Allocation of searching time within the soybean canopy by parasitoids attacking the green cloverworm

We examined the searching behavior of a guild of primary parasitoids which attack the green cloverworm,Plathypena scabra (Fabricius), as well as that of an associated hyperparasitoid. We hypothesized that self-superparasitism is an important constraint on the foraging behavior of primary parasitoids, and therefore these parasitoids should avoid portions of the soybean canopy where parasitized caterpillars accumulate. Conversely, we hypothesized that the hyperparasitoid preferentially searches parts of the canopy where parasitized caterpillars accumulate. In a greenhouse experiment, we found that exposure to parasitoids [eitherCotesia marginiventris (Cresson) orDiolcogaster facetosa Ashmead] resulted in the accumulation of caterpillars lower in the canopy. In a field experiment, we measured the amount of time parasitoids spent searching in each of three strata (upper, middle, bottom) of the soybean canopy. Leaf area in each stratum was used to calculate expected values for search effort. The time spent searching each of the strata was proportional to leaf area for all primary parasitoids, exceptD. facetosa, which spent significantly more time searching the top stratum of plants than predicted by leaf area in that stratum. The hyperparasitoidMesochorus discitergus (Say) tended to search the bottom stratum of the canopy. Thus only one of the three primary parasitoids appears to search in a manner that would reduce its rate of encounter with previously parasitized green cloverworms. The hyperparasitoid searching pattern may increase its probability of encountering parasitized caterpillars, thereby increasing its foraging success.     

A simple rule for the costs of vigilance: empirical evidence from a social forager

It is commonly assumed that anti-predator vigilance by foraging animals is costly because it interrupts food searching and handling time, leading to a reduction in feeding rate. When food handling does not require visual attention, however, a forager may handle food while simultaneously searching for the next food item or scanning for predators. We present a simple model of this process, showing that when the length of such compatible handling time Hc is long relative to search time S, specifically Hc/S > 1, it is possible to perform vigilance without a reduction in feeding rate. We test three predictions of this model regarding the relationships between feeding rate, vigilance and the Hc/S ratio, with data collected from a wild population of social foragers (samango monkeys, Cercopithecus mitis erythrarchus). These analyses consistently support our model, including our key prediction: as Hc/S increases, the negative relationship between feeding rate and the proportion of time spent scanning becomes progressively shallower. This pattern is more strongly driven by changes in median scan duration than scan frequency. Our study thus provides a simple rule that describes the extent to which vigilance can be expected to incur a feeding rate cost.     

Beyond diffusion: Modelling local and long-distance dispersal for organisms exhibiting intensive and extensive search modes

The distribution of foragers on the landscape has important consequences to, for example, the spread rate of an invasive species or the outcrossing levels between neighbouring crops. Since forager distribution can be difficult to measure directly, mathematical models are often used to predict the population density of dispersing foragers on the landscape. We model organism movement using a diffusion framework in which the foraging population is divided into two subpopulations engaged in intensive and extensive search modes respectively. Movement in the intensive search mode (ISM) is modeled by diffusion, and movement in the extensive search mode (ESM) is modeled by advection. We show that our model provides a superior fit to organism movement data than more traditional diffusion or diffusion-advection models in which the forager population is considered homogeneous. Our results have implications for the understanding of dispersal in a wide variety of applications.     

How a simple adaptive foraging strategy can lead to emergent home ranges and increased food intake

Animals often alternate between searching for food locally and moving over larger distances depending on the amount of food they find. This ability to switch between movement modes can have large implications on the fate of individuals and populations, and a mechanism that allows animals to find the optimal balance between alternative movement strategies is therefore selectively advantageous. Recent theory suggests that animals are capable of switching movement mode depending on heterogeneities in the landscape, and that different modes may predominate at different temporal scales. Here we develop a conceptual model that enables animals to use either an area‐concentrated food search behavior or undirected random movements. The model builds on the animals’ ability to remember the profitability and location of previously visited areas. In contrast to classical optimal foraging models, our model does not assume food to be distributed in large, well‐defined patches, and our focus is on animal movement rather than on how animals choose between foraging patches with known locations and value. After parameterizing the fine‐scale movements to resemble those of the harbor porpoise Phocoena phocoena we investigate whether the model is capable of producing emergent home ranges and use pattern‐oriented modeling to evaluate whether it can reproduce the large‐scale movement patterns observed for porpoises in nature. Finally we investigate whether the model enables animals to forage optimally. We found that the model was indeed able to produce either stable home ranges or movement patterns that resembled those of real porpoises. It enabled animals to maximize their food intake when fine‐tuning the memory parameters that controlled the relative contribution of area concentrated and random movements.     

Intraspecific variation in movement patterns: modeling individual behaviour in a large marine predator

In large marine predators, foraging entails movement. Quantitative models reveal how behaviours can mediate individual movement, such that deviations from a random pattern may reveal specific search tactics or behaviour. Using locations for 52 grey seals fitted with satellite-linked recorders on Sable Island; we modeled movement as a correlated random walk (CRW) for individual animals, at two temporal scales. Mean move length, turning angle, and net squared displacement (R²_n: the rate of change in area over time) at successive moves over 3 to 10 months were calculated. The distribution of move lengths of individual animals was compared to a Lévy distribution to determine if grey seals use a Lévy flight search tactic. Grey seals exhibited three types of movement as determined by CRW model fit: directed movers – animals displaying directed long distance travel that were significantly underpredicted by the CRW (23% of animals); residents – animals remaining in the area surrounding Sable Island that were overpredicted by the model (29% of animals); and correlated random walkers – those (48% of animals) in which movement was predicted by the CRW model. Kernel home range size differed significantly among all three movement types, as did travel speed, mean move length, mean R²_n and total distance traveled. Sex and season of deployment were significant predictors of movement type, with directed movers more likely to be male and residents more likely to be female. Only 30% of grey seals fit a Lévy distribution, which suggests that food patches used by the majority of seals are not randomly distributed. Intraspecific variation in movement behaviour is an important characteristic in grey seal foraging ecology, underscoring the need to account for such variability in developing models of habitat use and predation.     

Agonistic Tactics and Size Asymmetries between Opponents in Blattella germanica (L.) (Dictyoptera: Blattellidae)

In this paper we investigated the influence of size differences between opponents on the occurrence of different agonistic attack and response tactics displayed during encounters between Blattella germanica (L.) cockroaches of all developmental stages, competing for a limited food source. The two main results were: 1) attack and response tactics varied during development and 2) size asymmetries between contestants influenced agonistic tactics. Developmental stage of initiator influenced the frequency distribution of the three types of attack patterns (Bite, Kick and Jump) and of the four types of response patterns (Retaliation, Escape, Resettle and No Response). The proportion of kicks increased gradually with developmental stage of the initiator whereas the proportion of bites declined during development. In addition, cockroaches adapted their attack tactics to the developmental stage of their opponent. Similarly the response displayed varied in relation to the developmental stage of the attacked individual, the developmental stage of the initiator and the type of attack displayed. During an interaction, animals appeared to be able to evaluate the relative size of their opponent and to evaluate the consequences of the behavioural pattern they displayed. Larger animals tended to minimize the intensity of the agonistic act they initiated. Escalation in interactions was rare and smaller individuals tended to escape when attacked by larger ones. Wrong estimations of the relative size of opponent, when a smaller animal retaliated after being attacked by a larger animal or when a larger animal fled after being attacked by a smaller one, represented only 6% of the records.     

The effect of habitat complexity on the functional response of a seed-eating passerine

Recent population declines of seed-eating farmland birds have been associated with reduced overwinter survival due to reductions in food supply. An important component of predicting how food shortages will affect animal populations is to measure the functional response, i.e. the relationship between food density and feeding rate, over the range of environmental conditions experienced by foraging animals. Crop stubble fields are an important foraging habitat for many species of seed-eating farmland bird. However, some important questions remain regarding farmland bird foraging behaviour in this habitat, and in particular the effect of stubble on farmland bird functional responses is unknown. We measured the functional responses of a seed-eating passerine, the Chaffinch Fringilla coelebs , consuming seeds placed on the substrate surface in three different treatments: bare soil, low density stubble and high density stubble. Stubble presence significantly reduced feeding rates, but there was no significant difference between the two stubble treatments. Stubble reduced feeding rates by reducing the maximum attack distance, i.e. the distance over which an individual food item is targeted and consumed. The searching speed, handling time per seed, proportion of time spent vigilant, duration of vigilance bouts and duration of head-down search periods were unaffected by the presence of stubble. The frequency of vigilance bouts was higher in the bare soil treatment, but this is likely to be a consequence of the increased feeding rate. We show the influence of a key habitat type on the functional response of a seed-eating passerine, and discuss the consequences of this for farmland bird conservation.     

Experimental evidence of density dependent activity pattern of a large herbivore in an alpine ecosystem

Density dependent processes affecting foraging strategies may in turn influence vital rates and population regulation in large herbivores. Increased competition may lower both forage availability and quality, but whether the main activity constraint at high density is increased searching time or increased digestion time is poorly investigated. In a fully replicated landscape‐scale experiment, we used long‐term data (2003–2009) from domestic sheep grazing at high and low density (80 and 25 sheep km^–2, respectively) on alpine summer ranges to test density dependence in allocation of time to feeding (moving) vs digestion (resting) activities and how this in turn affected body growth. Sheep at high density spent more time actively feeding than sheep at low density, but sheep moved shorter distances while foraging at high density. Increased activity levels at high density suggest that the main activity constraint at high density was availability of high‐quality food increasing searching time and possibly reducing intake rates. Increased movement distances at low density is consistent with a higher selection for more productive vegetation types since high‐quality patches are dispersed in the landscape. The alternative hypothesis, that food processing time increased at high density was not supported as it would have reduced overall activity levels. Individual activity levels increased body growth, but this was not sufficient to fully compensate for lower habitat quality leading to an overall reduced body growth at high density. Our experiment clearly documents changes in activity budgets and movement distances of a large herbivore at high population density, providing one potential behavioural mechanism of density dependent responses observed in vital rates.     

Is It Worth the Risk? Food Deprivation Effects on Tadpole Anti-Predatory Responses

Predation risk affects foraging behavior and, particularly, the amount of time devoted to the search for food. When exposed to predation risk, food deprived animals should be risk prone and relax behavioral defenses to a wider extent than well fed individuals. To test for this prediction, we used a 2?×?2 factorial design experiment, manipulating both the energetic state (fed vs. fasted) and exposure to an attack-released cue (injured vs. uninjured conspecifics) of common water frog tadpoles (Pelophylax kl. esculentus). Contrary to expectations, food deprivation significantly lowered the activity level of predator-exposed tadpoles. As in this experiment no food resource was added to test containers, energy conserving behavior might have both delayed starvation and lowered the probability of encountering the potential predator. To test for the effect of food availability on behavioral responses, we performed a second experiment, using the same protocol and procedures, except for adding food to all test containers. All tadpoles showed similar levels of activity, while fed tadpoles exposed to alarm cues tended to swim farther from the cage containing the stimulus than in the first experiment. As many anuran larvae can feed on dead conspecifics, prey-borne cues may have been interpreted as the potential presence of both a food source and a predator, fed tadpoles possibly being more confident than fasted tadpoles in their ability to escape predation in case an actual attack occurs.     

First Passage Time Analysis of Animal Movement and Insights into the Functional Response

Movement plays a role in structuring the interactions between individuals, their environment, and other species. Although movement models coupled with empirical data are widely used to study animal distribution, they have seldom been used to study search time. This paper proposes first passage time as a novel approach for understanding the effect of the landscape on animal movement and search time. In the context of animal movement, first passage time is the time taken for an animal to reach a specified site for the first time. We synthesize current first passage time theory and derive a general first passage time equation for animal movement. This equation is related to the Fokker–Planck equation, which is used to describe the distribution of animals in the landscape. We illustrate the first passage time method by analyzing the effect of territorial behavior on the time required for a red fox to locate prey throughout its home range. Using first passage time to compute search times, we consider the effect of two different searching modes on a functional response. We show that random searching leads to a Holling type III functional response. First passage time analysis provides a new tool for studying how animal movement may influence ecological processes.     

Food Searching Strategy of Amoeboid Cells by Starvation Induced Run Length Extension

Food searching strategies of animals are key to their success in heterogeneous environments. The optimal search strategy may include specialized random walks such as Levy walks with heavy power-law tail distributions, or persistent walks with preferred movement in a similar direction. We have investigated the movement of the soil amoebae Dictyostelium searching for food. Dictyostelium cells move by extending pseudopodia, either in the direction of the previous pseudopod (persistent step) or in a different direction (turn). The analysis of ∼4000 pseudopodia reveals that step and turn pseudopodia are drawn from a probability distribution that is determined by cGMP/PLA2 signaling pathways. Starvation activates these pathways thereby suppressing turns and inducing steps. As a consequence, starved cells make very long nearly straight runs and disperse over ∼30-fold larger areas, without extending more or larger pseudopodia than vegetative cells. This ‘win-stay/lose-shift’ strategy for food searching is called Starvation Induced Run-length Extension. The SIRE walk explains very well the observed differences in search behavior between fed and starving organisms such as bumble-bees, flower bug, hoverfly and zooplankton.     

Time‐to‐kill: measuring attack rates in a heterogenous landscape with multiple prey types

Although spatial heterogeneity of prey and landscapes are known to contribute to variation around predator‐prey functional response models, few studies have quantified these effects. We illustrate a new approach using data from winter movement paths of GPS‐collared wolves in the Rocky Mountains of Canada and time‐to‐event models with competing risks for measuring the effect of prey and landscape characteristics on the time‐to‐kill, which is the reciprocal of attack rate (aN) in a Holling's functional response. We evaluated 13 a priori models representing hypothesized mechanisms influencing attack rates in a heterogeneous landscape with two prey types. Models ranged from variants on Holling's disc equation, including search rate and prey density, to a full model including prey density and patchiness, search rates, satiation, and landscape features, which were measured along the wolf's movement path. Movement rates of wolves while searching explained more of the variation in time‐to‐kill than prey densities. Wolves did not compensate for low prey density by increasing movement rates and there was little evidence that spatial aggregation of prey influenced attack rates in this multi‐prey system. The top model for predicting time‐to‐kill included only search rate and landscape features. Wolves killed prey more quickly in flat terrain, likely due to increased vulnerability from accumulated snow, whereas attack rates were lower when wolves hunted near human‐made features presumably due to human disturbance. Understanding the sources of variation in attack rates provides refinements to functional response models that can lead to more effective predator–prey management in human‐dominated landscapes.     

Time sharing between host searching and food searching in parasitoids: state-dependent optimal strategies

By varying the time spent searching for food, parasitoids modifytheir expected lifespan, and therefore their total lifetimereproductive success. Using a stochastic dynamic approach, wedefine the best choice between searching for food and searchingfor hosts as a function of the state of the parasitoid and theavailability of food when hosts and food are found in differentparts of the environment A first model deals with the influenceof food availability and survivorship conditions on the behaviorof a single parasitoid. Our results suggest that under conditionsof very low food availability, parasitoids should never searchfor food. When food availability is moderate, parasitoids shouldnot wait until their reserves are low before searching for food.When food is abundant and survivorship is independent of foodconsumption, parasitoids should search for food only when theirreserves are almost exhausted. They should not wait so longif survivorship depends on the energy reserves. By finding thestate-dependent ideal free distribution for a population ofparasitoids, we are able to predict their distribution betweenthe feeding area and the host living area at equilibrium. Theproportion of parasitoids in each area is altered by the numberof competitors and interference. Finally, the model predictsthat optimal time sharing between food searching and host searchingmay promote the stability of the host-parasitoid system.     

Prey selection and foraging period of the predaceous rocky intertidal snail,Acanthina punctulata

Summary The diet and foraging period of the neogastropod Acanthina punctulata were investigated in order to test various aspects of recent optimal foraging strategy models. This intertidal snail is an actively searching predator which preys on snails and barnacles by boring a hole in the shell and rasping out the flesh. Unlike many gastropod predators, Acanthina drill its gastropod prey at a very specific location on the columella, the thickest portion of the shell. Acanthina's foraging period can be interpreted as a compromise between maximizing the energy obtained by feeding and minimizing risk of mortality from exposure to wave action. That foraging period minimizing risk of being dislodged by waves appears to be during low tide when the predators can be in shallow pools. However, prey cannot be captured and consumed during one low tide. Thus Acanthina must be exposed during some high tides, and its strategy appears to be to restrict movement while exposed. Thus search is not initiated during high tide, but drilling and prey consumption are continued during that time. A snail not drilling or consuming prey seeks the protection of crevices or large anemones during high tide. A model is presented to indicate the relative amounts of risk and net energy for Acanthina at successive low and high tides. Predictions from the model, e.g., minimizing search time to avoid being exposed for an additional high tide and no movement during high tide are supported by field data. Acanthina commences foraging at the beginning of low tide, searches initially for preferred prey, but if unsuccessful, settles for a less preferred prey and begins drilling this prey before the end of low tide. Drilling and ingestion of prey occur during the following high and sometimes low tides. These handling times take 95% of the total foraging time in the field, while search time takes only 5% (pursuit time is negligible). Drilling alone accounts for 48–70% of the total drilling and eating time. In the laboratory, drilling and eating time for littorine food ranged from 15–60 hrs per item. The time to drill and eat a littorine increases exponentially with prey length. Since handling and processing prey items represents such a large investment of time, Acanthina would be expected to be very selective with respect to choice of prey items. Electivity coefficients from field data suggest that littorines are preferred over barnacles. Acanthina in the laboratory optimizes the amount of biomass ingested per time by choosing larger littorines over smaller ones and by preferring the more readily drilled species.It is suggested that Acanthina obtains information about the range of prey available initially by encountering and evaluating quite a few prey before making a selection, but usually by comparing an item of prey encountered to the prey it recently ingested. This latter method should provide a basis for evaluating prey encountered and has the advantage of reducing search time, the total amount of time spent feeding and thus the high-tide time exposed to wave action.In a similar manner, the decrease in the level of acceptability of prey as search time increases represents a compromise between maximizing energy obtained and minimizing risk from mortality.     

Producer–Scrounger Games in a Spatially Explicit World: Tactic Use Influences Flock Geometry of Spice Finches

Group-foraging animals can either search for their food (producer) or search for opportunities to join the food discoveries of others (scrounger). To maximize food returns, producers should distance themselves from potential competitors whereas scroungers should increase proximity to potential producers. We investigated the extent to which playing one or the other tactic affected an individual's location in captive flocks of ground-feeding spice finches ( Lonchura punctulata ) as they foraged for hidden clumps of food on an aviary floor. We constrained some individuals to use the producer tactic by pre-training them to find food hidden under lids. Constrained producers foraged significantly further from the center of flocks than constrained scroungers. Flocks with many scroungers were significantly more compact than flocks with fewer scroungers. The results are consistent with published simulations of spatially explicit producer–scrounger models and suggest that the use of producer and scrounger foraging tactics be included as a factor that affects an individual's position within foraging groups.     

Efficient harvesting of renewing resources

Many foraging animals return to feeding sites to harvest replenishingresources, but little is known about efficient tactics for doingthis. Can animals with adequate cognitive abilities increasetheir efficiency by modifying their behavior according to memoriesof past experience at particular sites? We developed a simulationmodel of animals harvesting renewable resources from isolatedpatches in undefended, competitive situations. We compared fourforaging tactics: (1) moving stochastically without using anyinformation from past experiences (random searching); (2) movingstochastically, but going longer distances after encounteringlower reward (area-restricted searching); (3) repeatedly movingalong a fixed route (complete traplining); and (4) traplining,but sampling and shifting to neighboring rewarding patches afterencountering low reward (sample-and-shift traplining). FollowingPossingham, we tracked both the resources actually harvestedby a focal forager (i.e., rewards) and the standing crops ofresources that accumulated at patches. Complete traplining alwaysproduces less variation in elapsed time between visits thanrandom searching or area-restricted searching, which has threebenefits: increasing the reward crop harvested, if resourcerenews nonlinearly; reducing resource standing crop in patches;and reducing variation in reward crop per patch. Moreover, thesystematic revisitation schedule produced by complete trapliningmakes it more competitive, regardless of resource renewal scheduleor competitor frequency. By responding to their past experiences,using sample-and-shift traplining, foragers benefit only whenmany patches are left unvisited in the habitat. Otherwise, theexploratory component of sample-and-shift traplining, whichincreases the movement distance and the variation in elapsedtime between visits, makes it more costly than complete traplining.Thus, traplining will usually be beneficial, but foragers shouldswitch between "impatient" (sample-and-shift traplining) and"tenacious" (complete traplining) traplining, according to temporalchanges in surrounding situations.     

Route finding by rats in an open arena

Rats were repeatedly exposed to an open arena containing two depletable food sources in a discrete-trials procedure. Their movement patterns were recorded and compared to adaptive foraging tactics such as minimizing distance or energy expenditure, thigmotaxis, and trail following. They were also compared to the predictions of the associative route-finder model of Reid and Staddon [Reid, A.K., Staddon, J.E.R., 1998. A dynamic route finder for the cognitive map. Psychol. Rev. 105 (3), 585-601]. We manipulated the presence/absence of food, goal cups, and a wooden runway to determine the influence of local and distal stimuli (visual, olfactory, and tactile) on movement patterns. Increased experience in the arena produced decreases in travel distance and time to the food sources. Local and distal stimuli influenced movement patterns in ways compatible with visual beacons and trail following. The route-finder model accurately predicted movement patterns except those that were influenced by local and distal stimuli. These results show how certain stimuli influence movement and provide a guide for the incorporation of local and distal stimuli in a future version of the dynamic route-finder model.     

Experimental evidence that group foragers can converge on predicted producer-scrounger equilibria

When foraging together, animals are often observed to feed from food discoveries of others. The producer-scrounger (PS) game predicts how frequently this phenomenon of food parasitism should occur. The game assumes: (1) at any moment all individuals can unambiguously be categorized as either playing producer (searching for undiscovered food resources) or scrounger (searching for exploitation opportunities), and (2) the payoffs received from the scrounger tactic are negatively frequency dependent; a scrounger does better than a producer when the scrounger tactic is rare, but worse when it is common. No study to date has shown that the payoffs of producer and scrounger conform to the game's assumptions or that groups of foragers reach the predicted stable equilibrium frequency (SEF) of scrounger, whereby both tactics obtain the same payoff. The current study of three captive flocks of spice finches, Lonchura punctulata, provides the first test of the PS game using an apparatus in which both assumptions of the PS game are met. The payoffs to the scrounger, measured as feeding rate (seeds/s), were highly negatively frequency dependent on the frequency of scrounger. The feeding rate for scrounger declined linearly while the rate for producer either declined only slightly or not at all with increasing scrounger frequency. When given the opportunity to alternate between tactics, the birds changed their use of each, such that the group converged on the predicted SEF of scrounger after 5-8 days of testing. Individuals in this study, therefore, demonstrated sufficient plasticity in tactic use such that the flock foraged at the SEF of scrounger. Copyright 2000 The Association for the Study of Animal Behaviour.