Is Leadership a Reliable Concept in Animals? An Empirical Study in the Horse

Leadership is commonly invoked when accounting for the coordination of group movements in animals, yet it remains loosely defined. In parallel, there is increased evidence of the sharing of group decisions by animals on the move. How leadership integrates within this recent framework on collective decision-making is unclear. Here, we question the occurrence of leadership in horses, a species in which this concept is of prevalent use. The relevance of the three main definitions of leadership – departing first, walking in front travel position, and eliciting the joining of mates – was tested on the collective movements of two semi-free ranging groups of Przewalski horses (Equus ferus przewalskii). We did not find any leader capable of driving most group movements or recruiting mates more quickly than others. Several group members often displayed pre-departure behaviours at the same time, and the simultaneous departure of several individuals was common. We conclude that the decision-making process was shared by several group members a group movement (i.e., partially shared consensus) and that the leadership concept did not help to depict individual departure and leading behaviour across movements in both study groups. Rather, the different proxies of leadership produced conflicting information about individual contributions to group coordination. This study discusses the implications of these findings for the field of coordination and decision-making research.     

Decision-Making in Przewalski Horses (Equus ferus przewalskii) is Driven by the Ecological Contexts of Collective Movements

We addressed decision-making processes in the collective movements of two groups of Przewalski horses ( Equus ferus przewalskii ) living in a semi free-ranging population. We investigated whether different patterns of group movement are related to certain ecological contexts (habitat use and group activity) and analysed the possible decision-making processes involved. We found two distinct patterns; 'single-bout' and 'multiple-bout' movements occurred in both study groups. The movements were defined by the occurrence of collective stops between bouts and differed by their duration, distance covered and ecological context. For both movements, we found that a preliminary period involving several horses occurred before departure. In single-bout movements, all group members rapidly joined the first moving horse, independently of the preliminary period. In multiple-bout movements, however, the joining process was longer; in particular when the number of decision-makers and their pre-departure behaviour before departure increased. Multiple-bout movements were more often used by horses to switch habitats and activities. This observation demonstrates that the horses need more time to resolve motivational conflicts before these departures. We conclude that decision-making in Przewalski horses is based on a shared consensus process driven by ecological determinants.     

Decision-making Processes in Group Departures of Cattle

To keep social cohesiveness, group-living animals have to reach consensus decisions through recruitment processes. This implies that decision-making depends on the behaviours and social relationships of several group members at different stages of movements. We tested these assumptions in a group of fifteen 18-mo-old Charolais heifers ( Bos taurus ) at pasture, in which two observers continuously videotaped social interactions and group departures after resting periods. These departures were preceded by a phase of preparation characterized by an increase in activity. The number of heifers participating to a movement increased with the number of group members oriented in the direction of the movement before departure. The first moving animal also recruited a higher number of mates when it had a greater number of close neighbours, the first individuals to follow being mainly its preferential partners. Coercive interactions such as pressing behaviours were observed within the 5 min preceding or following departure. After departure, the numbers of walks and restarts of the first two movers were still operative in recruiting others. The frequency of pauses of the first mover was significantly higher when it was not followed, meaning that it adjusted its behaviour to that of other group members. Decision-making was distributed among group members, with any individual being liable to move first. The behaviour of cows and their spatial distribution before departure, at departure and after departure significantly affected the number of participants in the movement, demonstrating that decision-making was time-distributed in the studied cattle group.     

The Organization of Collective Group Movements in Wild Barbary Macaques (Macaca sylvanus): Social Structure Drives Processes of Group Coordination in Macaques

Social animals have to coordinate activities and collective movements to benefit from the advantages of group living. Animals in large groups maintain cohesion by self-organization processes whereas in smaller groups consensus decisions can be reached. Where consensus decisions are relevant leadership may emerge. Variation in the organization of collective movements has been linked to variation in female social tolerance among macaque species ranging from despotic to egalitarian. Here we investigated the processes underlying group movements in a wild macaque species characterized by a degree of social tolerance intermediate to previously studied congeneric species. We focused on processes before, during and after the departure of the first individual. To this end, we observed one group of wild Barbary macaques (Macaca sylvanus) in the Middle Atlas, Morocco using all-occurrence behaviour sampling of 199 collective movements. We found that initiators of a collective movement usually chose the direction in which more individuals displayed pre-departure behavior. Dominant individuals contributed to group movements more than subordinates, especially juveniles, measured as frequencies of successful initiations and pre-departure behaviour. Joining was determined by affiliative relationships and the number of individuals that already joined the movement (mimetism). Thus, in our study group partially shared consensus decisions mediated by selective mimetism seemed to be prevalent, overall supporting the suggestion that a species’ social style affects the organization of group movements. As only the most tolerant species show equally shared consensus decisions whereas in others the decision is partially shared with a bias to dominant individuals the type of consensus decisions seems to follow a stepwise relation. Joining order may also follow a stepwise, however opposite, relationship, because dominance only determined joining in highly despotic, but not in intermediate and tolerant species.     

The influence of social organisation on leadership in brown lemurs (Eulemur fulvus fulvus) in a controlled environment

Studies on leadership during group movements in several lemur species showed that females were responsible for the travelling choices concerning time and direction. Interestingly, in these species females are dominant over males. We investigated the influence of social organisation upon leadership processes by studying a lemur species in which social organisation is characterized by the absence of female dominance: the brown lemur (Eulemur fulvus fulvus). The study was conducted on a semi-free ranging group of 11 individuals and the analysis performed on 69 group movements showed that all the individuals could initiate a group movement. In 34 cases, the whole group moved. There was no significant difference in the number of start attempts or in the number of group members involved from one initiator to another. Moreover, there was no effect of sex or age of the initiator on the number of individuals following it or on the speed of the joining process. Therefore, the leadership observed is widely distributed to all group members. These results support the hypothesis of an influence of social organisation upon the decision-making processes but still remain to be studied in a more relevant ecological context.     

Information transfer during shell exchange in the hermit crab Clibanarius antillensis

The shell exchange behaviour of the hermit crab Clibanarius antillensis was examined for evidence that the non-initiating crab used information about the initiator's shell. Manipulation of the internal volume of the initiator's shell altered the outcome of exchanges in a manner consistent with the negotiations model of resource exchange. It is suggested that the fundamental frequency of shells is detected by crabs during rapping behaviour when the initiator raps its shell against that of the non-initiator.     

Quorum responses and consensus decision making

Animal groups are said to make consensus decisions when group members come to agree on the same option. Consensus decisions are taxonomically widespread and potentially offer three key benefits: maintenance of group cohesion, enhancement of decision accuracy compared with lone individuals and improvement in decision speed. In the absence of centralized control, arriving at a consensus depends on local interactions in which each individual''s likelihood of choosing an option increases with the number of others already committed to that option. The resulting positive feedback can effectively direct most or all group members to the best available choice. In this paper, we examine the functional form of the individual response to others'' behaviour that lies at the heart of this process. We review recent theoretical and empirical work on consensus decisions, and we develop a simple mathematical model to show the central importance to speedy and accurate decisions of quorum responses, in which an animal''s probability of exhibiting a behaviour is a sharply nonlinear function of the number of other individuals already performing this behaviour. We argue that systems relying on such quorum rules can achieve cohesive choice of the best option while also permitting adaptive tuning of the trade-off between decision speed and accuracy.     

Moving together: Incidental leaders and naïve followers

Elucidating whether common general mechanisms govern collective movements in a wide range of species is a central issue in the study of social behaviour. In this paper, we describe a new experimental paradigm for studying the dynamic of collective movements. Some sheep (Ovis aries) were first trained to move towards a coloured panel, in response to a sound cue. We present data comparing the behaviour of test groups composed of one of the trained sheep and 3 naïve sheep, and control groups composed of 4 naïve sheep. In the tests, for both test and control groups, sheep were observed for 20 min before the sound cue was delivered and the panel made visible. Before the sound, trained and naïve sheep were similar in terms of activity budgets, spatial distribution, social behaviour and spontaneous movement initiation. After the sound, trained sheep moved toward the panel and systematically triggered a collective movement in all test groups. The results suggest that any individual moving away from the group can elicit a collective movement. Our experimental protocol provides an opportunity to quantify mechanisms involved in group movements, and to investigate differences between species and the effect of social context on collective decision-making.     

How many for dinner? Recruitment and monitoring by glances in capuchins

Group members present physical and physiological differences according to their age, sex or social status, which could generate motivation differences among individuals during travel. In spite of these divergences of interest among individuals, the group succeeds more often than not in making a collective decision about departure time and which direction to take. To reach a consensus decision, animals should exchange information relating to characteristics of group movement through different communication channels. The main purpose of this study is to understand the function of behaviour patterns displayed during movements of white-faced capuchins (Cebus capucinus). We designed experiments in which we provoked collective movements involving a binary choice. During experiments, a video camera recorded the behaviour of each capuchin, which enabled us to determine which individuals displayed a behavioural pattern during travel and how this behaviour influenced the other group members. We found that looking backwards seemed to permit the recruitment of group mates during collective movement. This behaviour also seemed to allow the quantification of the number of followers, since the emitter modified its locomotion speed according to this number. In this preliminary study, we showed that visual behaviour was used to recruit and monitor group mates during collective movements and provided information on mechanisms involved in maintaining cohesion and coordination among group members during travel.     

Leadership in Moving Human Groups

How is movement of individuals coordinated as a group? This is a fundamental question of social behaviour, encompassing phenomena such as bird flocking, fish schooling, and the innumerable activities in human groups that require people to synchronise their actions. We have developed an experimental paradigm, the HoneyComb computer-based multi-client game, to empirically investigate human movement coordination and leadership. Using economic games as a model, we set monetary incentives to motivate players on a virtual playfield to reach goals via players'' movements. We asked whether (I) humans coordinate their movements when information is limited to an individual group member''s observation of adjacent group member motion, (II) whether an informed group minority can lead an uninformed group majority to the minority''s goal, and if so, (III) how this minority exerts its influence. We showed that in a human group – on the basis of movement alone – a minority can successfully lead a majority. Minorities lead successfully when (a) their members choose similar initial steps towards their goal field and (b) they are among the first in the whole group to make a move. Using our approach, we empirically demonstrate that the rules of swarming behaviour apply to humans. Even complex human behaviour, such as leadership and directed group movement, follow simple rules that are based on visual perception of local movement.     

Elucidating the significance of spatial memory on movement decisions by African savannah elephants using state–space models

Spatial memory facilitates resource acquisition where resources are patchy, but how it influences movement behaviour of wide-ranging species remains to be resolved. We examined African elephant spatial memory reflected in movement decisions regarding access to perennial waterholes. State–space models of movement data revealed a rapid, highly directional movement behaviour almost exclusively associated with visiting perennial water. Behavioural change point (BCP) analyses demonstrated that these goal-oriented movements were initiated on average 4.59 km, and up to 49.97 km, from the visited waterhole, with the closest waterhole accessed 90% of the time. Distances of decision points increased when switching to different waterholes, during the dry season, or for female groups relative to males, while selection of the closest waterhole decreased when switching. Overall, our analyses indicated detailed spatial knowledge over large scales, enabling elephants to minimize travel distance through highly directional movement when accessing water. We discuss the likely cognitive and socioecological mechanisms driving these spatially precise movements that are most consistent with our findings. By applying modern analytic techniques to high-resolution movement data, this study illustrates emerging approaches for studying how cognition structures animal movement behaviour in different ecological and social contexts.     

Group personality during collective decision-making: a multi-level approach

Collective decision-making processes emerge from social feedback networks within a group. Many studies on collective behaviour underestimate the role of individual personality and, as a result, personality is rarely analysed in the context of collective dynamics. Here, we show evidence of sheltering behaviour personality in a gregarious insect (Periplaneta americana), which is characterized by a collective personality at the group level. We also highlight that the individuals within groups exhibited consistent personality traits in their probability of sheltering and total time sheltered during the three trials over one week. Moreover, the group personality, which arises from the synergy between the distribution of behaviour profiles in the group and social amplifications, affected the sheltering dynamics. However, owing to its robustness, personality did not affect the group probability of reaching a consensus. Finally, to prove social interactions, we developed a new statistical method that will be helpful for future research on personality traits and group behaviour. This approach will help to identify the circumstances under which particular group compositions may improve the fitness of individuals in gregarious species.     

Distributed leadership in semifree-ranging white-faced capuchin monkeys

We investigated the initiation of group movements in white-faced capuchin monkeys, Cebus capucinus, with the aim of determining whether a single individual with high dominance status consistently leads movements or whether leadership is distributed between group members. The group studied was reared in semifree-ranging conditions. A multivariate analysis followed by univariate analyses demonstrated that leadership was not concentrated on a single individual in this species. All individuals could initiate a collective movement. Nearly half of group members regularly succeeded in recruiting at least three followers. Although both sexes had similar rates of start attempts, females succeeded more frequently than males. We found no significant effect of the dominance status on the percentage of successful attempts. The use of a slow speed, looking back towards the other group members, or trills by the initiator heightened the likelihood of success in group movement initiation. An initiator starting from a core position in a clumped group was more successful than one starting from an edge position in a clumped group or from a dispersed group. Furthermore, the probability of successful start attempts was higher when the group remained stationary for a long period. Leadership in white-faced capuchins appears to be distributed between group members rather than exclusively concentrated on high-ranking individuals. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.      

Moving calls: a vocal mechanism underlying quorum decisions in cohesive groups

Members of social groups need to coordinate their behaviour when choosing between alternative activities. Consensus decisions enable group members to maintain group cohesion and one way to reach consensus is to rely on quorums. A quorum response is where the probability of an activity change sharply increases with the number of individuals supporting the new activity. Here, we investigated how meerkats (Suricata suricatta) use vocalizations in the context of movement decisions. Moving calls emitted by meerkats increased the speed of the group, with a sharp increase in the probability of changing foraging patch when the number of group members joining the chorus increased from two up to three. These calls had no apparent effect on the group's movement direction. When dominant individuals were involved in the chorus, the group's reaction was not stronger than when only subordinates called. Groups only increased speed in response to playbacks of moving calls from one individual when other group members emitted moving calls as well. The voting mechanism linked to a quorum probably allows meerkat groups to change foraging patches cohesively with increased speed. Such vocal coordination may reflect an aggregation rule linking individual assessment of foraging patch quality to group travel route.     

Personality and Affinities Play a Key Role in the Organisation of Collective Movements in a Group of Domestic Horses

Understanding how groups of individuals with different motives come to daily decisions about the exploitation of their environment is a key question in animal behaviour. While interindividual differences are often seen only as a threat to group cohesion, growing evidence shows that they may to some extent facilitate effective collective action. Recent studies suggest that personality differences influence how individuals are attracted to conspecifics and affect their behaviour as an initiator or a follower. However, most of the existing studies are limited to a few taxa, mainly social fish and arthropods. Horses are social herbivores that live in long‐lasting groups and show identifiable personality differences between individuals. We studied a group of 38 individuals living in a 30‐ha hilly pasture. Over 200 h, we sought to identify how far individual differences such as personality and affinity distribution affect the dynamic of their collective movements. First, we report that individuals distribute their relationships according to similar personality and hierarchical rank. This is the first study that demonstrates a positive assortment between unrelated individuals according to personality in a mammal species. Second, we measured individual propensity to initiate and found that bold individuals initiated more often than shy individuals. However, their success in terms of number of followers and joining duration did not depend on their individual characteristics. Moreover, joining process is influenced by social network, with preferred partners following each other and bolder individuals being located more often at the front of the movement. Our results illustrate the importance of taking into account interindividual behavioural differences in studies of social behaviours.     

Individual-level personality influences social foraging and collective behaviour in wild birds

There is increasing evidence that animal groups can maintain coordinated behaviour and make collective decisions based on simple interaction rules. Effective collective action may be further facilitated by individual variation within groups, particularly through leader–follower polymorphisms. Recent studies have suggested that individual-level personality traits influence the degree to which individuals use social information, are attracted to conspecifics, or act as leaders/followers. However, evidence is equivocal and largely limited to laboratory studies. We use an automated data-collection system to conduct an experiment testing the relationship between personality and collective decision-making in the wild. First, we report that foraging flocks of great tits (Parus major) show strikingly synchronous behaviour. A predictive model of collective decision-making replicates patterns well, suggesting simple interaction rules are sufficient to explain the observed social behaviour. Second, within groups, individuals with more reactive personalities behave more collectively, moving to within-flock areas of higher density. By contrast, proactive individuals tend to move to and feed at spatial periphery of flocks. Finally, comparing alternative simulations of flocking with empirical data, we demonstrate that variation in personality promotes within-patch movement while maintaining group cohesion. Our results illustrate the importance of incorporating individual variability in models of social behaviour.     

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.     

Shared or unshared consensus decision in macaques?

Members of a social group have to make collective decisions in order to synchronise their activities. In a shared consensus decision, all group members can take part in the decision whereas in an unshared consensus decision, one individual, usually a dominant member of the group, takes the decision for the rest of the group. It has been suggested that the type of decision-making of a species could be influenced by its social style. To investigate this further, we studied collective movements in two species with opposed social systems, the Tonkean macaque (Macaca tonkeana) and the rhesus macaque (Macaca mulatta). From our results, it appears that the decision to move is the result of the choices and actions of several individuals in both groups. However, this consensus decision involved nearly all group members in Tonkean macaques whereas dominant and old individuals took a prominent role in rhesus macaques. Thus, we suggest that Tonkean macaques display equally shared consensus decisions to move, whereas in the same context rhesus macaque exhibit partially shared consensus decisions. Such a difference in making a collective decision might be linked to the different social systems of the two studied species.     

‘Captivity bias’ in animal tool use and its implications for the evolution of hominin technology

Animals in captive or laboratory settings may outperform wild animals of the same species in both frequency and diversity of tool use, a phenomenon here termed ‘captivity bias’. Although speculative at this stage, a logical conclusion from this concept is that animals whose tool-use behaviour is observed solely under natural conditions may be judged cognitively or physically inferior than if they had also been tested or observed under controlled captive conditions. In turn, this situation creates a potential problem for studies of the behaviour of extinct members of the human family tree—the hominins—as hominin cognitive abilities are often judged on material evidence of tool-use behaviour left in the archaeological record. In this review, potential factors contributing to captivity bias in primates (including increased contact between individuals engaged in tool use, guidance or shaping of tool-use behaviour by other tool-users and increased free time and energy) are identified and assessed for their possible effects on the behaviour of the Late Pleistocene hominin Homo floresiensis. The captivity bias concept provides one way to uncouple hominin tool use from cognition, by considering hominins as subject to the same adaptive influences as other tool-using animals.     

Grunt to go—Vocal coordination of group movements in redfronted lemurs

To remain cohesive as a group, individuals must coordinate their movements between resources. In many species, vocalisations are used in this context. While some species have specific movement calls, others use calls which are also employed in different contexts. The use of such multicontextual calls has rarely been studied quantitatively, especially during both the pre‐departure and departure period associated with collective decisions. We thus investigated the use of close calls (“grunts”) for the coordination of collective movements in four groups of wild redfronted lemurs (Eulemur rufifrons) in Kirindy Forest, Western Madagascar. Group movements are started by an initiator, who moves away from the group and is joined by followers setting out in the same direction. We observed collective movements and recorded vocalisations from 18 focal individuals (54 movements recorded for followers, 21 for initiators). The grunt rate of both initiators and followers was higher in the pre‐departure period than in a control context (i.e., during foraging). Initiators of collective movements grunted more often than followers in the pre‐departure period as well as at individual departure. The latter difference was due to the initiators’ grunt rates increasing earlier than the followers’ and remaining at an elevated level for longer. These observations suggest that grunts serve to coordinate the departure by indicating the individual's readiness to move. The pre‐departure period, in which both initiators and followers showed an elevated grunt rate, may provide the basis for a shared decision on departure time. The difference in initiator and follower call rates suggests that grunts may have a recruitment function, but playback experiments are required to test this potential function. Overall, our study describes how multicontextual close calls can function as movement calls, with changes in call rate providing a potential feedback mechanism for the timing of group movements. This study thus contributes to a more detailed understanding of the mechanisms of group coordination and collective decision‐making.