Aggregations from using inadvertent social information: a form of ideal habitat selection

Social information in breeding site selection has received extensive study; however, few attempts have been made to link this process to pre‐existing models. We examine the importance of social information to three pertinent models of habitat selection that describe breeding aggregations and spatial patterns: 1) the ideal despotic distribution (IDD) which considers conspecific competition and habitat availability, 2) the perceptual constraints model which accounts for patch selection when animals experience a threshold of undetectable difference in quality, and 3) the “neighbourhood model” which predicts concordance between resources and settlers can be disrupted by conspecific attraction when resources are patchy. These models all predict initial settlers will select a high quality patch first. However, their predictions of subsequent settlement behaviour in remaining patches differ: the IDD predicts subsequent settlers will be distributed regularly, the perceptual constraints model predicts a random distribution, and the neighbourhood model predicts clustering from conspecific attraction. We examined which model best described settlement patterns of bobolink Dolichonyx oryzivorus and savannah sparrow Passerculus sandwichensis, in the context of social information. We observed settlement timing, quantified available resources, and determined where they occurred in the highest (local population “core”) and lowest densities (local population “periphery”). We then assessed whether individuals in the periphery settled in greater concordance with resources or conspecific presence. Core territories were clustered strongly on relevant resources, and these territory holders were older than in the periphery. Peripheral territories were likewise clustered but did not always co‐occur with the best available resources, matching the neighbourhood model prediction that social information may not always direct them to the best sites available. This suggests older individuals used their own experience to locate ideal habitat, whereas younger individuals attempted to aggregate on seemingly ideal habitat by using conspecific location; such information asymmetry due to age can be viewed as an “ideal aggregative distribution”.     

Bayesian foraging with only two patch types

I model the optimal Bayesian foraging strategy in environments with only two patch qualities. That is, all patches either belong to one rich type, or to one poor type. This has been a situation created in several foraging experiments. In contrast, previous theories of Bayesian foraging have dealt with prey distributions where patches may belong to one out of a large range of qualities (binomial, Poisson and negative binomial distributions). This study shows that two‐patch systems have some unique properties. One qualitative difference is that in many cases it will be possible for a Bayesian forager to gain perfect information about patch quality. As soon as it has found more than the number of prey items that should be available in a poor patch, it “knows” that it is in a rich patch. The model generates at least three testable predictions. 1) The distribution of giving‐up densities, GUDs, should be bimodal in rich patches, when rich patches are rare in the environment. This is because the optimal strategy is then devoted to using the poor patches correctly, at the expense of missing a large fraction of the few rich patches available. 2) There should be a negative relation between GUD and search time in poor patches, when rich patches are much more valuable than poor. This is because the forager gets good news about potential patch quality from finding some food. It therefore accepts a lower instantaneous intake rate, making it more resistant against runs of bad luck, decreasing the risk of discarding rich patches. 3) When the energy gains required to remain in the patch are high (such as under high predation risk), the overuse of poor patches and the underuse of rich increases. This is because less information about patch quality is gained if leaving at high intake rates (after short times). The predictions given by this model may provide a much needed and effective conceptual framework for testing (both in the lab and the field) whether animals are using Bayesian assessment.     

A game theoretical model of kleptoparasitism with incomplete information

Kleptoparasitism, the stealing of food from one animal by another, is a common natural phenomenon that has been modelled mathematically in a number of ways. The handling process of food items can take some time and the value of such items can vary depending upon how much handling an item has received. Furthermore this information may be known to the handler but not the potential challenger, so there is an asymmetry between the information possessed by the two competitors. We use game-theoretic methods to investigate the consequences of this asymmetry for continuously consumed food items, depending upon various natural parameters. A variety of solutions are found, and there are complex situations where three possible solutions can occur for the same set of parameters. It is also possible to have situations which involve members of the population exhibiting different behaviours from each other. We find that the asymmetry of information often appears to favour the challenger, despite the fact that it possesses less information than the challenged individual. The research was supported by EPSRC grant EP/E043402/1 and NSF 0634182.     

Species difference in adaptive use of public information in sticklebacks

Animals foraging on variable food sources can refine their estimates of patch quality by monitoring the success of others (i.e. collect 'public information'). Here, we show that both three-spined sticklebacks (Gasterosteus aculeatus) and nine-spined sticklebacks (Pungitius pungitius) use past cues provided by others to locate food but only nine-spined sticklebacks use prior public information to assess patch quality, regardless of whether demonstrators were conspecifics or heterospecifics. Moreover, nine-spined but not three-spined sticklebacks preferentially hid in vegetation during the demonstration, a position from which they could observe both patches simultaneously and collect public information. We conclude that species differences in the use of public information can be explained by variations in habitat choice and response to predation. Our findings expand current understanding of the scope of public-information use in animals by showing that fishes can use public-information in a foraging context and from heterospecifics. The study suggests that public-information use is an adaptation that allows animals vulnerable to predation to acquire valuable foraging information at low risk.     

Comparative experimental studies on the feeding of chaetognaths and ctenophores

There is very little quantitative experimental information onfeeding in chaetognaths and ctenophores because of the difficultyof obtaining undamaged specimens. As important plankton predators,a knowledge of their daily rations as a function of food concentrationis vital to ecosystem models of water column trophodynamics.At specified food concentrations, three ctenophore species alwaysingested higher specific daily rations than three species ofchaetognaths. Food consumption of ctenophores increased linearlythroughout the food concentration range (exceeding 10⁶ µgC/m³ Although food consumption of chaetognaths reached an upperlimit at very high food concentrations, it seemed unlikely thatsatiation would occur under environmental conditions. Below10,000 food items/m³ chaetognaths were unable to obtain l% oftheir specific daily ration, although the few environmentalstudies based on gut contents suggest substantial feeding evenbelow this concentration. The apparent threshold may reflectthe environmental patch density necessary for feeding to begin.At high food concentrations daily rations of newly-hatched ctenophoresand chaetognaths can exceed well over 10 times their body carbon.In ctenophores even adults can consume these extremely largerations although chaetognaths follow the more usual pattern,where ration decreases throughout life. Gut residence time offood (where single items were ingested) increased with age andsize of food item selected in Sagitta hispida from one to twohours, levelling off after animals reached 6 mm. Mnemiopsismccradyi maintained a uniform digestion time of one hour asit grew, except for a brief period when it first selected larger-sizedfood. In older animals of both species, multiple ingestion becamemore common, where gut residence time was directly proportionalto number of food items consumed, ranging up to 5 hours forfive or more food items. Chaetognaths showed more variabilityof gut residence time than did ctenophores. It was concludedthat these animals are much better at catching food in the environmentthan laboratory experiments would suggest.     

Titrating the cost of plant toxins against predators: determining the tipping point for foraging herbivores

1. Foraging herbivores must deal with plant characteristics that inhibit feeding and they must avoid being eaten. Principally, toxins limit food intake, while predation risk alters how long animals are prepared to harvest resources. Each of these factors strongly affects how herbivores use food patches, and both constraints can pose immediate proximate costs and long-term consequences to fitness. 2. Using a generalist mammalian herbivore, the common brushtail possum (Trichosurus vulpecula), our aim was to quantitatively compare the influence of plant toxin and predation risk on foraging decisions. 3. We performed a titration experiment by offering animals a choice between non-toxic food at a risky patch paired with food with one of five toxin concentrations at a safe patch. This allowed us to identify the tipping point, where the cost of toxin in the safe food patch was equivalent to the perceived predation risk in the alternative patch. 4. At low toxin concentration, animals ate more from the safe than the risky patch. As toxin concentration increased at the safe patch, intake shifted until animals ate mainly from the risky patch. This shift was associated with behavioural changes: animals spent more time and fed longer at the risky patch, while vigilance increased at both risky and safe patches. 5. Our results demonstrate that the variation in toxin concentration, which occurs intraspecifically among plants, can critically influence the relative cost of predation risk on foraging. We show that herbivores quantify, compare and balance these two different but proximate costs, altering their foraging patterns in the process. This has potential ecological and evolutionary implications for the production of plant defence compounds in relation to spatial variation in predation risk to herbivores.     

Bayes' theorem and its applications in animal behaviour

Bayesian decision theory can be used to model animal behaviour. In this paper we give an overview of the theoretical concepts in such models. We also review the biological contexts in which Bayesian models have been applied, and outline some directions where future studies would be useful. Bayesian decision theory, when applied to animal behaviour, is based on the assumption that the individual has some sort of "prior opinion" of the possible states of the world. This may, for example, be a previously experienced distribution of qualities of food patches, or qualities of potential mates. The animal is then assumed to be able use sampling information to arrive at a "posterior opinion", concerning e.g. the quality of a given food patch, or the average qualities of mates in a year. A correctly formulated Bayesian model predicts how animals may combine previous experience with sampling information to make optimal decisions. We argue that the assumption that animals may have "prior opinions" is reasonable. Their priors may come from one or both of two sources: either from their own individual experience, gained while sampling the environment, or from an adaptation to the environment experienced by previous generations. This means that we should often expect to see "Bayesian-like" decision-making in nature.     

A simple model of information use in the exploitation of patchily distributed food

In this paper we consider a simple model of an environment in which prey are distributed in patches. It is assumed that each patch contains at most one item, but items may vary in the ease with which they can be found. The time spent in unsuccessful search on a patch gives information of whether a patch contains an item, and if it does, how difficult that item is to find. We show how this information can be used to find the policy which maximizes the mean rate of reward for the environment. The analysis is illustrated by two examples.     

An evolutionarily stable joining policy for group foragers

For foragers that exploit patchily distributed resources thatare challenging to locate, detecting discoveries made by otherswith a view to joining them and sharing the patch may oftenbe an attractive tactic, and such behavior has been observedacross many taxa. If, as will commonly be true, the time takento join another individual on a patch increases with the distanceto that patch, then we would expect foragers to be selectivein accepting joining opportunities: preferentially joining nearbydiscoveries. If competition occurs on patches, then the profitabilityof joining (and of not joining) will be influenced by the strategiesadopted by others. Here we present a series of models designedto illuminate the evolutionarily stable joining strategy. Weconfirm rigorously the previous suggestion that there shouldbe a critical joining distance, with all joining opportunitieswithin that distance being accepted and all others being declined.Further, we predict that this distance should be unaffectedby the total availability of food in the environment, but shouldincrease with decreasing density of other foragers, increasingspeed of movement towards joining opportunities, increased difficultyin finding undiscovered food patches, and decreasing speed withwhich discovered patches can be harvested. We are further ableto make predictions as to how fully discovered patches shouldbe exploited before being abandoned as unprofitable, with discoveredpatches being more heavily exploited when patches are hard tofind: patches can be searched for remaining food more quickly,forager density is low, and foragers are relatively slow intraveling to discovered patches.     

Time's crooked arrow: optimal foraging and rate-biased time perception

Time perception is critical to animal behaviours requiring anticipation of future events based on present information about the environment. Most models of animal foraging assume that animals are capable of measuring absolute time despite evidence that animals measure time with predictable biases in mean and variance. We incorporate the evidence for a rate-biased subjective time perception into a classic model of optimal foraging, the marginal value theorem. If acceleration of the clock rate is proportional to food intake rate and time is perceived similarly when in transit between patches as it is while waiting in a patch following eating, organisms are predicted to follow the predictions of the marginal value theorem exactly. However, a nonlinear relationship between clock rate and food intake rate, unequal wait and transit time perception, or any lag in the clock predicts characteristic suboptimal behaviour. We discuss how this mechanism for suboptimal behaviour compares with others in the literature and how it can be recognized in experiments. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

People's study time allocation and its relation to animal foraging

In this article we suggest a relation between people's metacognitively guided study time allocation strategies and animal foraging. These two domains are similar insofar as people use specific metacognitive cues to assist their study time allocation just as other species use cues, such as scent marking. People decline to study items that they know they already know, just as other species use a win-shift strategy - avoiding already visited and depleted patches - in foraging. People selectively study the easiest as-yet-unlearned items first, before turning to more difficult items just as other species take the ‘just right’ size and challenge of prey—the so-called Goldilocks principle. People use a stop rule by which they give up on one item and turn to another when the returns diminish just as others species use a stop rule that guides shifting from one patch to another. The value that each item is assigned on the criterion test, if known during study, influences which items people choose to study and how long they study them just as knowledge of the nutritional or energy value of the food influences choices and perseverance in foraging. Finally, study time allocation strategies can differ in their effectiveness depending upon the expertise of the student just as some species forage close to optimally while others do not.     

The asymmetric war of attrition

The paper, which has an informal discussion at the end, provides a game theoretical analysis of the asymmetric “war of attrition” with incomplete information. This is a contest where animals adopt different roles like “owner” and “intruder” in a territorial conflict, and where the winner is the individual prepared to persist longer. The term incomplete information refers to mistakes in the identification of roles. The idea by Parker & Rubenstein (1981) is mathematically worked out and confirmed that there exists only a single evolutionarily stable strategy (ESS) for the model with a continuum of possible levels of persistence and no discontinuities in the increase of cost during attrition. The ESS prescribes to settle the conflict according to “who has more to gain or less to pay for persistence”. The only evolutionarily stable convention is thus to give the player access to the resource who has the role which is favoured with respect to payoffs. By contrast, it was shown earlier (Hammerstein, 1981) for various asymmetric versions of the “Hawks-Doves” model that an ESS can exist which appears paradoxical with respect to payoffs. The nature of this contrast is further analyzed by introducing elements of discreteness in the asymmetric war of attrition. It turns out that some conditions must be satisfied in order to have the possibility of an alternative ESS which is not of the above simple commonsense type. First, a decision to persist (or escalate) further in a contest must typically commit a contestant to go on fighting for a full “round”, before he can give up without danger. Second, such a “discontinuity” must occur at a level of persistence where the contest is still cheap, and, finally, errors in the identification of roles must be rare.     

Prior knowledge about spatial pattern affects patch assessment rather than movement between patches in tactile-feeding mallard

1. Heterogeneity in food abundance allows a forager to concentrate foraging effort in patches that are rich in food. This might be problematic when food is cryptic, as the content of patches is unknown prior to foraging. In such case knowledge about the spatial pattern in the distribution of food might be beneficial as this enables a forager to estimate the content of surrounding patches. A forager can benefit from this pre-harvest information about the food distribution by regulating time in patches and/or movement between patches. 2. We conducted an experiment with mallard Anas platyrhynchos foraging in environments with random, regular, and clumped spatial configurations of full and empty patches. An assessment model was used to predict the time in patches for different spatial distributions, in which a mallard is predicted to remain in a patch until its potential intake rate drops to the average intake rate that can be achieved in the environment. A movement model was used to predict lengths of interpatch movements for different spatial distributions, in which a mallard is predicted to travel to the patch where it expects the highest intake rate. 3. Consistent with predictions, in the clumped distribution mallard spent less time in an empty patch when the previously visited neighbouring patch had been empty than when it had been full. This effect was not observed for the random distribution. This shows that mallard use pre-harvest information on spatial pattern to improve patch assessment. Patch assessment could not be evaluated for the regular distribution. 4. Movements that started in an empty patch were longer than movements that started in a full patch. Contrary to model predictions this effect was observed for all spatial distributions, rather than for the clumped distribution only. In this experiment mallard did not regulate movement in relation to pattern. 5. An explanation for the result that pre-harvest information on spatial pattern affected patch assessment rather than movement is that mallard move to the nearest patch where the expected intake rate is higher than the critical value, rather than to the patch where the highest intake rate is expected.     

Learning to forage in a regenerating patchy environment: Can it fail to be optimal?

The behaviour of animals foraging along closed traplines of regenerating patches of food has been simulated using a learning rule that determines when an animal should leave the patch at which it is currently feeding to search for another one. The rule causes the animal to stay at the patch as long as it is feeding faster than it remembers doing. The foraging behaviour of one animal, and of two or more animals together, feeding in traplines containing patches of the same and of differing types has been simulated, and in all cases the foraging behaviour generated by the rule allowed the animals to exploit the food very efficiently. The learning model is also responsible for indirect social interactions among animals sharing the same trapline because the feeding of each animal reduces the availability of food for the others. This causes a population of animals to disperse themselves, on average, among patches of food according to the ideal free distribution. The relationship between the learning model and conventional optimal foraging models is examined and it is shown that it is pointless to try to account for learned behaviour in the context of optimal foraging theory.     

Optimal movement between patches under incomplete information about the spatial distribution of food items

If the food distribution contains spatial pattern, the food density in a particular patch provides a forager with information about nearby patches. Foragers might use this information to exploit patchily distributed resources profitably. We model the decision on how far to move to the next patch in linear environments with different spatial patterns in the food distribution (clumped, random, and regular) for foragers that differ in their degree of information. An ignorant forager is uninformed and therefore always moves to the nearest patch (be it empty or filled). In contrast, a prescient forager is fully informed and only exploits filled patches, skipping all empty patches. A Bayesian assessor has prior knowledge about the content of patches (i.e. it knows the characteristics of the spatial pattern) and may skip neighbouring patches accordingly by moving to the patch where the highest gain rate is expected. In most clumped and regular distributions there is a benefit of assessment, i.e. Bayesian assessors achieve substantially higher long-term gain rates than ignorant foragers. However, this is not the case in distributions with less strong spatial pattern, despite the fact that there is a large potential benefit from a sophisticated movement rule (i.e. a large penalty of ignorance). Bayesian assessors do also not achieve substantially higher gain rates in environments that are relatively rich or poor in food. These results underline that an incompletely informed forager that is sensitive to spatial pattern should not always respond to existing pattern. Furthermore, we show that an assessing forager can enhance its long-term gain rate in highly clumped and some specific near-regular food distributions, by sampling the environment in slightly larger spatial units.     

Optimal movement between patches under incomplete information about the spatial distribution of food items

If the food distribution contains spatial pattern, the food density in a particular patch provides a forager with information about nearby patches. Foragers might use this information to exploit patchily distributed resources profitably. We model the decision on how far to move to the next patch in linear environments with different spatial patterns in the food distribution (clumped, random, and regular) for foragers that differ in their degree of information. An ignorant forager is uninformed and therefore always moves to the nearest patch (be it empty or filled). In contrast, a prescient forager is fully informed and only exploits filled patches, skipping all empty patches. A Bayesian assessor has prior knowledge about the content of patches (i.e. it knows the characteristics of the spatial pattern) and may skip neighbouring patches accordingly by moving to the patch where the highest gain rate is expected. In most clumped and regular distributions there is a benefit of assessment, i.e. Bayesian assessors achieve substantially higher long-term gain rates than ignorant foragers. However, this is not the case in distributions with less strong spatial pattern, despite the fact that there is a large potential benefit from a sophisticated movement rule (i.e. a large penalty of ignorance). Bayesian assessors do also not achieve substantially higher gain rates in environments that are relatively rich or poor in food. These results underline that an incompletely informed forager that is sensitive to spatial pattern should not always respond to existing pattern. Furthermore, we show that an assessing forager can enhance its long-term gain rate in highly clumped and some specific near-regular food distributions, by sampling the environment in slightly larger spatial units.     

Development of a picture‐scale‐questionnaire to assess liking for fatty,salty, sweet,and umami seasonings compared to a sensory evaluation for Japanese consumers

PrefQuest is a web‐based questionnaire that measures the liking for sweet, fatty‐sweet, salty, and fatty‐salty sensations using combinations of images of French food items. We adapted the original PrefQuest questionnaire for Japanese respondents (J‐PrefQuest). J‐PrefQuest comprises 17 Japanese food items categorized into four sensations: fatty, salty, sweet, and umami. Participants responded by indicating their preferred level of seasoning for each food item on a 6‐point scale. Segmentation of 161 respondents identified groups who preferred: (a) high amount of umami seasonings; (b) low amount of umami seasonings; (c) high amount of fatty and low amount of salty seasonings. To validate the questionnaire, 70 female university students were recruited for a sensory evaluation of five levels of seasoning used for six food items on a 9‐point hedonic scale before completing the questionnaire. The validity of J‐PrefQuest questionnaire was then analyzed by comparing the two results.

Practical applications

Measuring personal preference or “liking” for the taste of food items is important as such information may contribute to the prevention of various diseases and improve the quality of life of consumers. We developed a simple questionnaire with pictures to evaluate the liking for fatty, salty, sweet, and umami sensations of Japanese respondents. This questionnaire focuses on typical seasonings added to different Japanese food items and provides objective information about liking different food sensations. Thus, it has application as an important screening tool to identify respondents who may need to make their food consumption behavior more healthy. Moreover, following minor modifications, this questionnaire could also be used in other Asian countries where plain white rice is the stable food.     

动物觅食行为对捕食风险的反应

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

The conflict between optimisation and orderliness in building behaviour of the caddis fly,Chaetopteryx villosa Fabr.(Limnephilidae: Trichoptera), larvae

The Optimal Foraging Theory predict that an animal should restrict its searching activity to those patches of environment, where the ratio of gain to time and energy spent is maximal. Experimenters usually verify the prediction against the null hypothesis of random activity distribution between patches. As animals always prefer profitable patches to some degree, experimental results can always be interpreted as confirmation of the theory. In opposite to this approach, we put forward the "regularity hypothesis". According to this hypothesis, the finding of a profitable item in some patch makes an animal to stay and test more items within the patch. The readiness to test decreases if the profitability of these items is low, and the animal eventually leaves the patch. It also searches in other patches less carefully. As a result, positive and negative responses to items are repeated by ordered series. In general, this regularity of responses helps animals to choose profitable patches. However, animals may also ignore some profitable patches because of the regularity, so that the behavioural optimazation fails. The regularity hypothesis have been tested in experiment with the search of building material and patch choice by caddisfly larvae. The first of two experimental patches contained egg shell fragments (profitable building material) mixed with a sand (unprofitable). The second patch contained sand only. Larva stayed within the first patch after a shell had been found, so that the probability to find more shells increased. However, larvae started walking after they had found several sand particles. Once starting to walk, they found new shell fragments and tested them, but tended to reject them and eventually leave the patch. Moreover, upon returning to the first patch, larvae also might reject fragments and leave the patch again. Rejections are accounted for by the fact that the duration of testing was too short to identify fragments correctly. As a result, negative responses were repeated to a certain degree independently of the profitability of building material, and the patch choice was not optimal. These results are agreed with the regularity hypothesis. It is argued that the hypothesis can be used as an alternative to the Optimal Foraging Theory.     

Self-Improvement for Team-Players: The Effects of Individual Effort on Aggregated Group Information

By putting effort into behaviours like foraging or scanning for predators, an animal can improve the correctness of its personal information about the environment. For animals living in groups, the individual can gain further information if it is able to assess public information about the environment from other group members. Earlier work has shown that consensus group decisions based upon the public information available within the group are more likely to be correct than decisions based upon personal information alone, given that each individual in a group has a fixed probability of being correct. This study develops a model where group members are able to improve their personal likelihood of making a correct decision by conducting some level of (costly) effort. I demonstrate that there is an evolutionarily stable level of effort for all the individuals within the group, and the effort made by an individual should decrease with increasing group size. The relevance of these results to social decision making is discussed: in particular, these results are similar to standard theoretical predictions about the amount of vigilance shown by individuals decreasing with increasing group size. However, this model suggests that these results could come about where individuals are coordinating their effort within the group (unlike standard models, which assume that all individual effort is independent of the actions of others). This ties in with experimental findings where individuals have been shown to monitor the efforts of others.