungulate species increased with the probability of encountering lion. Group composition differed between the two species and was driven by habitat selection as well as predation risk. For both species, composition of groups was, however, dominated by males in open scrub habitats, irrespective of the probability of encountering lion.

Conclusions/Significance

Distribution patterns of wildebeest and zebra groups at the landscape level directly support the theoretical and empirical evidence from a range of taxa predicting that grouping is favored in open habitats and when predation risk is high. Group composition reflected species-specific social, physiological and foraging constraints, as well as the importance of predation risk. Avoidance of high resource open scrub habitat by females can lead to loss of foraging opportunities, which can be particularly costly in areas such as KGR, where this resource is limited. Thus, landscape-level grouping dynamics are species specific and particular to the composition of the group, arising from a tradeoff between maximizing resource selection and minimizing predation risk.     

Multi-Scale Foraging Habitat Use and Interactions by Sympatric Cervids in Northeastern China

Abstract: Moose (Alces alces) and roe deer (Capreolus pygargus) are sympatric in the forest region of northeastern China. Using univariate analyses of feeding sign data, we found the 2 species were positively associated, but there were distinctions between their use of forage resources across landscape, patch, and microhabitat scales. We used resource selection function models to predict the influence of environmental covariates on moose and roe deer foraging; we detected covariate effects at the landscape and microhabitat scales but not at the patch scale. Forage resources used by the 2 species were similar, but moose used wetter areas and more low-visibility habitats than did roe deer, which strongly avoided areas with sparse vegetation. Both species were influenced by forage abundance and distribution at the microhabitat scale but exhibited differences in intensity of use of plant species and microhabitats. Moose used areas with deeper snow and avoided hiding cover; roe deer avoided areas with higher total basal areas of tree stems and preferred areas with high plant species richness. For moose, there was a trade-off in the use of concealment cover between the landscape and microhabitat scales. We detected avoidance by moose of roads where roe deer occurred. Roe deer exhibited more capacity for coping with human disturbance and interspecific interaction. In areas similar to our study area, road closures and suppression of roe deer near roads within 3–5 years postlogging may benefit moose. Furthermore, a mosaic of areas with different logging intervals may contribute to spatial separation of moose and roe deer and promote their coexistence.     

Foraging Patch Selection in Winter: A Balance between Predation Risk and Thermoregulation Benefit

In winter, foraging activity is intended to optimize food search while minimizing both thermoregulation costs and predation risk. Here we quantify the relative importance of thermoregulation and predation in foraging patch selection of woodland birds wintering in a Mediterranean montane forest. Specifically, we account for thermoregulation benefits related to temperature, and predation risk associated with both illumination of the feeding patch and distance to the nearest refuge provided by vegetation. We measured the amount of time that 38 marked individual birds belonging to five small passerine species spent foraging at artificial feeders. Feeders were located in forest patches that vary in distance to protective cover and exposure to sun radiation; temperature and illumination were registered locally by data loggers. Our results support the influence of both thermoregulation benefits and predation costs on feeding patch choice. The influence of distance to refuge (negative relationship) was nearly three times higher than that of temperature (positive relationship) in determining total foraging time spent at a patch. Light intensity had a negligible and no significant effect. This pattern was generalizable among species and individuals within species, and highlights the preponderance of latent predation risk over thermoregulation benefits on foraging decisions of birds wintering in temperate Mediterranean forests.     

非致命性捕食风险对鸟类的影响

非致命性捕食风险是捕食者对猎物产生的间接的、非致命性的捕食压力,对维持捕食者-猎物种群数量的相对稳定具有重要意义。鸟类拥有较复杂的反捕食对策,是研究非致命性捕食风险的理想对象。本文综述了非致命性捕食风险对鸟类影响的研究进展。研究者通过多种方法模拟非致命性捕食风险,发现不同鸟类的耐受性不仅会影响个体适合度,也会影响其种群数量动态。鸟类对非致命性捕食压力的行为响应受诸多因素影响,包括鸟类自身状态,栖树高度,集群大小等。研究鸟类对非致命性捕食风险的响应,对完善捕食风险权衡理论,建立种群数量预测模型,以及探讨人类与鸟类共存机制具有重要意义,同时可为生态旅游中提出合理的接近距离、制定有效的保护管理措施提供科学指导。     

The Influence of Pedipalp Autotomy on the Courtship and Mating Behavior of Pardosa milvina (Araneae: Lycosidae)

Male Pardosa milvina wolf spiders use their pedipalps both for copulation and courtship. Pedipalp loss is significantly more common among adult males compared to females. We measured the courtship and mating effects associated with the loss of one or both pedipalps among adult male P. milvina. Pedipalp loss significantly reduced courtship intensity, but had no influence on mounting success. Intact males were less likely to be cannibalized and suffered fewer predatory attacks by females than autotomized males. Loss of the left pedipalp resulted in significantly less intense courtship, higher female aggression levels, and delayed onset of courtship whereas loss of the right pedipalp minimally affected male and female behavior relative to intact males. Pedipalp displays may function in reducing female aggression rather than increasing female receptivity.     

Predation Risk and Opportunity Cost of Fleeing While Foraging on Plants Influence Escape Decisions of an Insular Lizard

Cost‐benefit models of escape behaviour predict how close a prey allows a predator to approach [flight initiation distance (FID)] based on cost of not fleeing (predation risk) and cost of fleeing (loss of opportunities). Models for FID have been used with some success to predict distance fled (DF). We studied effects of foraging opportunity cost of fleeing and examined differences between age‐sex groups in the omnivorous Balearic Lizard, Podarcis lilfordi. Balearic lizards forage on the ground for invertebrate prey and climb the thistle Carlina corymbosa to forage on its inflorescences. We studied escape behaviour in three experimental groups, with human beings as simulated predators: lizard foraging above ground on C. corymbosa, foraging on the ground away from thistles and on the ground with cut inflorescences. Flight initiation distance was shorter for lizards with cut inflorescences than for (1) lizards above ground due to the greater risk above ground due to conspicuousness of black lizards on yellow flowers; and (2) lizards on ground away from flowers due to the cost of leaving while feeding. The only age‐sex difference was slightly greater FID for adult males than subadults, presumably because larger adult males are more likely to be attacked by predators. Other potential factors affecting this difference are discussed. Experimental group and age‐sex group did not interact for FID or DF. Because lizards foraging on inflorescences above ground fled to the base of the plants to refuge provided by spiny thistle leaves, their DF was shorter than in the other groups, which fled across the ground, usually without entering refuge. DF did not differ between groups on the ground or among age‐sex groups. The predicted shorter DF for lizards with cut inflorescences than on ground without inflorescences did not occur. We hypothesize that the opportunity cost was small due to the abundance of blooming thistles and that DF may be less sensitive to opportunity cost than FID.     

Effects of Predation Risk and Food Availability on the Activity,Habitat Use,Feeding Behavior and Mating Behavior of a Pond Water Strider,Gerris marginatus (Hemiptera)

By experimentally manipulating predation risk and food availability, we explored the effects of these factors on refuge use, general activity, feeding activity, and mating activity of adults of the pond water strider Gerris marginatus. Predation risk (from adult backswimmers Notonecta undulata) profoundly influenced the behavior of G. marginatus, reducing mating activity, general activity in males and females, decreasing feeding activity in females and increasing the use of refuge by both sexes. Surprisingly, variations in food availability failed to produce significant effects on any of these behavior patterns. Risk x food interactions were also not statistically significant for any gerrid behavior. The significant responses to predators all changed in directions that should decrease the risk of predation for G. marginatus.     

Predation Risk Perception,Food Density and Conspecific Cues Shape Foraging Decisions in a Tropical Lizard

When foraging, animals can maximize their fitness if they are able to tailor their foraging decisions to current environmental conditions. When making foraging decisions, individuals need to assess the benefits of foraging while accounting for the potential risks of being captured by a predator. However, whether and how different factors interact to shape these decisions is not yet well understood, especially in individual foragers. Here we present a standardized set of manipulative field experiments in the form of foraging assays in the tropical lizard Anolis cristatellus in Puerto Rico. We presented male lizards with foraging opportunities to test how the presence of conspecifics, predation-risk perception, the abundance of food, and interactions among these factors determines the outcome of foraging decisions. In Experiment 1, anoles foraged faster when food was scarce and other conspecifics were present near the feeding tray, while they took longer to feed when food was abundant and when no conspecifics were present. These results suggest that foraging decisions in anoles are the result of a complex process in which individuals assess predation risk by using information from conspecific individuals while taking into account food abundance. In Experiment 2, a simulated increase in predation risk (i.e., distance to the feeding tray) confirmed the relevance of risk perception by showing that the use of available perches is strongly correlated with the latency to feed. We found Puerto Rican crested anoles integrate instantaneous ecological information about food abundance, conspecific activity and predation risk, and adjust their foraging behavior accordingly.     

Hamsters Use Predator Odors as Indirect Cues of Predation Risk

Golden hamsters (Mesocricetus auratus) use olfactory cues to assess traits of conspecifics such as kinship, individual identity, and reproductive status. The environment, however, is full of a wide variety of other olfactory information such as signals emitted by some of the hamster’s primary predators. Given this, we hypothesized that hamsters use odors from predators as an indirect sign of increased predation risk in the environment. In addition, based on data that show that wild hamsters are diurnal while laboratory hamsters are nocturnal, we hypothesized that if golden hamsters did respond to the predator odors, perceived predator risk might influence daily activity patterns in hamsters. We tested male and female hamsters over 5 d with scent gland secretion from domestic ferrets (Mustela putorius furo) and compared their behavior to that observed when they were exposed to a clean arena. In response to the predator odor, subjects significantly decreased the amount of time active outside of their burrow, returned to their burrow more quickly, and spent less time near the predator odor than the clean control stimulus. These results strongly support our hypothesis that hamsters, like other species of small mammals, avoid predator odors. The results did not, however, support our second hypothesis that exposure to predator odors during the dark phase of the light cycle would elicit a switch to a more diurnal pattern of activity. More work is needed to understand how environmental cues and internal mechanisms interact to shape activity patterns.     

Seasonal Effects of Habitat on Sources and Rates of Snowshoe Hare Predation in Alaskan Boreal Forests

Survival and predation of snowshoe hares (Lepus americanus) has been widely studied, yet there has been little quantification of the changes in vulnerability of hares to specific predators that may result from seasonal changes in vegetation and cover. We investigated survival and causes of mortalities of snowshoe hares during the late increase, peak, and decline of a population in interior Alaska. From June 2008 to May 2012, we radio-tagged 288 adult and older juvenile hares in early successional and black spruce (Picea mariana) forests and, using known-fate methods in program MARK, evaluated 85 survival models that included variables for sex, age, and body condition of hares, as well as trapping site, month, season, year, snowfall, snow depth, and air temperature. We compared the models using Akaike’s information criterion with correction for small sample size. Model results indicated that month, capture site, and body condition were the most important variables in explaining survival rates. Survival was highest in July, and more generally during summer, when alternative prey was available to predators of hares. Low survival rates coincided with molting periods, breeding activity in the spring, and the introduction of juveniles to the sample population in the fall. We identified predation as the cause of mortality in 86% of hare deaths. When the source of predation could be determined, hares were killed more often by goshawks (Accipiter gentilis) than other predators in early successional forest (30%), and more often by lynx (Lynx canadensis) than other predators in black spruce forest (31%). Great horned owls (Bubo virginianus) and coyotes (Canis latrans) represented smaller proportions of hare predation, and non-predatory causes were a minor source (3%) of mortality. Because hares rely on vegetative cover for concealment from predators, we measured cover in predation sites and habitats that the hares occupied and concluded that habitat type had a greater influence on the sources of predation than the amount of cover in any given location within a habitat. Our observations illustrate the vulnerability of hares to predators in even the densest coniferous habitat available in the boreal forest, and indicate strong seasonal changes in the rates and sources of predation.     

Habitat Use by Weddell Seals and Emperor Penguins Foraging in the Ross Sea, Antarctica

The only apex predators that live year-round at high latitudesof the Ross Sea are the Weddell seal and emperor penguin. Theseasonal distribution, foraging depths, and diet of these twospecies appear to overlap. What makes it possible for emperorpenguins and Weddell seals to co-exist at high latitude throughoutthe winter when other marine tetrapods apparently cannot? Bothspecies have similar adaptations for exploitation of the deep-waterhabitat, forage on the same species, and routinely make longand deep dives. Yet, despite these similarities, there is probablylittle trophic overlap between the adults of both species dueto geographical and seasonal differences in habitat use. Forexample, during the winter months while female emperor penguinsare ranging widely in the pack ice, adult seals are foragingand fattening for the upcoming summer fast, literally beneaththe feet of the male penguins. However, there is more extensiveoverlap between juvenile seals and adult penguins, and shiftsin prey abundance and/or distribution would likely affect thesetwo groups similarly. In contrast, juvenile penguins appearto avoid inter- and intra- specific competition by leaving theRoss Sea once they molt.