The response of a selfish herd to an attack from outside the group perimeter

According to the selfish herd hypothesis, animals can decrease predation risk by moving toward one another if the predator can appear anywhere and will attack the nearest target. Previous studies have shown that aggregations can form using simple movement rules designed to decrease each animal's Domain of Danger. However, if the predator attacks from outside the group's perimeter, these simple movement rules might not lead to aggregation. To test whether simple selfish movement rules would decrease predation risk for those situations when the predator attacks from outside the flock perimeter, we constructed a computer model that allowed flocks of 75 simulated fiddler crabs to react to one another, and to a predator attacking from 7 m away. We attacked simulated crab flocks with predators of different sizes and attack speeds, and computed relative predation risk after 120 time steps. Final trajectories showed flight toward the center of the flock, but curving away from the predator. Path curvature depended on the predator's size and approach speed. The average crab experienced a greater decrease in predation risk when the predator was small or slow moving. Regardless of the predator's size and speed, however, predation risk always decreased as long as crabs took their flock-mates into account. We conclude that, even when flight away from an external predator occurs, the selfish avoidance of danger can lead to aggregation.     

Simulated evolution of selfish herd behavior

Single species aggregations are a commonly observed phenomenon. One potential explanation for these aggregations is provided by the selfish herd hypothesis, which states that aggregations result from individual efforts to reduce personnel predation risk at the expense of group-mates. Not all movement rules based on the selfish herd hypothesis are consistent with observed animal behavior. Previous work has shown that herd-like aggregations are not generated by movement rules limited to local interactions between nearest neighbors. Instead, rules generating realistic herds appear to require delocalized interactions. To date, it has been an open question whether or not the necessary delocalization can emerge from local interactions under natural selection. To address this question, we study an individual-based model with a single quantitative genetic trait that controls the influence of neighbors as a function of distance. The results indicate that predation-based selection can increase the influence of distant neighbors relative to near neighbors. Our results lend support for the idea that selfish herd behavior can arise from localized movement rules under natural selection.     

Geometry for mutualistic and selfish herds: the limited domain of danger

We present a two-dimensional individual-based model of aggregation behaviour in animals by introducing the concept of a "limited domain of danger", which represents either a limited detection range or a limited attack range of predators. The limited domain of danger provides a suitable framework for the analysis of individual movement rules under real-life conditions because it takes into account the predator's prey detection and capture abilities. For the first time, a single geometrical construct can be used to analyse the predation risk of both peripheral and central individuals in a group. Furthermore, our model provides a conceptual framework that can be equally applied to aggregation behaviour and refuge use and thus presents a conceptual advance on current theory that treats these antipredator behaviours separately. An analysis of individual movement rules using limited domains of danger showed that the time minimization strategy outcompetes the nearest neighbour strategy proposed by Hamilton's (J. Theor. Biol. 31 (1971) 295) selfish herd model, whereas a random strategy confers no benefit and can even be disadvantageous. The superior performance of the time minimization strategy highlights the importance of taking biological constraints, such as an animal's orientation relative to its neighbours, into account when searching for efficient movement rules underlying the aggregation process.     

Mechanisms for aggregation in animals: rule success depends on ecological variables

Under the threat of predation, animals often group tightly together,with all group members benefiting from a reduction in predationrisk through various mechanisms, including the dilution, encounter-dilution,and predator confusion effects. Additionally, the selfish herdhypothesis was first put forward by Hamilton (1971). He proposedthat in order to reduce its risk of predation, an individualshould approach its nearest neighbor, reducing its risk at theexpense of those around it. Despite extensive empirical support,the selfish herd hypothesis has been criticized on theoreticalgrounds: approaching the nearest neighbor does not result inthe observed dense aggregations, and the nearest neighbor inspace is not necessarily the one that can be reached fastest.Increasingly complex movement rules have been proposed, successfullyproducing dense aggregations of individuals. However, no studyto date has made a full comparison of the different proposedmovement rules within the same modeling environment. Further,ecologically relevant parameters, such as the size and densityof a population or group and the time it takes a predator toattack, have thus far been ignored. Here, we investigate thereduction in risk for animals aggregating using different strategiesand demonstrate the importance of ecological parameters on riskreduction in group-living animals. We find that complex rulesare most successful at reducing risk in small, compact populations,whereas simpler rules are most successful in larger, low-densitypopulations, and when predators attack quickly after being detectedby their prey.     

The temporal selfish herd: predation risk while aggregations form

The hypothesis of the selfish herd has been highly influential to our understanding of animal aggregation. Various movement strategies have been proposed by which individuals might aggregate to form a selfish herd as a defence against predation, but although the spatial benefits of these strategies have been extensively studied, little attention has been paid to the importance of predator attacks that occur while the aggregation is forming. We investigate the success of mutant aggregation strategies invading populations of individuals using alternative strategies and find that the invasion dynamics depend critically on the time scale of movement. If predation occurs early in the movement sequence, simpler strategies are likely to prevail. If predators attack later, more complex strategies invade. If there is variation in the timing of predator attacks (through variation within or between individual predators), we hypothesize that groups will consist of a mixture of strategies, dependent upon the distribution of predator attack times. Thus, behavioural diversity can evolve and be maintained in populations of animals experiencing a diverse range of predators differing solely in their attack behaviour. This has implications for our understanding of predator–prey dynamics, as the timing of predator attacks will exert selection pressure on prey behavioural responses, to which predators must respond.     

Quantitative analysis of fiddler crab flock movement: evidence for ‘selfish herd’ behaviour

The selfish herd hypothesis predicts that aggregations form because individuals move towards one another, and that this movement will minimize predation risk as measured by the domain of danger. To test the predictions of the selfish herd hypothesis in the field, we videotaped the movements of sand fiddler crab, Uca pugilator, flocks being attacked by predators. After recording 12 attacks on crabs by shorebird and human attackers, we digitized the video, and determined the positions of crabs before and after being frightened. We estimated the time of panic initiation by the rapid increase in the crabs' velocity. Crab flocks became more cohesive after panic initiation. The frequency distribution of the crabs' domains of danger shifted significantly towards smaller domains after panic initiation. The median domain of danger was significantly lower after panic initiation than beforehand. Two other indices of aggregation also showed statistically significant increases in flock cohesion following panic initiation. We conclude that fiddler crab behaviour is consistent with the selfish herd hypothesis. Therefore, our results support the selfish herd hypothesis as an explanation for gregarious behaviour. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

‘Selfish herds’ of guppies follow complex movement rules,but not when information is limited

Under the threat of predation, animals can decrease their level of risk by moving towards other individuals to form compact groups. A significant body of theoretical work has proposed multiple movement rules, varying in complexity, which might underlie this process of aggregation. However, if and how animals use these rules to form compact groups is still not well understood, and how environmental factors affect the use of these rules even less so. Here, we evaluate the success of different movement rules, by comparing their predictions with the movement seen when shoals of guppies (Poecilia reticulata) form under the threat of predation. We repeated the experiment in a turbid environment to assess how the use of the movement rules changed when visual information is reduced. During a simulated predator attack, guppies in clear water used complex rules that took multiple neighbours into account, forming compact groups. In turbid water, the difference between all rule predictions and fish movement paths increased, particularly for complex rules, and the resulting shoals were more fragmented than in clear water. We conclude that guppies are able to use complex rules to form dense aggregations, but that environmental factors can limit their ability to do so.     

Sharks shape the geometry of a selfish seal herd: experimental evidence from seal decoys

Many animals respond to predation risk by forming groups. Evolutionary explanations for group formation in previously ungrouped, but loosely associated prey have typically evoked the selfish herd hypothesis. However, despite over 600 studies across a diverse array of taxa, the critical assumptions of this hypothesis have remained collectively untested, owing to several confounding problems in real predator–prey systems. To solve this, we manipulated the domains of danger of Cape fur seal (Arctocephalus pusillus pusillus) decoys to provide evidence that a selfish reduction in a seals'' domain of danger results in a proportional reduction in its predation risk from ambush shark attacks. This behaviour confers a survival advantage to individual seals within a group and explains the evolution of selfish herds in a prey species. These findings empirically elevate Hamilton''s selfish herd hypothesis to more than a ‘theoretical curiosity’.     

Escaping parasitism in the selfish herd: age, size and density-dependent warble fly infestation in reindeer

It has been suggested that animals may escape attack from mobile parasites by aggregating in selfish herds. A selfish herd disperses the risk of being attacked among its members and the per individual risk of parasite infection should therefore decrease with increasing animal density through the encounter–dilution effect. Moreover, in a selfish herd, dominant and agile animals should occupy the best positions and thereby receive fewer attacks compared to lower ranked animals at the periphery. We tested these predictions on reindeer ( Rangifer tarandus tarandus ) parasitized by warble flies ( Hypoderma tarandi ). Warble flies oviposit their eggs on reindeer during summer and induce strong anti-parasitic behavioural responses in the herds. In this period, reindeer are sexually segregated; females and calves form large and dense herds while males are more solitary. After hatching, the warble fly larvae migrate under the skin of their host where they encyst. In the present study encysted larvae were counted on newly slaughtered hides of male calves and 1.5 year old males from 18 different reindeer herds in Finnmark, northern Norway with large contrasts in reindeer density. In reindeer, body mass is correlated with fitness and social status and we hypothesized that individual carcass mass reflected the animal's ability to occupy the best positions within the herd. Larval abundance was higher among the 1.5 year old males than among the calves. For calves we found in accordance with the selfish herd hypothesis a negative relationship between larval abundance and animal density and between larval abundance and body mass. These relationships were absent for the 1.5 year old males. We suggest that these differences were due to different grouping behaviour where calves and females, but not males, aggregated in selfish herds where they escaped parasitism.     

Predator-driven fragmentation of fiddler crab droves into selfish miniherds of biased composition

An explanation for animal groups is the selfish herd, characterized by aggregation as each member tries to shield itself from a predator by moving into a tight gap between other members. We test the hypotheses that: (1) droves, the large feeding groups of fiddler crabs, are selfish herds; (2) the miniherds that form when droves fragment on approach of a large predator are selfish herds; (3) selfish herds form when refugia are unlikely to be reached before an approaching predator arrives; and (4) the composition of selfish miniherds is biased toward individuals most vulnerable to predation. The study was conducted in South Carolina (USA) by videotaping the movements of sand fiddler crabs Uca pugilator when approached by a human predator. In both droves and miniherds, interindividual distance decreases with predator approach, consistent with behavior in a selfish herd. However, two other expectations for selfish herds—herd cohesion and sacrificing distance from the predator in order to get closer to other herders—are only met in miniherds. Crabs farther from refugia are more likely to form and remain in miniherds, indicating that selfish herding is only favored when refugia cannot be quickly reached. The composition of the smallest miniherds, consisting of 2-18 crabs, is biased toward females and small males. These individuals may be more vulnerable to predation because they lack the enlarged claw of large males that deters some predators. The small miniherds are relatively homogeneous with respect to the size and sex of their members, which may enhance cohesion and effectiveness as selfish herds. Miniherds will be effective selfish groups when predator attack has a significant vertical component and when the strike distance is large relative to both the size of the prey and the distance between group members. Droves are not selfish herds but permit crabs to flee feeding grounds as members of selfish miniherds.     

Evolving the selfish herd: emergence of distinct aggregating strategies in an individual-based model

From zebra to starlings, herring and even tadpoles, many creatures move in an organized group. The emergent behaviour arises from simple underlying movement rules, but the evolutionary pressure which favours these rules has not been conclusively identified. Various explanations exist for the advantage to the individual of group formation: reduction of predation risk; increased foraging efficiency or reproductive success. Here, we adopt an individual-based model for group formation and subject it to simulated predation and foraging; the haploid individuals evolve via a genetic algorithm based on their relative success under such pressure. Our work suggests that flock or herd formation is likely to be driven by predator avoidance. Individual fitness in the model is strongly dependent on the presence of other phenotypes, such that two distinct types of evolved group can be produced by the same predation or foraging conditions, each stable against individual mutation. We draw analogies with multiple Nash equilibria theory of iterated games to explain and categorize these behaviours. Our model is sufficient to capture the complex behaviour of dynamic collective groups, yet is flexible enough to manifest evolutionary behaviour.     

On the evolution of group-escape strategies of selfish prey

The phenomenon of group escape cannot be explained by an argument of risk dilution, applied to gregarious behaviour of passive prey whose risk of predation is equally shared by all group members (Hamilton, 1971). Instead, individuals at the tail of an escaping group suffer the bulk of the group’s predation risk, and thus have the highest incentive to desert it. Just because of this, desertion, in this case, may serve as a signal of vulnerability for the pursuing predator. Under wide conditions, it is therefore shown that the predator is always expected to prefer the chasing of a deserter, whenever it is observed. Consequently, an individual who finds himself at the tail of the herd must compare the risk of remaining there with that of deserting the herd and thereby becoming a likely target for predation. If the first risk is higher than the latter, the herd disperses; if the latter is higher, the herd cohesively follows the fastest individuals in its lead (we deal also with cases in which only part of the herd disperses). We see, however, that the question which risk is higher depends not only on the terrain, but also on the route of escape that is decided by the fastest members at the lead of the herd, those that are least likely to be caught. Concentrating on herds without family structure, we assume that the route of escape is selfishly chosen by these ad hoc leaders to minimize their own predation risk, regardless of the others’ welfare. However, the predation risk of the leader depends very much on the willingness of other herd members to follow him, thus providing a buffer between him and the pursuing predator. Consequently, when choosing an escape route, the leader has also to consider the cohesion of the herd, i.e., the reaction of slower individuals to his choice. Under some plausible conditions, this choice may force the herd to follow, while other conditions may lead to its dispersal. In some cases the leader may choose a route that serves the needs of the entire group, and sometime only those of its more vulnerable members. In other cases the leader may choose a route that sacrifices the weakest members, thereby improving the survival probability of the others.We employ a model of a k+1 players game, a single predator, and k heterogeneous prey individuals. The predator aims to maximize the probability of a successful catch, and each individual aims to minimize his probability of being caught.     

Scalable rules for coherent group motion in a gregarious vertebrate

Individuals of gregarious species that initiate collective movement require mechanisms of cohesion in order to maintain advantages of group living. One fundamental question in the study of collective movement is what individual rules are employed when making movement decisions. Previous studies have revealed that group movements often depend on social interactions among individual members and specifically that collective decisions to move often follow a quorum-like response. However, these studies either did not quantify the response function at the individual scale (but rather tested hypotheses based on group-level behaviours), or they used a single group size and did not demonstrate which social stimuli influence the individual decision-making process. One challenge in the study of collective movement has been to discriminate between a common response to an external stimulus and the synchronization of behaviours resulting from social interactions. Here we discriminate between these two mechanisms by triggering the departure of one trained Merino sheep (Ovis aries) from groups containing one, three, five and seven naïve individuals. Each individual was thus exposed to various combinations of already-departed and non-departed individuals, depending on its rank of departure. To investigate which individual mechanisms are involved in maintaining group cohesion under conditions of leadership, we quantified the temporal dynamic of response at the individual scale. We found that individuals'' decisions to move do not follow a quorum response but rather follow a rule based on a double mimetic effect: attraction to already-departed individuals and attraction to non-departed individuals. This rule is shown to be in agreement with an adaptive strategy that is inherently scalable as a function of group size.     

The Role of Neighbours Selection on Cohesion and Order of Swarms

We introduce a multi-agent model for exploring how selection of neighbours determines some aspects of order and cohesion in swarms. The model algorithm states that every agents'' motion seeks for an optimal distance from the nearest topological neighbour encompassed in a limited attention field. Despite the great simplicity of the implementation, varying the amplitude of the attention landscape, swarms pass from cohesive and regular structures towards fragmented and irregular configurations. Interestingly, this movement rule is an ideal candidate for implementing the selfish herd hypothesis which explains aggregation of alarmed group of social animals.     

Antipredator defense as a limited resource: unequal predation risk in broods of an insect with maternal care

The allocation of parental investment is a potential sourceof conflict within broods whenever offspring are able obtaindifferential access to the parental resource. Unlike the provisioningof food, parental antipredator behavior is usually considereda resource that benefits all offspring simultaneously. In thethornbug treehopper (Umbonia crassicornis), offspring formaggregations in exposed positions on host-plant stems. Theyare subject to intense predation, and maternal defense is theirprimary means of protection. I examined the distribution ofrisk within these offspring groups, using natural variationin the outcome of more than 500 predation attempts (324 recordedon videotape) by vespid wasps (Pseudopolybia compressa) on18 U. crassicornis aggregations. I found three influences onan individual offspring's risk of predation. The first wasthe presence of a defending female: as expected, offspringwere much more likely to survive contact with a wasp if thefemale was present than if the female had disappeared. Thesecond influence was position relative to other offspring: when wasps were successful in removing an individual, they almostalways removed it from the edge of the group. The third influencewas distance from the female: the closer an offspring was tothe female at the time it was contacted by a wasp, the higherits likelihood of survival. The distribution of risk is determinedlargely by the behavior of defending females and the prey-searchingbehavior of wasps. The nature of risk within these aggregations sets the stage for two forms of sibling rivalry: selfish herdbehavior and competition for access to maternal defense. Italso raises the question of how a parent should allocate defenseamong offspring when it is unable to defend them all simultaneously.     

Negotiation may lead selfish individuals to cooperate: the example of the collective vigilance game

Game-theoretical models have been highly influential in behavioural ecology. However, these models generally assume that animals choose their action before observing the behaviour of their opponents while, in many natural situations, individuals in fact continuously react to the actions of others. A negotiation process then takes place and this may fundamentally influence the individual attitudes and the tendency to cooperate. Here, I use the classical model system of vigilance behaviour to demonstrate the consequences of such behavioural negotiation among selfish individuals, by predicting patterns of vigilance in a pair of animals foraging under threat of predation. I show that the game played by the animals and the resulting vigilance strategies take radically different forms, according to the way predation risk is shared in the pair. In particular, if predators choose their target at random, the prey respond by displaying moderate vigilance and taking turns scanning. By contrast, if the individual that takes flight later in an attack endures a higher risk of being targeted, vigilance increases and there is always at least one sentinel in the pair. Finally, when lagging behind its companion in fleeing from an attacker becomes extremely risky, vigilance decreases again and the animals scan simultaneously.     

Communication and Cognition in Primate Group Movement

We here review the communicative and cognitive processes underpinning collective group movement in animals. Generally, we identify 2 major axes to explain the dynamics of decision making in animal or human groups or aggregations: One describes whether the behavior is largely determined by simple rules such as keeping a specific distance from the neighbor, or whether global information is also factored in. The second axis describes whether or not the individual constituents of the group have overlapping or diverging interests. We then review the available evidence for baboons, which have been particularly well studied, but we also draw from further studies on other nonhuman primate species. Baboons and other nonhuman primates may produce specific signals in the group movement context, such as the notifying behavior of male hamadryas baboons at the departure from the sleeping site, or clear barks that are given by chacma baboons that have lost contact with the group or specific individuals. Such signals can be understood as expressions of specific motivational states of the individuals, but there is no evidence that the subjects intend to alter the knowledge state of the recipients. There is also no evidence for shared intentionality. The cognitive demands that are associated with decision making in the context of group coordination vary with the amount of information and possibly conflicting sources of information that need to be integrated. Thus, selective pressures should favor the use of signals that maintain group cohesion, while recipients should be selected to be able to make the decision that is in their own best interest in light of all the available information.     

Dynamical modeling of collective behavior from pigeon flight data: flock cohesion and dispersion

Several models of flocking have been promoted based on simulations with qualitatively naturalistic behavior. In this paper we provide the first direct application of computational modeling methods to infer flocking behavior from experimental field data. We show that this approach is able to infer general rules for interaction, or lack of interaction, among members of a flock or, more generally, any community. Using experimental field measurements of homing pigeons in flight we demonstrate the existence of a basic distance dependent attraction/repulsion relationship and show that this rule is sufficient to explain collective behavior observed in nature. Positional data of individuals over time are used as input data to a computational algorithm capable of building complex nonlinear functions that can represent the system behavior. Topological nearest neighbor interactions are considered to characterize the components within this model. The efficacy of this method is demonstrated with simulated noisy data generated from the classical (two dimensional) Vicsek model. When applied to experimental data from homing pigeon flights we show that the more complex three dimensional models are capable of simulating trajectories, as well as exhibiting realistic collective dynamics. The simulations of the reconstructed models are used to extract properties of the collective behavior in pigeons, and how it is affected by changing the initial conditions of the system. Our results demonstrate that this approach may be applied to construct models capable of simulating trajectories and collective dynamics using experimental field measurements of herd movement. From these models, the behavior of the individual agents (animals) may be inferred.     

The influence of flock size and geometry on the scanning behaviour of spotted turtle doves,Streptopelia chinensis

Abstract The negative correlation between the time individuals spend scanning the environment for predators and group size is usually explained by the benefit of corporate vigilance. However, this negative correlation may be explained additionally in terms of the ‘dilution effect’ and ‘selfish herd geometry’. Our experimental investigation of the scanning behaviour of free-living spotted turtle doves foraging at different shaped feeders revealed that flock geometry influenced individual scanning rates. The time spent scanning declined with group size less rapidly among birds foraging in linear flocks than among those foraging in more two-dimensional flocks. These results were not confounded by aggressive behaviour, and indicate that the benefits of foraging in groups include the so-called selfish herd geometry.     

Cooperative sentinel calling? Foragers gain increased biomass intake

Many foraging animals face a fundamental tradeoff between predation and starvation. In a range of social species, this tradeoff has probably driven the evolution of sentinel behavior, where individuals adopt prominent positions to watch for predators while groupmates forage. Although there has been much debate about whether acting as a sentinel is a selfish or cooperative behavior, far less attention has focused on why sentinels often produce quiet vocalizations (hereafter known as "sentinel calls") to announce their presence. We use observational and experimental data to provide the first evidence that group members gain an increase in foraging success by responding to these vocal cues given by sentinels. Foraging pied babblers (Turdoides bicolor) spread out more, use more exposed patches, look up less often, and spend less time vigilant in response to sentinel calling. Crucially, we demonstrate that these behavioral alterations lead to an increase in biomass intake by foragers, which is likely to enhance survival. We argue that this benefit may be the reason for sentinel calling, making it a truly cooperative behavior.