Variation in foraging success among predators and its implications for population dynamics

The effects of the expected predation rate on population dynamics have been studied intensively, but little is known about the effects of predation rate variability (i.e., predator individuals having variable foraging success) on population dynamics. In this study, variation in foraging success among predators was quantified by observing the predation of the wolf spider Pardosa pseudoannulata on the cricket Gryllus bimaculatus in the laboratory. A population model was then developed, and the effect of foraging variability on predator–prey dynamics was examined by incorporating levels of variation comparable to those quantified in the experiment. The variability in the foraging success among spiders was greater than would be expected by chance (i.e., the random allocation of prey to predators). The foraging variation was density‐dependent; it became higher as the predator density increased. A population model that incorporates foraging variation shows that the variation influences population dynamics by affecting the numerical response of predators. In particular, the variation induces negative density‐dependent effects among predators and stabilizes predator–prey dynamics.     

Predator foraging success and habitat complexity: quantitative test of the threshold hypothesis

Summary Numerous studies have demonstrated a negative relationship between increasing habitat complexity and predator foraging success. Results from many of these studies suggest a non-linear relationship, and it has been hypothesised that some threshold level of complexity is required before foraging success is reduced significantly. We examined this hypothesis using largemouth bass (Micropterus salmoides) foraging on juvenile bluegill sunfish (Lepomis macrochirus) in various densities of artificial vegetation. Largemouth foraging success differed significantly among the densities of vegetation tested. Regression analysis revealed a non-linear relationship between increasing plant stem density and predator foraging success. Logistic analysis demonstrated a significant fit of our data to a logistic model, from which was calculated the threshold level of plant stem desity necessary to reduce predator foraging success. Studies with various prey species have shown selection by prey for more complex habitats as a refuge from predation. In this stydy, we also examined the effects of increasing habitat complexity (i.e. plant stem density) on choice of habitat by juvenile bluegills while avoiding predation. Plant stem density significantly effected choice of habitat as a refuge. The relationship between increasing habitat complexity and prey choice of habitat was found to be positive and non-linear. As with predator foraging success, logistic analysis demonstrated a significant fit of our data to a logistic model. Using this model we calculated the threshold level of habitat complexity required before prey select a habitat as a refuge. This density of vegetation proved to be considerably higher than that necessary to significantly reduce predator foraging success, indicating that bluegill select habitats safe from predation.Implications of these results and various factors which may affect the relationships described are discussed.     

Positive complexity-stability relations in food web models without foraging adaptation

May's [1972. Will a large complex system be stable? Nature 238, 413-414] local stability analysis of random food web models showed that increasing network complexity leads to decreasing stability, a result that is contradictory to earlier empirical findings. Since this seminal work, research of complexity-stability relations became one of the most challenging issues in theoretical ecology. We investigate conditions for positive complexity-stability relations in the niche, cascade, nested hierarchy, and random models by evaluating the network robustness, i.e., the fraction of surviving species after population dynamics. We find that positive relations between robustness and complexity can be obtained when resources are large, Holling II functional response is used and interaction strengths are weighted with the number of prey species, in order to take foraging efforts into account. In order to obtain these results, no foraging dynamics needs to be included. However, the niche model does not show positive complexity-stability relations under these conditions. By comparing to empirical food web data, we show that the niche model has unrealistic distributions of predator numbers. When this distribution is randomized, positive complexity-stability relations can be found also in the niche model.     

Food limitation of population density in the orb-wed spider,Nephila clavata

Field studies were conducted to clarify whether variation in food availability among habitats influences population density, and whether population density has a negative effect on foraging success in the orb-web spider, Nephila clavata. Lifetime food consumption per individual (i.e., foraging success) strongly correlated with mean body size of adult females and mean fecundity in populations. Also, there was a positive correlation between foraging success and population density. Since foraging success reflected potential prey availability in the habitat, food resource appeared to be a limiting factor for populations in this spider. Mean fecundity per individual correlated with population density of the following year, suggesting that decreased reproduction is a major component of food limitation on population density. Consistent defferences in mean body size between particular sites were observed over years, while such difference was less obvious in density. Thus, ranking of food abundance among habitats seems to be predictable between years. A field experiment revealed that an artificial increase in population density had no negative effect on the feeding rate of individuals, suggesting that intraspecific competition for food is not important in this species.     

Life-history plasticity and inbreeding depression under mate limitation and predation risk: cumulative lifetime fitness dissected with a life table response experiment

Environmental effects on the evolution of mating systems are increasingly discussed, but we lack many examples of how environmental conditions affect the expression and consequences of alternative mating systems. Variation in mate availability sets up a trade-off between reproductive assurance and inbreeding depression, but the consequences of both mate limitation and inbreeding may depend on other environmental conditions. Predation risk is common under natural conditions, and known to affect allocation to reproduction, but we know little about the effects of isolation and inbreeding under predation risk. We reared selfed and outcrossed hermaphroditic freshwater snails (Physa acuta) in four environments (predator cues present or absent crossed with mating partners available or not) and quantified life-history traits and cumulative lifetime fitness. Our results confirm that isolation from mates can increase longevity and growth, resulting in higher lifetime fecundity. Thus, we observed no evidence for mate limitation of reproduction. However, reproduction under isolation (i.e., selfing) resulted in inbreeding depression, which should counteract the benefits of selfing. Inbreeding depression in fitness occurred in both predator and no-predator environments, but there was no overall change in inbreeding depression with predator cues. This represents, to our knowledge, the first empirical estimate of the effect of predation risk on inbreeding depression in an animal. Cumulative fitness was most influenced by early survival and especially early fecundity. As predation risk and inbreeding (both ancestral and due to a lack of mates) reduced early fecundity, these effect are predicted to have important contributions to population growth under natural conditions. Therefore life-history plasticity (e.g., delayed reproduction) is likely to be very important to overall fitness.     

Response of a complex foraging phenotype to artificial selection on its component traits

In nature, where predators must often track dynamic and dispersed prey populations, predator consumption rate, conversion efficiency, dispersal, and prey finding are likely to be important links between foraging and predator–prey population dynamics. Small differences in predator foraging caused by variation in any of the abovementioned traits might lead to significant differences in predator success as well as population dynamics. We used artificial selection to create lines of the predatory mite, Phytoseiulus persimilis in order to determine the potential for or constraints on the evolution of predator foraging behaviors. All four foraging traits demonstrated considerable phenotypic variation. They also exhibited significant realized heritabilities after artificial selection, except that prey finding did not respond to downward selection. Lines that responded to selection did so rapidly, and high-consumption, high-conversion efficiency, and high- and low-dispersal were stable for at least four generations after artificial selection was relaxed. There were some indirect responses to selection among the foraging traits. For example, there was positive correlation between consumption and dispersal. However, none of the correlated responses were of the magnitude of the direct responses we measured on the same trait. We also observed some correlations between foraging traits and life-history traits such as low-consumption and development time (negative), high-consumption and fecundity (positive), and high-conversion efficiency and fecundity (positive), but these were more likely to represent non-genetic constraints. Intrinsic rates of increase in low-consumption and low-conversion efficiency lines were lower than in their respective high lines and the unselected control, whereas rates of increase in dispersal and olfactory response lines did not differ from the unselected control. Thus, traits that make up foraging share partially overlapping genetic architectures with highly heritable phenotypic components, suggesting that each foraging trait will be able to respond rapidly to changes in the density and distribution of resources.     

Frequency-dependent predation, crypsis and aposematic coloration

Frequency-dependent predation may maintain or prevent colour pattern polymorphisms in prey, and can be caused by a variety of biological phenomena, including perceptual processes (search images), optimal foraging and learning. Most species are preyed upon by more than one predator species, which are likely to differ in foraging styles, perceptual and learning abilities. Depending upon the interaction between predator vision, background and colour pattern parameters, certain morphs may be actively maintained in some conditions and not in others, even with the same predators. More than one kind of predator will also affect stability, and only slight changes in conditions can cause a transition between polymorphism and monomorphism. Frequency-dependent selection is not a panacea for the explanation of variation in animal colour patterns, although it may be important in some systems.     

Do seasonal patterns of rat snake (Pantherophis obsoletus) and black racer (Coluber constrictor) activity predict avian nest predation?

Avian nest success often varies seasonally and because predation is the primary cause of nest failure, seasonal variation in predator activity has been hypothesized to explain seasonal variation in nest success. Despite the fact that nest predator communities are often diverse, recent evidence from studies of snakes that are nest predators has lent some support to the link between snake activity and nest predation. However, the strength of the relationship has varied among studies. Explaining this variation is difficult, because none of these studies directly identified nest predators, the link between predator activity and nest survival was inferred. To address this knowledge gap, we examined seasonal variation in daily survival rates of 463 bird nests (of 17 bird species) and used cameras to document predator identity at 137 nests. We simultaneously quantified seasonal activity patterns of two local snake species (N = 30 individuals) using manual (2136 snake locations) and automated (89,165 movements detected) radiotelemetry. Rat snakes (Pantherophis obsoletus), the dominant snake predator at the site (~28% of observed nest predations), were most active in late May and early June, a pattern reported elsewhere for this species. When analyzing all monitored nests, we found no link between nest predation and seasonal activity of rat snakes. When analyzing only nests with known predator identities (filmed nests), however, we found that rat snakes were more likely to prey on nests during periods when they were moving the greatest distances. Similarly, analyses of all monitored nests indicated that nest survival was not linked to racer activity patterns, but racer‐specific predation (N = 17 nests) of filmed nests was higher when racers were moving the greatest distances. Our results suggest that the activity of predators may be associated with higher predation rates by those predators, but that those effects can be difficult to detect when nest predator communities are diverse and predator identities are not known. Additionally, our results suggest that hand‐tracking of snakes provides a reliable indicator of predator activity that may be more indicative of foraging behavior than movement frequency provided by automated telemetry systems.     

Effects of age,sex, and ENSO phase on foraging and flight performance in Nazca boobies

Age‐related changes in survival and reproduction are common in seabirds; however, the underlying causes remain elusive. A lack of experience for young individuals, and a decline in foraging performance for old birds, could underlie age‐related variation in reproduction because reproductive success is connected closely to provisioning offspring. For seabirds, flapping flight during foraging trips is physiologically costly; inexperience or senescent decline in performance of this demanding activity might cap delivery of food to the nest, providing a proximate explanation for poor breeding success in young and old age, respectively. We evaluated the hypothesis that young and old Nazca boobies (Sula granti), a Galápagos seabird, demonstrate deficits in foraging outcomes and flight performance. We tagged incubating male and female adults across the life span with both accelerometer and GPS loggers during the incubation periods of two breeding seasons (years), during the 2015 El Niño and the following weak La Niña. We tested the ability of age, sex, and environment to explain variation in foraging outcomes (e.g., mass gained) and flight variables (e.g., wingbeat frequency). Consistent with senescence, old birds gained less mass while foraging than middle‐aged individuals, a marginal effect, and achieved a slower airspeed late in a foraging trip. Contrary to expectations, young birds showed no deficit in foraging outcomes or flight performance, except for airspeed (contingent on environment). Young birds flew slower than middle‐aged birds in 2015, but faster than middle‐aged birds in 2016. Wingbeat frequency, flap–glide ratio, and body displacement (approximating wingbeat strength) failed to predict airspeed and were unaffected by age. Sex influenced nearly all aspects of performance. Environment affected flight performance and foraging outcomes. Boobies'' foraging outcomes were better during the extreme 2015 El Niño than during the 2016 weak La Niña, a surprising result given the negative effects tropical seabirds often experience during extreme El Niños.     

Relation between complexity and stability in food webs with adaptive behavior

We investigate the influence of functional responses (Lotka-Volterra or Holling type), initial topological web structure (randomly connected or niche model), adaptive behavior (adaptive foraging and predator avoidance) and the type of constraints on the adaptive behavior (linear or nonlinear) on the stability and structure of food webs. Two kinds of stability are considered: one is the network robustness (i.e., the proportion of species surviving after population dynamics) and the other is the species deletion stability. When evaluating the network structure, we consider link density as well as the trophic level structure. We show that the types of functional responses and initial web structure do not have a large effect on the stability of food webs, but foraging behavior has a large stabilizing effect. It leads to a positive complexity-stability relationship whenever higher "complexity" implies more potential prey per species. The other type of adaptive behavior, predator avoidance behavior, makes food webs only slightly more stable. The observed link density after population dynamics depends strongly on the presence or absence of adaptive foraging, and on the type of constraints used. We also show that the trophic level structure is preserved under population dynamics with adaptive foraging.     

Flexible nest-site selection under anthropogenic habitat change in an Afrotropical understorey insectivore

Human activities impact upon natural habitats used by birds for breeding and foraging, and lead to changes in the composition and spatial distribution of predator communities, mainly through loss, fragmentation and disturbance of formerly pristine habitat. Yet possible fitness consequences of such changes through impacts on bird nest-site selection remain poorly known. Here we study nest-site selection and reproductive success of Placid Greenbuls Phyllastrephus placidus in the Taita Hills, southeast Kenya. We show that habitat features associated with nest-site selection by this insectivorous, open-cup-nesting bird species vary among forest fragments that are exposed to different levels of habitat disturbance. Such differences in sites selected for breeding result from a plastic response to fragment-specific conditions or may be driven by fragment-specific variation in the distribution and availability of certain habitat features. Given the overall high nest predation rates in our study area, we expected variation in nest-site selection to correlate with reproductive success and nestling condition, but detected no such relationship. Because predator density and nest predation rates may vary strongly in space and time, a better understanding of spatio-temporal variation in predator communities is needed to assess the possible adaptive value of nest-site selection strategies for reducing the high predation rates that are typical for this and many other open-cup-nesting tropical passerines.     

Masting in Buxus balearica: assessing fruiting patterns and processes at a large spatial scale

Masting consists of the synchronous highly variable seed production among years by a plant population. We studied spatiotemporal variation in fruit production in ten populations of Buxus balearica (six in the Balearic Islands and four in the Iberian Peninsula) from 2001 to 2004 in the light of masting. In some of them we assessed, by means of both observational and experimental data, the relationship between fruit production and some abiotic variables, the role of previous reproduction, the "pollination efficiency" and the "predator satiation" hypotheses, as well as the consequences for seedling density and survival. Fruit production in B. balearica showed substantial between-year variation, especially in island compared to mainland populations. Correlative evidence indicated that this variation and its geographic pattern were related to differences between regions in rainfall variability, cost of reproduction and the degree of ambophily. We found no indication of predator satiation. However, experimental tests failed to support many of our results, namely a negative effect of previous reproduction on future flowering in island populations and lower pollen limitation with increasing flower production. We therefore warn against exclusive reliance on correlations when testing hypotheses related to masting. In addition, seedling recruitment increased after some episodes of high fruit production but probably additional factors had a role in recruitment, suggesting that mast events not always translate into increased reproductive success. Although a limited time series only allows considering B. balearica as showing 'putative' masting, weather and pollination-related processes are good candidates for further exploration of fruiting patterns and processes at a large spatial scale.     

Seasonal and temporal variations in colony-level foraging activity of a queenless ant, <Emphasis Type="Italic">Diacamma</Emphasis> sp., in Japan

We investigated colony-level foraging activities of Diacamma sp., a queenless ponerine ant, in the field. Our aim was to elucidate the presence of any pattern in foraging activity in field colonies in relation to: (1) circadian rhythm, (2) physical environmental conditions such as extreme temperatures, (3) seasonality, and (4) short-term foraging efficiency (i.e. the success ratio in obtaining food per foraging trip). Colony-level foraging activity tended to be diurnal throughout the year, as more foraging trips were observed in the daytime. Although temperature had no linear effect on overall foraging activity, lower temperature precluded foraging at night. Overall, foraging was more frequent at times of day when foraging efficiency was high, but this relationship was weak and varied seasonally. Interestingly, we found that hourly foraging efficiency and hourly foraging activity were negatively correlated in autumn, the season when the average foraging efficiency peaked, whereas they were positively correlated in winter and spring.     

Foraging to balance conflicting demands: novel insights from grasshoppers under predation risk

Animal foraging may be influenced by multiple demands simultaneously(e g., nutrient gain and predator avoidance). Conventional approachesto understand the trade-offs between these demands require crammingthem in similar currencies, which is impractical in many fieldsituations. We introduce a new method, called multiobjectiveprogramming, as a framework to explore how animals balance conflictingdemands. Multiobjective programming allows one to explore theinfluence of foraging demands directly, without explicit assumptionsabout how they enter into fitness and without conversion tosome common currency. Using multiobjective programming, we showthat, as foraging demands change, animals may adaptively adjusttheir behavior, even if the constraints on feasible behaviorare unaffected (contrary to the predictions of the conventionalmodels). Hence, we may see a variable response in foraging thatis consistent with adaptive behavior. We used an empirical testwith herbivore grasshoppers and predator spiders to evaluatethe utility of multiobjective programming Our experiments showthat grasshoppers are able to optimally balance the foragingobjectives of energy intake and vigilance under changing levelsof predation risk. The multiobjective model is used both toevaluate the biological significance of the broad variationthat was observed in the grasshoppers' foraging behavior andto quantify explicitly the trade-off between energy intake andpredator avoidance.     

Should "handled" prey be considered? Some consequences for functional response, predator-prey dynamics and optimal foraging theory

Predator-prey models consider those prey that are free. They assume that once a prey is captured by a predator it leaves the system. A question arises whether in predator-prey population models the variable describing prey population shall consider only those prey which are free, or both free and handled prey together. In the latter case prey leave the system after they have been handled. The classical Holling type II functional response was derived with respect to free prey. In this article we derive a functional response with respect to prey density which considers also handled prey. This functional response depends on predator density, i.e., it accounts naturally for interference. We study consequences of this functional response for stability of a simple predator-prey model and for optimal foraging theory. We show that, qualitatively, the population dynamics are similar regardless of whether we consider only free or free and handled prey. However, the latter case may change predictions in some other cases. We document this for optimal foraging theory where the functional response which considers both free and handled prey leads to partial preferences which are not observed when only free prey are considered.     

An experimental design and a statistical analysis separating interference from exploitation competition

Previous experimental studies of competition among foragers rarely distinguished between exploitation and interference competition. In many systems this separation is experimentally impossible without interfering with the natural behavior of the animals. Consequently, these studies can only demonstrate the combined effect of interference and exploitation on the forager’s feeding rate, namely, it usually decreases in a decelerating rate as a function of density. We suggest here a simple experimental and statistical procedure that facilitates the separation of the effects of interference from those of exploitation. This procedure includes manipulation of both predator density and the foraging experiment duration. The statistical analysis is based on multiple linear regression. The working assumption is that exploitation can be neglected at the beginning of the foraging experiment because, initially, predators do not experience diminishing returns in prey capture rates. Using both the results of an individual-based simulation and a field experiment dataset of gerbils foraging for seeds in an artificial food patch located in the field, we demonstrate that our procedure can successfully detect and separate the effect of interference from the combined overall effect of competition (i.e., interference plus exploitation). Inon Scharf and Ido Filin contributed equally to this paper.     

Perspectives on the ecomorphology of bony fishes

Synopsis The field of ecomorphology has a long history with early roots in Europe. In this half of the century the application of ecomorphology to the biology of fishes has developed in the former Soviet Union, Poland and Czechoslovakia, The Netherlands, and in North America. While the specific approaches vary among countries, many North American studies begin by comparing morphological variation with variation in ecological characteristics at the intra or interspecific levels. These initial correlative studies form the ground work for hypotheses that explore the mechanistic underpinnings of the observed ecomorphological associations. Supporting these mechanistic hypotheses are insights from functional studies which demonstrate the limits to potential resource use resulting from a particular morphology; however, the actual resource use is likely to be more limited due to additional constraints provided by internal (e.g., behavior, physiology) and external (e.g., resource abundance, predator distribution) factors. The results from performance studies in the laboratory or field can be used to test specific ecomorphological hypotheses developed from the initial correlational and functional studies. Such studies may, but rarely do, incorporate an ontogenetic analysis of the ecomorphological association to determine their effect on performance. Finally, input from phylogenetic analyses allow an investigator to examine the evolution of specific features and to assess the rates and directionality of character evolution. The structural and ecological diversity of fishes provides a fertile ground to investigate these interactions. The contributions in this volume highlight some of the specific directions for ecomorphological research covering a variety of biological processes in fishes. These include foraging, locomotion, reproduction, respiration, and sensory systems. Running through these papers are new insights into universal ecomorphological issues, i.e., the relationships between form and ecological role and the factors that modify these relationships.     

Individual variation in prey choice in a predator-prey community

One predator-two prey community models are studied with an emphasis on individual variation in predator behavior. The predator behaves according to a well-known prey choice model. The behavioral model predicts that predators should always attack the primary prey (more profitable prey of the two), but only attack the alternative prey (less profitable prey of the two) when the density of the primary prey is below a threshold density. The predator that accepts the alternative prey does not discriminate between the primary and alternative prey (all-or-nothing preference for the alternative prey). However, empirical studies do not result in clear all-or-nothing responses. Previous models examined the relaxation of the all-or-nothing response by assuming partial preference (e.g., predators preferentially forage on the primary prey even when they also attack the alternative prey). In this study, I consider individual variation in two predator traits (prey density perception and handling time) as the sources of the variation in the threshold density, which can make empirical data appear deviated from the expectation. I examine how community models with partial preference and individual variation differ in their dynamics and show that the differences can be substantial. For example, the dynamics of a model based on individual variation can be more stable (e.g., stable in a wider parameter region) than that of a model based on partial preference. As the general statistical property (Jensen’s inequality) is a main factor that causes the differences, the results of the study have general implications to the interpretation of models based on average per-capita rates.     

Detecting predator–prey relationships in the sea

Understanding the strength and diversity of predator‐prey interactions among species is essential to understand ecosystem consequences of population‐level variation. Directly quantifying the predatory behaviour of wild fishes at large spatial scales (>100 m) in the open sea is fraught with difficulties. To date the only empirical approach has been to search for correlations in the abundance of predators and their putative prey. As an example we use this approach to search for predators of the keystone crown‐of‐thorns starfish. We show that this approach is unlikely to detect predator–prey linkages because the theoretical relationship is non‐linear, resulting in multiple possible prey responses for single given predator abundance. Instead we suggest some indication of the strength and ecosystem importance of a predator–prey relationship can be gained by using the abundance of both predators and their putative prey to parameterize functional response models.     

Seasonal Trophic Niche Shift and Cascading Effect of a Generalist Predator Fish

Few studies have examined how foraging niche shift of a predator over time cascade down to local prey communities. Here we examine patterns of temporal foraging niche shifts of a generalist predator (yellow catfish, Pelteobagrus fulvidraco) and the abundance of prey communities in a subtropical lake. We predicted that the nature of these interactions would have implications for patterns in diet shifts and growth of the predator. Our results show significant decreases in planktivory and benthivory from late spring to summer and autumn, whereas piscivory increased significantly from mid-summer until late autumn and also increased steadily with predator body length. The temporal dynamics in predator/prey ratios indicate that the predation pressure on zooplankton and zoobenthos decreased when the predation pressure on the prey fish and shrimps was high. Yellow catfish adjusted their foraging strategies to temporal changes in food availability, which is in agreement with optimal foraging theory. Meanwhile the decrease in planktivory and benthivory of yellow catfish enabled primary consumers, such as zooplankton and benthic invertebrates, to develop under low grazing pressure via trophic cascading effects in the local food web. Thus, yellow catfish shifts its foraging niche to intermediate consumers in the food web to benefit the energetic demand on growth and reproduction during summer, which in turn indirectly facilitate the primary consumers. In complex food webs, trophic interactions are usually expected to reduce the strength and penetrance of trophic cascades. However, our study demonstrates strong associations between foraging niche of piscivorous fish and abundance of prey. This relationship appeared to be an important factor in producing top-down effects on both benthic and planktonic food webs.