The effect of microhabitat feeding site selection on aphid foraging and predation risk

Organisms attempt to optimize foraging by maximizing resource acquisition while minimizing predation risk. Aphids (group-living, phloem-feeding insects) routinely change feeding positions and interact with predators and parasites at the single-leaf scale. Here, we assess the life history and predation risk consequences of within-leaf feeding site choices in pea aphids in response to different natural enemies. First, three-chambered clip cages were used to isolate first instar aphids anterior and posterior to a centrally feeding adult on the underside of a single broad bean leaf. Development time to adulthood did not differ between feeding sites, nor did fecundity within the first 24 h of reproduction. Second, we recorded the frequency and latency of natural enemy attacks on aphids adhered to three leaf sites, matching those of the clip cage experiment, on the underside of a single leaf. Aphids feeding nearest the leaf petiole were at greatest risk of predation by a foliar foraging coccinellid predator, Hippodamia convergens, but not by a parasitoid wasp, Aphidius ervi. Thus, feeding nearer the leaf petiole provided no individual life history benefits and exposes the aphid to increased predation risk. We further discuss the notion that feeding at these sites may provide inclusive fitness benefits for colony mates via alarm signaling and subsequent decreased predation success.     

The scale-dependent impact of wolf predation risk on resource selection by three sympatric ungulates

Resource selection is a fundamental ecological process impacting population dynamics and ecosystem structure. Understanding which factors drive selection is vital for effective species- and landscape-level management. We used resource selection probability functions (RSPFs) to study the influence of two forms of wolf (Canis lupus) predation risk, snow conditions and habitat variables on white-tailed deer (Odocoileus virginianus), elk (Cervus elaphus) and moose (Alces alces) resource selection in central Ontario's mixed forest French River-Burwash ecosystem. Direct predation risk was defined as the frequency of a predator's occurrence across the landscape and indirect predation risk as landscape features associated with a higher risk of predation. Models were developed for two winters, each at two spatial scales, using a combination of GIS-derived and ground-measured data. Ungulate presence was determined from snow track transects in 64 16- and 128 1-km(2) resource units, and direct predation risk from GPS radio collar locations of four adjacent wolf packs. Ungulates did not select resources based on the avoidance of areas of direct predation risk at any scale, and instead exhibited selection patterns that tradeoff predation risk minimization with forage and/or mobility requirements. Elk did not avoid indirect predation risk, while both deer and moose exhibited inconsistent responses to this risk. Direct predation risk was more important to models than indirect predation risk but overall, abiotic topographical factors were most influential. These results indicate that wolf predation risk does not limit ungulate habitat use at the scales investigated and that responses to spatial sources of predation risk are complex, incorporating a variety of anti-predator behaviours. Moose resource selection was influenced less by snow conditions than cover type, particularly selection for dense forest, whereas deer showed the opposite pattern. Temporal and spatial scale influenced resource selection by all ungulate species, underlining the importance of incorporating scale into resource selection studies.     

Ontogenetic shifts in competitive interactions and intra-guild predation between two wolf spider species

Abstract. 1. The wolf spiders (Araneae: Lycosidae) Hogna helluo (Walckenaer) and Pardosa milvina Hentz co-occur in soybean fields of south-west Ohio, U.S.A. As adults, Hogna is the larger species and has the competitive advantage in most interactions; due to differing phenologies, however, their size-classes frequently overlap and as such there is potential for shifts in competitive ability and intra-guild predation. The hypothesis that competitive interactions and intra-guild predation will favour Pardosa when Pardosa is similar-sized, or has a size advantage over Hogna , was tested in laboratory and field experiments.
2. Studies in laboratory arenas, pairing similar-sized individuals of these species and Hogna spiderlings with larger spiders of both species, revealed that intra-guild predation seldom occurs with similar-sized Hogna and Pardosa , however Pardosa will consume small Hogna individuals in laboratory arenas.
3. Field experiments involved stocking high densities (50 m^{– 2} ) of Pardosa and Hogna in enclosures placed in soybean fields. In experiments with spiders of similar size, no interspecific effects were uncovered, but an intraspecific effect was found for Pardosa as its survival and weight gain were lower in the presence of more conspecifics. Large Hogna or Pardosa had no effect on the survival or weight gain of Hogna spiderlings.
4. Although Hogna is a better competitor as an adult, it has no advantage over Pardosa when their size-classes overlap, and Pardosa effects on Hogna may be inconsequential under field conditions. Therefore, the co-existence of these species is fostered by the fact that there are few negative interspecific interactions during their ontogeny.     

Effect of adult females' predation risk on oviposition site selection in a water strider

In many egg‐laying species, females avoid ovipositing at sites where the predation risk is high. Previous studies have mainly focused on the risk for offspring. The effect of predation risk for the females has been considered in some taxa in which parents spend much time at an oviposition site for parental care or mating (e.g., birds, amphibians). In species in which females do not perform activities other than oviposition at sites, the effect of predation risk for females on oviposition site selection has been rarely investigated. We examined whether the predation risk for ovipositing females affects the decision on oviposition in the water strider Aquarius paludum insularis (Motschulsky) (Heteroptera: Gerridae). Adult A. paludum are preyed upon by a backswimmer, Notonecta triguttata (Motschulsky) (Heteroptera: Notonectidae), but N. triguttata does not prey upon eggs or early instars of A. paludum. We allowed female A. paludum to oviposit under one of three conditions: in presence of N. triguttata, in presence of its chemicals, and in absence of the predator or its cues (control). Female A. paludum less frequently oviposited in presence of N. triguttata than in its absence. Oviposition frequency did not differ between females in presence of chemicals of N. triguttata vs. those in the control. Female A. paludum recognized the predation risk upon themselves from the presence of N. triguttata and avoided ovipositing. This study is the first to directly show that the predation risk upon ovipositing females changes oviposition site selection in species in which the time spent at an oviposition site is short.     

The benefits of stealing from a predator: foraging rates, predation risk, and intraspecific aggression in the kleptoparasitic spider Argyrodes antipodiana

In the trade-off between food and safety, the role of aggressiveintraspecific interactions has not been extensively examined.Here I present information on this system using a kleptoparasiticspider, Argyrodts antipodiana, and its host spider and potentialpredator, Eriophora pustulosa. A. antipodiana can feed eitherat a potentially dangerous site (the hub of its host's web withthe host), or at a relatively safe site (on food bundles aroundthe edge of the host's web). I found that A. antipodiana cangain food very quickly when feeding with the host, apparentlyby exploiting the host's ability to digest the prey. Thus A.antipodiana follows predictions based on foraging models inthat it accepts a higher predation risk at the hub because ofthe higher food payoff. A. antipodiana also aggressively competesfor access to more food. However, aggressive competition increasesthe predation risk from the host, especially at tile hub wherethe host is very close. Consequently, A. antipodiana modifiesits level of intraspecific aggressiveness in accordance withits position on the web: at the hub, where the cost of aggressionis high (due to predation risk), A. antipodiana reduces itsaggressiveness, but it is aggressive away from the hub whencompeting for food bundles. The ability of A. antipodiana tochange interaction intensity as a function of its position onthe web enables it to exploit a rich, but risky, food sourceand provides a new angle for examining food and safety tradeoffs in light of intraspecific competition for food     

The effects of predation risk from crab spiders on bee foraging behavior

Recent studies have suggested that top–down effects ofpredation on plant–pollinator interactions may not be,as previously thought, rare and/or weak. In this paper, we explorethe effects of crab spiders (Araneae: Thomisidae) on the behaviorof 2 species of bee (Hymenoptera: Apidae) foraging for nectarand pollen on 3 different plant species in central Portugal.In 2 experiments, we found that the eusocial bee Apis melliferawas significantly less likely to inspect and accept a floweror inflorescence if it harbored a spider. In contrast, we foundno such effects of spiders on the behavior of the solitary beeEucera notata. Further experiments showed that the effects ofenvironmental cues associated with predators on flower visitationby A. mellifera were detectable even when no spider was presentat the moment a flower was encountered. Such indirect effectswere only identified, however, in bees foraging on 1 of 2 plantspecies studied. In a final experiment, A. mellifera was shownto respond negatively to the presence of the corpses of conspecificsglued to flowers. This suggests that prey corpses left exposedon petals or bracts by spiders provide an obvious cue that beescan use to avoid predators. These results add to a growing bodyof evidence that plant–pollinator interactions are notimmune to the effects of predation and suggest that the strengthof such effects vary both between and within species.     

The effects of predation risk on the use of social foraging tactics

The effects of predation on the use of social foraging tactics, such as producing and scrounging, are poorly known in animals. On the one hand, recent theoretical models predict increased use of scrounging with increasing predation risk, when scroungers seeking feeding opportunities also have a higher chance of detecting predators. On the other hand, there may be no relation between tactic use and predation when antipredator vigilance is not compatible with scanning flockmates. We investigated experimentally the effects of predation risk on social foraging tactic use in tree sparrows, Passer montanus. We manipulated predation risk in the field by changing the distance between shelter and a feeder. Birds visited the feeder in smaller flocks, spent less time on it and were somewhat more vigilant far from shelter than close to it. Increased predation risk strongly affected the social foraging tactic used: birds used the scrounger tactic 30% more often far from cover than close to it. Between-flock variability in scrounging frequency was not related to the average vigilance level of the flock members, and within-flock variability in the use of scrounging was negatively related to the vigilance of birds. Our results suggest that in tree sparrows, the increased frequency of scrounging during high predation risk cannot simply be explained by an additional advantage of increasing antipredator vigilance. We propose alternative mechanisms (e.g. increased stochasticity in food supply, and that riskier places are used by individuals with lower reserves) that may explain increased scrounging when animals forage under high predation risk.     

A test of the hypothesis that a pathway of intraguild predation limits densities of a wolf spider

Abstract.  1. The high number of potential predatory interactions among the many arthropod generalist predators in terrestrial food webs makes exhaustive testing of interaction strengths by field experiments unfeasible. Thus, correlative patterns and laboratory observations of behaviour often form the basis of inferences about the strength of interaction pathways involving generalist predators (intraguild predation).
2. Previous research has revealed a negative correlation between survival of juvenile wolf spiders of the genus Schizocosa (Lycosidae) and densities of another abundant spider family, the Gnaphosidae.
3. Feeding trials in laboratory microcosms with a leaf-litter substrate revealed that gnaphosids prey on juvenile Schizocosa in a structurally complex habitat.
4. Gnaphosid densities were manipulated in two different field experiments, each conducted in a different year, in order to test directly the hypothesis that intraguild predation by gnaphosids limits densities of juvenile Schizocosa .
5. Reducing numbers of gnaphosids, and doubling their numbers to two times the mean natural density, had no impact on the survival of juvenile Schizocosa in either field experiment. This finding suggests that correlative patterns in nature and feeding trials in the laboratory may at times provide deceptively simple and potentially misleading generalisations about the strengths of interaction pathways in complex networks of generalist predators.     

State-dependent decisions in nest site selection by a web-building spider

Foraging decisions may reflect a trade-off between food intake and safety and can also be influenced by the animal's internal state. Foraging in the web-building spider Stegodyphus lineatus depends on a capture web associated with a retreat (the nest). The relocation of nests, including the take-over of conspecific nests, may be viewed as a foraing decision, which depends on risk of exposure, cost of silk production and hunger state. We investigated a possible state-dependent trade-off in nest site choice of S. lineatus spiderlings. The philopatric nature ofS. lineatus implies a high risk of encounters with potentially cannibalistic conspecifics and a potential loss of inclusive fitness because encountered conspecifics are likely to be kin. To test for state dependence of foraging decisions, we compared preferences of well-fed and hungry spiders for their own nests and those of siblings and nonsiblings. We expected satiated spiders to prefer their own to conspecific nests and hungry spiders to choose the risky option of a conspecific nest. SinceS. lineatus is less aggressive towards kin, we tested the ability of spiders to discriminate kin by silk-bound cues. Because of this reduced aggression, preference for kin nests should be safer than preference for nonkin nests. A strong preference for self-nests demonstrated self-recognition in well-fed spiders. However, neither well-fed nor hungry spiders discriminated between nests of siblings and nonsiblings. Well-fed spiders preferred self-nests to empty chambers, but showed no discrimination between nonself-nests and empty chambers. Hungry spiders showed a reduced preference for self-nests, suggesting that hunger elicits a more risky foraging strategy. A tendency of hungry spiders to adopt the nests of conspecific spiders may reflect a silk-saving strategy. We conclude that S. lineatus spiders show state-dependent decision making in nest site selection. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

How avian nest site selection responds to predation risk: testing an 'adaptive peak hypothesis'

1. Nest predation limits avian fitness, so birds should favour nest sites that minimize predation risk. Nevertheless, preferred nest microhabitat features are often uncorrelated with apparent variation in predation rates. 2. This lack of congruence between theory-based expectation and empirical data may arise when birds already occupy 'adaptive peaks'. If birds nest exclusively in low-predation microhabitats, microhabitat and nest predation may no longer be correlated even though predation ultimately shaped microhabitat selection. 3. This 'adaptive peak hypothesis' was tested for a population of Yellow Warblers (Dendroica petechia) focusing on two nest microhabitat features: concealment and height. Experimental nests measured relative predation risk both within and outside the microhabitat range typically occupied by natural nests to examine whether nest site choices made by birds restricted our ability to detect microhabitat effects on predation. 4. Within the natural range (30-80% concealment, >75 cm height), microhabitat-predation relationships were weak and inconsistent, and similar for experimental and natural nests. Over an extended range, however, experimental predation rates were elevated in exposed sites (<30% concealed), indicating a concealment-related 'adaptive plateau'. 5. Clay egg bite data revealed a concealment effect on avian predators, and the abundance of one avian predator group correlated with nest concealment among years, suggesting these predators may cue birds to modulate nest concealment choices. 6. This study demonstrates how avian responses to predation pressure can obscure the adaptive significance of nest site selection, so predation influences may be more important than apparent from published data.     

Temporal variation in site fidelity: scale-dependent effects of forage abundance and predation risk in a non-migratory large herbivore

Large herbivores are typically confronted by considerable spatial and temporal variation in forage abundance and predation risk. Although animals can employ a range of behaviours to balance these limiting factors, scale-dependent movement patterns are expected to be an effective strategy to reduce predation risk and optimise foraging opportunities. We tested this prediction by quantifying site fidelity of global positioning system-collared, non-migratory female elk (Cervus canadensis manitobensis) across multiple nested temporal scales using a long-established elk–wolf (Canis lupus) system in Manitoba, Canada. Using a hierarchical analytical approach, we determined the combined effect of forage abundance and predation risk on variation in site fidelity within four seasons across four nested temporal scales: monthly, biweekly, weekly, daily. Site fidelity of female elk was positively related to forage-rich habitat across all seasons and most temporal scales. At the biweekly, weekly and daily scales, elk became increasingly attached to low forage habitat when risk was high (e.g. when wolves were close or pack sizes were large), which supports the notion that predator-avoidance movements lead to a trade-off between energetic requirements and safety. Unexpectedly, predation risk at the monthly scale increased fidelity, which may indicate that elk use multiple behavioural responses (e.g. movement, vigilance, and aggregation) simultaneously to dilute predation risk, especially at longer temporal scales. Our study clearly shows that forage abundance and predation risk are important scale-dependent determinants of variation in site fidelity of non-migratory female elk and that their combined effect is most apparent at short temporal scales. Insight into the scale-dependent behavioural responses of ungulate populations to limiting factors such as predation risk and forage variability is essential to infer the fitness costs incurred.     

On the economics of sit-and-wait foraging: site selection and assessment

We present two models of optimal resource exploitation for sit-and-waitforagers. The first model assumes immediate recognition of sitequality and that site quality does not change over time. Thismodel predicts a forager's minimum acceptable site quality.We present a graphical analysis to show how (1) the distributionof site qualities, (2) the travel time between sites, (3) costof search, and (4) expected duration of the foraging processinfluence the minimum acceptable rate. Our second model allowssite qualities to change and relaxes the assumption of immediaterecognition. This model defines conditions of (1) state duration,(2) recognition time, (3) site abundance, and (4) cost of searchwhere the optimal policy is to stay put in a site regardlessof experience. We discuss the implications of these models forthe design and interpretation of field experiments of site useand habitat selection.     

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.     

Acquired food aversion of a wolf spider to three cereal aphids: Intra- and interspecific effects

Naive spiders usually accept aphids though these are low quality food, but they subsequently develop aversions as a result of experience. An aversion induced by one species of cereal aphid will be effective with the same strength also against two other species, no matter which of the three species induces the aversions; thus, it plays no role which species the spider encounters. However, the strength of the aversions is significantly dependent on which species induces them. With the three species of aphids tested, it turned out that the aphid of lowest quality as food for the spider created the weakest aversions, while the relatively higher quality aphids produced stronger aversions. Hypothetical explanations for these findings are discussed.     

The effects of prey lipid on female mating and reproduction of a wolf spider

As predators, the macronutrients spiders extract from their prey play important roles in their mating and reproduction. Previous studies of macronutrients on spider mating and reproduction focus on protein, the potential impact of prey lipid content on spider mating and reproduction remains largely unexplored. Here, we tested the influence of prey varying in lipid content on female mating, sexual cannibalism, reproduction, and offspring fitness in the wolf spider Pardosa pseudoannulata. We acquired 2 groups of fruit fly Drosophila melanogaster that differed significantly in lipid but not protein content by supplementing cultural media with a high or low dose of sucrose on which the fruit flies were reared (HL: high lipid and LL: low lipid). Subadult (i.e., 1 molt before adult) female spiders that fed HL flies matured with significantly higher lipid content than those fed LL flies. We found that the mated females fed with HL flies significantly shortened pre-oviposition time and resulted in a significantly higher fecundity. However, there was no significant difference in female spiders varying in lipid content on other behaviors and traits, including the latency to courtship, courtship duration, mating, copulation duration, sexual cannibalism, offspring body size, and survival. Hence, our results suggest that the lipid content of prey may be a limiting factor for female reproduction, but not for other behavioral traits in the wolf spiders P. pseudoannulata.     

Habitat selection in a large orb‐weaving spider: vegetational complexity determines site selection and distribution

1. The distribution of the large orb‐weaving spider Argiope trifasciata in old field habitats of North America and the habitat selection process this species used was studied for 2 years. 2. Because web spiders have limited dispersal abilities and an energetically costly prey capture device, they do not have the ability to sample potential foraging sites. Structural complexity of the vegetation to which the web must be attached is relatively easy to assess. The hypothesis that the structural complexity is a primary factor in determining initial web site selection was tested both by relating the natural distribution of the spiders across habitats to vegetational complexity and by manipulating the complexity of the habitats in a series of experiments. 3. Argiope trifasciata was not distributed evenly among three old field vegetation types. Habitat complexity was related to spider density in both years although no measure of insect activity, prey capture, or prey consumption was correlated with spider distribution. 4. Three experimental manipulations were conducted to test the impact of habitat structure on spider establishment: (1) the amount of natural vegetation was reduced, (2) structures were added to a simple habitat, and (3) the complexity of the structures added was varied. In each case, spiders were introduced and establishment of webs was monitored. In all manipulations, spider establishment was related to the complexity of the substrate available. 5. These results are important for understanding the cues that influence foraging site selection and therefore provide insight into the distribution of species with limited dispersal abilities and high site investment requirements.     

Risk allocation and competition in foraging groups: reversed effects of competition if group size varies under risk of predation

Animals often feed more quickly when in larger groups. This group-size effect is often explained by safety advantages for groups but an alternative explanation is that animals feed faster in larger groups because of greater scramble competition for limited food. We show that predation risk enhances the group-size effect if groups vary in size. By contrast, competition leads to the group-size effect only when individuals feed in groups of constant size. When individuals feed in groups that vary in size, the best strategy for dealing with competition is to feed intensely when in smaller groups and feed little when in larger (more competitive) groups. In all situations, the effects of competition interact with the effects of predation risk in a simple multiplicative way. Our results suggest that scramble competition is not a general explanation for the group-size effect on vigilance in situations where group size changes relatively rapidly.