Choosing where to feed: the influence of competition on feeding behaviour of cod, Gadus morhua L.

Groups of cod, Gadus morhua L, presented with two feeding patches with a food abundance ratio of 2:1, distributed themselves between the patches in a ratio of 2.5:1. This is slightly higher than the 2:1 ratio predicted by the ideal free distribution theory. Large differences were observed in competitive ability between individual fish. A strong correlation was found between feeding success of individuals and time spent in a feeding patch. The more successful competitors caught about 2.5 times as many food items in the rich patch as in the poor patch. The less successful competitors caught an equal amount of food in both patches. All competitors, however, spent significantly more time in the rich patch. These results suggest that hunting success is the most important factor in assessing patch quality. However, it is not the only parameter which cod use in deciding where to feed.     

Patch Choice Decisions among Ifaluk Fishers

Studies of patch choice decisions among human foragers have failed to explain why foragers do not exclusively exploit the patch with the highest mean profitability. One possible explanation is that profitability rankings are likely to vary daily; however, this instability is not captured when profitabilities are calculated as a sampled average over a longer time span. Here I present data on the patch choice decisions of Ifaluk fishers to evaluate whether men are responding to daily variation in the profitability of their primary fishing patch. Results show that men choose to fish most frequently in the patch with the highest mean profitability. Men fish in alternative patches (alternative from the most profitable patch) when, on that morning or the previous day, return rates in the most profitable patch are lower than the overall mean per capita return rate of alternative patches. Results also indicate that when fishers pursue alternative patches after fishing in the patch with the highest profitability, their mean per capita return rates are generally higher in the alternative patches exploited. However, variance in the profitability of the most profitable patch cannot explain why men exploit two patches, the Nine-mile reef and the dogtoothed tuna patch, which on average have very low profitability. These results and directions for future research are discussed. [Keywords: human behavioral ecology, patch choice decisions, Micronesia]     

Do cattle egrets gain information from conspecifics when foraging?

Summary We examined whether individual cattle egrets (Bubulcus ibis) base their decisions of where to forage, and how long to stay in a patch, on the behavior of other flock members. Cattle egrets commonly forage in flocks associated with cattle and capture prey at higher rates when they do not share a cow with another egret. Foraging egrets provide cues of the location of prey and their success in capturing prey. Therefore, there is the possibility of information transfer between egrets in a flock. We predicted that egrets should only move to occupied patches when the resident was capturing enough prey that it is profitable for the invader to share the patch or take over the patch. However, egrets did not seem to decide where to forage based on neighbors' rates of energy intake, but rather on the presence or absence of conspecifics in a patch. We also predicted that an egret should remain in a patch until its rate of energy intake dropped to or below the average rate for other egrets within the flock. However, egrets that were foraging more efficiently than the average rate for the flock switched patches sooner than less efficient foragers. Egrets did not appear to increase foraging success by gaining information on patch quality from neighbors.     

Sex differences in foraging behaviour and oviposition site preference in an insect predator, Orius sauteri

The effects of patch quality on the foraging behaviour of an anthocorid predator Orius sauteri (Poppius) were compared between sexes. Prior experience in patches was also studied to determine whether this was a factor affecting oviposition decisions. Patch quality affected patch residence time differently for the two sexes; females stayed much longer in a patch with prey (60 Thrips palmi larvae) than a patch without prey, while males did not remain in any patch for extended periods. Most of the females remained in or moved to patches with prey, whereas males dispersed, irrespective of patch quality. Both females released in patches with prey and females released in patches without prey deposited more eggs per hour in patches with prey than in patches without prey. Females released in patches without prey laid eggs in patches with prey at higher rates than did females released in patches with prey. Causes for the sex difference in patch residence time and allocation are discussed in relation to optimal foraging theory. The significance of selective oviposition and the role of experience in oviposition decisions within heterogeneous environments are also discussed.     

Response of predators to prey abundance: separating the effects of prey density and patch size

We tested the relative and combined effects of prey density and patch size on the functional response (number of attacks per unit time and duration of attacks) of a predatory reef fish (Cheilodactylus nigripes (Richardson)) to their invertebrate prey. Fish attacked prey at a greater rate and for longer time in large than small patches of prey, but large patches had naturally greater densities of prey. We isolated the effects of patch size and prey density by reducing the density of prey in larger patches to equal that of small patches; thereby controlling for prey density. We found that the intensity at which fish attacked prey (combination of attack rate and duration) was primarily a response to prey density rather than the size of patch they occupied. However, there was evidence that fish spent more time foraging in larger than smaller patches independent of prey density; presumably because of the greater total number of prey available. These experimental observations suggest that fish can distinguish between different notions of prey abundance in ways that enhance their rate of consumption. Although fish may feed in a density dependent manner, a critical issue is whether their rate of consumption outstrips the rate of increase in prey abundance to cause density dependent mortality of prey.     

Ideal free distributions when individuals differ in competitive ability: phenotype-limited ideal free models

A series of prospective models is developed to investigate ideal free distributions in populations where individuals differ in competitive ability. The models are of three types. In the continuous-input models, there is continuous arrival of food or mates into each habitat patch, and competitors scramble to obtain as large a share as possible. In the interference models, the prey density in a particular patch stays constant but the presence of competitors slows down the rate at which prey are captured. In the kleptoparasitism model, individuals have food or females stolen from them by competitors higher in the dominance hierarchy, and in turn steal items from subordinates. A general result of the continuous-input and interference models is that the population of competitors can be truncated between patches so that the individuals with the highest competitive ability occur in the best patches, or in the patches where competitive differences are greatest. Individuals of lowest competitive ability occur in the poorest patches or where competitive differences are least, and intermediate phenotypes are ranked between these two extremes. Thus the ideal free prediction that all individuals will achieve equal fitness will not apply. However, in continuous-input cases where competitive differences between phenotypes remain constant across patches, this solution is only neutrally stable, and forms only one element of a set of equilibrium distributions. The fact that many empirical studies of continuous-input have found approximately equal mean fitness across patches may relate to this finding. Most interference studies contradict the simple ideal free solution by having different mean intake rates across patches; this may relate to the predicted positive correlation of competitive ability with patch quality. The kleptoparasitism model usually generated continuous cycling of individuals between habitat patches, though some correlation could be found between competitive ability and patch quality.     

On the role of patch density and patch variability in central-place foraging

Existing models of the foraging behavior of single-prey loaders in patchy environments differ on whether the optimal forager is predicted to stay in a patch until a prey is found, or to leave a patch for a next one if a prey is not found by a certain “deadline.” This article examines conditions on the probability distribution of prey density across patches that are necessary or sufficient for the existence of a finite, optimal deadline. It is shown that, for environments in which prey density is variable but never falls below some strictly positive level, a finite, optimal deadline exists when and only when the spatial density of patches is “high.” Also, a characterization is given of a large class of distributions (including the gamma distribution) for which a finite, optimal deadline exists for all levels of spatial density of patches.     

Minimax strategies for a predator—Prey game

Suppose prey are distributed in patches. The predator knows the fraction of patches containing 0, 1, 2,… prey, but not how many prey a particular patch contains. It searches each patch randomly, at constant speed. It leaves a patch when the intercapture times satisfy a formula designed to maximize the number of prey eaten per unit time. We show that, if the prey distribution is Poisson, the predator should stay in each patch for the same time, regardless of what happens there. Accordingly, the prey can minimize the predator's maximum intake by choosing the Poisson distribution, and the predator can maximize its minimum intake (against a “smart” prey) by choosing the constant-time strategy.     

On optimal diet in a patchy environment

Optimal foraging theory has dealt with the following questions independently: (1) On what prey types should an individual predator feed (optimal diet)? (2) How long should a predator stay in each patch if prey is patchily distributed (optimal allocation of time to patches) ? This paper explores optimal foraging in patches containing several different kinds of prey. Results obtained by simulation show that deviations from recent predictions are to be expected, particularly for long interpatch travel times and rapid depletion of profitable prey types. In these situations the tactics of feeding as either specialist or as a generalist can be inferior to a tactic which starts as a specialist and then expands the diet after some time in the patch. Furthermore, predators should not necessarily stay longer in a patch if interpatch travel time increases. Some experimental tests of these new predictions are proposed.     

Foraging Rules for Group Feeders: Area Copying Depends upon Food Density in Shoaling Goldfish

The decision rules governing forage area copying behaviour were investigated in shoaling fish. Shoaling goldfish were offered two equal food patches, one of which was adjacent to an equal-sized shoal feeding behind a transparent barrier. When food was low, goldfish foraged according to an area copying rule, but under high and zero food area copying disappeared. Only under high food density did equal numbers of fish feed at both sites as predicted by foraging theory. Under zero food the fish were less certain about where to forage. Precise visual cues from feeding fish were required: non-feeders did not attract area copiers. Furthermore, area copying was task-dependent since it reappeared strongly if fish were not able to forage on patches like their fellows. Control experiments eliminated an increase in group size for anti-predator advantage as an explanation. Two sequential decisions: to stay or move, and to join or leave may explain the results, which are not accommodated by simple optimality models. These decisions may be based on a comparison of current food intake with the anticipation of a higher reward by foraging socially.     

Optimal Bayesian foraging policies and prey population dynamics-some comments on Rodriguez-Girones and Vasquez

In this paper we show the density-dependent harvest rates of optimal Bayesian foragers exploiting prey occurring with clumped spatial distribution. Rodríguez-Gironés and Vásquez (1997) recently treated the issue, but they used a patch-leaving rule (current value assessment rule) that is not optimal for the case described here. An optimal Bayesian forager exploiting prey whose distribution follows the negative binomial distribution should leave a patch when the potential (and not instantaneous) gain rate in that patch equals the best long-term gain rate in the environment (potential value assessment rule). It follows that the instantaneous gain rate at which the patches are abandoned is an increasing function of the time spent searching in the patch. It also follows that the proportion of prey harvested in a patch is an increasing sigmoidal function of the number of prey initially present. In this paper we vary several parameters of the model to evaluate the effects on the forager's intake rate, the proportion of prey harvested per patch, and the prey's average mortality rate in the environment. In each case, we study an intake rate maximizing forager's optimal response to the parameter changes. For the potential value assessment rule we find that at a higher average prey density in the environment, a lower proportion of the prey is taken in a patch with a given initial prey density. The proportion of prey taken in a patch of a given prey density also decreases when the variance of the prey density distribution is increased and if the travel time between patches is reduced. We also evaluate the effect of using predation minimization, rather than rate maximization, as the currency. Then a higher proportion of the prey is taken for each given initial prey density. This is related to the assumption that traveling between patches is the most risky activity. Compared to the optimal potential value assessment rule, the current value assessment rule performs worse, in terms of long-term intake rate achieved. The difference in performance is amplified when prey density is high or highly aggregated. These results pertain to the foraging patch spatial scale and may have consequences for the spatial distribution of prey in the environment.     

How does hunger affect convergence on prey patches in a social forager?

Internal state, in this case hunger, is known to influence both the organisation of animal groups and the social foraging interactions that occur within them. In this study, we investigated the effects of hunger upon the time taken to locate and converge upon hidden simulated prey patches in a socially foraging fish, the threespine stickleback (Gasterosteus aculeatus). We predicted that groups of food‐deprived fish would find and recruit to prey patches faster than recently fed groups, reasoning that they might search more rapidly and be more attentive to inadvertent social information produced by other foragers. Instead we saw no difference between the two groups in the time taken to find the patches and found that in fact, once prey patches had been discovered, it was the recently fed fish that converged on them most rapidly. This finding is likely due to the fact that recently fed fish tend to organise themselves into fewer but larger subgroups, which arrived at the food patch together. Hunger has a significant impact upon the social organisation of the fish shoals, and it appears that this has a stronger effect upon the rate at which they converged upon the food patches than does internal state itself.     

Optimal Bayesian Foraging Policies and Prey Population Dynamics—Some Comments on Rodríguez-Gironés and Vásquez

In this paper we show the density-dependent harvest rates of optimal Bayesian foragers exploiting prey occurring with clumped spatial distribution. Rodríguez-Gironés and Vásquez (1997) recently treated the issue, but they used a patch-leaving rule (current value assessment rule) that is not optimal for the case described here. An optimal Bayesian forager exploiting prey whose distribution follows the negative binomial distribution should leave a patch when the potential (and not instantaneous) gain rate in that patch equals the best long-term gain rate in the environment (potential value assessment rule). It follows that the instantaneous gain rate at which the patches are abandoned is an increasing function of the time spent searching in the patch. It also follows that the proportion of prey harvested in a patch is an increasing sigmoidal function of the number of prey initially present. In this paper we vary several parameters of the model to evaluate the effects on the forager's intake rate, the proportion of prey harvested per patch, and the prey's average mortality rate in the environment. In each case, we study an intake rate maximizing forager's optimal response to the parameter changes. For the potential value assessment rule we find that at a higher average prey density in the environment, a lower proportion of the prey is taken in a patch with a given initial prey density. The proportion of prey taken in a patch of a given prey density also decreases when the variance of the prey density distribution is increased and if the travel time between patches is reduced. We also evaluate the effect of using predation minimization, rather than rate maximization, as the currency. Then a higher proportion of the prey is taken for each given initial prey density. This is related to the assumption that traveling between patches is the most risky activity. Compared to the optimal potential value assessment rule, the current value assessment rule performs worse, in terms of long-term intake rate achieved. The difference in performance is amplified when prey density is high or highly aggregated. These results pertain to the foraging patch spatial scale and may have consequences for the spatial distribution of prey in the environment.     

The effects of perceived competition and parasitism on the foraging behaviour of the upland bully (Eleotridae)

Prey choice by fish is subject to many constraints, some of which may interact to determine the relative preference of fish for prey with different profitabilities. The constraining effects of parasitism and perceived competition on foraging behaviour were examined in the upland bully, Gobiomorphus breviceps . In the laboratory, bullies faced with a choice of prey items of two different sizes chose the larger prey significantly more often than the smaller ones. The presence of a conspecific fish near the source of large prey significantly reduced the bullies'preference for larger prey. Neither the size of the test fish, nor the number of digenean metacercarial cysts they harboured, had any influence on their relative preference for larger prey, or on how that preference was dampened by the presence of a competitor. The threat of competition appears to be a more important constraint on prey choice in upland bullies than parasitism.     

Foraging on patchily distributed prey by a cichlid fish (Teleostei,Cichlidae): A test of the ideal free distribution theory

Cichlid fish (Aequidens curviceps) distributed themselves and allocated their foraging time between two drift food patches in close approximation to the patch profitability ratio, as predicted by the ideal free distribution theory. The fish thereby achieved similar average feeding rates in the two patches, in two of three patch profitability ratio experiments. However, one major assumption of the ideal free model was violated, since individual fish differed in their competitive abilities for limited food resources, which resulted in unequal payoffs among individuals within each patch. Individual variation in feeding rates, and thus in competitive ability, was not related to despotism, but perhaps rather to individual differences in perceptual ability and in the ability to learn which patch was currently the more profitable. The strategy used by the fish to assess patch profitability included sampling available patches. However, individual fish switched (sampled) patches with varying frequency. Sampling had an associated cost, since high-frequency switchers had lower feeding rates on average than low-frequency switchers. Differences in foraging strategy among the fish therefore contributed to the observed in-equality in individual payoffs within patches.     

Sharing, consumption, and patch choice on Ifaluk atoll

Anthropological tests of patch choice models from optimal foraging theory have primarily employed acquisition rates as the currency of the model. Where foragers share their returns, acquisition rates may not be similar to consumption rates and thus may not be an appropriate currency to use when modeling foraging decisions. Indeed, on Ifaluk Atoll the distribution patterns of fish vary by fishing method and location. Previous analyses of Ifaluk patch choice decisions suggested that if Ifaluk fishers are trying to maximize their production rates they should rarely torch fish for dogtoothed tuna. However, some men do spend considerable time and energy exploiting the dogtoothed tuna patch. To improve our understanding of the constraints and motivations influencing men’s decisions to exploit this patch, here I use per capita consumption rates as a currency, rather than production rates, to evaluate predictions generated from a patch choice model. Results indicate that although fish caught in other patches are more widely distributed than fish caught in the dogtoothed tuna patch, the consumption rates of torch fishers and their kin are still considerably lower than the consumption rates of men pursuing fish in other patches. Although these results are unable to explain why Ifaluk men exploit the dogtoothed tuna patch, an important explanatory hypothesis is eliminated.     

Patch use in time and space for a meso-predator in a risky world

Predator–prey studies often assume a three trophic level system where predators forage free from any risk of predation. Since meso-predators themselves are also prospective prey, they too need to trade-off between food and safety. We applied foraging theory to study patch use and habitat selection by a meso-predator, the red fox. We present evidence that foxes use a quitting harvest rate rule when deciding whether or not to abandon a foraging patch, and experience diminishing returns when foraging from a depletable food patch. Furthermore, our data suggest that patch use decisions of red foxes are influenced not just by the availability of food, but also by their perceived risk of predation. Fox behavior was affected by moonlight, with foxes depleting food resources more thoroughly (lower giving-up density) on darker nights compared to moonlit nights. Foxes reduced risk from hyenas by being more active where and when hyena activity was low. While hyenas were least active during moon, and most active during full moon nights, the reverse was true for foxes. Foxes showed twice as much activity during new moon compared to full moon nights, suggesting different costs of predation. Interestingly, resources in patches with cues of another predator (scat of wolf) were depleted to significantly lower levels compared to patches without. Our results emphasize the need for considering risk of predation for intermediate predators, and also shows how patch use theory and experimental food patches can be used for a predator. Taken together, these results may help us better understand trophic interactions.     

Failure of simple optimal foraging models to predict residence time when patch quality is uncertain

Blue jays (Cyanocitta cristata) were presented with a foragingsituation in which half of the patches they encountered containedno prey and half contained a single prey item. Experimentallydetermined probability distributions controlled prey arrivaltimes in those patches that contained prey. Patch residencein empty patches was studied during four experiments. In thefirst, prey arrival was exponentially distributed. Residencetimes increased with travel time as predicted by a rate-maximizationmodel, but the bird stayed in empty patches much longer thanpredicted. During the second experiment, prey arrival was uniformlydistributed. The jays again stayed longer than optimal, andpatch residence times increased as travel time increased, althoughthe residence time that maximized rate of intake was independentof travel time under these conditions. In the third experiment,exponential and uniform patches were randomly intermixed. Thejays showed larger travel-time effects in the exponential thanin the uniform patch. However, the travel-time effect in theuniform patch was contrary to rate-maximization predictions,and the birds again overstayed in both patch types. In the fourthexperiment, prefeeding at the start of each foraging bout slightlyincreased overstaying rather than decreasing overstaying, aswould be expected if overstaying were due to underestimatingenvironmental quality. Consistent and dramatic overstaying anda travel-time effect under conditions where travel time hasno effect on optimal residence times suggest that the rate-maximizationapproach does not apply to foraging problems involving patchuncertainty.     

Allochthonous prey subsidies provide an asymmetric growth benefit to invasive bluegills over native cyprinids under the competitive conditions in a pond

Asymmetry in the competition abilities between invasive and native consumers can potentially influence the colonization success by invasive species. We tested whether a subsidy of allochthonous prey enhanced an asymmetric competition between invasive bluegill (Lepomis macrochirus) and two native cyprinid fish, that is, stone moroko (Pseudorasbora parva) and tamoroko (Gnathopogon elongatus elongatus). A field experiment was conducted using enclosures wherein the strength of interspecific competition and the presence/absence of allochthonous prey were manipulated. The experiment revealed that allochthonous prey alleviated the limitation of fish growths caused by a severe competition for aquatic prey resources. However, the importance of allochthonous prey differed considerably between invasive bluegill and the two native cyprinids. Individual bluegills grew faster when the allochthonous prey was supplied, whereas no difference in growth was observed in the two cyprinids whether or not allochthonous prey was supplied. Interestingly, the importance of allochthonous prey on the total amount of bluegill growth varied depending on the numerical abundance of native cyprinid competitors, and this importance increased when the native cyprinids were abundant. These findings indicated that allochthonous prey provides an asymmetric growth benefit to invasive bluegills over the two native cyprinids by alleviating asymmetrically the competition strength in a Japanese pond, especially under the conditions of severe interspecific resource competition and a limitation in the utilization of in situ prey resources.     

Foraging strategy of a non-omniscient predator in a changing environment (I) model with a data window and absolute criterion

1. Almost all the models so far presented assume that predators are omniscient in the sense that they always have complete information about the spatial distribution of prey abundance and its change over time. But this type of model cannot cover the situation where the prey abundance in each patch changes over time due to factors other than predation. The model with a data window and absolute criterion (SAC) here enables us to treat such situations.
2. The strategy of non-omniscient predators can be generally devided into four procedures; collection of information, its memorization, decision of tactics and its execution. SAC involves only two tactics; to stay another time period in the patch the predator is staying presently or to move to another patch chosen at random. The choice of either one of the two tactics is made by comparing the profitability of the current patch estimated by the data window with a pre-determined absolute criterion.
3. Three changing patterns of prey abundance are considered. In the most general pattern good patches have a higher mean profitability than poor patches, but the profitability changes cyclically in each of patches.
4. There are only two possibilities for an optimal strategy; the “patch choice strategy” in which once the predator has taken a good patch, it tries to stay there even when the state becomes poor, and the ‘state choice strategy” in which the predator seeks for only good states in good patches. The condition for which either of the two foraging strategies is superior to the other is specified analytically.