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What Animals Follow The Leader Of The Group

  • Journal List
  • Commun Integr Biol
  • five.ii(ii); Mar-April 2009
  • PMC2686370

Commun Integr Biol. 2009 Mar-Apr; two(ii): 147–150.

Leaders, followers and group controlling

Andrew J Rex

aneConstitute of Zoology; Zoological Society of London; London, United kingdom of great britain and northern ireland

2Department of Anthropology; University College London; London, United kingdom

Guy Cowlishaw

1Institute of Zoology; Zoological Society of London; London, UK

Received 2008 Dec vii; Accepted 2008 Dec eight.

Abstract

Social animals have to brand a multitude of group decisions on a daily basis. At the almost bones level, this will involve coordination of activities and travel directions. In groups of insects, birds and fish, much of this 'coordination' can be the effect of relatively unproblematic interaction patterns amongst grouping members. Such systems are self-organizing, and ofttimes do non require specific leaders, or followers. However, in more than socially complex groups, achieving commonage grouping action—a consensus—may not be achieved past simple rules alone. Instead, a consensus may be reached by the averaging of preferences (democracy), or by post-obit the choices of specific leaders (despotism). In this mini-review, we discuss the atmospheric condition necessary for despotism in creature groups, and focus upon new studies investigating coordinated actions in primates. We ask how specific leaders arise and why others follow them—providing new insight into the mechanisms of effective leadership in groups characterized by potent social relationships.

Central words: decisions, leaders, followers, social relationships, group-living

Introduction

Colonies, schools, flocks, herds and troops—the formation of groups is a universal phenomenon in the Animate being Kingdom. Such incredible diverseness of sociality has intrigued behavioral and evolutionary biologists, and there is a rich and various literature that strives to explain the origins and maintenance of group living. We now know much about the relative costs and benefits of grouping in animals,one even so for individuals to maximize the benefits and minimize the costs of grouping, they are required to be at to the lowest degree partially coordinated in their activities and travel directions. At the most fundamental level, this requires that different group members do not undertake such divergent activities equally to compromise group cohesion; for example, past initiating a foraging journey while the rest of the group remains at a sleeping site.2 Group-living animals must therefore according to act in unison.3 , 4 Interim in unison tin all the same be costly to individuals if it requires them to postpone an action that would be personally more than profitable in lodge to practise what the rest of the group is doing.2 This scenario will be more mutual in more stable social groups which can be especially heterogeneous as a consequence of private variation in authorization,5 , 6 relatedness,vii , 8 internal land9 xi and levels of information.12 14 In such groups and so, coordinated behavior and group cohesion requires individuals to be constantly balancing their desired actions and behaviors with that of their neighbors.13 , fifteen Ultimately, it is expected that the resulting cost of such 'balancing' will influence an individual's decision to remain in the group. Where between-individual variation in the timing of activities becomes likewise large then animals may not be able to reach a consensus on their activities and their coordination breaks down.xvi , 17 So how exercise group-living animals coordinate themselves under such conflicts of interest, and make (group) decisions?

Coordinated Behavior from Simple Rules

Examining the trouble of coordinating periods of foraging and resting, models by Rands and colleagues9 , 18 provide a straightforward resolution to the problem of group coordination where individuals' interests differ. They utilize a game-theoretic, state-dependent, individual-based approach to model the foraging behavior of a pair of animals. Their models predict that differences in the energetic reserves of the two players spontaneously develop, as a upshot of stochastic processes, leading them to adopt different behavioral roles. The individual with lower reserves tends to emerge as the leader, since the private with the college reserves will always adopt to minimize predation risk by foraging only when the other role player is doing so. All the same, this arroyo only considers minor groups (two animals), and while the furnishings of the decision rule derived from this model have been explored in larger groups,6 , nineteen these studies have non specifically addressed the question of how groups reach a consensus, on the timing of activities and travel directions for example. For this, alternative approaches are needed.

Much of the coordination in the timing of activities and travel directions evident in biological systems can exist the upshot of relatively unproblematic interaction patterns amidst group members.20 22 In such 'self-organizing systems'4 , 23 , 24 multiple individuals following unproblematic movement rules can produce complex collective behaviours.25 Such emergent collective behaviors can therefore be explained without invoking complex controlling abilities at the level of the private.26 , 27 But whilst self-organizing models tin be usefully practical to a variety of group behaviors and in many study systems,22 , 24 , 25 such models tend to work all-time where groups are composed of individuals with identical interests, and which only communicate locally (i.e., between proximate neighbors). Typical examples are decisions made by eusocial insects about choosing a new nest site,twenty , 28 , 29 or by navigating birds about travel routes.30 , 31 For many of the groups that we see in nature, withal, individuals and their interests will differ greatly, equally already discussed above.

Decision-Making under Conditions of Conflict

To specifically address the problem of conflicts of involvement, Conradt and Roper32 examined consensus decisions: when the members of a grouping choose between two or more mutually exclusive actions, resulting in a consensus. They specifically addressed the issue of 'consensus costs', which are the costs (in terms of reduced fitness) of animals forgoing their own optimal action to comply with the group consensus.33 Thus, if there is a large disharmonize of involvement involved in a consensus decision, the consensus costs will be as big. They modeled two alternative decision processes. First, decisions may be made in a 'democratic' manner, where the average behavior of individuals is adopted. Second, decisions may exist made by a single animal or minority of animals in a more 'despotic' manner.16 , 32 , 33 Conradt and Roper's models show that both democratic and despotic decision-making tin evolve through, and be maintained past, individual pick.16 Nevertheless, they predict that under most conditions the costs to subordinate group members, and to the group as a whole, are considerably higher for despotic than for democratic decisions. As a consequence, they suggest that democratic decisions are more likely to evolve. Conradt and Roper's models further signal that democratic decisions can fifty-fifty evolve when groups are heterogeneous in limerick; 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.16

So does empirical show support these recent models? Tests of consensus decision-making in vertebrate groups have largely concentrated on decisions about travel routes or the timing of activities.33 Inside this torso of enquiry, testify for both democratic and despotic decision-making has been presented, eastward.g., primates,34 37 ungulates,38 42 and birds.13 , 43 , 44 Why despotism appears at to the lowest degree every bit often as commonwealth in nature—contrary to theoretical predictions—remains unclear. In fact, explaining the profusion of despotic controlling in nature presents us with a significant claiming to our understanding of sociality.

Agreement Leadership

New insights into the emergence of despotic grouping decisions in animate being groups may exist caused by agreement how leaders arise and why others follow them. In that location are conceivably several different types of animate being that might emerge as a leader. In eusocial insects, information technology has been shown that very few individuals within a group may actually possess pertinent information with respect to the decision in hand,45 , 46 and thus get crucial to coordinating behavior and the decision process. In vertebrates as well, a minority of informed individuals (often elders) are seen to guide entire groups to specific resources. These include golden shiner fish Notemigonus crysoleucas,47 elephants Loxodonta africana,48 ravens Corvus corax,49 and broad-winged hawks Buteo platypterus.50 Specific animals may also lead groups on the basis that they are the hungriest, or because of the feeding benefits they derive from leading groups to nutrient resource.51 53

Just the incentive or information required to create leaders does not necessarily generate post-obit, and both processes are necessary for effective leadership. Consider long-lived and cognitively complex organisms, like primates, that display intricate social interactions. These create college-order properties of groups that can be studied and quantified as authorisation hierarchies and social networks.5 , 54 Given that such higher-society properties tin can modify private behavior, should nosotros expect all individuals to have an 'equal say' where group coordination and decision-making is concerned sensu Conradt and Roper?32 Concerning authorization, high-ranking individuals are known to hold a particularly strong influence over the behaviors of group-mates.55 , 56 Where members of families (or matrilines) coexist together, specific individuals may too take a larger influence according to the relative number of kin relations (i.e., size of matriline).57 59 Similarly, given the corporeality of time invested in social relationships, and the established importance of social networks to individual fettle (for case encounter Silk et al.lx), individuals with stronger and/or more social bonds inside groups may be in a meliorate position to generate follower behavior. The influence of social relationships can therefore not be ignored, and their critical part has been highlighted by a number of contempo studies on primates.

Looking for Leaders

Sueur and Petit61 used network metrics (Fig. 1) to assess what rules may underlie follower behavior in two macaque species: rhesus macaques (Macaca mulatta) and Tonkean macaques (Macaca tonkeana) living in semi costless-ranging conditions. Rhesus macaques are a highly hierarchical and nepotistic species, whereas Tonkean macaques are often more than tolerant and egalitarian in nature.62 Sueur and Petit examined the organization of group members when joining a move initiated by a commencement individual, and found the fashion macaques joined a movement reflected the 2 species' differing social systems. Older and more dominant male rhesus macaques were more often at the front of the movement. In contrast, Tonkean macaques exhibited no specific order. Interestingly the researchers also found that rhesus macaques preferred to follow high-ranking or related individuals, whereas Tonkean macaques' follower behavior reflected only male-female sexual relationships. These observations suggest that ascendant rhesus macaques, which were at the front of the movements observed, have an especially strong influence over group-mates beliefs, and can exist described as leaders that elicit follower behaviour.32 , 35

An external file that holds a picture, illustration, etc.  Object name is cib0202_0147_fig001.jpg

Network of associations during collective movements for rhesus macaques provided courtesy of C. Sueur and O. Petit (adjusted from Sueur & Petit61). Nodes represent individuals. The number indicates the authorization rank. Females are in black, males in grey (two males: rank 1 for the alpha male person and rank 2 for a peripheral male). The distances between individuals are 'half-weight indices' which, in this instance, represent individuals that are more oft associated during group movements. The size of a node is directly related to the private eigenvector axis coefficient; the higher the centrality coefficient, the greater the importance of the individual in the joining of group members. Male 1 (alpha male) and female person vi (who connects two matrilines of females, see Sueur and Petit61) are thus key to eliciting follower behavior. Meet Sueur and Petit61 for further details on the adding of these network associations.

A recent study by King et al.63 ostend Sueur and Petit's prediction that social relationships tin have a large influence upon an private'southward ability to human action every bit a leader, and dictate the behavior of grouping mates. Rex et al. presented two wild baboon groups with experimental food patches within their habitation ranges. In the experimental patches, food intake amidst group members was highly skewed so that a minority of (ascendant) group members acquired a lot of food, while the majority of (subordinate) group members obtained very little (if any). In dissimilarity, in natural patches, private food intake was relatively evenly spread beyond group members. Thus, King et al. predicted that simply if ascendant individuals were able to act as leaders, and elicit follower beliefs from subordinates, would groups choose an experimental over a natural nutrient patch. This is because the bulk of group members would incur substantial consensus costs (see to a higher place) from this decision. If even so, the dominant individuals were unable to dictate group decisions—and act as leaders—then groups were predicted to choose natural patches over experimental patches. Retrieve that theory predicts that nether these circumstances groups should movement to the natural patch, which benefits the majority of group members and minimizes overall consensus costs.xvi , 32 What Rex et al. observed was that their baboon groups consistently visited experimental patches in preference to natural patches. What is more, the dominant male person consistently led his group to the experimental patches (Fig. two), and the individuals that followed him most closely were those with whom he shared the strongest social bonds (as indexed by grooming interactions). They besides noted that coercion by dominants did not play a office in this choice.

An external file that holds a picture, illustration, etc.  Object name is cib0202_0147_fig002.jpg

A wild baboon troop in Namibia, studied by Rex et al.,63 travelling in single file toward a known food source. King et al. showed that the dominant male—when provided with incentives—will lead groups to experimental nutrient patches, fifty-fifty though the group equally a whole do worse than if they foraged elsewhere. See text for more details. Photo courtesy of Hannah Peck/ZSL Tsaobis Baboon Project.

Such leadership behavior is puzzling, because choice is predicted to favor equally shared decisions over dominant decisions under a wide diverseness of conditions.xvi What Sueur and Petit observed, and King et al. showed experimentally, was that social ties appear key to such patterns of beliefs. So why should the desire to follow an private of high social status be so potent? King et al. suggest that for their baboons, shut clan with the dominant male person may provide females and their dependent offspring with direct fettle benefits, such as increased babe survival and protection from predators. In event, it is worth experiencing the short-term costs that might ensue from following, because the long-term benefits of association with the leader should outweigh these. Thus far, theoretical models take not considered the importance of social ties in group decision-making and in the emergence of leaders in groups. These recent studies of primates, and observations of other taxa too, east.yard., zebra,38 suggest that this may be a fruitful avenue of investigation in the future.

Outlook

Theoretical developments tackling the topic of leadership and group decision-making have blossomed in contempo years,ix , 12 , 32 and whilst empirical studies of insects tackling the subject have ofttimes matched these developments,45 , 46 , 64 work on vertebrate groups has been less productive. There are several reasons for this. One is that information technology tin take many months or even years to investigate the complex aspects of leadership and group decision-making that these theoretical models tackle. Another is that such studies are often express by pocket-sized sample sizes (in terms of the number of groups they are able to written report), frequently due to the difficulty of studying vertebrate groups, which may be shy of observers and range over large areas. However, the outlook for futurity research in this field looks promising. For example, we believe that studies of captive and semi-wild populations tin can provide enormous scope for investigating the office of certain individuals in group-level behavior. Flack et al.65 carried out experiments in captive pigtailed macaques (Macaca nemestrina), in which high-ranked individuals were temporarily removed. They institute that their removal was associated with dramatic reductions in the size and connectivity of social networks, which finer de-stabilized the social groups. It might well be possible to blueprint similar experiments to investigate the role of certain individuals in maintaining grouping coordination and directing collective actions. Field-based studies in this subject field will at present surely advance too, given technological developments. For example, Global Positioning System (GPS) tracking of individuals volition allow continuous sampling of multiple individuals' movements in real-time.66 Combining these information with social network analyses54 volition let researchers to examine the office of social interactions in shaping the spatial properties of groups and the importance of specific individuals in leading groups to resources in their environment.

What nosotros accept learned is that in that location is a need to understand how social interactions tin can shape group-level patterns of behavior in vertebrate groups, and how they regulate the basic coordination of multiple individuals. We hope that this mini-review will help stimulate other researchers to explore these issues in a diverseness of new species and written report systems, and nosotros eagerly anticipate their findings.

Acknowledgements

Andrew J. King gratefully acknowledges support from an AXA Postdoctoral Fellowship and a Natural Environment Inquiry Council (NERC) Studentship.

Footnotes

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