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.     

Do Pied Flycatchers Use Personal or Social Information for Replacement Clutch Decisions? A Field Experiment

Life is a continuous stream of decisions: organisms are repeatedly confronted with multiple options from which to choose. To make the best decisions and maximise their fitness, individuals need to collect information: from direct interactions with the environment (personal information) or from observing other individuals interacting with the environment (social information). Yet, very little is known about the details of decision‐making by animals in their natural environments, including the sources of information they use in the process. We set out to determine the relative importance of social and personal information in breeding habitat decisions in the migratory pied flycatcher, Ficedula hypoleuca. In a field experiment in Finland, we manipulated flycatcher breeding success and then allowed individuals to choose among breeding sites labelled with artificial cues (arbitrary symbols) associated with their own (manipulated) and neighbour's reproductive success, respectively, for laying a replacement clutch. Birds did not appear to distinguish among the symbols representing their own, their neighbour's or a neutral symbol, indicating that they did not use either information source when deciding about a replacement clutch. Also, the choice did not influence subsequent investment into breeding (clutch size). This may suggest that pied flycatchers utilise other cues (e.g. breeding success or predation risk on larger scales) when having to make a decision about replacement clutches. We discuss our results in light of constraints imposed by a short breeding season of migratory birds.     

Swarm Intelligence in Animal Groups: When Can a Collective Out-Perform an Expert?

An important potential advantage of group-living that has been mostly neglected by life scientists is that individuals in animal groups may cope more effectively with unfamiliar situations. Social interaction can provide a solution to a cognitive problem that is not available to single individuals via two potential mechanisms: (i) individuals can aggregate information, thus augmenting their 'collective cognition', or (ii) interaction with conspecifics can allow individuals to follow specific 'leaders', those experts with information particularly relevant to the decision at hand. However, a-priori, theory-based expectations about which of these decision rules should be preferred are lacking. Using a set of simple models, we present theoretical conditions (involving group size, and diversity of individual information) under which groups should aggregate information, or follow an expert, when faced with a binary choice. We found that, in single-shot decisions, experts are almost always more accurate than the collective across a range of conditions. However, for repeated decisions - where individuals are able to consider the success of previous decision outcomes - the collective's aggregated information is almost always superior. The results improve our understanding of how social animals may process information and make decisions when accuracy is a key component of individual fitness, and provide a solid theoretical framework for future experimental tests where group size, diversity of individual information, and the repeatability of decisions can be measured and manipulated.     

Frequency-dependent payoffs and sequential decision-making favour consistent tactic use

Although natural selection should have favoured individuals capable of adjusting the weight they give to personal and social information according to circumstances, individuals generally differ consistently in their individual weighting of both types of information. Such individual differences are correlated with personality traits, suggesting that personality could directly affect individuals' ability to collect personal or social information. Alternatively, the link between personality and information use could simply emerge as a by-product of the sequential decision-making process in a frequency-dependent context. Indeed, when the gains associated with behavioural options depend on the choices of others, an individual's sequence of arrival could constrain its choice of options leading to the emergence of correlated behaviours. Any factor such as personality that affects decision order could thus be correlated with information use. To test this new explanation, we developed an individual-based model that simulates a group of animals engaged in a game of sequential frequency-dependent decision: a producer-scrounger game. Our results confirm that the sequence of decision, in this case enforced by the order in which animals enter a foraging area, consistently influences their mean tactic use and their individual plasticity, an outcome reminiscent of the correlation reported between personality and social information use.     

Exploring the costs and benefits of social information use: an appraisal of current experimental evidence

Research on social learning has focused traditionally on whether animals possess the cognitive ability to learn novel motor patterns from tutors. More recently, social learning has included the use of others as sources of inadvertent social information. This type of social learning seems more taxonomically widespread and its use can more readily be approached as an economic decision. Social sampling information, however, can be tricky to use and calls for a more lucid appraisal of its costs. In this four-part review, we address these costs. Firstly, we address the possibility that only a fraction of group members are actually providing social information at any one time. Secondly, we review experimental research which shows that animals are circumspect about social information use. Thirdly, we consider the cases where social information can lead to incorrect decisions and finally, we review studies investigating the effect of social information quality. We address the possibility that using social information or not is not a binary decision and present results of a study showing that nutmeg mannikins combine both sources of information, a condition that can lead to the establishment of informational cascades. We discuss the importance of empirically investigating the economics of social information use.     

The higher the better: sentinel height influences foraging success in a social bird

In all social species, information relevant to survival and reproduction can be obtained in two main ways: through personal interaction with the environment (i.e. ‘personal’ information) and from the performance of others (i.e. ‘public’ information). While public information is less costly to obtain than personal information, it may be inappropriate or inaccurate. When deciding how much to rely on public information, individuals should therefore assess its potential quality, but this possibility requires empirical testing in animals. Here, we use the sentinel system of cooperatively breeding pied babblers (Turdoides bicolor) to investigate how behavioural decisions of foragers are influenced by potential variation in the quality of anti-predator information from a vigilant groupmate. When sentinels moved to a higher position, from where their probability of detecting predators is likely to be greater, foragers reduced their vigilance, spread out more widely and were more likely to venture into the open. Consequently, they spent more time foraging and increased their foraging efficiency, resulting in a profound increase in biomass intake rate. The opposite behavioural changes, and consequent foraging outcomes, were found when sentinels moved lower. A playback experiment demonstrated that foragers can use vocal cues alone to assess sentinel height. This is the first study to link explicitly a measure of the potential quality of public information with a fitness measure from those relying on the information, and our results emphasize that a full understanding of the evolution of communication in complex societies requires consideration of the reliability of information.     

Individuals in foraging groups may use vocal cues when assessing their need for anti-predator vigilance

Many studies of social species have reported variation in the anti-predator vigilance behaviour of foraging individuals depending on the presence and relative position of other group members. However, little attention has focused on how foragers assess these variables. It is commonly assumed that they do so visually, but many social species produce frequent calls while foraging, and these 'close' calls might provide valuable spatial information. Here, we show that foraging pied babblers (Turdoides bicolor) are less vigilant when in larger groups, in the centre of a group and in closer proximity to another group member. We then show that foragers are less vigilant during playbacks of close calling by more individuals and individuals on either side of them when compared with calls of fewer individuals and calls on one side of them. These results suggest that foragers can use vocal cues to gain information on group size and their spatial position within a group. Future studies of anti-predator vigilance should consider the relative importance of both visual and vocal monitoring of group members.     

Wood warblers copy settlement decisions of poor quality conspecifics: support for the tradeoff between the benefit of social information use and competition avoidance

Social information use in songbird habitat selection commonly involves a conspecific attraction strategy. Individuals copy the breeding‐site choices of conspecifics, that is, bias their own settlement decisions towards sites (tracts of spatially limited habitat with similar structure) already occupied by others. In order to be adaptive, social information use has to be discriminative. Especially the decisions of good quality individuals, i.e. measuring high at observable fitness correlates, should be copied more frequently than those of poor quality individuals. It is unknown, however, whether songbirds discriminatively use conspecific presence by evaluating the quality of information providers in habitat selection. We experimentally tested whether wood warblers Phylloscopus sibilatrix selectively copied settlement decisions of conspecifics in relation to the quality of observed individuals. We also tested whether the use of social cues was influenced by the population density at a particular site in the preceding year. We found that wood warblers selectively used intraspecific social information, but in a pattern opposite to that expected based on existing hypotheses. Wood warblers copied breeding‐site choices of poor quality conspecifics and despite temporary attraction to sites where the presence of good quality individuals was simulated, they did not ultimately settle near these individuals. Population density in the preceding year did not influence settlement patterns. We argue that when making settlement decisions, wood warblers assessed the expected level of local intraspecific competition and selectively copied breeding‐site choices of conspecifics or refused to settle, depending on competitive abilities of observed individuals. This adds a novel aspect to the patterns and processes of social information use proposed thus far, and provides support for the predicted negative effect of intraspecific competition on benefit of information. Moreover, it seems that habitat selection in wood warblers is a complex decision‐making process, in which initial decisions are adjusted after acquiring more accurate information. Synthesis Social information use in songbird habitat selection commonly involves copying the breeding‐site choices of conspecifics (so‐called conspecific attraction). To be adaptive, this strategy has to be discriminative, but almost no empirical studies have tested this assertion. Our study shows that birds may selectively use social information by copying settlement decisions of poor quality conspecifics, but avoid settling near good quality individuals, likely because of their high competitive abilities. This decision‐making pattern supports the predicted, yet not experimentally tested, tradeoff between information value and cost of competition in social information use. Our study highlights also that the use of social cues in settlement decisions may be both positively and negatively biased.     

Dynamic choices are most accurate in small groups

According to the classic results of Galton and Condorcet, as well as in modern decision-making models, accuracy in groups increases with group size. However, these studies do not consider the naturally occurring situation in which individuals dynamically re-evaluate their decision with a possible change of opinion. The dynamics of re-evaluation in groups are very different to individual re-evaluation because individuals influence the group and the group influences the individual. We find that individual accuracy in a group is higher when individuals re-evaluate because all members have more access to social information, while in single decisions, those deciding first have less. This improvement is smaller in large groups as in this case errors can cascade across the members of the group before re-evaluation can correct them. The net result is a maximal accuracy at a small group size. We also analyzed the case in which individuals are influenced only by a small number of the other individuals. In this case, cascading errors affect the interacting subgroups but are very unlikely to reach the whole group. This results in a local optimum at a small group size but also an optimum at a very large size. We thus suggest that re-evaluation dynamics can make small and very large groups optimal. Also, features that may be seen as limitations, like an influence from only a small number of individuals, may turn to be beneficial when considering local animal interactions, here filtering out cascading of errors in the group when reconsideration dynamics takes place.     

Motion-guided attention promotes adaptive communications during social navigation

Animals are capable of enhanced decision making through cooperation, whereby accurate decisions can occur quickly through decentralized consensus. These interactions often depend upon reliable social cues, which can result in highly coordinated activities in uncertain environments. Yet information within a crowd may be lost in translation, generating confusion and enhancing individual risk. As quantitative data detailing animal social interactions accumulate, the mechanisms enabling individuals to rapidly and accurately process competing social cues remain unresolved. Here, we model how motion-guided attention influences the exchange of visual information during social navigation. We also compare the performance of this mechanism to the hypothesis that robust social coordination requires individuals to numerically limit their attention to a set of n-nearest neighbours. While we find that such numerically limited attention does not generate robust social navigation across ecological contexts, several notable qualities arise from selective attention to motion cues. First, individuals can instantly become a local information hub when startled into action, without requiring changes in neighbour attention level. Second, individuals can circumvent speed–accuracy trade-offs by tuning their motion thresholds. In turn, these properties enable groups to collectively dampen or amplify social information. Lastly, the minority required to sway a group''s short-term directional decisions can change substantially with social context. Our findings suggest that motion-guided attention is a fundamental and efficient mechanism underlying collaborative decision making during social navigation.     

Variation in social information use: the influences of information reliability and mass on decision making in a group‐living fish Gambusia affinis

The effect of the reliability of available social information was assessed by examining whether the age of social information changes its effects on a foraging decision in a group‐living fish Gambusia affinis. Individuals switched their patch preference when faced with social information that conflicted with personal information in general; the age of the social information, however, did not significantly influence preference for feeding patch. The mass of decision makers was positively correlated with their use of available social information, with heavier individuals exhibiting a greater difference in patch preference than lighter individuals, suggesting that large and small G. affinis trade‐off the benefits of information acquisition and the costs of competition from conspecifics differently.     

The role of conformity in foraging when personal and social information conflict

There is currently considerable interest in the interplay betweenpersonal and social information in decision-making processes.Two experiments are presented exploring the relative use ofprior personal information and subsequent social informationin foraging decisions of guppies. Experiment 1 tested the assumptionthat when the use of information acquired through personal experienceis not costly, conflicting social information will be ignored.The assumption was confirmed because, when given a choice betweenfeeding at two food patches, at one of which they had previouslyseen conspecifics feed, individual fish with prior experienceof feeding at the alternative site chose the alternative, whereasfish with no prior experience chose the site at which theirconspecifics had fed. Experiment 2 tested theoretical predictionsthat when the use of information acquired through personal experienceis potentially costly, conflicting social information will beweighed more heavily than will personal information. The predictionwas confirmed because, when given a choice between feeding attwo food patches, one at which they had previously seen conspecificsfeed and one behind a visual barrier, individual fish with priorexperience of feeding behind the barrier chose the site at whichtheir conspecifics had fed. These findings suggest that conformitycan promote social learning in naïve individuals, but priorexperience can insulate individuals from conformity providedthe costs of relying on that experience are small. In addition,the experiments highlight the fact that personal and socialinformation are not always weighed equally.     

Democracy in animals: the evolution of shared group decisions

A 'consensus decision' is when the members of a group choose, collectively, between mutually exclusive actions. In humans, consensus decisions are often made democratically or in an 'equally shared' manner, i.e. all group members contribute to the decision. Biologists are only now realizing that shared consensus decisions also occur in social animals (other than eusocial insects). Sharing of decisions is, in principle, more profitable for groups than accepting the 'unshared' decision of a single dominant member. However, this is not true for all individual group members, posing a question as to how shared decision making could evolve. Here, we use a game theory model to show that sharing of decisions can evolve under a wide range of circumstances but especially in the following ones: when groups are heterogeneous in composition; when alternative decision outcomes differ in potential costs and these costs are large; when grouping benefits are marginal; or when groups are close to, or above, optimal size. Since these conditions are common in nature, it is easy to see how mechanisms for shared decision making could have arisen in a wide range of species, including early human ancestors.     

Spying on your neighbours? Social information affects timing of breeding and stress hormone levels in a colonial seabird

A good overlap between offspring energetic requirements and availability of resources is required for successful reproduction. Accordingly, individuals from numerous species fine-tune their timing of breeding by integrating cues that predict environmental conditions during the offspring period. Besides acquiring information from their direct interaction with the environment (personal information), individuals can integrate information by observing the behaviours or performance of others (social information). The use of social information is often beneficial because the accumulated knowledge of conspecifics may represent a source of information more reliable than the intrinsically more limited personal information. However, although social information constitutes the major source of information in a wide range of contexts, studies investigating its use in the context of timing of breeding are scarce. We investigated whether black-legged kittiwakes (Rissa tridactyla) used social information to adjust the timing of egg-laying. We manipulated social information using a food-supplementation experiment, known to advance kittiwakes' reproductive phenology. We expected food-supplemented and unsupplemented pairs to delay and advance, respectively, their timing of laying when surrounded by a majority of neighbours from the opposite food-treatment. However, both unsupplemented and food-supplemented kittiwakes delayed egg-laying when surrounded by a higher proportion of neighbours from the opposite food-treatment. This result shows that kittiwakes use social information to time egg-laying, but that it is not used to match the seasonal peak of food availability. We suggest that when social and personal cues give contradictory environmental information, individuals may benefit from delaying laying to gather more information to make better decisions about investment into eggs. Further, we explored a potential proximate mechanism for the pattern we report. We show that baseline corticosterone, known to mediate reproductive decisions, was lower in unsupplemented females facing a higher proportion of food-supplemented neighbours. Altogether, our results suggest that to fine-tune their timing of laying, kittiwakes use complex decision-making processes in which social and personal information interplay.

     

Personal experience and reputation interact in human decisions to help reciprocally

There is ample evidence that human cooperative behaviour towards other individuals is often conditioned on information about previous interactions. This information derives both from personal experience (direct reciprocity) and from experience of others (i.e. reputation; indirect reciprocity). Direct and indirect reciprocity have been studied separately, but humans often have access to both types of information. Here, we experimentally investigate information use in a repeated helping game. When acting as donor, subjects can condition their decisions to help recipients with both types of information at a small cost to access such information. We find that information from direct interactions weighs more heavily in decisions to help, and participants tend to react less forgivingly to negative personal experience than to negative reputation. Moreover, effects of personal experience and reputation interact in decisions to help. If a recipient''s reputation is positive, the personal experience of the donor has a weak effect on the decision to help, and vice versa. Yet if the two types of information indicate conflicting signatures of helpfulness, most decisions to help follow personal experience. To understand the roles of direct and indirect reciprocity in human cooperation, they should be studied in concert, not in isolation.     

Nine-spined sticklebacks exploit the most reliable source when public and private information conflict

Social foragers can potentially use private information gained from personal experience and public information gained from observing the foraging success of others to determine the profitability of a foraging patch. We investigated how nine-spined sticklebacks use conflicting public and private information of variable reliability to make foraging decisions. In a first experiment, when private information was reliable, sticklebacks ignored public information and based their foraging decision on private information. However, when private information was less reliable, sticklebacks tended to use public rather than private information. A second experiment investigated how the time since experiencing private information affected sticklebacks' use of this information when it conflicted with recent public information. Fish based their foraging decisions on recently acquired private information, but reliance on private information diminished as the period since experiencing it increased. Fish used public information if 7 days had elapsed since updating their private information. Our findings suggest that nine-spined sticklebacks flexibly adjust their decision making to exploit the most reliable information available, be it public or private, and that animals will weight private and public information appropriately depending on circumstances.     

Potential disadvantages of using socially acquired information

The acquisition and use of socially acquired information is commonly assumed to be profitable. We challenge this assumption by exploring hypothetical scenarios where the use of such information either provides no benefit or can actually be costly. First, we show that the level of incompatibility between the acquisition of personal and socially acquired information will directly affect the extent to which the use of socially acquired information can be profitable. When these two sources of information cannot be acquired simultaneously, there may be no benefit to socially acquired information. Second, we assume that a solitary individual's behavioural decisions will be based on cues revealed by its own interactions with the environment. However, in many cases, for social animals the only socially acquired information available to individuals is the behavioural actions of others that expose their decisions, rather than the cues on which these decisions were based. We argue that in such a situation the use of socially acquired information can lead to informational cascades that sometimes result in sub-optimal behaviour. From this theory of informational cascades, we predict that when erroneous cascades are costly, individuals should pay attention only to socially generated cues and not behavioural decisions. We suggest three scenarios that might be examples of informational cascades in nature.     

Group decision making in fission-fusion societies: evidence from two-field experiments in Bechstein's bats

Group decisions are required when group coordination is beneficial, but individuals can choose between alternatives. Despite the increased interest in animal group decision making, there is a lack of experimental field studies that investigate how animals with conflicting information make group decisions. In particular, no field studies have considered the influence of fission-fusion behaviour (temporary splitting into subgroups) on group decisions. We studied group decision making in two wild Bechstein's bat colonies, which are fission-fusion societies of stable individual composition. Since they frequently switch communal roosts, colony members must regularly make group decisions over where to roost. In the two-field experiments, we provided marked individuals with conflicting information about the suitability of potential roosts. We investigated whether conflicting information led to group decisions that followed a 'unanimous' or a 'majority' rule, or increased colony fission. Individual behaviour suggests that bats considered both their own information and the behaviour of others when deciding where to roost. Group decisions about communal roosts reflected the information available to a majority of the bats roosting together, but conflicting information led to an increased fission in one colony. Our results suggest that fission-fusion societies allow individuals to avoid majority decisions that are not in their favour.     

Number of neighbors instead of group size significantly affects individual vigilance levels in large animal aggregations

The group size effect states that animals living in groups gain anti‐predator benefits through reducing vigilance levels as group size increases. A basic assumption of group size effect is that all individuals are equally important for a focal individual, who may adjust its vigilance levels according to social information acquired from them. However, some studies have indicated that neighbors pose greater influences on an individual's vigilance decisions than other group members, especially in large aggregations. Vigilance has also been found to be directed to both predators (anti‐predation vigilance) and conspecifics (social vigilance). Central individuals might rely more on social vigilance than peripheral individuals. To test these hypotheses, we examined the effects of flock size, number of neighbors and position within a flock on vigilance and competition of greater white‐fronted goose Anser albifrons that form large foraging flocks in winter, controlling the effects of other variables (group identity, winter period and site). We found that individual vigilance levels were significantly affected by number of neighbors and position within a flock, whereas flock size showed no effect. Individuals devoted a large component of vigilance to nearby flock mates. Central individuals directed a relatively larger proportion of vigilance to monitor neighbors than peripheral ones, indicating that central individuals more relied on social information acquired from neighbors, possibly caused by the more blocked visual field of central individuals. Moreover, some social vigilance may function as conducting or preventing agonistic interactions since competition intensity was positively correlated with number of neighbors. Our study therefore demonstrate that the number of neighbors is more important than group size in determining individual vigilance in large animal groups. Further studies are still needed to unravel which neighbors pose greater influence on individual vigilance, and the factors that influence individuals to acquire information from their neighbors to adjust vigilance behaviors.     

To follow or not? How animals in fusion–fission societies handle conflicting information during group decision‐making

When group members possess differing information about the environment, they may disagree on the best movement decision. Such conflicts result in group break‐ups, and are therefore a fundamental driver of fusion–fission group dynamics. Yet, a paucity of empirical work hampers our understanding of how adaptive evolution has shaped plasticity in collective behaviours that promote and maintain fusion–fission dynamics. Using movement data from GPS‐collared bison, we found that individuals constantly associated with other animals possessing different spatial knowledge, and both personal and conspecific information influenced an individual's patch choice decisions. During conflict situations, bison used group familiarity coupled with their knowledge of local foraging options and recently sampled resource quality when deciding to follow or leave a group – a tactic that led to energy‐rewarding movements. Natural selection has shaped collective behaviours for coping with social conflicts and resource heterogeneity, which maintain fusion–fission dynamics and play an essential role in animal distribution.