evolution and leadership 1 running head: evolution and

EVOLUTION AND LEADERSHIP 1

Running Head: EVOLUTION AND LEADERSHIP

What Evolution Teaches Us About Leadership:

Some Lessons From The Past

Mark Van Vugt

University of Kent

Robert Hogan

Hogan Assessment Systems

Invited paper prepared for end-of-year theory issue of Leadership Quarterly (2005)


Abstract

Taking a Darwinian perspective, this paper suggests that leadership is the product of an

evolved design for within-group coordination. Group coordination involves collective

decision-making about what to do, how, and when to do it. Leader-follower distinctions

emerge when two or more individuals adopt different but complementary decision rules in

order to engage in joint action. These decision rules are adaptive in that they are the product

of an evolutionary process, selecting for individuals who were better at solving problems of

social organization. This perspective produces various novel insights into the origins of

leadership and followership and offers an agenda for future research. Furthermore, an

evolutionary approach to the study of leadership serves to integrate a dispersed set of theories

and findings.


What Evolution Teaches Us About Leadership:

Some Lessons From The Past

“The UP [Universal People] have leaders, though they may be ephemeral or situational. The

UP admire, or profess to admire, generosity and this is particularly desired in a leader. No

leader of the UP ever has complete power lodged in himself alone. UP leaders go beyond the

limits of UP reason and morality. Since the UP never have complete democracy, and never

have complete autocracy, they always have a de facto oligarchy” (Brown, 1991; p. 138).

In “Human Universals” Donald Brown investigated what all people, societies,

cultures, and languages have in common and identified leadership as a human universal.

Similarly, in a review of modern and traditional hunter-gatherer societies, Boehm (1999)

concluded that, even in strictly egalitarian bands and tribes, band members exercise

differential influence on group decision-making. This is supported by the experimental social

psychological literature. When individuals form groups to work on collective tasks, leader-

follower patterns emerge quickly and spontaneously even when groups initially lack a leader

(Bales, 1951; Mann, 1959). Leadership functions might not even be restricted to the human

species. In studies of nonhuman primates and other social mammals, there have been

numerous observations of social interactions that resemble leader-follower patterns in

humans (e.g., Boehm, 1999; De Waal, 1996; Wilson, 1975).

For example, De Waal (1996) observed the following in a captive chimpanzee colony

at Arnhem Zoo in the Netherlands: “On one occasion, a quarrel between Mama and Spin got

out of hand and ended in fighting and biting. Numerous apes rushed up to the warring

females and joined in the fray. A huge knot of fighting, screaming apes rolled around in the


sand until Luit [the alpha male] leapt in and literally beast them apart. He did not choose

sides in the conflict, like others; instead anyone who continued received a blow from him”(p.

129).

Given the vast literature on leadership, accumulated by scientists in disciplines

ranging from psychology and management to political science and primatology, one might

expect there would be an effort to make sense of all the findings. Currently, however, there is

no such grand theory. As Hogan and Kaiser (2005) stated recently “The academic tradition

[in the study of leadership] is a collection of dependable empirical nuggets, but it is also a

collection of decontextualized facts that do not add up to a persuasive account of leadership”

(p. 171). This is partly due to the fact that the research is fragmented across different

disciplines, and there is little communication among them. What seems to be lacking is a

larger theoretical context in which to interpret and integrate findings across disciplines

(Chemers, 2000; Rost, 1991; Yukl, 1989).

Moreover, such as theory could also provide a more efficient guide for future research

This article presents a theory of leadership, inspired by Darwin’s theory of evolution via

natural selection and adaptation (Buss, 1999; Darwin, 1859; Dawkins, 1976; Van Vugt &

Van Lange, in press). This perspective offers a new way of looking at an old problem.

Evolutionary thinking can help integrate a massive body of empirical data into a coherent

conceptual framework and develop non-trivial hypotheses on leadership and followership

processes. Moreover, evolutionary theory is a good place to start because other models of

leadership drawn from economics, psychoanalysis, personality, social, cognitive, or cultural

theory must ultimately be consistent with evolutionary theory to be viable (Barrett, Dunbar &

Lycett, 2002; Buss, 1999). For example, any trait-based theory of leadership (Stogdill, 1974)

must ultimately account for how the particular traits evolved in the human species. Any

cognitive model, such as leader categorization theory (Lord & Maher, 1991), must explain


how leader prototypes evolved and why they are so influential. Finally, cultural theories of

leadership (House et al., 1999) must ultimately account for the human capacity for culture—

and explain how leaders in different cultures learn which style of leadership is effective

(Boyd & Richerson, 2005).

A Darwinian Perspective on Leadership: Asking the Why-question

Why do people defer to leaders? Tinbergen (1963) described four ways to answer the

why-question, each representing a different level of analysis from an evolutionary

perspective. First, one might ask what motivates a person to follow or lead another person at

any particular point in time. This is a question about the proximate or immediate causes of

behavior, and it is the question in which most psychologists are interested. Answers to this

question would refer to the motivation of individuals to lead or follow, and how they

perceive their relationships. For example, people might defer to a leader because they expect

doing so will be instrumental in securing such benefits as money, prestige, safety, or sexual

favors (Hollander, 1985). Alternatively, they might follow a leader because they are attracted

to and identify with the person (Bass, 1997; Hogg, 2001).

Second, one might ask if there is something about a person’s developmental history that

prompts him/her to lead or follow in a particular context? This question concerns the

developmental or ontogenetic causes of leadership and would interest developmental,

personality, and educational psychologists studying leadership (Ilies, Gerhardt, & Le, 2004).

For example, do leaders and followers have different personalities, and, if so, are their

differences based on heritable factors or different experiences in childhood or adolescence?

A third question concerns the evolutionary history of leadership. When and why did

this behavior arise? Through cross-species comparisons, one could determine whether

leadership can be found in other social species including the nonhuman primates. In addition,

studies of modern hunter-gatherer societies (Barrett et al., 2002; Boehm, 1999; Diamond,


1997) might reveal the development of leader-follower relations in humans. This is the

phylogenetic or historical cause of behavior, which concerns anthropologists and students of

animal behavior.

The final question concerns the functional or ultimate causes of leadership, which

interests evolutionary psychologists and evolutionary biologists. This perspective is linked to

the concept of natural selection. Does leadership confer reproductive advantages on leaders

and followers and does leadership contribute to the survival of these individuals and their

groups? If so, we might have an ultimate explanation of leadership.

Each of these questions analyzes leadership from a different perspective and these

perspectives are fairly independent from each other (although they are also interconnected).

For example, a developmental or ontogenetic understanding of why some people with

particular personalities and life histories are more likely to emerge as leaders tells us little

about whether or not leadership contributes to the reproductive success of individuals.

However, if we investigate leadership from different levels of analysis, we will derive a

better account of this phenomenon. For example, finding a proximate mechanism – that

people readily and strongly identify with their leaders (Bass, 1997; Hogg, 2001) – that

produce the ultimate reproductive benefits of followership – being a loyal follower enhances

one’s survival and reproduction abilities – strengthens an evolutionary account of leadership.

What must be avoided at all costs, however, is confusing one level of analysis with

another. For example, it is pointless to argue about whether people become leaders because

they are ambitious and energetic people, or because they want to enhance their reproductive

success. Both views may be right, but they concern a different level of explanation. Hence,

they complement each other, but they are not alternative explanations.

Finally, we should note that a Darwinian account of leadership and followership does

not assume that the decisions and cognitions associated with these roles are always


conscious. People may often not consciously be aware of why they take up a position of

leader or follower in their social groups. This is presumably because evolution has done the

thinking for us in such situations. Based on millions of years of evolutionary engineering,

humans are designed to act in certain ways in response to particular environmental stimuli

(Buss, 1999; Dawkins, 1976; Pinker, 2002; Wilson, 1975). Of course, there are individual

differences in the strengths of these inclinations, which accounts for why some people

emerge as leaders while others become followers. But these dispositions and their associated

cognitive, motivational, and behavioral elements may not be consciously acknowledged by

individuals. Their response is driven by the interaction between situational stimuli and

tendencies latent in a person’s genes.

Natural Selection and Adaptations for Group Life

How might natural selection account for the emergence of leadership in social

interaction? Natural selection operates via three principles (Darwin, 1859). First, individuals

in a species differ in terms of behavioral, physical, and physiological traits – this is the

Principle of Variation. Second, part of the variations in phenotype is heritable and will

therefore be passed on from one generation to the next (Principle of Inheritance). Third, some

of these variants allow individuals to compete more effectively for scarce resources

(Principle of Adaptation). As a consequence of these premises, some individuals enjoy

greater reproductive success because their traits give them an advantage: They are more

successful at mating, finding food, or avoiding predators. Their offspring will inherit their

successful traits and natural selection will have taken place. Through this process, organisms

thus become “adapted” to their environment (Barrett et al., 2002; Buss, 1999).

A crucial element of the human environment is other human beings (Alexander, 1979;

Sober & Wilson, 1998). Through natural selection, humans have acquired behaviors that

enable them to interact with other humans, sometimes in groups containing relatives and non-


relatives (Fehr & Fishbacher, 2003). Natural selection has produced a variety of adaptations

that facilitate group living. These include, for example, parental investment, food sharing,

collective defense, and behaviors that minimize the costs of group life, for example, conflict

and violence, struggles for status and dominance, and jealousy (Kenrick, Li, & Butner, 2003;

Kurzban & Leary, 2001; Van Vugt & Van Lange, in press). Leadership might have emerged

in human history as a way to maximize the benefits of group life while minimizing its costs.

Evolution, Leadership and Group Life

Many adaptive group challenges faced by the ancestors of modern humans involved the

coordination and organization of actions of smaller and larger social units. Human ancestors

lived in semi-nomadic groups of 30-50 members for about 95% of human history (Barrett et

al., 2002). They necessarily had to make collective decisions about what to do, when and

how to do it. For example, they would have needed to move frequently between waterholes

(Gamble, 1993). Given the risks of being attacked by predators or rival groups, it would pay

to stay together on the move. Yet, how would a group decide where and when to go? This

decision required some form of leadership, whereby someone would take initiative, and the

rest would follow. Similarly, there would be occasional conflicts between individual

members that threatened to spill over to the rest of the group (Chagnon, 1997; Diamond,

1997). Again, it would be useful for someone with the support of the rest to step in, resolve

the conflict and restore order in the group.

Problems of coordination and conflict resolution may have paved the way for the

emergence of a primordial distinction between leadership and followership. The evolution of

these social roles would have been unlikely unless there were clear benefits for all involved,

for example, in terms of improved group mobility, hunting success, food distribution, and

group defense and security. Especially in the case of followership, it is hard to argue why

people would voluntarily defer to another individual unless there were clear rewards. Once


these roles were “invented” they presumably became fine-tuned over time by being subjected

to natural selection forces (Boyd & Richerson, 2005). Furthermore, specific problems would

presumably have given rise to the emergence of different types of leaders. For example, an

intragroup conflict might persuade followers to endorse a neutral and impartial leader,

whereas an imminent attack from a rival group might lead to the endorsement of an

aggressive, coercive style leader such as in Big Man societies (Chagnon, 1997; McCann,

1997). We now turn to the literature to present a more detailed evolutionary account of

leadership.

Leadership: Lessons From a Distant Past

To indicate how an evolutionary perspective contributes to understanding leadership,

we offer six propositions (Table 1). We support each proposition with relevant theory and

research. Our list of evolutionary-based ideas about leadership only focuses on the main

issues. Some of our propositions can be derived from frameworks other than evolutionary

theory. However, it would be hard to find an alternative theory that accounts for this broad

range of leadership and followership phenomena.

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Insert Table 1 here

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Proposition 1: Leadership and Followership Are Adaptive Decision Rules

To determine the adaptive significance of leadership and followership as two separate but

complementary decision rules, we might consider two pieces of evidence (Buss, 1999;

Schmitt & Pilger, 2004). First, are there indications for differences in reproductive success of

individuals or, for that matter groups, that varied in the ability to establish good leader-

follower relationships? We believe that there is evidence that leader-follower patterns confer


certain benefits to people that would be otherwise unattainable. Second, is there evidence that

leadership is designed specifically to solve a set of group problems that would otherwise

remain unsolved? Again, we believe the evidence is there.

Reproductive Benefits Associated With Leadership and Followership

Historical data show that, in traditional societies, higher ranked males had more

offspring (Betzig, 1986). For example, one emperor of Morocco had more than 1000 children

with many different wives, and his harem was arranged in such a way as to maximize his

reproductive success (Betzig, 1986). There are also many proximal benefits associated with

leadership in contemporary society that might enhance reproductive success. For example,

leaders are generally wealthier than followers (Bass, 1990) and enjoy better health (Marmot,

2004). Experimental social psychological research shows that putting people into positions of

leadership enhances their mood and optimism and reduces their stress levels (Hardy & Van

Vugt, 2005; Keltner, Gruenfeld, & Anderson, 2003).

What about followers? An evolutionary account of followership requires reproductive

benefits for this role to emerge as well. Although followers are likely to extract fewer

benefits from group cooperation and coordination than leaders do, members of groups with

leaders should do better, overall, than members of groups without leaders. For example, if a

group had no leadership structure, it would have trouble moving to a new waterhole in time

when the old one dried up (Boehm, 1999). Similarly, when a group is attacked by rival

groups, deferring to a leader might be preferable to not having a leader at all (Diamond,

1997). Although there is no firm evidence to date to suggest that leaders enhance the

reproductive success of followers, case studies of work teams and army units clearly show

the benefits of good leadership in terms of enhancing performance, satisfaction and rewards

for followers (Hogan, Curphy & Hogan, 1994). Leadership also increases the contributions to

public goods from which all group members profit (Van Vugt & De Cremer, 1999).


The above suggests that leader and follower roles might have evolved through natural

selection forces operating at the individual level as well as the group level – the differential

success rates of groups competing with each other. Although group selection models have

been criticized by evolutionary scientists in the past, recent treatments of group selection like

D. S. Wilson’s multilevel selection theory (Sober & Wilson, 1998) and Boyd and

Richerson’s (2005) cultural group selection model have been endorsed and are gaining

influence in the scientific community. This is not the place, however, to discuss these

models.

Evidence of Special Design

A second line of evidence suggesting that leader and follower relations are adaptive

concerns whether these behaviors show evidence of “special design” (Dawkins, 1976;

Williams, 1966). This judgment is based on criteria involving the complexity and efficiency

of the attribute or behavior and its appropriateness for solving problems that humans have

faced through evolutionary time. Consider, for example, humans’ highly specialized tool

making ability. Making axes and spears allowed our ancestors to hunt large animals and

defend against large predators in a hostile savannah environment (Gamble, 1993). It is an

ability that all humans possess. In contrast, gorillas who live in regions without predators and

eat plants, do not need tools and therefore have no tool making ability.

Is there any evidence that leadership is the product of special design? To answer this

requires integrating evidence from many different disciplines (Schmitt & Pilcher, 2004).

First, we have noted that leadership is a universal attribute of human societies (Brown, 1991),

including modern hunter-gatherers whose living arrangements resemble those in which our

ancestors lived (Boehm, 1999; Diamond, 1997). Second, there are patterns of leader-follower

relations among chimpanzees, our closest living relative (De Waal, 1996), baboons (Dunbar,

1983) and other social mammals like dolphins (Packer, 1983). Third, adaptive challenges in


human groups require individuals to coordinate their activities, for organizing a hunt or group

defense. Moreover, evidence from game theoretical simulations show that stable leader-

follower arrangements emerge when two (or more) individuals coordinate their actions but

have slightly different preferences for what activity to take, for example, to rest or eat

(Browning & Colman, 2004; Van Vugt, 2005).

Fourth, demographic studies show a correlation between leadership and reproductive

success (Betzig, 1986) health and wealth (Marmot, 2004). Fifth, evidence from psychological

experiments suggests that, in groups of strangers, leader-follower differences quickly emerge

(Hemphill, 1950). Sixth, people process information about leaders automatically and have

prototypes of leadership that are readily available and difficult to change (Lord & Maher,

1991). Finally, evidence from behavior genetics research suggests that traits such

extraversion, ambition, and boldness, which are related to leadership emergence, show some

degree of heritability (Ilies et al., 2004; Rushton et al., 1986)

Summary. Obviously we need more research to establish the adaptive significance of

leadership and followership behaviors and to examine the specific evolutionary trajectories

of these social roles. But the evidence so far is encouraging. Leadership and followership

might have evolved as special designs for facilitating group action and coordination. Yet we

still know very little about the evolutionary advantages of becoming a follower nor about the

role of group selection forces in the emergence of leadership.

Proposition 2: Leadership and Followership are Strategies for Group Coordination

Evolutionary game theory (Maynard-Smith, 1982) suggests how leadership might have

emerged in our species’ history. Evolutionary game theory is like classic economic game

theory (Luce & Raiffa, 1957), only the pay-offs are in terms of reproductive rather than


economic costs and benefits (although they might be related, of course). Evolutionary game

theory has been used to model a range of social behaviors like cooperation and competition,

and suggest how these behaviors evolved (Maynard-Smith, 1982). The goal is to determine

which strategies succeed when pitted against others as well as themselves (Axelrod, 1984).

Successful strategies spread through a population by virtue of the superior decision rules they

adopt in relevant situations, whereas unsuccessful strategies become extinct—and this

resembles the process of natural selection. How do leadership and followership fare as game

strategies?

The Leader Game

Because leadership involves coordination, initiative-taking, risk, and responsibility

(Bass, 1990), it is appropriate to use a group coordination game as a model for the evolution

of leadership (Rapoport, 1967). The simplest version of the leader game involves two

players, 1 and 2, each with two strategies, say to Wait or Move (Figure 1). There are four

cells in Figure 1, each with two outcomes (pay-offs in reproductive terms). The first pay-off

is always for player 1 and the second for player 2. For example, if both choose to wait, they

receive a zero pay-off, and if both choose to move simultaneously, they receive a negative

pay-off of -100 – for example, they could bump into each other if they move at the same

time. They can solve this problem only if they coordinate their actions such that one player

waits for the other to move and then follows. Moving is the most attractive option (a pay-off

of 200), provided that the other waits (who then gets 100). However, moving is also the risky

option because if both move at the same time, both players receive -100. If the players

coordinate their actions, they can both win, but the player who makes the risky move (200)

does better and emerges as the leader and the player playing the cautious option (100), to

wait, becomes follower.


As a real-world example, imagine two individuals on a hostile savannah who need to

stay close together for safety. At some point, they must decide to stay where they are, or

move on for food. They will differ in how hungry they are; the hungriest individual might

make a move at some point and thus obtains the best pay-off because he eats when he is

hungry. This means that the less hungry individual must follow, otherwise they end up at

different locations and lose each other’s protection. Another example is a village that is being

attacked by men of a neighboring clan. The village members must decide whether to defend

themselves or run away to hide. Given that they must stay together for protection, the best

option is for one of the members to decide what to do, fight or flight, and for the rest to

support that decision. The first person then emerges as the group leader, and receives a

higher pay-off because he can impose his preference on the others.

In game theory, (Move, Wait) and (Wait, Move) are referred to as the equilibriums of

this game. Thus, once the players’ strategies settle into either of these cells, neither player has

an incentive to choose a different strategy. Consequently, the strategies become stabilized in

a population, because they cannot be improved by any alternative strategies. From this, one

would expect behavioral adaptations for leadership and followership to become increasingly

complementary and fine-tuned. Furthermore, the logic of the leader game dictates that a

leader-follower pair (Move, Wait) would always receive a higher pay-off than any dyad of

leaders (Move, Move) or followers (Wait, Wait). This implies that these roles may have co-

evolved over time, producing a stable mixture of leaders and followers in a population at

large (Frank, 1988; Maynard-Smith, 1982; D.S. Wilson et al., 1996).

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Insert Figure 1 here

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Evolutionary Hypotheses


Conceptualizing leadership and followership as complementary strategies in a

coordination game leads to a number of testable predictions about leadership. We have done

that elsewhere in greater detail (Van Vugt, 2005). Table 1 summarizes the main predictions

about leadership from a game theoretical perspective. For example, we predict that

individuals who are dispositionally more inclined to take the initiative, are more likely to

emerge as leaders in any given situation. This is clearly supported in the psychological

literature (Gough, 1984). Also, emergent leadership should correlate with intelligence

because individuals would want to wait until a wise person makes a first move and then

follow him or her (Bass, 1990). Leaders, according to this analysis, are also expected to be

wealthier because their relative pay-off is expected to be higher (in Figure 1 it is 2x the

followers’ pay-off). This is supported overwhelmingly in the literature (Bass, 1990). Yet

signs of generosity should and do matter as well (De Cremer & Van Knippenberg, 2002),

because before people become followers, they need to be sure that a leader is willing to share

the benefits with them (according to Figure 1, they should expect at least 100 in pay-offs).

________________________

Insert Table 2 here

________________________

Summary

Evolutionary game theory provides an account of how strategies producing leadership

and followership behaviors might have evolved and spread through the population (at the

expense of alternative strategies) and become increasingly fine-tuned to each other. Also, it

makes a number of predictions, some of which deserve further testing, about the likelihood

that people with particular personalities emerge as either leaders or followers. Furthermore, it

explains what particular set of attributes followers might be looking for in their leaders.


Proposition 3: Leadership Emerges in Response to Group Threats and Opportunities

As indicated earlier, there are certain costs associated with following another person, because

followers give up their preferred outcome (200 in Figure 1) in order to coordinate their

actions with leaders. To offset the costs, there must be substantial benefits associated with

group coordination before a leader-follower structure emerges. From an adaptionist

framework (Cosmides & Tooby, 1992; Schmitt & Pilcher, 2004), designs for leadership and

followership should be context-specific. Because these behaviors have evolved to solve

specific problems associated with group life, they should be elicited only when organisms

perceive situations as relevant to these problems.

What are these situations? Kenrick et al. (2003) identified six social problem domains

in response to which group-relevant cognitive and behavioral adaptations might have

evolved: (1) coalition formation – forming and maintaining alliances, (2) self-protection --

protecting oneself against threats to survival and reproduction, (3) status – gaining respect

from and power over others, (4) mate choice – obtaining a partner to insure one’s fitness, (5)

relationship maintenance – maintaining a bond with a partner, and (6) parental care –

promoting the survival and success of one’s off-spring. The first three domains are

particularly relevant to the evolution of leader and follower behaviors.

Coalitions

Several authors have discussed the adaptive value functions of cooperative alliances

(Baumeister & Leary, 1995; Diamond, 1997). For example, hunting large game provides the

ingredients for a healthy and nutritious diet, and sharing food provides an insurance policy

for survival through bad times. Trade with benevolent others provides access to otherwise

unavailable resources (food, tools, status objects). Finally, political coalitions facilitate

gaining and holding power in a group as well as dominating other groups (Sidanius & Pratto,


1999). Coalitional behavior requires that some individuals take the lead and others follow.

For example, to organize a group to hunt large animals, it would be useful if one individual

would tell the rest what positions they should take and follow his orders. By engaging in

collective action, everyone would benefit, although some individuals would clearly benefit

more than others, for example, because they are more instrumental in organizing the hunt.

Self-protection

Self-protection is another problem domain in response to which leader-follower

adaptations might have emerged. Here it is useful to distinguish between intra- and

intergroup threats (Van Vugt, 2004). The social life of our hunter gatherer ancestors involved

constant conflict and violence and homicide was a leading cause of death (Buss, 1999;

Chagnon, 1997). To minimize conflict in small bands, peacemakers are needed to prevent

fights, particularly when they threaten to incorporate the rest of the group. Because such

interventions depend on the support of other group members, this would have encouraged

followership behavior. Groups were in competition with one another. Minimizing conflict

within the group improves its viability vis a vis other groups. Interestingly, peacemaking

behavior is found in most of the nonhuman primates, suggesting that it has ancient origins

(Boehm, 1999; DeWaal, 1996; Wilson, 1975).

Intergroup threats would have also created selection pressures for the evolution of

leadership and followership. The main threat here would be attacks from outgroup members

on oneself or other members of one’s ingroup. Alexander (1979) has suggested that

intergroup rivalry and warfare acted as a strong selection pressure for the evolution of a

whole range of social behaviors, including altruism towards ingroup members, conformity,

morality, and perhaps even religion (D.S Wilson, 2002). Leadership may have emerged

through intergroup pressures as well. Because defeat in an intergroup battle would mean

almost a certain death for oneself and one’s relatives, it would be functional to act as a unit to


fend off such threats (Boehm, 1999; Chagnon, 1997; Van Vugt, 2004). Good leadership and

disciplined followership would be vital to group and personal survival and thus may have

evolved for this particular purpose.

Again, this adaptation might have an ancient origin. Boehm (1999) observes an

encounter between rival groups of chimpanzees in the colony at Gombe: “Goblin [the alpha

male and leader] moves forward quickly to a vantage spot to peer across the valley and

Mustard now emulates him. As Goblin (number one), Satan (number two) and Evered

(number three) scan the valley, they break off several times to look at one another quickly.

After nearly 60 seconds, Goblin suddenly makes his decision and begins to vocalize and

display, The entire group, which includes adolescents Freud and Beethoven, immediately

follows suit. And the result is the usual one: Both groups vocalize and display ferociously

the slowly retreat into their home ranges.” (p. 28).

Status

The adaptive problems, coalition formation and self-protection, explain why individuals

might be keen to follow another individual. But what is in it for the leaders? Why would they

take on dangerous roles as peacekeepers or warriors? Their benefits derive presumably from

the emergence of a hierarchy in human groups, whereby those at the top enjoy greater

rewards than those at the bottom. Although human hierarchies are much flatter than those of

even the most egalitarian non-human primates, high ranking individuals still have more

respect and status (Boehm, 1999; DeWaal, 1996). As a result, humans are almost certainly

programmed to seek status gains and avoid status losses (Buss, 1999; Kenrick et al., 2003).

Leadership confers status, which allows access to such resources as wealth, health, and,

ultimately, more and better mates (Chagnon, 1997; Perusse, 1993).

Opportunities for Leadership


Is there any evidence that leadership emerges in response to the various threats and

opportunities of group life? The answer appears to be yes. First, group members are more

willing to endorse a leader when their shared resource is being depleted (Samuelson &

Messick, 1986; Van Vugt & De Cremer, 1999). Second, people show leader and follower

behaviors when faced with natural crises such as earthquakes (Baumeister et al., 1989). The

Robbers Cave experiment shows that when a group faces competition, one of its first

activities is to select a leader (Sherif, 1966). Finally, leaders sometimes strengthen their

position by inducing intergroup competition – for example, Margaret Thatcher and the

Falkland War (Rabbie & Bekkers, 1978).

An evolutionary perspective also suggests that different types of leadership emerge in

response to different group challenges. Protection against aggressive outgroup members

encourages followers to endorse aggressive and coercive leaders, presumably because they

are more effective warriors. McCann’s (1997) US-presidential study confirms this. The

public voted for more aggressive candidates when faced with an external threat. Anecdotal

evidence also supports this conclusion. The tough-minded Winston Churchill only became a

viable candidate for the role of Prime Minister of Britain when war broke out, and when it

ended, the public no longer supported him. Thus, aggressive and coercive forms of leadership

may be endorsed in crisis situations, but may not be tolerated otherwise.

Leaderless groups

An evolutionary analysis suggests that leadership is unlikely to emerge when groups

face relatively simple coordination problems, there are no conflicts of interests, and there are

no pressures to solve the problem quickly. This hypothesis is supported by the literature on

leadership substitutes (Kerr & Jermier, 1978). For example, when group activity is routine

and predictable, no leadership structure is likely to emerge (Comstock & Scott, 1977).

Friendship groups and other small cohesive units usually do better without a leader, because


power differences undermine group cohesion (Haslam et al., 1998). Finally, improvements in

technology and communication, such as the internet, may make leadership irrelevant because

group members can coordinate their actions without central control (Kerr & Jermier, 1978;

Wright, 2000).

Summary

An evolutionary perspective assumes that leadership and followership are associated

with specific problems of group life - coalition formation to acquire valuable resources, and

self protection against ingroup and outgroup threats. In combination with a universal drive

for status, individuals emerge as group leaders when there is a need for direction and

coordination. Still, simple coordination problems can be solved without leadership

sometimes.

Proposition 4: Leadership Styles Are Flexible

There are phylogenetic similarities in the social behavior of human and nonhuman primates.

There are important phylogenetic differences as well. Leadership in gorillas, chimpanzees

and bonobos is tyrannical, but human leadership varies from dictatorship to egalitarianism

(Boehm, 1999). The alpha male chimpanzee, for example, bullies the lower ranked males and

harasses the females in the group. However, this behavior is less extreme than that of Stalin,

Hitler, or Mao, who were willing to kill anyone they disliked. Thus, a theory of leadership

needs to explain the variability in leadership styles found in human populations as well as the

unique nature of human leadership.

Compared to other social species, humans show great behavioral flexibility (D. S.

Wilson et al., 1996). Being flexible is adaptive in environments that are relatively unstable in

terms of climate, risks of predation, and availability of resources such as food and water. For


example, when food is scarce, it would be useful for a group to fission into smaller groups,

whereas when food is abundant, it is useful to merge into a single larger group. Fission-

fusion tendencies are characteristic of human social life, and they require flexibility in

organizational structure (Diamond, 1997; Hart & Van Vugt, in press). Similarly, different

leader structures should emerge in response to different group challenges. To understand

variations in leadership, we must look at the specific contextual dynamics in which

leadership structures emerge. This can be clarified by using a game theoretical account of

leadership.

Autocratic versus Democratic Leadership Styles

The distinction between autocratic and democratic leadership is an old one. Autocratic

leaders decide group matters without consulting their followers, whereas democratic leaders

involve followers in decision-making and try to build consensus (Bass, 1990). Followers

endorse autocratic leaders when they are desperate, for example, when faced with war,

famine, or drought when the group must move together quickly (Vroom & Yetton, 1973).

Furthermore, the support for an autocratic leader should be greater when deviants threaten to

undermine group unity and group unity is necessary to fend off threats (Van Vugt & De

Cremer, 1999). Of course, once leaders have such powers, it is difficult to take them away

because, for leaders, they make life so much easier (Kipnis, 1972).

When individuals are reluctant followers and they have options elsewhere, leaders must

generally become more democratic. To compensate for the losses that followers incur in

autonomy and wealth, leaders must share the decision-making with them so that they can

influence the direction of the group action or decision. The democratic leader is more like a

“primus inter pares.” We would expect democratic structures to emerge when two conditions

are met. First, there should be obvious benefits associated with group coordination so that

everyone, leader and followers, has a stake in acting together to achieve their goals. Second,


individuals must have alternatives to following a particular leader, for example, following a

different person, acting independently, or joining a different group. For example, in medieval

times, noblemen involved mercenary soldiers in their decision-making, because they much

needed their support and did not want them to join a rival army (Betzig, 1986). Exit

opportunities give followers some power in negotiating with their leader (Van Vugt et al,

2004). The result is a shift towards a more democratic structure. Nonetheless, leaders can

stop the democratic drift by creating group threats—e.g., wars—which foreclose the options

for group members and strengthen the case for an autocratic system (Rabbie & Bekkers,

1978).

Figures 2a and 2b show the pay-off structures associated with the emergence of

autocratic or democratic leadership, respectively. In Figure 2a, to Move is the dominant

strategy for player 1. He is not dependent on the strategy of player 2 to achieve good

outcomes for himself (either receiving a pay-off of 200 or 100 by choosing to move).

Therefore he should always make a first move and by doing so leaves player 2 no other

option but following. This is essentially what autocratic leadership is about. In contrast,

Figure 2b shows a situation, like in Figure 1, in which the players have to coordinate their

actions to achieve good outcomes for both of them. But in contrast to Figure 1, the costs of

not coordinating are smaller, putting pressure on leaders to share more with followers. This is

essentially a model of democratic leadership

____________________________

Insert Figures 2a and 2b here.

____________________________

Task or Relationship Leadership

From a Darwinian framework, different adaptive problems underlie the difference

between task-oriented and relationship-oriented leadership (Fiedler, 1967; Hemphill, 1950).


Task leaders focus on group goal achievement, whereas relationship leaders are concerned

with establishing and maintaining good relations within the group. This difference reflects

the extent to which the interests of leaders and followers overlap (Figure 3a) or conflict

(Figure 3b). For example, in organizing a small group for a collective hunt, individual goals

should be congruent. Hence, the leader primarily faces a problem of coordinating activities

so as to maximize the success of the group but there is no conflict between leader and

followers about what to do (i.e., each party receives a pay-off of 150). Yet, when a band of

nomads must decide where to move next but they disagree about the direction of the move –

thus following is costly (see Figure 3b) -- then a leader who has good relations with followers

should be more effective. This is corroborated by research with Fielder’s (1967) contingency

model which suggests that in situations that are moderately but not dismally unfavorable, a

relationship leader is most persuasive.

___________________________

Insert Figures 3a and 3b here

___________________________

Transactional versus Transformational Leadership

Transformational leaders inspire followers with charisma and vision, and persuade

them to move beyond their immediate self-interest (Bass, 1990). Transactional leaders, in

contrast, appeal to followers’ self interest by providing them with favorable outcomes in

return for tangible and symbolic rewards (social exchange; Hollander, 1985). Again, it is

instructive to ask what adaptive problems might give rise to these two forms of leadership.

Transactional relationships (Figure 4a) follow pretty closely the pay-off matrix of the Leader

game (Figure 1). In exchange for their commitment to the leader, followers get rewards and

the higher the rewards relative to those of non-followers or leaders, the more dedicated

followers will be.


Transformational relationships change the pay-off matrix so that individuals believe

they are better off by following the leader. Thus, followers may think that their outcomes are

actually better than what leaders receive. Leaders seemingly turn followers into leaders

(Figure 4b), which explains the enthusiasm of followers supervised by transformational

leaders (Bass, 1997). Bass (1997) suggests that “the followers experience a magnetic

attention that transcends their usual experience. They become zealots and leaders in their

own right” (p. 40).

Certain conditions must be met for transformational leadership to occur. First, the

benefits associated with group coordination must be such that there is no attractive

alternative course of action for individuals. This might explain why transformational

leadership is more likely to emerge when a group is threatened and followers are uncertain

about what to do (Bass, 1990; Yul, 1999). Hence, they become willing to follow someone

with the vision and willingness to take the initiative.

Second, the leader must persuade followers that there are no costs, only benefits accrue

to coordinating their actions with the leader. Not surprising, such leaders are great actors

(Gardner & Avioli, 1998), and probably score high on a Machiavellian scale. One strategy is

to invent an ideology that justifies following a particular leader. Religions and national

identities provide such justifications; consequently, charismatic leaders often emerge in

religious groups, political settings, and the army (Bass, 1990; Burns, 1978). A second

strategy is for leaders to display self-sacrificial behaviors, for example giving up status,

giving away wealth, or even risking their lives so that followers think they are as well or

better off than their leader (De Cremer & Van Knippenberg, 2003). Not surprising, Burns

(1978) labels this as heroic leadership and the game depicted in Figure 4b is known as the

Hero game (Rapoport, 1967). In this game, the first to move – the leader – actually gets a

lower pay-off (100) than the people who wait – the followers (200). Indeed, no one is more


charismatic than a leader who suffers for his/her people, think of Jesus of Nazareth, Thomas

Beckett, Joan of Arc, or, a more recent example, Nelson Mandela.

______________________________

Insert Figures 4a and 4b here

_____________________________

Summary

An evolutionary perspective illuminates the variability in leadership styles in human

groups. Different problems give rise to different strategies for both leaders and followers. For

example, autocratic leadership is associated with a dominance game, in which leaders can

play their preferred strategies with little regard for the interests of followers. Democratic

leadership is associated with a game in which people are reluctant followers because they

have reasonably attractive alternatives beyond coordination with a leader. This enhances their

negotiation power with the leader. Transformational leadership in essence transforms the

leader game such that followers believe that their outcomes are as good as if not better than

what the leader gets. Thus, they turn into dedicated followers.

Proposition 5: Leadership Can Be Both a State and Trait

A common misconception about evolutionary explanations of behavior is that they imply

genetic determinism (Kenrick & Simpson, 1997). This is wrong. For example, a boy might

have genes that make him a potential leader, but if he is dominated by his older brothers or

bullied by his class mates, this inclination might not materialize. Similarly, a person

somebody with no predisposition for leadership might emerge as leader in an emergency,

when he is the first to notice a danger (Baumeister et al., 1989). All that can be said is that if


there is a genetic basis for leadership, then an individual with the right combination of genes

is statistically more likely to emerge as leader.

Furthermore, a key adaptive feature of human behavior is its flexibility or, to use the

proper term, phenotypic plasticity (Barrett et al., 2002; D.S. Wilson et al., 1996). This

flexibility enables humans to cope with a variety of environments. The complexities of group

life suggest that individuals need many strategies for social interactions, from aggression to

altruism, from conformity to deviance, and from leading to following. Phenotypic plasticity

makes social life possible. Without it, human groups would be like beehives, with each

individual assigned by birth to a specific role in the group (Wilson, 1975).

There might be a genetic basis for leadership, but environmental factors determine

whether this inclination results in the phenotypic expression of leadership. Ultimately,

empirical research must choose among personality, situational or interaction models of

leadership. Although there is a lot of anecdotal evidence (Simonton, 1994), there are only a

handful of studies on the heritability of leadership traits (Ilies et al., 2004; Rusthon et al.,

1986) but none on followership traits as far as we know. Twin studies are the appropriate tool

for evaluating the genetic basis of leadership. Recent studies show strong heritability

coefficients for traits associated with leadership, such as extraversion, intelligence, ambition,

and empathy (Rushton et al., 1986). Furthermore, there is a relationship between children’s

desire for leadership roles and the ambition levels of their parents (Bass, 1990; Klonsky,

1983). In addition, people who have been rated as leaders at college are more likely to

emerge as leaders later on in life (Russell et al., 1986). Finally, when group composition and

group tasks change, the same individuals still emerge as leaders (Kenny & Zaccoro, 1983;

Zaccoro, Foti, & Kenny, 1991).

Ontogenetic factors moderate the influence of traits on leadership. For example, in

children between the age of 3 and 8, the physically stronger children exercise more influence.


By the age of 12, friendly children are more likely to be leaders of their peer group (Barner-

Berry, 1982). Both physique and friendliness are influenced by genes, but they predict

leadership in a different age range. Trait models of leadership are further qualified by cultural

factors. For example, Hofstede (1980) presents evidence for a more hierarchical leadership

style among East-Asian and Middle-East managers than among Northern European

managers. This suggests that an authoritarian person is more likely to be a leader in Japan

than in the United Kingdom. Even within the same culture, perceptions of leadership

potential can change. In an analysis of US-elections, McCann (1992) found that the

electorate voted for a more aggressive and conservative president when the US faced either a

domestic (economy) or international crisis (war).

Summary

To summarize, the evidence suggests that leadership is determined by both heritable

and non-heritable factors, but the exact weighting of the two is yet unclear. More research is

needed to establish the relative importance of these two sources of influence on leadership.

We know relatively little about the determinants of followership, however. The evidence also

suggests that flexibility or adaptability in leadership style may be key to leadership

effectiveness, and that preferences in leadership style can change as a function of age,

culture, and the specific environment that requires leadership.

Proposition 6: Leader-Follower Relationships are Fundamentally Ambivalent

Most theories assume that leaders and followers work together to pursue shared goals

(Chemers, 2000; Fiedler, 1967; Hogg, 2001). However, a Darwinian perspective suggests

that, in reality, there are frequent conflicts between leaders and followers. The criteria in

evolutionary theory is relative reproductive success, that is, how well individuals do,


compared to others, in terms of propagating their genes into subsequent generations (Barrett

et al., 2002). Consequently, in nature, competition is likely to be the rule and cooperation the

exception. Humans are slightly odd in that the benefits associated with cooperation are so

substantial that we have evolved into an ultrasocial species. Nevertheless, human nature is at

best ambivalent, torn between cooperation and competition (Boehm, 1999; Hogan, 1982).

Leader-follower relationships are an example. These adaptations have evolved to reap

the benefits of coordinated action. At the same time, coordinated actions frequently produce

unequal outcomes such that leaders benefit more than followers (Figure 1). This creates

constant tension between leaders and followers. Leaders want to maintain or increase their

relative benefits, whereas followers want to reduce their relative losses. This creates selection

pressures for self-serving leadership behaviors as well as selection pressures for undermining

the self-serving activities of leaders.

Staying in Power

There are at least five strategies that leaders use to maintain their privileged positions

(Diamond, 1997). The first two are prosocial, because they benefit both leaders and

followers. First, leaders can strengthen their position by redistributing the gains of collective

action generously and fairly among followers. Anthropological research suggests that, across

many different societies, generosity is one of the most important traits in a leader (Boehm,

1993; Brown, 1991). Interestingly, in hunter-gatherer societies the head man gets to eat fist

when large game is killed but all members of the hunting party eat equivalent portions

(Boehm, 1999). Social psychological studies have also shown that promoting fair outcomes

increases the legitimacy of and support for leaders (Thibaut & Walker, 1975).

A second prosocial strategy involves providing security for followers. Group members

are always at risk for attack from hostile outgroups and unhappy ingroup members (Chagnon,

1997). Leaders who effectively reduce these threats tend to receive greater respect and


endorsement. For example, in a recent survey, Winston Churchill was voted the Greatest

Briton of all times, primarily for his contribution as Prime Minister during the Second World

War. In post-war France, Charles De Gaulle, the leader of the French resistance, won this

election (Hardy & Van Vugt, 2005). In addition, leaders must also be seen to deal effectively

with violence, cheating, and other antisocial behaviors (De Cremer & Van Vugt, 2002; Van

Vugt & De Cremer, 1999). Because conflict resolution is so important for the cohesion of a

group, a leader’s impartiality, honesty and neutrality are seen as essential leader traits (Tyler

& Lind, 1992).

Using evidence from many hunter-gatherer societies, Boehm (1993) concluded that “a

desirable leader is likely to be of high social standing, generous, wise, experienced,

successful in what he does, and self-assertive in general. It also helps if he is fair minded,

tactful, reliable,, morally upright, apt at resolving disputes and a competent speaker.”

Nonetheless, it is often easier for leaders to maintain their privileged position by adopting a

more selfish strategy. The most obvious strategy is to distribute the outcomes from

coordinated action to their allies. Although leaders’ corruption and nepotism undermine

societies, there is little systematic research on the matter. Anecdotal evidence from

investigative journalists suggests that the practice is widespread in some countries. Pitt

(1985) reports that over 90 % of company managers in South Africa had observed colleagues

taking bribes from suppliers.

Another more obvious exploitative strategy is to physically dominate and coerce

followers. Once people emerge as leaders and gain a privileged position, it is easy to

strengthen this position by limiting the choices of followers. This is often justified by

appealing to civil disorder, which enhances the perceived need for coercive leadership (Bass,

1990). In essence, tyrants do what they want with little regard for the needs and concerns of

followers (Figure 3a), and those that do not comply with their dictates are punished. In an


ingenious laboratory experiment, Kipnis (1972) showed that when leaders of small groups

were given coercive powers (penalizing poor performing members), they were more likely to

use this power, regardless of the performance of the group members. This finding is

consistent with the old saying that “power corrupts, but absolute power corrupts absolutely.”

A more subtle but selfish strategy is to impose an ideology that justifies one’s position

as leader. Throughout history, leaders have adopted religions which give them and their

descendants privileged access to resources -- the divine right of kings. In traditional

chiefdoms, the chief either combines the offices of political leader and priest, or he supports

an elite priesthood who justify the chief’s position (Diamond, 1997). Modern monarchies use

the same methods to justify hereditary leadership. Furthermore, charismatic leaders gain

influence by their connection to a religious or political ideology. Why do followers accept

this farce? The evolutionary anthropologists Tiger and Fox (1971) note that “We are so

committed to the notion of hierarchy that even our mightiest leaders must subserve to

something or someone” (p. 52).

Leveling The Power of Leaders

Our ambivalence model suggests that, in order to reap the benefits of group

coordination, followers would have developed a range of counteracting strategies to monitor

their leaders, control them, and, if necessary, reduce their power position. These strategies –

which can be referred to as “leveling” mechanisms (Boehm, 1999) -- presumably have an

evolutionary basis as well, because they enable individuals to reap the benefits of coordinated

action without paying excessive costs (e.g., losing autonomy, being exploited ). As in a

nuclear arms race, follower strategies have co-evolved with leader strategies in order to

undermine the exploitative intentions of leaders. The result is an evolutionary stable

equilibrium that balances the benefits to leaders and followers. Yet, in any particular

environment, the balance can shift in favor of the leaders or the followers, and this may


explain the variability in leadership structures that are found across human groups and

societies today.

What strategies do followers use to curtail the power of leaders? Drawing on Boehm

(1999), we suggest the following. The first tactic involves undermining the authority of

leaders by gossiping privately about them or by holding meetings in which the leader’s

decisions are scrutinized publicly. Leaders’ decisions in hunter-gatherer and Big Man

societies are largely overridden by group decision-making, often in public meetings and

through the manipulation of minorities by the majority (Meggitt, 1977). Human societies

across the world show remarkable consistency in the use of private gossip and public debate

to control the decision-making of leaders.

A second control strategy is to criticize and ridicule leaders openly. In African and

South American tribes, for example, people shout down leading hunters who become

overassertive (Maybury-Lewis, 2000). In the Philippines, when a chief misbehaves, he may

be publicly criticized and if he tries to give commands, he is rebuffed (Freeman, 1970). In

modern society, it is largely the role of the media (TV, newspapers, and internet) to criticize

self-serving leaders, who are often caricatured in cartoons.

Third, group members may simply disobey self-aggrandizing leaders. Among the

Bedouin, leaders who tried to impress European visitors by exercising their authority were

simply disobeyed. Freeman (1970) reports that Phillipine chiefs who issue commands as

opposed to suggestions are ignored. In traditional African societies, leaders cannot order

people to do things that they do not want to do. Disobeying leaders can be effective, because

they are sanctioned without being replaced.

Fourth, followers ostracize or shun exploitative leaders. Moore (1972) reports that,

when aggressive tribal leaders start a feud, the tribesmen distance themselves by no longer

regarding them as one of theirs. This allows rival groups to kill the ostracized leader with


impunity because he is no longer defended by his group. The social psychological literature

confirms that ostracism is a severe group sanction with implications for the mental and

physical well-being of the ostracized (Williams & Sommer, 1997).

Fifth, followers can depose selfish leaders and turn them into ordinary group members.

Field research shows that chiefs are deposed for attempting gross injustices or for failing to

consult about important decisions. Leaders who become violent or behave immorally may

also be replaced. In some African tribes, stingy leaders are also routinely removed from

office (Lewis, 1974), suggesting the importance of sharing. Sometimes followers simply

desert an exploitative leader. This reaction has been observed in both African and South

American societies (Boehm, 1999). An experimental study in social psychology found that

attrition rates in groups supervised by autocratic leaders are four times higher than in groups

supervised by a democratic or laissez-faire style leader (Van Vugt et al., 2004). As a

consequence, many of the autocratically led groups collapsed before the experiment was

over.

Finally, selfish leaders may be assassinated. The chief of a South American tribe was

killed for making private deals with outsiders (Boehm, 1993). In New Guinea, a Big man

who seriously oversteps his prerogatives can be killed by the community (Hogbin, 1951).

Political murders also occur in modern society. In US-history, several presidents have been

murdered and some survived attempts on their lives, making it one of the most dangerous

jobs in the world.

Ambivalence Model of Leadership

Gossip, criticism, ridicule, disobedience, ostracism, desertion, and violence are key

strategies for reducing the exploitative tendencies of leaders. Both the self-serving strategies

of leaders and the leveling strategies of followers reflect our evolutionary history, creating

leader-follower relationships that are fundamentally ambivalent. This ambivalence model


may explain why human leadership structures are so varied, compared to other social species

(Vehrencamp, 1983).

Under normal conditions, one would expect that the self-serving tendencies of leaders

and the leveling tendencies of followers would balance out, resulting in a structure that is

somewhere between a dictatorship and a true democracy, a de-facto oligarchy perhaps (see

our opening quote about the Universal People; Brown, 1991). Constitutional monarchies, for

example, are a compromise between the interests of monarchs, hereditary leaders, and the

lower ranking members of a society (Tiger & Fox, 1971). However, this balance could be

changed if forces push a group towards the autocratic or democratic end. For example, Hitler

moved a democratic society in an autocratic direction and Germans with authoritarian

attitudes supported the trend (Altemeyer, 1988). In contrast, a democratic leader with a group

of assertive and self-confident followers would push a group towards the democratic end of

the scale.

In addition, external or internal conditions can shift the equilibrium. For example,

during times of war, where quick decisions are often needed, conditions create opportunities

for leaders to exploit followers. Furthermore, under these conditions, it becomes difficult to

criticize, ridicule or desert leaders, thereby firmly shifting the balance of power toward the

leader (Van Vugt et al., 2004). In contrast, in situations where followers can stand up to,

criticize, desert, or take up arms against their leaders, the leadership structure should become

more democratic. Obviously leaders will be tempted to oppose these democratic movements

by restricting criticism (limiting free press), limiting exit opportunities for followers

(selectively granting passports), or disarming them (creating laws against gun possession).

Summary

By viewing leadership as an exchange relationship—the result of strategic interactions

between individuals engaged in coordinated action—a Darwinian perspective can account


for the variation in leadership styles observed in societies. Based on the assumption that

leader-follower relationships are inherently ambivalent, this analysis makes various

predictions about the type of leadership found in groups, depending upon the interaction

between external and internal conditions as well as the personalities of the actors involved.

Furthermore, this model specifies how leadership changes might come about and what

leadership structures become stabilized in groups over time (all else being equal). These

predictions can be tested using archived data, survey data, and small scale experimental

research.

Conclusions

This article puts forward a Darwinian perspective on the study of leadership, based on

the principles of natural selection and adaptation. Leadership and followership are evolved

designs for solving recurrent coordination problems among individuals. Across evolutionary

time, these complimentary behaviors have become increasingly synchronized and fine-tuned,

becoming part and parcel of human functioning in groups.

We have reviewed the evidence for six key propositions about leadership, emanating

from an evolutionary perspective. First, there is ample evidence that leadership and

followership are adaptive solutions to problems associated with group life, in particular

problems involving group coordination. Second, these decision making problems can be

modeled as coordination games that have presumably been played out across human

evolutionary history. Third, the design for leader and followership is extremely context-

specific. These behaviors are elicited only when there is a need to form coalitional alliances

to achieve certain goals like access to valuable resources or the protection of oneself or one’s

group. Fourth, the interplay between genetic and situational factors accounts for the

variability in human leadership styles, which can range from a despotic and autocratic style

to more benign, democratic styles. Fifth, leadership is based on a complex, dynamic


interaction between both heritable (genetic) and non-heritable factors. Finally, relationships

between leaders and followers are fundamentally ambivalent. There is a constant tension

between cooperating with each other to achieve mutual goals and competition to stay ahead

of each other in the reproductive game.

Theoretical Issues

Rather than offering a definite account of the evolutionary history of leadership, the

main aim of this article is to build a bridge between various behavioral science disciplines

studying leadership such as anthropology, biology, psychology, and management. As various

researchers have pointed out, there is a wealth of data on leadership but a lack of integration

into a universal theory of leadership (Bass, 1990; Chemers, 2000; Hogan & Kaiser, 2005).

Evolutionary theory is a good candidate for such an overarching theory, first, because any

middle-level social, psychological or cultural theory of leadership must ultimately be

consistent with it to be viable. For example, Freud’s psycho-analytic theory of leadership can

ultimately be understood in terms of a fundamental conflict between the reproductive

interests of leaders versus followers (Hogan, 1982). Second, an evolutionary approach

requires researchers to take an interdisciplinary approach by analyzing leadership from many

different angles, including examining the proximal, ultimate, ontogenetic and phylogenetic

causes of leadership.

Misconceptions. There are certain misconceptions about the evolutionary perspective

that might prevent it being incorporated in the study of leadership (Kenrick & Simpson,

1997). The first is that genes cannot influence complex social behaviors like leadership. Yet

there are a host of equally complex actions that, according to experts, have deep evolutionary

roots such as altruism, group identity, political, and religious behaviors (Boyd & Richerson,

2005; Van Vugt & Van Lange, in press; D. S. Wilson, 2002). A second misconception that

we have refuted is that evolutionary explanations imply genetic determinism and ignore the


role of the environment. But genes associated with leadership can only become manifest in

the context of a particular environment, for example, when leadership is required due to an

intergroup or intragroup threat (Sherif, 1966; Van Vugt & De Cremer, 1999).

A related misconception is that culture is more important than genes. Yet this is a false

dichotomy in that culture, or at least the capacity for social learning, is ultimately also a

product of evolution (Barrett et al., 2002). Moreover, cultural innovations that are successful

can spread through a population in much the same way as genes – so-called memes (Boyd &

Richerson, 2005; Dawkins, 1976). An evolutionary analysis of leadership memes seems

therefore quite appropriate. Finally, cultural improvements ultimately affect gene frequencies

in a population via the process of natural selection and adaptation (Boyd & Richerson, 2005).

For example, leadership in proto-human groups may have started as a cultural innovation

initially, but by virtue of its success on group life, it influenced the genetic make-up of early

humans.

A final misconception is that leadership researchers can ignore evolutionary

explanations because they operate at a different level of analysis. This would be a mistake. If

we want to understand leadership, we simply cannot ignore studying the cross-cultural, cross-

special and ontogenetic similarities and differences in the manifestation of leader and

followership.

Why an evolutionary perspective? There are a number of reasons why we think

leadership researchers should seriously consider a Darwinian perspective. The first is to

avoid saying things about leadership that are clearly wrong. For example, various

psychological theories of leadership assume that the goals of leaders and followers are

interchangeable, and that leaders always work for the good of the group. An evolutionary

perspective makes clear, however, that there is a fundamental tension in the relationship

between leaders and followers. As each is trying to pursue their reproductive goals, there will


be frequent conflicts of interests, for example, in the distribution of the benefits or burdens of

collective action.

A second reason is to avoid incomplete explanations for leadership phenomena. An

evolutionary analysis delineates that there is little point in studying leadership without

examining the motives and desires of followers (Yukl, 1999). In many ways, followership is

the most interesting of the two roles from an evolutionary perspective, because the benefits

of following are not always clear-cut. A more specific example is research on leader

categorization theory (Lord et al., 1986; Lord & Maher, 1991). This body of research shows

that followers have fairly stable impressions regarding the desired attributes in their leaders --

so-called leader prototypes. Examples of leader prototypes are intelligence, determination,

and honesty. An evolutionary account can help to understand where these intuitive beliefs

about leadership come from, why they are likely to be universal, and why they are so readily

accessible. Evolutionary theory can thus provide the underpinnings for such social-cognitive

models of leadership.

A third reason for adopting an evolutionary approach is to account for findings that are

difficult to explain with traditional leadership theories. A major criticism of leadership

research is that it has failed to account adequately for the relationship between leadership and

a diversity of factors such as age, height, weight, health, appearance, and speech fluency

(Bass, 1990; Stogdill, 1974). Such patterns begin to make more sense if one assumes that

evolution has selected for attendance to cues that signal leadership potential or ability. In

ancient history (and sometimes today) it would be potentially lethal to follow the wrong

person, for example, in moving a long distance towards a fresh water hole (Gamble, 1993).

Information about the health status and physical attributes of a leader would thus be

important to consider.

Practical Matters


Our Darwinian analysis suggests various new directions for leadership research. There

is already considerable evidence for the proximate benefits of leadership for those who

occupy such positions, but future research might want to address whether this is paid out in

reproductive success (Betzig, 1986; Perusse, 1993). Furthermore, what are the benefits of

leadership for those that follow? According to our analysis, loyal followers should be better

off under leaders that are good, but they should suffer under bad forms of leadership

compared to more opportunistic followers. Future studies could use data from businesses and

other organizations to test whether this is true.

Laboratory research is also needed to examine decision making in experimental games

like leader (Figure 1) to find out if there are personality differences that can meaningfully

predict whether some people are more likely to make a risky first move. For example, do

risky choices correlate with indices of empathy, extraversion, risk-taking, and

Machiavellianism? Furthermore, do choices in the Leader game correspond to displays of

leadership in the real-world? To our knowledge, there have been very few studies on the

Leader Game (Kremer & Mack, 1983) in contrast to, for example, the Prisoner’s Dilemma

Games (Van Vugt & Hart, 2004).

Finally, cross-cultural data could be gathered to examine the similarities and

differences in leadership and followership style across diverse societies. Although we believe

that the human potential for engaging in leader-follower relationships is universal, there

might be cultural factors determining the exact nature of this relationship. Differences in

climate, food provision, kin relations, intergroup rivalries, and status hierarchies within

groups might determine the type of leadership that is most likely to emerge in a particular

culture. For instance, the hierarchical style leadership and the importance of seniority

leadership that one finds in collectivistic cultures in East Asia (Hofstede, 1980) might well be

the product of a less mobile society in which people tend to stay in one place longer. Because


exit options are limited, power differences between superiors and subordinates are enhanced.

Similarly, we expect more aggressive and coercive forms of authority and leadership in

cultures in which conflicts between neighboring communities are frequent (Chagnon, 1997).

Before closing we should not some implications of a Darwinian perspective on

leadership. One implication is derived from the assumption that leadership is the product of

the particular phylogenetic history of humans as a highly social and highly organized species.

This unique history puts constraints on the way humans organize their societies. For

example, unlike gorillas or even chimpanzees, a leadership system based exclusively on

dominance, coercion and threat is not sustainable. Human leaders can not ignore the wishes

and preferences of followers for too long, otherwise they are ridiculed, replaced, put in

prison, or sometimes even killed. As Boehm (1999) states “it only takes one little arrow.”

Dictatorship is the right theory, but simply for the wrong species.

A second implication from a Darwinian account is that the relationship between leaders

and followers is always likely to contain elements of an Arms race in which both parties have

an incentive to cooperate as well as compete. While leaders will look for ways to extend their

power base, followers must try to keep leaders in place through a combination of subtle and

sometimes aggressive tactics. From the perspective of followers, their best outcome is to give

leaders the opportunity to strengthen their privileged position through prosocial means like

honesty, generosity, and offering safety and protection. Yet at the same time, leaders will

always be tempted to exploit their position for selfish means by benefiting themselves and

their close relatives (nepotism). Recognizing the fundamental ambivalence in the relationship

between leaders and followers is perhaps the greatest merit of a Darwinian model of

leadership.


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Author Notes

Correspondence regarding this manuscript may be sent to Mark van Vugt, University of

Kent, Canterbury CT2 7NP, England, [email protected].

Portions of this paper were presented at the EAESP leadership small group meeting in

Amsterdam, the Netherlands, June 2003 and the Human Behavior and Evolution conference

in Austin, US, June 2005


Tables

Table 1. Propositions about leadership and followership from an evolutionary perspective

Propositions Descriptions

___________________________________________________________________________

1 Leadership and followership are adaptive decision rules

2 Leadership and followership are strategies for group

coordination

3 Leadership emerges in response to group threats and

opportunities

4 Leadership styles are flexible

5 Leadership can be both a state and trait

6 Leader-follower relationships are fundamentally

ambivalent

_____________________________________________________________________


Table 2. Results Supporting the Evolutionary Game Hypothesis of Leadership

Hypothesis Rationale Result Reference

1. Leadership

correlates with

initiative taking

Those who take the

initiative are more

likely to emerge as

leaders

Leadership

correlates with the

following trait

measures:

- activity and

energy levels

- ambition

- assertiveness

- extraversion

- shyness (negative)

- active

participation in

group discussion

- initiating structure

is one of the

primary leadership

activities

Bray & Howard

(1983)

Gough (1984)

Sorrentino &

Boutilllier (1975)

(Hemphill, 1950)

2. Leadership

correlates with (social)

intelligence

More (socially)

intelligent persons

know better what

group action is

- Leadership

correlates with

general measures of

intelligence

Bass (1990)


needed and what

followers want

- Verbal ability

correlates most

strongly with

leadership

- Sociability

predicts leadership

- Individuals who

appear more

intelligent (high

Machs) emerge

more often as

leaders

Korman (1968)

Kenny & Zaccaro

(1983)

Christie & Geiss

(1970)

3. Leaders Show

Competence

Individuals would

only follow a leader

if they can be sure

the leader achieves

his goals

- Leadership in a

particular domain

correlates with

expertise in that

domain

- The successful

head of an

accounting

department is a

better accountant

than most

- Group members

are more willing to

Aidar (1989)

(Tsui, 1984)

Hollander (1985)


follow directions

from those with

task ability

4. Leadership

correlates with

generosity

Followers are more

dedicated to leaders

that show a

willingness to share

- Leadership

correlates with self-

sacrifice in social

dilemmas

- Leaders score

high on trust scale

- Leaders who are

procedurally and

distributively fair

get more support

- Stingy leaders are

replaced

De Cremer & Van

Knippenberg

(2002) Hardy &

van Vugt (2005)

Sgro, Worchel,

Pence, & Orban

(1980)

Thibaut & Walker

(1975)

Tyler & Lind

(1992)

Boehm (1999)

5. Leadership

correlates with health

Individuals expect

their leaders to

deliver long-term

benefits

- Leaders appear to

possess above-

average health

- prospective

leaders (e.g.,

presidential

candidates) must

appear healthy to be

elected

Bass (1990)

Marmot (2004)

Simonton (1994)


6. Leaders are

wealthier than

followers

Leaders benefit

more from

coordinated action

than followers

Leadership

correlates with all

kinds of socio-

economic indices of

wealth:

- income

- class

- education

- property

- Village leaders are

of higher caste and

have more property

Switzer (1975)

Roy et al. (1974)


Figure Captions

Figure 1. Leader Game; When Leader-Follower Patterns Emerge;

Figure 2a. Dominance Game; When Leaders Dictate the Outcomes of Followers;

Figure 2b. Democratic Game; When Leaders Must Share Power With Followers;

Figure 3a. Coordination Game; When Leaders and Followers Want the Same Thing;

Figure 3b. Conflict Game; When Followers Are Reluctant;

Figure 4a. Leader Game;

Figure 4b. Hero Game; When Followers Become Leaders;

Notes with Figures: The first value is for Player 1, and the second for Player 2; The pay-

offs represent fitness benefits (+) or costs (-) for the actors involved.


Figure 1

Player 2

Player 1

Wait

Move

Wait Move

0,0

200,100

100,200

-100,-100


Figure 2a

Player 2

Player 1

Wait

Move

Wait Move

0,0

200,100

100,200

100,-100


Figure 2b

Player 2

Player 1

Wait

Move

Wait Move

50,50

150,50

50,150

-50,-50


Figure 3a

Player 2

Player 1

Wait

Move

Wait Move

0,0

150,150

150,150

-100,-100


Figure 3b

Player 2

Player 1

Wait

Move

Wait Move

0,0

300,0

0,300

-100,-100


Figure 4a

Player 2

Player 1

Wait

Move

Wait Move

0,0

200,100

100,200

-100,-100


Figure 4b

Player 2

Player 1

Wait

Move

Wait Move

0,0

100,200

200,100

-100,-100

evolution and leadership 1 running head: evolution and

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