Kesebir 2012

Personality and Social Psychology
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The Superorganism Account of Human Sociality : How and When Human Groups Are Like Beehives
Selin Kesebir
Pers Soc Psychol Rev 2012 16: 233 originally published online 27 December 2011
DOI: 10.1177/1088868311430834
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PSRXXX10.1177/1088868311430834
The Superorganism Account of

Human Sociality: How and When
Human Groups Are Like Beehives
Personality and Social Psychology Review

16(3) 233261
2012 by the Society for Personality
and Social Psychology, Inc.
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DOI: 10.1177/1088868311430834
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Selin Kesebir1
Abstract
Biologists call highly cooperative and socially integrated animal groups like beehives and ant colonies superorganisms. In
such species, the colony acts like an organism despite each animals physical individuality. This article frames human sociality
through the superorganisms metaphor by systematically reviewing the superorganismic features of human psychology. These
features include (1) mechanisms to integrate individual units, (2) mechanisms to achieve unity of action, (3) low levels of
heritable within-group variation, (4) a common fate, and (5) mechanisms to resolve conflicts of interest in the collectives
favor. It is concluded that human beings have a capacity to partly and flexibly display each of these superorganismic properties.
Group identification is a key mechanism that activates human superorganismic properties, and threats to the group a key
activating condition.This metaphor organizes diverse aspects of human psychology (e.g., normative conformity, social identity
processes, religion, and the rally around the flag reflex) into a coherent framework.
Keywords
evolutionary psychology, group processes, interdependence, intergroup relations, interpersonal processes, morality, social
identity
What is it about beehives that captures the social imagination of so many? From Aristotle to Shakespeare, beehives
have been invoked repeatedly as a metaphor and an ideal for
human groups (Farrell, 1985). Ralph Waldo Emerson (1904)
relied on the beehive image to refer to the futility of selfinterested pursuits in human groups: No matter how you
seem to fatten on a crime, there can never be good for the
bee which is bad for the hive (p. 189). Mormons, who see
beehives as the symbol of societal harmony, nicknamed
Utah the Beehive State and put a beehive on the states
flag. This article looks at human sociality through the lens of
the beehive metaphor and systematically reviews its hivelike features.
Beehives are quintessential examples of the many coexisting in harmony. Despite each bees physical individuality,
a colony acts like an organism unto itself: The bee colony is
a mammal in many bodies (Tautz, 2008, p. 3). The queen
and drones represent the female and male reproductive
organs, respectively, and worker bees carry out the remaining bodily functions. The mammalian bodys division of
labor among organs is paralleled by the division of labor
among cleaner bees, builder bees, brood care bees, guard
bees, and foragers. The beehives functional unity is also
apparent when it collects food: The foraging activity of
20,000 or so bees in a colony is coordinated such that surrounding areas are surveyed most efficiently for nectar and
pollen. The nutrients are collected both in the precise quantities needed by the colony and in the nutritionally optimal
mix. In short, the hive is an integrated unit of function
(Seeley, 1995) that demonstrates a correlation and cooperation of parts (Wheeler, 1911, p. 324). Biologists call such
highly integrated entities like beehives superorganisms
(Hlldobler & Wilson, 2009).
Superorganismic species are rare emergences in the history of life, yet when they emerge they are extremely successful. This is because solitary individuals and uncoordinated
groups are no match for them. Given their size advantage,
ability to aggregate information, and efficiency that comes
from a massive division of labor, superorganismic colonies
gain the ecological upper hand. As a result, in the more than
100 million years over which they have existed, no family of
ultrasocial insects is known to have become extinct (E. O.
Wilson, 1990). Even though such insects make up just 2% of
the approximately 900,000 known insect species, they are
believed to compose more than half of the total insect mass.
Ants alone are estimated to equal human beings in mass
1
University of Virginia, Charlottesville,VA, USA
Corresponding Author:
Selin Kesebir, University of Virginia, Darden School of Business, P.O. Box
6550, Charlottesville,VA 22906-6500
Email: selin.kesebir@gmail.com
234
Personality and Social Psychology Review 16(3)
(Hlldobler & Wilson, 2009). Beyond sheer numbers, social

insects have pushed solitary insects (e.g., cockroaches and
grasshoppers) away from the best nest sites on vegetation
and ground, to remote and less stable sites, such as peripheral twigs, leaf surfaces, and mudflats (E. O. Wilson, 1990).
Are Human Groups

Superorganisms?
The human ecological dominance is reminiscent of the
superorganismic species. Recently, a number of biologists
claimed that human groups also are (at least partially) superorganisms (Foster & Ratnieks, 2005; Stearns, 2007; D. S.
Wilson & Wilson, 2007). They argued that human beings
have made an evolutionary transition from primate colonies
to a superorganismic existence. If human groups are superorganisms, this should be evident first and foremost in their
social behavior. Social psychology is thus in a privileged
position to evaluate whether human groups display any
superorganismic traits. Some social psychologists have
indeed suggested that a superorganism perspective can help
explain various social-psychological phenomena (Brewer,
2004; Brewer & Caporael, 2006; D. S. Wilson, van Vugt, &
OGorman, 2008). This article systematically reviews how
and when human groups display superorganismic properties.
Based on a wealth of psychological evidence, I conclude that
human psychology is characterized by numerous mechanisms that make superorganismic social structures possible.
Identification with a group is the switch that activates these
mechanisms, and superorganismic features are magnified
under threat to the group. Human groups thus have superorganismic potential that is manifested opportunistically and
flexibly. This account of the human superorganismic capacity is neither a denial nor a rejection of human traits that are
not superorganism-like. Selfishness, conflict, and competition are facts of many human groups. Without question,
human groups are nothing like superorganisms in many
ways. Still, I argue that superorganismic capacity is part of
human psychology and that the superorganism metaphor is a
useful heuristic to explain human sociality.
This article is organized as follows: I first revisit the history of social psychology to discuss how the idea of human
groups as superorganismic structures was lost in psychology.
I then review ideas from evolutionary science on superorganisms to set the stage for the psychological evidence on
human superorganisms. In the articles major part, I discuss
how human psychology and social phenomena resonate with
the design of superorganismic species.
The History of the

Superorganism Idea
The idea that human groups resemble organisms can be
found in ancient India, where Hinduisms four major castes
were described as having descended from different body
parts of a god-created primal giant (OFlaherty, 1981). The

implication is that all castes must work together, as do the
parts of one body. A similar idea occurs in Platos Republic,
where Socrates made the extended analogy that a city-state
is like a man (A. Bloom, 1991). He described justice in an
ideal city-state (which involves each caste doing its job well
and for the common good) to segue into his discussion of
justice in a man (in which each part of the psyche does its
job well and for the common good).
As these examples show, superorganismic conceptions of
human society are ancient. Moreover, numerous social thinkers have conceived of human societies as organisms that need
to be understood on their own terms rather than as a mere
collection of individuals (for a review, see MacLay, 1990).
The term superorganism, which was later appropriated by
biologists, was first used by 19th-century thinker Herbert
Spencer (1876/1969) to describe human groups. Like the
Hindu sages and Plato, he thought that human groups interdependence is similar to a bodys functional integration:
When we see that in a mammal, arresting the lungs
quickly brings the heart to a stand; that if the stomach
fails absolutely in its office all other parts by-and-by
cease to act; that paralysis of its limbs entails on the
body at large death from want of food, or inability to
escape; that loss of even such small organs as the eyes,
deprives the rest of a service essential to their preservation; we cannot but admit that mutual dependence is
an essential characteristic. And when, in a society, we
see that the workers in iron stop if the minders do not
supply the materials; that makers of clothes cannot
carry on their business in the absence of those who
spin and weave textile fabrics; that the manufacturing
community will cease to act unless the food-producing
and food-distributing agencies are acting...we are
obliged to say that this mutual dependence of parts is
similarly rigorous. (p. 12)
Spencer was a sociologist, and the idea that groups are
more than their parts sum is foundational to sociology
(Durkheim, 1895/1950). Psychology, in contrast, seldom
goes beyond the individual unit of analysis (Oishi, Kesebir,
& Snyder, 2009). And yet the history of psychology also has
included organismic conceptions of human groups. Wilhelm
Wundt divided psychology into individual psychology,
which would rely on experimentation, and Vlkerpsychologie,
which would study psychological processes in historical and cultural context (Wundt, 1912/1916). He defined Vlkerpsychologie
as the study of those mental products which are created by a
community of human life and are, therefore, inexplicable in
terms merely of individual consciousness since they presuppose the reciprocal action of many (Wundt, 1912/1916, p. 3).
His 10-volume Elements of Folk Psychology covered such
topics as language, religion, cults, customs, morals, and
myth. Wundt believed that these collective phenomena
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Kesebir
would not fully lend themselves to experimentation, and the
experimental project of psychology was thus a limited one
(Farr, 1996). Soon after these volumes were published, however, Vlkerpsychologie was declared outdated and too philosophical to be relevant to psychology (e.g., Eliot, 1917).
Wundts deviation from experimentalism was not received
kindly and prompted the positivist repudiation of Wundt
(Danziger, 1979). This backlash was a harbinger of the
reductionism and methodological individualism that would
soon dominate social psychology (Cahan & White, 1992;
Farr, 1983; Jahoda, 2007).
Another organismic brand of psychology, the group
mind, appeared around the turn of the 20th century, when
large collectives such as labor unions and socialist organizations began to play significant roles in public life. Scholars
responded to the social and political changes of the times.
Some believed that there was a distinct psychology of group
behavior that could not be fully captured through the psychology of the isolated individual (Freud, 1921; Le Bon,
1896/1920; McDougall, 1920). Gustave Le Bon (1896/1920)
specifically invoked the society-as-organism metaphor to
argue this point:
The psychological group is a provisional being formed
of heterogeneous elements, which for a moment are
combined, exactly as the cells which constitute a living body form by their reunion a new being which
displays characteristics very different from those possessed by each of the cells singly. (p. 30)
William McDougall (1920), the author of social psychologys first textbook, similarly believed that the disciplines
proper subject was the general principles of collective mental life which are incapable of being deduced from the laws
of the mental life of isolated individuals (pp. 7-8).
But this organismic view of groups was also rejected. In
his seminal 1924 textbook, Floyd Allport wrote, There is
no psychology of groups which is not essentially and entirely
a psychology of individuals.... Psychology in all its
branches is a science of the individual. To extend its principles to larger units is to destroy their meaning (p. 4). He
rejected any metaphysical notion of the group mind, noting
that the actions of all are nothing more than the sum of the
actions of each taken separately (p. 5). Historians of psychology have remarked that this was a non sequitur:
Rejecting the metaphysical aspect of the group mind concept does not necessitate the rejection of a group psychology
that emerges from the interactions of individuals and is irreducible to individual behavior (Greenwood, 2004). Yet this
rejection was the next critical step toward the reign of methodological individualism in social psychology (Pepitone,
1981; Sampson, 1977; Semin, 1986).
Particularly since World War II, American social psychology has distanced itself from sociological approaches (Oishi
et al., 2009). As a result, group studies have relied heavily on
small groups of strangers, and group behavior has often been

modeled as the aggregate of individual processes. Group
cohesiveness, for example, has been operationalized as the
average attraction of group members to each other (Hogg,
1992). This reductionist treatment and neglect of emergent
collective phenomena was a major trigger of the crisis in
social psychology during the 1970s (Cartwright, 1979;
McGuire, 1973; Ring, 1967; Sherif, 1977; Steiner, 1974).
The crisis has since abated, but individualism and reductionism remain dominant (Rodrigues & Levine, 1999) despite
such notable exceptions as research on social identity that
will be reviewed below (Tajfel & Turner, 1979), research on
cultural psychology (Shweder, 1991), and research on
dynamic social psychology (Latan & Bourgeois, 1996; A.
Nowak & Vallacher, 1998).
As a discipline, social psychology is located between the
individual and social levels of analysis. This position presents
the challenge of bridging the two levels. As we have seen,
social psychology opted for methodological individualism
early in its history, which led to the neglect of emergent social
phenomena and reciprocal determination between individual
and social levels. Given the trends just described, social psychology todayparticularly in the United Statesis theoretically and methodologically disinclined to look at individuals
as units of a superorganism. Current perspectives from evolutionary science, however, provide some analytical tools that
are conducive to conceptualizing human sociality in such
terms. I next review these evolutionary ideas to set the stage
for the discussion of superorganismic human features.
Major Transitions in Evolution and

the Emergence of Superorganisms
Life forms are organized in nested clusters. Genes are bundled in chromosomes that occur in cells. Cells are joined
together in multicellular organisms, and some multicellular
organisms, such as bees and ants, live in societies. This hierarchical organization strongly suggests that the amazing
diversity of life forms is partly the result of the grouping of
biological units into higher-level units. Although this idea
has been endorsed since the end of the 19th century, it has
not been part of the mid-20th-century evolutionary synthesis, most likely because it has lacked a strong theoretical
underpinning (Bourke, 2011). Since then, however, a modern view of lifes hierarchical organization has emerged
(Bonner, 1974; Buss, 1987). The dynamic underlying the
hierarchical organization of life forms has been called major
transitions in evolution (Maynard Smith & Szathmry,
1995). A major transition in evolution occurs when individual organisms become so integrated that they transform into
a higher-level organism in their own right. There are multiple obstacles to such integration: Integration requires communication among individual units, capabilities for
coordination, and suppression of competition at the lower
level. This is why major transitions are rare.
236
Major transitions are among the most important events of

the last 4 billion years because they have been key propellers
in the history of life. At every level of biological organization,
a closer look reveals cooperative communities. Yesterdays
aggregates become todays integrates by joining into survival alliances and suppressing competition at the individual
level (G. O. Mackie, 1986). Self-replicating molecules have
given way to self-replicating complexes (e.g., chromosomes). The eukaryotic cell is a symbiotic community of
previously autonomous bacteria, some of which have given
up their reproductive independence to join the eukaryotic
cell (Margulis, 1981). Multicellular organisms are collections of individual eukaryotic cells that work together as a highly
cooperative unit. And finally, such multicellular organisms as
ants, termites, honeybees, and naked mole rats transitioned from
groups of solitary individuals to colonies that act much like one
organism: a superorganism. Superorganisms have a division of
labor resembling the organ systems in a body, and some individuals forgo reproduction, much like somatic cell lineages in
multicellular bodies that do not pass genetic material to offspring (Buss, 1987; Maynard Smith & Szathmry, 1995;
Michod, 1999). Major transitions demonstrate how at any
level of organization, consolidating power by cooperating with
other units at the same level may lead to a huge competitive
advantage (also see H. Bloom, 2000; Corning, 2005; Wright,
2000). As a result, life is full of such alliances.
This article explores how far the superorganism metaphor
can take us in capturing aspects of human sociality. It should
be noted at the outset that this metaphor is applied to specific
human groups, and not to humanity in general or such temporary aggregations as crowds. Here, a group is defined as
any collection of individuals that allows its members to
switch from me to us. In social scientific terminology,
this is any collection of individuals who share a social identity. As such, a group could be based on religion, nationality,
race, political party, or sports team. Among others, it could
take the form of a tribe, a street gang, a family, a troop, a
fraternity or sorority, a clique, or a work team.
I use two strategies to establish the analogy between
human groups and superorganisms. First, I ask what superorganismic features are evident in human psychology and
human groups. Second, I look at two major phenomena of
human sociality (religion and morality) to identify how they
embody superorganismic principles.
Superorganismic Aspects
of Human Psychology
Table 1 lists properties of a superorganism and how they are
manifested in honeybee colonies versus human groups.
These properties are:
1. Integration of lower-level units through communication
2. Unity of action
3. Low levels of heritable variation among the superorganisms units

4. A common fate
5. Mechanisms to resolve conflicts of interest in favor
of the collective
The following discussion identifies these superorganismic
features in human psychology and human groups.
1. Integration of Lower-Level
Units Through Communication
Integration is defined as the forging of connections among
disconnected parts. Connectedness among lower-level units
is an elemental requirement for any organism. In the words
of a prominent honeybee expert, [T]he formation of a
higher-level unit by integrating lower-level units will succeed only if the emerging organization acquires the appropriate technologies for passing information among its
members (Seeley, 1995, p. 247). All living organisms
have internal communication and coordination systems.
Human cells, for example, communicate through nervous
and hormonal systems in addition to cell-to-cell contact.
Unlike cells in a body, bees are physically dispersed yet
integrated through various chemical and tactile signals such
as the waggle dance (von Frisch, 1967). In human beings,
this function is fulfilled through the uniquely versatile
means of symbolic communication and communication
through the bodily expression of internal states.
Integration through symbolic communication. One means of
human communication is the use of symbols with shared
meanings. This ability allows us to access other minds so we
can offer information, make an argument, request something,
make a promise, give advice, challenge a claim, and engage
in countless other speech acts (Searle, 1969). As the biographies of feral children suggest (Newton, 2002), the human
brain requires immersion in a complex symbolic web to realize its potential. This human dependence on group culture is
symptomatic of a superorganism as it reflects the functional
integration of parts into the whole for their existence. Just as
a cell cannot exist outside the body, and a bee cannot exist
outside the colony, human beings cannot meaningfully exist
outside symbolic culture. Human cognition is instead distributed between the individual mind and the symbolic culture in
the sense that human cognitive processes heavily rely on cultural input (Donald, 1991; Hutchins, 1995; Semin, 2004).
Culture that is transmitted via shared symbols connects individuals to their societies cognitive models (Shore, 1996).
Language is a major medium of human communication
that deeply pervades human sociality. The remarkable generativity of language injects extraordinary versatility into
human interactions and unlocks collective possibilities not
available to species with less flexible communicational
means. The discussion of symbolic communication in this
section briefly touches on all human superorganismic aspects
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Kesebir
Table 1. Properties of a Superorganism and Their Manifestation in Human Groups Versus Beehives
Manifestations of these properties in
Properties of a superorganism
Human groups
1. Integration of lower-level units through

communication
2. Unity of action
3. Low levels of heritable variation among the
superorganisms units
4. A common fate
5. M
echanisms to resolve conflicts of interest
in favor of the collective
Symbolic communication
Synchronous activities
Shared intentionality
Social identity processes
Deference to legitimate authority
Phenotypical similarity through culture
Egalitarianism
Intergroup warfare
Individual self-restraint
Social control through prosocial
emotions, norms, and institutions
to be reviewed later, for symbolic communication is essential to each of them.

First, language aids coordination and planning, which, in
turn, facilitate cooperative joint action. Experimental studies
of social dilemmas show that participants are more likely to
commit to a cooperative strategy if they can talk to each
other (R. M. Dawes, 1980; Kerr & Kaufman-Gilliland,
1994). Moreover, because language allows people to transcend the here and now, it increases cooperation through the
social control device of reputation. Gossip, a very effective
means of reputation transmission, allows individuals to monitor group members, so they can selectively interact with
cooperators and withhold benefits from or punish noncooperators (Dunbar, 2004; Merry, 1984). Gossip not only
exposes noncooperators but also communicates norms and
values by revealing acceptable and unacceptable behavior
(Baumeister, Zhang, & Vohs, 2004; Sabini & Silver, 1982).
Like gossip, folktales, legends, and proverbs also convey
cultural values and norms. Anthropologists have noted that
however primitive its material technology, no society exists
without these folkloric forms that express group values and
norms (Bascom, 1953). In sum, language facilitates coordination, planning, and commitment; enables social control
through reputation transmission; and serves as a carrier of
cultural values and norms.
Symbols also promote group identity and cohesion by
defining who is in and who is out. In other words, they
draw the groups boundaries (Giles, Coupland, & Coupland,
1991; Scherer & Giles, 1979). Why are there so many languages? Why do languages keep changing? And why is
Esperanto not taking off as the language of the masses? A
clue to all these puzzles is that language variation serves
Beehives
Various chemical and tactile signals (e.g.,
the dance language)
Self-organization (decentralized control)
through genetically programmed
behaviors
Dependence of worker bees on the
queen for reproducing their genetic
material
Genotypical similarity
Shared reproductive fate
Self-restraint by worker bees
Social control (e.g., destruction of
worker-laid eggs by other worker bees)
group formation and maintenance by making it costly to

enter and exit groups (Nettle, 1999). Consciously or unconsciously, people modify their speech to expand and shrink
social boundaries: Positive and cooperative interactions are
associated with a convergence of speech styles (Giles &
Smith, 1979). In contrast, people whose identity is being
threatened rely more heavily on linguistic features that make
their speech distinctive (Bourhis & Giles, 1977; Labov,
1972). People also react more negatively to those who speak
different languages, dialects, or accents (for a review, see
Bradac, 1990). This preference for native speakers emerges
early in life. Infants who are 5 months old prefer to look at
native speakers, 10-month-old infants preferentially accept toys
from native speakers, and preschool children are more likely to
choose to befriend a native speaker. Moreover, all of these age
groups prefer speakers with a native accent over speakers with
foreign accents (Kinzler, Dupoux, & Spelke, 2007).
Symbolic group markers are so potent that they can
uphold very abstract groups such as nations. Nations are
in Benedict Andersons (1983) words imagined communities. They are imagined because never, not even in the
smallest nation, can citizens know all their compatriots.
Yet despite the impossibility of intimate personal bonds
with fellow citizens, many people have a strong sense of
allegiance to their nation. What gives power to the idea
of nation is an elaborate set of shared symbols and meanings. Research on nation building shows how newly independent nation-states immediately embark on massive
symbolic projects (Birch, 1989): Nation builders declare
an official language and create national paraphernalia
such as a flag, a national anthem, and new military uniforms. They rename towns, buildings, and capital cities.
238
They create and cultivate distinctive myths, memories of

an ethnic past, and traditions that, at times, may be more
invented than real (A. D. Smith, 1999). These symbolic
processes serve to create an identity and a sense of solidarity among people who will always remain strangers.
Nation-building cases from around the world illustrate
the extent to which symbolic processes can integrate
groups.
In sum, symbolic communication is foundational to human
sociality. Symbols play a crucial role in fulfilling each requirement for a superorganismic formation: They foster coordination and cooperation by facilitating planning, information
sharing, norm enforcement, and boundary marking. Clearly,
language is not an exclusively cooperative tool for it may be
used to lie, mislead, insult, and threaten. Still, symbols afford
enormous capabilities for joint action and collective problem
solving. Language multiplies what individuals can gain by
cooperating, rather than ignoring, competing with, or attacking each other, and allows the attainment of collective goals
that are beyond the reach of any single individual. It significantly amplifies the potential for non-zero-sum interactions
(E. A. Smith, 2003; Wright, 2000).
Integration through synchronous movement. Symbolic
communication is not the only way to bridge the gulf
between individual minds. Human beings also communicate through their bodily expressions. Mirror neurons are
activated when someone both does an action and observes
someone else perform it (Rizzolatti & Craighero, 2004).
Consequently, corresponding internal states can be
achieved between two people in the absence of any symbolic exchange. This phenomenon is held to precede symbolic communication evolutionarily and ontogenetically
(Semin, 2007). In this section, I review the evidence on
the role of bodily communication in human group formation and maintenance, and in particular the importance of
synchronous activities as a means of group integration.
One form of synchronous activity is dancing. Although
dancing does not figure prominently in modern societies
or in psychological theorizing, it has been a major cultural
element throughout human history. The anthropological
record shows that festive dancing was a very significant
aspect of prehistoric human life (Ehrenreich, 2006).
Archaeologist Garfinkel asserted that dancing scenes
were a most popular, indeed almost the only, subject used
to describe interaction between people in the Neolithic
and Chalcolithic periods (cited in Ehrenreich, 2006,
p. 22). Festive dancing is also documented to be ubiquitous
in the small-scale societies studied by anthropologists.
Participants report that submitting oneself bodily to the
music and dance creates an altered state of consciousness
a deeply satisfying sense of being incorporated into a community (Rappaport, 1999; V. Turner, 1969). Anthropologist
Radcliffe-Brown (1933/1948) wrote on dancing among
the Andaman Islanders,
As the dancer loses himself in the dance, as he

becomes absorbed in the unified community, he
reaches a state of elation in which he feels himself
filled with an energy of force immensely beyond his
ordinary state...finding himself in complete and
ecstatic harmony with all the fellow-members of his
community, experiences a great increase in his feelings
of amity and attachment towards them. (p. 252)
Dancing is often accompanied by drumming. Anthropological
works consistently refer to the ritualistic use of drumming
to alter consciousness and induce transcendent spiritual
states (Maxfield, 1992; Woodside, Kumar, & Pekala, 1997).
Clinical applications have also shown that drumming
reduces stress and creates a sense of connection to others
(Blackett & Payne, 2005; Friedman, 2000; Winkielman,
2000). In one study, participants in a small group reported
more attraction toward each other after a hand-drumming
session (C. D. Kaplan, 2000).
Military drill also creates ecstatic joy, similar to the one
experienced through ritualistic dance and drumming. Here is
how historian William McNeill (1995) described the blissful
feelings caused by military drill:
Words are inadequate to describe the emotion aroused by
the prolonged movement in unison that drilling involved.
A sense of pervasive well-being is what I recall; more
specifically, a strange sense of personal enlargement; a
sort of swelling out, becoming bigger than life, thanks to
participation in collective ritual. (p. 2)
The evidence, then, strongly suggests that rhythmic entrainment leads people to an ecstatic state of union with others,
wherein the self seems to merge with something larger.
Neurophysiologist Freeman (1995) described dance and
music as the biotechnology of group formation because
they are so effective in triggering a collective mental state
of we-ness. This sense of unity is not communicated in
symbols; rather, participants experience it physically as
they move in sync.
Despite the plethora of historical and anecdotal evidence on
the effects of synchronous activity in integrating groups,
empirical work is scant. We know that interactional synchrony is linked to rapport (Berneiri, 1988; Cappella, 1997),
and experimental tests of the role of synchronous activity have
found that singing and moving together lead to more cooperative behavior in adults (Wiltermuth & Heath, 2009) and children (Kirschner & Tomasello, 2010). The rich anthropological
evidence and partakers passionate accounts suggest that synchronous activity may be an ancient group-bonding technology as well as a fundamental human experience that is largely
missed by modern people and modern psychologists.
In sum, integration of units through various communicational means is a critical aspect of superorganisms. Human
239
Kesebir
groups have at least two means of integration, symbol use
and bodily communication.
2. Unity of Action: Shared Intentionality,

Social Identity Processes, and Deference to
Legitimate Authority
Integration of units is a necessary, but not a sufficient, condition for superorganism formation. Units may be highly interconnected, and yet they may not have the capacity to act as a
functional unit. The millions of computers connected through
the World Wide Web provide an example of dense interconnection without unity of action. The capacity to act as a functional unit is another key feature of a superorganism.
Beehives are functional units thanks to their capacity for
self-organization. Self-organization refers to decentralized
organization achieved without an external controller or an
internal control center (Seeley, 1995). It is common in biological systems made of numerous subunits that yet lack the
communicational or computational abilities required for centralized control (Seeley, 2002). In these systems, high-level
organization emerges as units respond to local cues. A magnolia tree, for example, efficiently allocates resources to
growing and maintaining its roots, trunk, branches, leaves,
and flowers, in the absence of central oversight by any of the
trees parts. The interactions of bees are similarly governed
by local environmental stimuli that activate genetically given
properties. Because these properties have been finely tuned
over evolutionary time, the emerging pattern is adaptive for
the hive (Camazine et al., 2001). As a result, the beehive acts
as a functional unit when it distributes its workforce among
the hives needs, collects its nutrients, regulates its temperature, selects a location to colonize, or builds honeycombs.
Self-organization also accounts for the emergence of
coordination, order, and social patterns in human groups
(Kenrick, Li, & Butner, 2003; A. Nowak & Vallacher, 1998).
The human capacities for sharing intentionality and identifying with a group create bottom-up functional integrity
through self-organization. In addition to these bottom-up
processes, human psychology also lends itself to establishing
functional integrity top-down, owing to the human willingness to submit to legitimate authority. I next review these
three capabilities that enable united action in human groups.
Shared intentionality. Human beings can engage in collaborative action because they can form joint goals and effectively coordinate the roles involved in goal pursuit. Consider
a joint action as deceptively simple as two people moving a
couch from one corner of the room to the other. To successfully complete this task, the two individuals must each form
an intention to move the couch, know that they share this
intention, be motivated to cooperate, and be able to initiate
collaborative action. Once they begin the task, they need to
adjust their behavior instantaneously to their partners actual
and anticipated behaviors as well as the task environment.

Although the origins of such collaborative undertakings are
evident in chimpanzees, our closest genetic relatives (Melis,
Hare, & Tomasello, 2006), human beings are extraordinary
collaborators, able to undertake collective actions involving
thousands or even millions of people. What accounts for this
exceptional capacity?
The ability to participate with others in collaborative
activities with shared goals is called shared intentionality
(Tomasello, 2009; Tomasello, Carpenter, Call, Behne, & Moll,
2005). Shared intentionality involves forming corresponding
mental states regarding aspects of the world (common ground)
along with mutual knowledge of this correspondence. What
is crucial here of course is not the correspondence of mental
states, which could be achieved incidentally by any two
organisms, but the mutual awareness of it. This mutual
awareness is the base from which communicative exchange
and joint action is launched.
Joint attention is the most elemental skill of human collaboration (Moore & Dunham, 1995). It has precedents in
nonhuman primates, but human beings take joint attention to
qualitatively new levels. Before 9 months of age, infants
dyadically interact with objects or people, such as when they
grasp and manipulate an object, or have proto-conversations
with a caregiver. In this period, attention is singularly focused
on one object or person at a time. At around 9 months something revolutionary happens: Infants develop a capacity for
triadic interactionsa capacity to coordinate their attention
between another person and an external object. They, for
example, start to follow an adults gaze by alternately looking at the adult and the target of the adults attention. Infants
also point to or hold up an object for an adult to see, thereby
attempting to direct the adults attention to it. A baby will, for
example, point to a flitting butterfly that dazzles her, and
check the adults face. She will want the adult to look at the
butterfly and then back at her, not being satisfied if the
adult looks only at her or only at the butterfly (Liszkowski,
Carpenter, Henning, Striano, & Tomasello, 2004). At around
this time, infants also start participating in joint engagements
involving an object, such as rolling a ball back and forth. All
of these activities involve a triadic coordination of attention
among the infant, a social partner, and an external object.
This capacity for triadic coordination of attention is the foundation of human cultural cognition that is based on symbolic
exchange (Tomasello, 1999). At the minimum, a symbol is a
triadic act involving two people and an external object: It
represents a mutually recognized convention between at
least two people on how to communicate about an object.
Chimpanzees who lack cultural cognition also lack the
capacity for joint attention. Like human beings, they can follow the gaze of other chimpanzees. But, they rely on head
movements, whereas human beings track eye direction
(Tomasello, Hare, Lehmann, & Call, 2007). This reliance on eye
direction is afforded by the human eyes unique morphology.
240
Human eyes seem to be designed to enhance the gaze signal,

whereas nonhuman primate eyes seem to be designed to
camouflage it. An analysis of primate eyes found that out of
a sample of 81, only human eyes showed a high color contrast with the surrounding skin. Of all species, human beings
had the highest ratio of exposed sclera in the eye outline and
the most horizontally elongated shape. Moreover, human
eyes were the only ones to have white sclera (Kobayashi &
Kohshima, 2001). These features all serve to more effectively communicate attentional focus to conspecifics: They
facilitate joint attention. Chimpanzees, unlike human infants,
also are not interested in establishing joint attention by pointing. This is not for a want of pointing capacitycaptive
chimpanzees do sometimes point to demand an object
(Leavens, Hopkins, & Bard, 2005). They just do not seem to
have any desire to use pointing to share an interest with a
conspecific. Joint attention thus sets human beings apart
from other primates.
Joint attention, the first step toward joint action, allows
individuals to start from a common ground with shared representations of objects and events. After all, how can people
jointly manipulate their environment if they cannot first
focus on the same aspect of the world? But that is not all, for
joint action also requires coordination of activities. Coordination
involves a division of labor and mutual awareness of each
partys different role and perspective. As mentioned above,
language is an extremely powerful coordination tool.
Interestingly, though, very young children can coordinate
roles before they can talk. Children 18 months old are able to
reverse their roles in a game involving an object (Carpenter,
Tomasello, & Striano, 2005). In one study, the experimenter
put a Big Bird toy on the floor and covered it with a cloth,
asking the child, Where is Big Bird? Can you find him?
After doing this three times, the experimenter gave the toy
and the cloth to the child saying, Its your turn now. For
this particular task, 18-month-olds were more likely than
12-month-olds to hide the Big Bird under the cloth and look
at the experimenter, presumably in anticipation of her role
fulfillment. This capacity to reverse roles suggests that the
infants had a global representation of the joint activity as
well as a sense of the roles involved. In childrens ability to
mentally represent an activity with its differentiated roles,
we find the developmental origins of collaboration and division of labor.
All of that shows that people are capable of joint attention
and engagement. But just as importantly, they take delight in
it. When children point to an object, they want others to
respond enthusiastically. When a mutual engagement is broken, they want to reestablish it. In one study, children aged
14 to 24 months and three human-raised juvenile chimpanzees were presented with instrumental tasks that required
collaboration with the experimenter (Warneken, Chen, &
Tomasello, 2006; Warneken & Tomasello, 2007). In the
midst of these joint tasks, the experimenter suddenly stopped
participating. Human children reacted to this disengagement
very differently from the chimpanzees: They waited for

reengagement, and when it was not forthcoming they actively
tried to reengage the experimenter by establishing eye contact, gesturing, and making sounds. No chimpanzee, on the
other hand, made any attempt to reengage the experimenter.
Once the experimenter disengaged, so did the chimpanzees.
Moreover, some human children apparently found collaboration so rewarding that they turned the instrumental task into
a social game: They placed the reward back into the apparatus to start the activity over.
This readiness to turn an instrumental task into a game
should not come as a surprise, for childrens vast appetite
for play reflects an eagerness for joint creation and joint
action. Play often involves establishing a shared universe
inhabited by play partners. The universe has distinctive
roles (Im the doctor and youre the patient), rules
(everybody will hide until it counts to ten), functions for
objects (the bed sheet is my cape and the umbrella is my
sword), and meanings for actions (wetting with a water
pistol means killing). Through play, children exercise their
capacity for building and functioning in shared social realities (Rakoczy, 2006). In other words, they exercise shared
intentionality. The ubiquity and appeal of such play
unequivocally illustrate the human ability and motivation
for binding oneself to shared social constructions. Without
this motivation to share emotions, activities, and conventions, collaborative action could not occur.
We saw that people differ from other primates in their
capacity for shared intentionality. Still, this comparison fails
to convey fully what it would mean for a human being to
lack this capacity. One group of human beings, unfortunately, exhibits exactly this condition. Autistic individuals
have low levels of skill and motivation for joint attention and
engagement. They are mostly unable to infer conclusions
from gaze direction, such as the referent of an uttered word
(Baron-Cohen, Baldwin, & Crowson, 1997) or an individuals mental state (Baron-Cohen, Campbell, Karmiloff-Smith,
Grant, & Walker, 1995). Autistic children are unlikely to
offer something to another persons attention for the sake of
sharing an interest (Mundy, Sigman, Ungerer, & Sherman,
1986) or to engage in pretend play (Baron-Cohen, 1987).
They also have difficulties with language development.
According to one estimate, about half of the autistic population never acquires functional linguistic skills (Bailey,
Phillips, & Rutter, 1996). Autistic individuals, in other
words, are not adequately in tune with their social environment and are short on the skills needed to respond to it adaptively. Their predicament highlights the significance of the
shared intentionality skill set for human sociality. Without
those skills, human beings could not show the initiative and
responsiveness required to act together.
In sum, human beings have a unique capacity for joint
action. This is possible because they are able and willing to
coordinate attention on aspects of the world with others and
take on roles with attached expectations and obligations.
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Kesebir
Division of labor is crucial for any superorganisms effective
functioning, and shared intentionality is the most elemental
building block underlying human division of labor in joint
goal pursuit.
Social identity processes. The previous section described
the micro foundations of the human capacity for joint action
that allow two people to collaborate. This section and the
next one describe how human beings achieve unified action
at larger scales of social organization. A group of people can
succeed in facing challenges and achieve goals to the extent
that they can merge their forces for a common purpose and
act one for all and all for one. In human groups, this happens through social identification processes, by which individuals adopt group goals and display preferential treatment
and loyalty to group members.
To understand what makes groups act in unison, consider
the times people do not take collective action even if the conditions call for it. Political scientists have noted the curious
lack of political cohesion in some highly visible groups such
as the Chinese in Indonesia, native people in Argentina, and
North Africans in Italy (Huddy, 2003). Why do these people
fail to work in solidarity for their groups benefit? The
answer is that political coalescing requires more than objective membership in a demographic category. In each of these
cases, the missing ingredient is a subjective sense of belonging to the group. Because they do not feel strong ties to their
group, these people do not form a united front. Identification
with a group is what allows individuals to adopt group-level
goals and get mobilized. Tajfel (1981) defined social identity
as an individuals knowledge of his membership in a social
group (or groups) together with the value and emotional significance attached to the membership (p. 255). Research
inspired by social identity theory (Tajfel, 1981; Tajfel &
Turner, 1979), and its subsequent elaboration social categorization theory (Hogg & Abrams, 1988; J. C. Turner, Hogg,
Oakes, Reicher, & Wetherell, 1987) explains the basic mechanisms that bind people to groups and coordinate their collective actions.
Social identity theory is based on the premise that group
processes cannot be reduced to interpersonal processes
among group members (J. C. Turner et al., 1987). When they
identify with a group, individuals not only like group members and feel similar to them, but they also incorporate the
group into their self-concept by self-categorizing as a group
member. Consequently, a shift happens towards the perception of the self as an interchangeable exemplar of some social
category and away from the perception of self as a unique
person (J. C. Turner et al., 1987, p. 50). This shift increases
peoples perceived similarity to their group members and
perceived dissimilarity to members of other groups (Simon,
Pantaleo, & Mummendey, 1995).
The incorporation of the group into the self-concept marshals the same cognitive, affective, and behavioral processes
associated with the self. Cognitively, groups become included
within peoples mental representation of themselves. For
example, people are faster to say that a particular trait

describes them if it applies to both themselves and their
group, compared to when there is no match (E. R. Smith &
Henry, 1996). Conversely, reporting on the ingroups characteristics is facilitated to the extent that those characteristics
match ones own (E. R. Smith, Coats, & Walling, 1999).
Notably, this representational self-group overlap is as strong
as the representational overlap with a spouse, as measured by
the response speed to traits describing oneself versus the
spouse (Aron, Aron, Tudor, & Nelson, 1991). People who
identify closely with a group are also more likely to take
extreme action on behalf of their group, when either their
personal or social identity is activated, suggesting that their
mental representations of themselves and of their group have
fused (Swann, Gmez, Seyle, & Morales, 2009).
Group identification has affective consequences, too.
According to intergroup emotions theory, identifying with a
group makes the group part of a persons self-concept and
thereby a significant source of emotions (D. M. Mackie,
Devos, & Smith, 2000; E. R. Smith, 1993, 1999). When a
group identity is salient, events concerning the group will
evoke emotions in accordance with the events implications
for the group and not necessarily its implications for the individual. Consistent with this theory, group-based emotional
reactions are sometimes distinct from reactions based on
individual concerns, such as when people feel guilt over their
nations colonial past or record of genocide, even though
they did not partake in these historical episodes (Doosje,
Branscombe, Spears, & Manstead, 1998; E. R. Smith, Seger,
& Mackie, 2007). As a result of the groups incorporation
into the self-concept, group-relevant events also affect emotions and cognitions about oneself. For example, male basketball and soccer fans experience a testosterone surge after
their teams victory and a testosterone drop after the teams
defeat (Bernhardt, Dabbs, Fielden, & Lutter, 1998). People
who strongly identify with a sports team also lose faith in
their own mental and social abilities after their team loses a
game (Hirt, Zillman, Erickson, & Kennedy, 1992).
Identification with a group also leads to preferential treatment of group members. The minimal group paradigm strikingly demonstrates the human readiness to identify with a
group and favor ones group over other groups (for reviews,
see Diehl, 1990; Tajfel & Turner, 1986). People discriminate
in favor of their ingroup even when their individual gains do
not depend on allocations by ingroup or outgroup members,
suggesting that this preferential treatment of ingroup members owes to more than considerations of reciprocity (Gagnon
& Bourhis, 1996). Further supporting the idea that group
identification cannot be fully explained by self-interest,
diverse lines of research find that political behavior is driven
by collective rather than individual interests. Research on
relative deprivation shows that peoples perception of their
groups deprivation, as opposed to their personal deprivation, best predicts their political attitudes and behavior
(Guimond & Dub-Simard, 1983; Kawakami & Dion, 1993;
242
Vanneman & Pettigrew, 1972). Southern black college students participation in the civil rights movement, for example,
was predicted more strongly by their feelings about the treatment of black Americans in general than by their discontent
with their own lives (Orbell, 1967). Voting and public opinion show a similar pattern. Reviewing decades of public
opinion research, Kinder (1998) concluded,
Interests, it is now clear, do have a part to play in public opinionsthat is, interests that are collective rather
than personal, group-centered rather than self-centered.
In matters of public opinion, citizens seem to be asking themselves not Whats in it for me? but rather
Whats in it for my group? (as well as Whats in it
for other groups?). (p. 808)
The stronger the identification with the group, the stronger
the groups grip on the individual. People who closely identify with a group are more likely to invest their own
resources in the group (Brewer & Kramer, 1986; De Cremer
& Van Vugt, 1999), more likely to engage in discretionary
cooperative behavior (Tyler & Blader, 2000), less likely to
exit the group when presented with an attractive option
(Abrams, Ando, & Hinkle, 1998; Ellemers, Spears, &
Doosje, 1997; Van Vugt & Hart, 2004), and more likely to
show ingroup bias and ingroup pride (Jackson & Smith,
1999). Social identity processes, in sum, constitute the basic
mechanism by which a group achieves psychological unity.
Status hierarchies and deference to legitimate authority.
Identifying with a group provides the basic internal mechanism that connects individuals with their group and motivates them to act on group interests. Whereas social identity
processes regulate group behavior by providing individuals
with emotional and cognitive incentives to think, feel, and
act in unison with their group, the human tendency to defer
to legitimate authority allows the top-down regulation of
lower-level units. Leadership is born out of a need for effective
collective action. Eminent leadership thinker John Gardner
(1993) wrote, The larger topic of which leadership is a subtopic is the accomplishment of group purpose (p. xvi).
Importantly, leadership is firmly tied to social identity processes. An effective leader is someone who can create a powerful sense of us, who is perceived as one of us, and who
is able and willing to advance our interests (Haslam,
Reicher, & Platow, 2010). A leaders reliance on coercive
means suggests a failure at mobilizing the social identity
processes that cause group members to follow a leader
voluntarily and enthusiastically.
Deference to legitimate authority is a fundamental aspect
of human psychology, as demonstrated by the Milgram
(1974) studies. Milgram wrote that even when the destructive effects of their work become patently clear, and they are
asked to carry out actions incompatible with fundamental
standards of morality, relatively few people have the
resources needed to resist authority (p. 6). This deference
may seem like a form of submission to dominance; however,

the Milgram studies did not involve the use or threat of force.
In general, within human groups deference is freely given
based on status rather than obtained through coercion
(Henrich & Gil-White, 2001). Human status hierarchies differ from the dominance hierarchies common in social primates and some nonprimate species. Whereas dominance
hierarchies determine the allocation of a fixed amount of
resources such as food and mates, human hierarchies often
also aim to accomplish group goals that might expand the
groups resources (Rubin, 2002).
Status hierarchies facilitate collective decision making and
intragroup coordination toward goal pursuit, as low-status
individuals defer to high-status individuals (De Kwaadsteniet
& Van Dijk, 2010; Van Vugt, Hogan, & Kaiser, 2008). This
differentiation of roles among group members is a form of
division of labor that parallels the division of labor within
organisms. But such status differentiation will expand the
resource pie and increase the groups efficiency only if status
is granted to those who exercise their influence for the groups
benefit. Research suggests that this is often the goal. High
status is usually bestowed on those who are expected to contribute most to valued group goals (J. Berger, Cohen, &
Zelditch, 1972; J. Berger, Fisek, Norman, & Zelditch, 1977;
Tyler, 1997; Willer, 2009). In one study, the groups most altruistic members were accorded the highest status. Moreover, as
the costs of altruism increased, the status rewards also
increased (Hardy & Van Vugt, 2006). As a result, individuals
pursue status by enhancing their apparent value to the group
(C. Anderson & Kilduff, 2009). Conversely, people detest
leaders who do not serve group goals and band together
against that leader (Boehm, 1993). By offering status and
respect to those who exert themselves for valued group goals,
well-functioning status hierarchies also serve to align individual goals with group goals (Magee & Galinsky, 2008).
Despite the potential benefits of hierarchies, it should also
be noted that hierarchies may under certain circumstances
hamper group performance and morale (for a review, see
C. Anderson & Brown, 2010). Status differences among
group members may suppress effective information transmission and decision making (Milanovich, Driskell, Stout, &
Salas, 1998; Thomas-Hunt & Phillips, 2004), groups may
fail to assign status to most competent members (C. Anderson
& Kilduff, 2009), and even when they do, power may corrupt those at the top of hierarchies or make them reckless
(C. Anderson & Galinsky, 2006; Kipnis, 1972). Nevertheless,
status hierarchies are functional for coordinating goal pursuit
and motivating individuals to contribute to the group
(Halevy, Chou, & Galinsky, 2011).
In sum, shared intentionality, social identity processes,
and deference to legitimate authority constitute the basic
mechanisms that allow human groups to undertake united
collective action. These psychological processes align individuals with the group, enable division of labor, and make
joint goal achievement possible.
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3. Low Levels of Heritable Variation

Among the Superorganisms Units
Barring mutations, cells in a body are genetically identical.
Bees in a hive are not genetically identical, but they are
highly related. As a result of this genetic homogeneity, the
cells of one organism are genetically far more similar to each
other than they are to the cells of a different organism, and
members of one hive are more similar to each other than they
are to members of other hives. Moreover, these shared
genetic properties are heritable. An organism is genetically
more similar to its parent organism(s) than to other members
of the same species. Similarly, a hive is genetically more
similar to its parent colony than to other bee colonies. In
other words, the level of heritable variation is low within
members of a hive. This section discusses aspects of human
psychology that reduce within-group heritable variation and
increase between-group variation.
Human groups do not have the genetic relatedness of ant
and bee colonies. Migration and intermarriage rates are high,
and human genes often mix across group boundaries (Rogers,
1990). It would thus be hard to make a case for genetic homogeneity among members of any human group. And yet human
beings do have means to reduce heritable variation within
groups, for, unlike other animals, phenotypical variation in
human groups is largely the result of culture. Human culture
thus functions like a social inheritance mechanism that promotes phenotypical similarity in somewhat the same way that
genetic inheritance promotes phenotypical similarity in bees
(Boyd & Richerson, 1985; Jablonka & Lamb, 2005).
Social learning is the mechanism underlying the emergence of between-group cultural differences (Henrich &
Boyd, 1998). Cultural differences arise and persist because
human beings readily absorb language, attitudes, skills, and
norms from members of their groups. Compliance with
social norms is a fundamental aspect of human psychology
(Cialdini & Trost, 1998). Young children become upset when
an experimenter breaks a norm, such as the rules of a game
or the word used to refer to an activity, even if the norm was
established minutes ago in the lab. When this happens, children protest, criticize the norm breaker, try to teach the correct norm, and sometimes tattle (Rakoczy, 2008; Rakoczy,
Warneken, & Tomasello, 2008). These behaviors illustrate
the early ontogenic origins of conformity to a socially constructed reality (P. L. Berger & Luckmann, 1966).
The power of normative influence was recognized in classical studies of social psychology (Asch, 1956; Deutsch &
Gerard, 1955; Sherif, 1936). More recent studies have shown
repeatedly that what other group members do, approve of,
and care about are vastly important determinants of human
behavior. The most reliable way of getting people to do
something is to make others around them do it: Telling people that their neighbors are conserving energy is a better way
of getting them to conserve energy than appealing to their
sense of social responsibility, desire to save money, or hope
to safeguard the Earth for future generations (Nolan, Schultz,

Cialdini, Goldstein, & Griskevicius, 2008). What people will
and will not do closely mirrors what they think others do and
do not do: Peoples perception of how much their friends
cheat on their partners is positively related to how much they
are willing to cheat themselves (Buunk & Bakker, 1995). In
addition, people care about what others around them care
about: The strongest predictor of eating disorders among
elementary and middle school girls is the importance their
peers place on weight, trumping factors such as the girls
self-confidence, actual weight, or desire to look like women
depicted in the media (Taylor et al., 1998).
The psychological evidence on the power of norms is corroborated by the anthropological record. In his analysis of 25
randomly picked cultures from the Human Relations Area
Files, David Sloan Wilson found that even though cultural
norms varied widely, every group had clearly defined and vigilantly enforced norms (Sober & Wilson, 1998). For example,
one record on the Mbuti people who live in the Congo region
of Africa said, Even the most insignificant and routine action
in the daily life of the family is potentially a major concern to
the band as a whole.... It is important that there should be a
pattern of behavior that is generally accepted (Turnbull, 1965,
p. 118, cited in Sober & Wilson, 1998, p. 165).
Part of the reason why people are so eager to follow
norms is that those who stand against the majority are perceived negatively, disliked, and rejected (Bassili & Provencal,
1988; J. M. Levine, 1989). Furthermore, resisting normative
influence feels uncomfortable as a study of brain activity
suggested. In this study, more activation was observed in
participants amygdala, which is associated with negative
emotions, when they disagreed with the confederates in an
Asch-type setting by giving the correct answer (Berns et al.,
2005). These adverse reactions to norm violators and norm
violation ensure that people will adhere to group norms lest
they suffer the internal and external consequences.
Importantly, people follow only their own groups norms
and not just any norms that come their way. The key determinant in norm compliance is identification with the group.
In line with the previous discussion of social identity processes, people who identify with a group conform more to
group attitudes and behavior. In one Asch-type study, psychology students conformed 58% of the time to other psychology students but only 8% of the time to ancient history
students (Abrams, Wetherell, Cochrane, & Hogg, 1990). In
another study, participants were much more responsive to
the canned laughter of ingroup members than the laughter
of outgroup members: They laughed more and for longer,
smiled more, and rated the material as more entertaining
when the canned laughter was attributed to students from
the same university (Platow et al., 2005). Participants were
also more likely to cheat to get money when they had seen
an ingroup member cheat blatantly. Yet when they had seen
an outgroup member cheat, they were less likely to cheat
(Gino, Ayal, & Ariely, 2009).
244
Along the same lines, the more people identify with a

group, the keener they are to observe group norms and
have other members observe the norms, too (Hyman &
Singer, 1968). In one study, the intention to exercise regularly was most strongly associated with the perceived
norms of a reference group for participants who identified
highly with that reference group but not for weakly identified participants (Terry & Hogg, 1996). In another study,
greater identification with a group was associated with
more positive emotions toward members who conformed
to group norms compared to members who violated them
(Christensen, Rothgerber, Wood, & Matz, 2004). People
also care more about group members norm compliance:
According to the black sheep effect, we are generally
less tolerant toward a norm transgressor when this person
belongs to an ingroup rather than an outgroup (Abrams,
Marques, Bown, & Henson, 2000; Marques & Yzerbyt,
1988). Norms, therefore, are primarily binding for group
members, and, in turn, adherence to them is an indicator of
commitment to the group.
In sum, cultural learning is the basis for low heritable
variation in human groups. The susceptibility to the normative influence of fellow group members reduces withingroup heritable variation and increases between-group
differentiation.
4. A Common Fate
Another property of superorganisms is the commonality of
individual fates. Common fate aligns the interests of individual subunits. In a multicellular body, for example, the
organisms reproductive fate coincides with the fitness interests of its cells. Since these cells all share the same genes,
the interests of one cell are identical to those of the next one.
Although bees in a hive are not genetically identical, a
strong alignment of individual interests is ensured by their
high levels of relatedness. In these cases, because individual
outcomes are highly correlated, what is good for one unit
is often also good for the other. In the absence of such an
alignment, a higher-level entity would either not emerge or
be unstable and collapse (Bourke, 2011). What, if anything,
ensures common fate in human groups?
Egalitarianism. Human groups, as noted, do not have the
genetic homogeneity of cells in a body or bees in a hive
reproductive interests do not overlap much within a group.
Do human beings have alternative means of leveling the
variation in individual fates? The answer is yes. To understand these means, we need to shift our gaze from major
transitions that involve related individuals to those that do
not. Some major transitions affected nonrelated parties that
subsequently retained their ability to reproduce. An example
of this is the emergence of meiosis, a form of cell division
whereby each daughter cell gets half the parent cells genetic
material. This is an egalitarian process in that every allele
has just about the same chance of ending up in the sperm or
egg. Because this process is mostly fair, there is not much

individual alleles can do to increase their chances of representation in future generationsexcept for cooperating to make
a finer organism together (Leigh, 1971; Maynard Smith &
Szathmry, 1995). These transitions involving nonkin are
known as egalitarian transitions (Queller, 2000), and they
ensure an alignment of interests through an egalitarian distribution of outcomes. I argue below that human groups also
have a preference for egalitarianism, and this preference helps
to even out the variation in individual fates to some extent.
The major factor sustaining egalitarian outcomes in
human groups is the sense of fairness. This sense runs so
deep that people might feel guilty for surviving catastrophes
when others have not (Niederland, 1961). This survivor
guilt syndrome has been observed among survivors of the
Holocaust and Hiroshima, workers whose colleagues have
been laid off (Brockner, Davy, & Carter, 1985), and sexually
active gay men who tested HIV negative (Wayment, Silver,
& Kemeny, 1995). In experimental setups, people are willing
to pay from their own pockets to secure egalitarian outcomes
for others, taking from the rich and giving to the poor (C. T.
Dawes, Fowler, Johnson, McElreath, & Smirnov, 2007). The
aversion to inequality is apparent in children, too, and it
seems to be stronger in older children than younger children
(Fehr, Bernhard, & Rockenbach, 2008). In one study children had to choose between getting two stickers for themselves versus getting one sticker for themselves whereas
their play partner would also get one sticker. By 5 years of
age, children were far more likely to opt for an egalitarian
distribution (Thompson, Barresi, & Moore, 1997).
Another emotion that sustains egalitarian outcomes is
envy, which generates ill will and hostility toward those perceived to have advantages (R. H. Smith & Kim, 2007). The
human propensity to envy helps level inequalities, as potential targets of envy try to deflect the evil eye by sharing
their fortunes or engaging in other acts of appeasement. A
study found that people provided more help to someone
whom they thought might be maliciously envying them (van
de Ven, Zeelenberg, & Pieters, 2010). Envy thus also plays a
role in evening out the distribution of outcomes.
Egalitarian emotions and cultural practices align individual fates within a group from the bottom up. Variation in
individual fates may also be reduced from the top down as
egalitarian principles get embodied in the practices by
which people govern themselves, such as when all citizens
are granted equal rights before the law and when these laws
mandate an egalitarian distribution of benefits and burdens
(e.g., universal suffrage and universal conscription).
Egalitarianism seems to be characteristic of huntergatherer
and horticulturalist societies that are living in conditions of economic uncertainty (Cashdan, 1980). Boehm (1993) described
several leveling mechanisms in huntergatherer societies that
keep leaders in check and prevent them from obtaining too
much power. For example, members of these societies accept
leadership only in areas of expertise, and chiefs are publicly
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Kesebir
criticized, ignored, or even ostracized if the group does not
approve of their actions (also see Boehm, 1999). One consequence of the preference for equality is that men often do not
acquire enough wealth to have more than one wife even when
polygamy is socially accepted. Anthropologist Murdock
(1949) concludes, An impartial observer employing the criterion of numerical preponderance...would be compelled to
characterize nearly every known human society as monogamous, despite the preference for and frequency of polygyny
in the overwhelming majority (pp. 27-28). Practices and
norms that level the fitness differences within a group, such as
monogamy, are called mechanisms of reproductive leveling
(Alexander, 1987; Bowles, 2006). Food sharing, another such
mechanism, is common among human huntergatherers and
many foragerhorticulturalists (H. Kaplan & Gurven, 2005).
Such egalitarian norms and practices serve to even out
inequalities in a group.
Of course, inequalities characterize past and current civilizations. Throughout history, complex societies with a more
specialized division of labor were as a rule richer and more
powerful. At the same time, these societies were more stratified and unequal (J. R. McNeill & McNeill, 2003). Does that
mean that egalitarian practices are dispensable for human
groups? The social patterns accompanying inequality suggest otherwise. A large body of evidence, for example, indicates that social harmony is threatened in societies with high
levels of income inequality. These societies do not have
much social capital: People trust each other less and participate less in community life (Kawachi, Kennedy, Lochner, &
Prothrow-Stith, 1997; Putnam, Leonardi, & Nanetti, 1993;
Uslaner, 2002). The low levels of social capital are also evident in higher levels of violent crime (Hsieh & Pugh, 1993).
In fact, the level of income inequality seems to be the strongest environmental predictor of homicide rates (Neapolitan,
1999). More equal societies, in contrast, are happier (Oishi,
Kesebir, & Diener, 2011) and healthier (Deaton, 2003;
Wilkinson & Pickett, 2006). These findings have been replicated within and across nations, cross-sectionally, and longitudinally (for a review, see Wilkinson, 2005).
Although correlational, these patterns are consistent with
the idea that an alignment of individual outcomes is more
conducive to social harmony in human groups. Deviations
from egalitarian arrangements in contrast, seem to disrupt
effective group functioning and divide communities.
Intergroup warfare. Common fate is a matter of degree in
human groups, and perhaps nothing aligns individual fates
more compellingly than warfare. Although bee colonies do not
fight, warfare among ant colonies can be ferocious (Hlldobler
& Wilson, 1994). At the battlefield, ants spray poisons over
their opponents, cut off their limbs, and stab them with their
stings. Some ant genera have soldier castes that are idle until
threat is impending, and others conduct cannibal wars. Survival is at stake since whole ant colonies can get annihilated
in the hands of another colony. Warfare may be similarly
consequential for human groups. A defeated group might
perish, lose the freedom of self-governance, or have to concede valuable resources such as territory and material wealth.
The groups women may be coerced to sexual intercourse.
To the extent that intergroup warfare has been part of the
human experience, we can claim that a high degree of common fate characterizes some human groups. Was warfare
historically a prevalent phenomenon?
Humanitys distinctive social nature is said to have
evolved within the context of a huntergatherer lifestyle
during the Pleistocene Epoch, which spans the period from
2 million years ago to about 11,000 years ago (Tooby &
Cosmides, 2005). Given that material evidence is rare and
ambivalent, archeologists are reluctant to make generalizations about warfare in this era (Gat, 2006). To study warfare
in the Pleistocene, researchers instead turn to anthropological evidence from contemporary huntergatherer societies.
This strategy, of course, is also fallible since the hunter
gatherer societies anthropologists study have had contact
with modernity and are marginalized groups today. As such,
they are hardly representative of the huntergatherer societies of the Pleistocene in which we are interested. A partial
work-around is studying Australian Aborigines, who had no
contact with modern civilization and had not been marginalized until the European arrival at the end of the 18th century. The picture of the huntergatherer lifestyle they
provide is thus as close to the Pleistocene lifestyle as we can
ever hope to get.
Before the Europeans arrived in Australia, about 300,000
huntergatherers lived in some 400 to 700 regional groups. It
turns out that these aboriginal groups were highly territorial
and lived in a constant state of warfare (Gat, 2006). Papua
New Guinean huntergatherers were similarly unexposed
until recently, and like the Australian Aboriginal groups,
they fought frequently. Regarding the Papua New Guinea
tribes, anthropologist LeBlanc (2003) wrote that this last
place on Earth to have remained unaffected by modern society was not the most peaceful but one of the most warlike
ever encountered (p. 151). Studies based on larger samples
of huntergatherers affirm these trends. A study of 99 hunter
gatherer societies found that almost all of them were engaged
in warfare at the time they were studied or had been so in
recent past (Divale, 1972). Evidence, although not airtight,
suggests that warfare was common during the Pleistocene.
Moving to the Holocene Epoch, which began about
11,000 ago when the last continental glaciers retreated, the
evidence for warfare is reliable and unequivocal. The
Holocene covers the period when agriculture took off and
human social structures grew far more complex. The evidence for warfare includes defensive location choices for
settlements such as hilltops surrounded by fortifications;
highly concentrated numbers of arrowheads outside these fortifications (particularly at the gates); mass graves with extraordinarily high rates of death by projectiles; evidence of burning
in entire communities; caches of weapons, shields, and armor;
and finally empty zones between clusters of communities,
246
which were presumably contested no-mans-lands too dangerous to inhabit (Keeley, 1996; LeBlanc, 2003).
The anthropological evidence from these two epochs suggests that warfare was common in human prehistory. In addition to the anthropological data, population genetics research
provides striking evidence on the occurrence and consequences of intergroup conflict. Studies show how groups of
men have reproductively replaced other groups of men over
time. One study mapped the genetic patterns of a Columbian
population established in the 16th to 17th centuries, comparing maternal and paternal lines of inheritance (CarvajalCarmona et al., 2000). This populations maternal ancestry
was more than 90% South American, yet the paternal line
was 94% European, suggesting that coalitions of European
men had replaced South American men. Similar, though less
stark, differences have been found for other populations
(e.g., Bortolini et al., 1999; Underhill et al., 2001). These
findings underscore the importance of intergroup competition for reproductive fate, especially for men (on this latter
point, see Geary, Byrd-Craven, Hoard, Vigil, & Numtee,
2003; Van Vugt, De Cremer, & Janssen, 2007). Overall, evidence strongly suggests that intergroup warfare was a stable
element of human history, with drastic consequences for the
group as a whole.
In sum, egalitarian emotions, norms, and practices serve
to bring individual fates closer together within a group.
Egalitarian institutional practices such as the rule of law also
equalize outcomes. A paradigmatic case of common fate is
intergroup warfare, where much is at stake for the group as a
whole. The evidence suggests that warfare was an integral
part of human history.
5. Mechanisms to Resolve Conflicts of

Interest in Favor of the Collective: Prosocial
Emotions, Norms, and Institutions
Collective enterprises can easily be undermined by indi
viduals who selfishly impose costs on the group or enjoy the
benefits of public goods without bearing their fair share of
the costs. For cooperative alliances to take off and endure,
free riding needs to be minimized. A common fate reduces
conflicts of interests by aligning them, yet such conflicts are
never fully eliminated. Although shared genes ensure a very
high coincidence of interests, conflicts of interests remain
even in multicellular organisms. For instance, cancer occurs
when mutations in some somatic cells cause them to multiply at the expense of other cell lineages and, eventually, at
the expense of the whole body. In honeybees, too, the coincidence of fitness interests is not complete, and worker bees
could increase their own fitness by laying eggs. But this
rarely happens because many worker bees restrain themselves from laying eggs. Moreover, they are coerced by the
policing behavior of other worker bees that eat worker-laid
eggs (Ratnieks & Visscher, 1989). Turning to human groups,
we similarly find various forms of both self-restraint and

coercion that help resolve interest conflicts in favor of the
collective. This section discusses how human groups counter the free-rider problem and promote within-group cooperation by building on the other capabilities discussed so far,
including language, a strong propensity to comply with
group norms, and an intense need for belonging and
approval.
Human interactions are coordinated by a variety of emotions that orchestrate social behavior. Of particular interest to
us are moral emotions that provide internal rewards for
cooperation and internal costs for noncooperation (Fessler &
Haley, 2003; Haidt, 2003; Tangney, Stuewig, & Mashek,
2007). For example, the anticipation of shame, embarrassment, and guilt makes individuals resist the temptation of
acting in nonnormative or noncooperative ways. Similarly,
distress at others suffering prevents us from harming others,
whereas gratitude motivates us to reciprocate benefits. The
importance of moral emotions to cooperation and social
order is highlighted by the portraits of those who lack them.
Psychopaths have a deficiency in feeling moral emotions
such as guilt, shame, and distress at others suffering (Blair,
1999). This deficiency makes them a major source of social
trouble. Even though they make up less than 1% of the population, they commit more than 50% of the most serious crimes
such as serial murder (Hare, 1993). Moral emotions thus
serve as self-restraint mechanisms.
Moral emotions also fulfill a coercive function. One
important emotion that keeps free riders in check is moralistic anger toward noncooperators. In economic games, people
are eager to punish defectors even if they have to pay for it
(Fehr & Gchter, 2002; Henrich et al., 2006). Moreover,
peoples dorsal striatum is activated when they punish those
who do not cooperate. This brain part is implicated in the
processing of rewards, suggesting that people derive satisfaction from punishing noncooperators (de Quervain et al.,
2004). This punishment, in turn, has been shown to uphold
cooperation levels in a public goods game, in the absence of
which cooperation quickly disappears (Boyd & Richerson,
1992; Fehr & Gchter, 2002).
A remarkable aspect of moral emotions is that they are
concerned with third parties. We are not just inclined to punish those who cheat us and reward those who cooperate with
us. We also have moral reactions to events that do not directly
concern us. For example, many people feel uplifted when
they hear about someone who risks his life to save a stranger
from being run over by a train or someone who dedicates her
life to providing medical care to the underprivileged (Haidt,
2003). People also react negatively to moral transgressions
that do not directly affect them (see M. A. Nowak & Sigmund,
2005). In one study, participants included as third parties in
dictator or prisoners dilemma games had the option of punishing other players by paying out of their own endowment.
Of the participants, 60% punished violators of fairness and
cooperation norms, even though these participants were not
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affected by the transaction and had to pay to punish (Fehr &
Fischbacher, 2004).
Sanctions may entail very little or no cost to the punisher
when they take the form of social disapproval or exclusion.
People are deeply averse to criticism, ridicule, and ostracism
and strongly motivated to get others to approve and like
them (Baumeister & Leary, 1995; Leary, 2001; Williams,
2001). Our sense of self-worth is contingent on our perception of how much we are socially accepted (Leary, Tambor,
Terdal, & Downs, 1995). Moreover, social exclusion activates brain areas that are also associated with physical pain,
suggesting that social pain may be just as compelling and
consequential as physical pain (Eisenberger, Lieberman, &
Williams, 2003; Lieberman & Eisenberger, 2009; MacDonald
& Leary, 2005; also see Zhong & Leonardelli, 2008).
Disapproval and exclusion are thus low-cost mechanisms to
encourage cooperation.
Given that people care about norm violations and are
willing to impose sanctions, concern with ones reputation is
critical to both individual social success and the sustenance of
cooperation (Milinski, Semmann, & Krambeck, 2002a, 2002b;
Sommerfeld, Krambeck, Semmann, & Milinski, 2007).
Information about others reputations is also highly valuable as
it allows us to navigate the social world more adroitly.
Reflecting this high premium on social information, people
keenly seek out gossip and sensational news. According to one
study covering eight countries and three centuries, cheating
and violence were the most common themes of sensational
front page stories (Davis & McLeod, 2003). This attention to
the social behavior of third parties, coupled with the willingness to impose sanctions on them, renders the pursuit of selfinterest at the expense of others an ineffective strategy and
serves the group as a whole.
All human groups value cooperative norms (e.g., honesty,
reciprocity, generosity, and hospitality) and teach them to
their children (Brown, 1991). Given the crucial role of normative influence in shaping human behavior, this prevalence
of cooperative norms provides another layer of incentives for
cooperation. Importantly, though, it has been repeatedly
shown that people are more likely to act cooperatively
toward ingroup members than outgroup members (Hornstein,
Masor, Sole, & Heilman, 1971; M. Levine, Cassidy, Brazier,
& Reicher, 2002). In one study, supporters of the Manchester
United soccer team were encouraged to think of themselves
as Manchester United fans. When they were next exposed to
a staged emergency, they helped an injured stranger wearing
a Manchester United shirt more than they helped an injured
stranger wearing a plain sports shirt or a fan wearing a
Liverpool shirt. But when they were first encouraged to think
of themselves as soccer fans, they helped the strangers wearing either teams shirt but not the stranger wearing a plain
shirt (M. Levine, Prosser, Evans, & Reicher, 2005). Like
normative conformity, then, cooperation is more common
and robust when ingroup members are involved.
In sum, human psychology is configured to make cooperation with group members rewarding and noncooperation distressing. In addition, cultural practices that build
on capabilities such as language and normative compliance serve to align individual interests with collective
interests. As a result, cooperation is often a win-win strategy for members of a group, whereas selfishness is often
self-defeating.
This concludes the section on the superorganismic
aspects of human psychology and human groups. The discussion indicates that each superorganismic feature is in
some form present in human beings. Moreover, different
superorganismic features act in concert and sustain each
other. For example, symbolic communication is crucial to
social identity processes, coordination and division of labor,
propagation of group norms, and reputation mechanisms
that keep free riders in check. Similarly, social identity
processes determine which norms will be followed and
enforced and who will be helped. Norms, in turn, often prescribe fairness and cooperation as well as loyalty to the
group. Human psychology thus presents an array of interlocking features that synergistically work to support superorganismic structures.
Superorganismic Aspects
of Human Social Phenomena
So far, I have discussed the features characterizing superorganisms and reviewed the evidence on their occurrence in
human beings. Each of these features was treated in isolation. This section presents two emergent social phenomena,
religion and morality, that present a confluence of multiple
superorganismic components.
Religion
Several social scientists have argued that religions primary
function is to foster group cooperation and solidarity
(Durkheim, 1915/1967; Rappaport, 1979, 1999; V. Turner,
1969). Durkheim noted that a religions symbols and beliefs
are secondary to its role in forging a community. According
to him, understanding religion requires looking beyond a
religions specific beliefs and zeroing in on what it does to a
group of peoplenamely, affirming and sanctifying its normative order:
[S]omething eternal in religion...is destined to survive all the particular symbols in which religious
thought has successively enveloped itself. There can
be no society which does not feel the need of upholding and reaffirming at regular intervals the collective
sentiments and the collective ideas which make
its unity and its personality. (Durkheim, 1915/1967,
pp. 474-475)
248
Anthropologist Rappaport (1999) similarly remarked on

religions unifying role:
[H]igh-order meaning...is grounded in identity or
unity, the radical identification or unification of self
with other. It is not so much, or even at all, intellectual
but is, rather, experiential. It may be experienced
through art, or in the acts of love, but is, perhaps, most
often felt in ritual and other religious devotions. (p. 71)
These accounts suggest that religion is a distinctly effective
means of binding groups (Graham & Haidt, 2010; D. S.
Wilson, 2002).
Empirical research supports the idea of religion as a
group-binding mechanism. A study of 19th-century American
communes shows that religious communes were more stable
and durable than their secular counterparts. Every year, utopian communes with religious convictions were 2 to 4 times
more likely to survive than secular communes (Sosis, 2000).
When religious people interact with fellow group members,
they achieve higher rates of cooperation than do members of
a matched secular group (Sosis & Ruffle, 2003).
Consistent with the idea of religion as a superorganismic
regulator, religious beliefs flexibly adjust to a societys
cooperation needs. Religiosity flourishes more readily under
conditions of threat and scarcity, when regulation of individual behavior is particularly crucial to the groups existence. In a sample of societies included in the Human
Relations Area Files, celestial deities were more common in
societies under resource stress (Hayden, 1987). Other
research shows that societies with water scarcity are more
likely to have moralizing gods, thereby promoting more prosocial behavior (Snarey, 1996). Deities also become more
powerful, punitive, and moralistic with increasing group
size, presumably because social regulation becomes harder
as groups grow (Roes & Raymond, 2003).
In sum, humanitys various religions seem to serve group
bonding through subordination to a sacred authority and/or
belief system that regulates individual action for the common
good. This authority demands cooperative behavior from
group members. Religion also provides people with a strong
social identity, prosocial norms, and incentives to abide by
those norms. As such, religions simultaneously strike multiple chords of the human superorganismic psychology.
Human Morality
Across cultures, superorganismic features are incorporated
into moral codes. Based on their anthropological work in
India, Shweder and colleagues proposed the ethics of autonomy, community, and divinity to account for different moral
systems across cultures (Shweder, Much, Mahapatra, &
Park, 1997). The ethics of autonomy, the moral code of
autonomous individuals, is concerned with harm, rights,
and justice. The ethics of community and divinity, in con-
trast, presume an individual who is deeply implanted in a

role-based social structure within a sacred order and whose
preferences are subordinated to communal obligations and
demands of the sacred domain. The moral codes of community and divinity capture issues such as proper gender roles,
observation of traditions, obedience to authority, and respect
for the natural order.
Haidt and his colleagues have extended Shweders theory
by identifying five domains of morality, corresponding to five
moral taste receptors (Haidt & Joseph, 2007). The first two,
harm/care and fairness/reciprocity, correspond to the ethics of
autonomy. The harm/care foundation derives from our evolutionary history as a mammalian species with an attachment
system. It underlines the basic moral capacity to feel and dislike the pain of others. This capacity, however, can be suspended when outgroup members are concerned, such as in
warfare (Bandura, 1999). The fairness/reciprocity foundation
underlies the human concern with egalitarianism.
The remaining three foundations are ingroup/loyalty,
authority/respect, and purity/sanctity. Each of these foundations expresses an affirmation of one or more superorganismic features and serves the cause of tighter groups (Haidt &
Graham, 2009). The ingroup/loyalty foundation is about
valuing ones group, putting it before personal interests, and
being hostile toward free riders and traitors. It is concerned
with sharpening group boundaries, prioritizing the group
over individuals, and creating a united front. A manifestation of this foundation is the outrage evoked by traitors
(Baumeister, 1999, pp. 194-197; Marques & Yzerbyt,
1988). The enmity bestowed on traitors far exceeds that
bestowed on enemies. In Dantes Inferno for example, traitors are found in the lowest level of hell, below even the
nonbelievers, the lustful, the violent, the murderous, and the
fraudulent. In experimental games, people inflict harsher
punishments on noncooperating group members than noncooperating outgroup members (Shinada, Yamagishi, &
Ohmura, 2004). This foundation is about placing the group
(the whole) above individuals (the parts). Such primacy of
the whole over the part is a major characteristic of any
superorganism.
The moral foundation of hierarchy/duty calls for respectful submission to legitimate authorities, institutions, traditions, and restraints. Even though Western liberals sometimes
equate hierarchy with subjugation and oppression, this is a
restricted vision of the authority/respect foundation (Fiske,
1992; Haidt & Graham, 2009). Legitimate authority creates
mutual obligations and is conceptually the opposite of
power, which is the one-sided exercise of muscle (Nisbet,
1966/1993). As we saw, respect for authority serves coordination and unity of action within a group. Respect for institutions and traditions also is a form of conformity to
time-honored group norms.
The purity/sanctity foundation is related to concerns
about sacredness and preservation of the natural order of creation. It is about respect for conventions defining the proper
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ways of living as prescribed by a sacredly held cultural system. The goal is to protect the sacred domain from any
encroachment of the profane. Any such encroachment would
violate the sacred authority to which the group has subordinated itself and thereby threaten the meaning system and the
unity of the people sharing the conceptions of the sacred.
Cultures and individuals vary in their endorsement of these
last three moral foundations. In modern societies, liberals are
less likely to endorse them, and conservatives are more likely
(Graham, Haidt, & Nosek, 2009). Although not uniformly recognized as virtues in modern societies, these three group-serving foundations were highly valued in many groups studied by
historians and anthropologists (Haidt & Graham, 2007; Haidt,
Koller, & Dias, 1993; Miller & Bersoff, 1992; Miller, Bersoff,
& Harwood, 1990; Shweder et al., 1997).
In sum, all five moral foundations are associated with superorganismic features. In particular, the foundations of ingroup/
outgroup, hierarchy/duty, and purity/sanctity are functional for
forming and sustaining tightly bound groups. They demand
loyalty to the group as well as respect for its values, traditions,
and institutions. They also demand cooperation from individuals to achieve group goals, even if doing so means that they
have to subordinate their own interests or sacrifice themselves.
Religion and morality thus both engage various superorganismic aspects of human psychology. Consequently, they
can be instrumental in the formation of superorganismic
social structures.
The Group Reflex

We have seen that superorganismic capabilities are part of
humanitys psychological repertoire and are evident in the
major social phenomena of religion and morality. At the
same time, their manifestation varies across time and place:
Human behavior reflects a constantly shifting balance
between individual and group concerns (Brewer & Caporael,
2006). We repeatedly saw, for example, how the activation
of superorganismic human traits is conditional on group
identification. This section describes the factors associated
with variations in the endorsement of superorganismic principles across time and place.
Immediate ecological conditions set an organisms motivational priorities, and many evolutionarily adaptive mechanisms need environmental triggers to become active
(Kenrick, Griskevicius, Neuberg, & Schaller, 2010). An
instructive case comes from slime molds: When food is
abundant, slime molds are invisibly dispersed on damp forest soils and wander around as single cells. But when the
conditions deteriorate, these cells secrete chemicals that
make the individual cells coalesce into a multicellular body.
This body then crawls as a single superorganism, and once a
better spot is found, cells are dispersed again and return to
their single-cell life (Bonner, 2008). Slime molds bear a
striking resemblance to human beings in the way they oscillate between individualistic and superorganismic behaviors.
Binding to a group is an optional strategy for human beings

that is prompted by external threats and relaxed under conditions of peace and plenty.
A threat to the group is a major trigger for social cohesion
(Branscombe, Ellemers, Spears, & Doosje, 1999; Brewer &
Campbell, 1976; Dion, 1979; LeVine & Campbell, 1972).
The threat may be realistic (e.g., a competing outgroup or
failing leadership), psychological (e.g., a challenge to the
groups status and prestige), or symbolic (e.g., challenges to
the groups normative order, shared values, beliefs, and morals). All these threats fuel ethnocentrism, conformity, outgroup discrimination, and stronger ingroup bias in those
who strongly identify with the group (Bobo, 1999; Riek,
Mania, & Gaertner, 2006; Stephan & Stephan, 2000). When
they feel that their groups status is under threat, people who
closely identify with their group become more likely to derogate outgroup members (Branscombe & Wann, 1994; Grant,
1993), to see their ingroup as homogeneous (Doosje, Ellemers,
& Spears, 1995), and to self-stereotype (Spears, Doosje, &
Ellemers, 1997). A review of the literature concluded,
Group-level threat combined with high group commitment is associated with perceptual, affective, and
behavioral reactions aimed at the group reasserting
itself in terms of either value or distinctiveness. This
may lead to a high degree of self-stereotyping, expressions of strong ingroup loyalty, and a readiness for
collective action. (Ellemers, Spears, & Doosje, 2002,
p. 178)
Note again, how group identification is a key moderator.
Venturing beyond social psychology, other disciplines
reach parallel conclusions on the solidarity-inducing role of
threat. Sociologist Nisbet (1953/1990) wrote, Society attains
its maximum sense of organization and community and its
most exalted sense of moral purpose during the period of
war (p. 35). Political scientists who have investigated the
effects of external threats on the public named the patriotic
reactions to external threats the rallying around the flag
reflex (Lee, 1977; Mueller, 1973). This reflex manifests itself
as increased support for leaders during times of war or international crisis. For example, according to Gallup polls, immediately after the terrorist attacks of 9/11, George W. Bushs
approval rating catapulted from 51% at the beginning of
September 2001 to his all-time high of 90% at the end of
the month. Moreover, between 2001 and 2004, his approval
rating went up every time the terror alert levels were raised
(Willer, 2004). This rallying around the president phenomenon in times of crisis is likewise a well-established empirical
pattern (Brody, 1991).
Perceived threats to the group often also increase authoritarian tendencies that seek to eliminate diversity and deviance. Authoritarianism refers to a concern with uniformity
and group authority over diversity and individual autonomy
(Duckitt, 1989; Stenner, 2005). According to political
250
scientist Stenner (2005), authoritarianism tends to produce

a characteristic array of stances, all of which have the effect
of glorifying, encouraging, and rewarding uniformity and of
disparaging, suppressing, and punishing difference (p. 16).
As such, authoritarianism is a marker of relative emphasis on
group concerns over individual concerns (Adorno, FrenkelBrunswik, Levinson, & Sanford, 1950; Altemeyer, 1981). In
her book The Authoritarian Dynamic, Stenner (2005) argued
that authoritarianism is best understood as a dynamic
response to societal conditions and not as a stable trait.
Perceptions of threat to the groups oneness and sameness
trigger the authoritarian response, and societal threats are
empirically associated with increases in authoritarianism
(McCann, 1997; Sales, 1973; Winter, 1996). Research in the
United States shows that periods of societal threats, such as
economic downturns, saw higher conversion rates to fundamentalist church dominations (Sales, 1972), more censorship attempts, and a higher willingness to express prejudiced
attitudes (Doty, Peterson, & Winter, 1991). Experimental
studies similarly find that exposure to normative threats such
as lack of social consensus increase authoritarian attitudes
and decrease support for democratic values (for a review, see
Stenner, 2005).
Conservatism is related to, but conceptually distinct from,
authoritarianism. As noted, conservatives are more concerned
with group-oriented moral foundations than are liberals
(Graham et al., 2009; Haidt & Graham, 2007). Therefore, if
threats to social stability foster a group-oriented mentality,
they may also move people to the political right. Indeed, people who were in or near the World Trade Center on 9/11 were
more likely to subsequently shift away from liberalism toward
conservatism (Bonanno & Jost, 2006). When reminded of
9/11, people also reported higher endorsement for George W.
Bush and his policies over his liberal opponent (Landau et al.,
2004). Similarly, a Spanish study conducted before and after
the 2004 Madrid terrorist attack found higher levels of prejudice against outgroups, support for authoritarianism, and
attachment to traditional conservative values after the attacks
(Echebarria-Echabe & Fernndez-Guede, 2006). More generally, a meta-analysis showed that one of the two strongest predictors of conservatism is perceptions of threatened social
stability, with a mean correlation coefficient of 0.47 (Jost,
Glaser, Kruglanski, & Sulloway, 2003).
As noted above, nowhere is threat to a groups existence
more acute than in intergroup combat, and that is exactly
where the highest amount of self-sacrifice occurs. Warfare
therefore represents the pinnacle of human superorganismic
potential. People put their lives at risksometimes with full
knowledge of their impending deathas in suicide attacks.
Such attacks are not an invention of our times. From the
Jewish sect of Zealots who organized suicide attacks on the
Roman Empire to the Japanese kamikaze pilots, they have
been part of intergroup conflict throughout history. Suicide
attackers are neither psychologically sick individuals nor
motivated by a sense of personal desperation (McCauley,
2007). Yet they not only sacrifice themselves for their group
but often do so with a sense of joy, as evinced by writings
and interviews they leave behind (Oliver & Steinberg, 2006).
A Hamas militant, for example, asked his family to receive
the news of his martyrdom with elation and chants to God
for it is a day of celebration (cited in Hafez, 2006, p. 177).
Two scholars who interviewed suicide bombers noted,
These guys knew theyre going to die.... At the same time,
we get the feeling that life has never seemed better to them
so intense, so exuberant, so full of meaning (Oliver &
Steinberg, 2006, p. 118). Apparently, suicide attackers derive
meaning and satisfaction from sacrificing themselves for
their group.
Similarly, at least some men report the ease of sacrificing
themselves for their comrades during combat. Comradeship
reaches its peak during battle, and attachment to ones buddies and refusal to let them down is a major motive in combat (Stouffer et al., 1949). Philosopher Glenn Gray (1959),
who fought in World War II, described the ease of selfsacrifice on the battlefield:
Such sacrifice seems hard and heroic to those who
have never felt communal ecstasy. In fact, it is not
nearly so difficult as many less absolute acts in peacetime and in civilian life, for death becomes in measure
unreal and unbelievable to one who is sharing his life
with his companions. (p. 46)
Whether in suicide attacks or during combat, when ones
group is under severe threat, self-sacrifice seems to come
easily, at least to some. Referring to the ease with which
brothers in arms are ready to sacrifice themselves, D. S.
Wilson and Sober (1994) wrote, Quibbles about the definition of altruism aside, nothing more is required to convert a
social group into an organism (p. 601). Just as cells are
expendable for a body and individual bees are expendable
for the hive, human beings, under some circumstances, are
expendable for their group.
Whereas threats to the group bond people together, the
lack of external threats weakens group solidarity. Writing on
the history of northern Africa, 14th-century Arab thinker Ibn
Khaldun noted that tribes would conquer states on the northern coast thanks to their high level of group solidarity. But
these tribes soon would be replaced by other conquering
tribes, as this solidarity declined after successive generations
of settled life (Ibn Khaldun, 1969). Building on this theory,
Turchin (2006) showed that empires almost always form on
frontiers between religious or ethnic groups. This is presumably because the frontiers are where constant threats from
other groups enable the buildup of strong solidarity necessary to displace other groups. However, this military success,
driven by the willingness to die for ones group, eventually
sets in motion processes that reduce the groups solidarity.
The lack of an external threat, combined with rising inequalities within empires, leads to higher within-group competition
251
Kesebir
and a decline in group solidarity. Solidarity thus seems to melt
away in the absence of external threats. Recently, psychological research has also shown that the loss of individual control
over ones fate leads to a heightened belief in the legitimacy of
controlling sociopolitical institutions or an interventionist
God (Kay, Gaucher, Napier, Callan, & Laurin, 2008). The
combined evidence thus strongly suggests that higher-level
regulatory systems lose their appeal to the extent that people
have control over their lives, such as when they are not constricted by intergroup threats, poverty, or natural disaster.
In sum, converging multidisciplinary evidence suggests
that threats to the group trigger group cohesiveness. Perceptions
of threat set off psychological processes that enable group
members to face the threat as a unified body. Whereas people
bond together in response to a threat, similar to the way slime
molds transition to a superorganismic state, the lack of adversity atrophies group cohesiveness.
Conclusion
This article has used the superorganism metaphor to reflect
on human sociality. I have reviewed the evidence for superorganismic aspects of human sociality and argued that multiple psychological and cultural mechanisms make human
groups resemble superorganisms: Human beings have versatile means of integration through communication as well as
mechanisms to achieve united action and to promote cooperation. Religions and morality present rich ensembles of
superorganismic features as well as the tensions intrinsic to
human groups.
Superorganismic conceptions of human psychology, as
noted, have not featured prominently in social psychology.
Unfavorable reactions to Wundts Vlkerpsychologie or to
the group mind idea, although partly justified, have resulted
in a psychology that has neglected the vision of the individual as an element within a larger whole. Looking at human
societies through a superorganism lens allows for a clearer
appreciation of the full scope of human existence. A unifying narrative emerges for phenomena that are treated piecemeal within an individualist paradigm. According to this
narrative, cultural meaning systems, shared intentionality,
norm compliance, deference to authority, social identity processes, religiosity, and morality can be understood parsimoniously as manifestations of the same dynamics that create
superorganism-like social structures. Superorganisms thus
offer a useful heuristic around which to organize our understanding of human sociality.
Despite the heuristics usefulness, it is important to be clear
on how human groups are not like beehives, or any other
superorganisms. Human groups are fuzzier and less solid than
beehives (Campbell, 1958). Unlike bees, most modern human
beings are not particles of one all-encompassing social structure. Human groups are fluid and human loyalties are multiple. People can simultaneously identify with various groups
and break their attachment to them at will. At any time, one of
multiple identities may be salient and shape behavior.

Honeybees do not have such flexibility. What characterizes
human beings is thus not membership in one discrete superorganism but a capacity to create and function in superorganismic structures.
Granting this human capacity for building superorganismic structures, there is also no question that most human
groups are much less superorganismic than a beehive. Selfish
motives are pervasive and conflict is frequent within groups,
as individuals compete over mates, resources, and power.
Subversion from within threatens all groups. Nepotistic tendencies and narrow interest groups distort political systems
and undermine the collective good. Individuals constantly
negotiate the tension between individuality and group membership (Brewer & Caporael, 2006; Stenner, 2005). On a
larger scale, this fundamental tension between individual
and group concerns is reflected in the culture wars between
conservatives and liberals or religious people and atheists.
The superorganism metaphor is thus obviously not perfect
its success is qualified.
The task of this article has been describing how and when
human groups are like superorganisms. The answers raise a
third question that I have not addressed: Why are human
groups like superorganisms? The why question invites an
evolutionary explanation. Specifically, we have to ask
whether the superorganism metaphor works because humans
actually have gone through a major evolutionary transition to
arrive at superorganismic capacity. Do we have in our hands
a case of convergent evolution rather than just a surface
resemblance? Even though this article has not sought to
make an evolutionary case for a major transition account, the
reviewed evidence speaks to the possibility of a major transition for two reasons. First and simply, the abundance of
superorganismic human features suggests that a major transition might have taken place. If human groups act like superorganisms in so many ways, we have to consider the
possibility of a major evolutionary transition.
Second, the existence of superorganismic human traits
might itself have facilitated a major evolutionary transition
(Bourke, 2011). Major biological transitions often involve
positive feedback loops (Crespi, 2004). The idea here is that
superorganismic traits might trigger dynamics that will lead
to their further proliferation and thus accelerate the transition. We can imagine a straightforward example involving
human warfare. Intergroup warfare would select for more
superorganismic groups that are better able to muster their
collective capacity to defeat an enemy (Turchin, 2006). At
the same time, higher levels of collective capacity could be
directed toward expropriating other groups resources, thus
promoting more intergroup warfare. An autocatalytic process would ensue by which superorganismic capacity leads
to warfare and warfare selects more superorganismic groups
over generations. Another example of a positive feedback
loop involving human superorganismic traits is the coevolution of cooperative cultural norms and genes for cooperation
252
(Henrich, 2004). Cultural norms that discourage cheating

would favor the proliferation of genes that make individuals
more cooperative as noncooperative individuals would be
excluded from valuable coalitions. Once these genes were
selected, the carriers of these cooperative genes would more
vigorously observe cooperative cultural practices, leading to
further proliferation of cooperative genes. As these two
examples indicate, superorganismic traits could set in motion
positive feedback loops. This is why the psychological evidence reviewed here is relevant to the evolutionary account of
human sociality yet to emerge in full shape.
As noted in the opening paragraph, beehives have often
been invoked as symbols of utter harmony and models for
the ideal human society. Not all artistic representations of
beehives were so favorable, though. Beehives have also
inspired chilling society portraits in science fiction, as in
Frank Herberts (1973) book Hellstroms Hive. Of course,
one need not consult science fiction to discern the dark side
of human superorganisms. The long history of warfare and
the fascist regimes of the 20th century in particular teach us
how ugly things get when superorganismic human features
rule the day. Although this article implies that the quickest
way to unite humanity would be an interstellar attack, there
is power in knowing the hivish minds intricacies. What sets
human groups apart from other superorganisms is the flexibility with which superorganismic potential is exercised.
Human moral imagination is not bound by narrowly constructed group identities, as evinced by the existence of organizations with universal moral commitments, such as
Amnesty International, Greenpeace, and Doctors Without
Borders. If we grant that hivishness is part of who we are, the
human challenge becomes converting it into a force for
good, based on what social scientists have uncovered about
its workings.
Acknowledgments
For their insights and thoughtful comments, I thank Jon Haidt,
Gary Sherman, Yoav Bar-Anan, Jesse Graham, Shigehiro Oishi,
Carlee Hawkins, Nicole Lindner, Simon Pearish, and N. Sriram.
Declaration of Conflicting Interests

The author(s) declared no potential conflicts of interest with
respect to the research, authorship, and/or publication of this
article.
Funding
The author(s) received no financial support for the research,
authorship, and/or publication of this article.
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Personality and Social Psychology

Society for Personality and Social Psychology

>> Version of Record - Jul 3, 2012

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The Superorganism Account of

Personality and Social Psychology Review

University of Virginia, Charlottesville,VA, USA

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Personality and Social Psychology Review 16(3)

(Hlldobler & Wilson, 2009). Beyond sheer numbers, social

Are Human Groups

The History of the

parts of a god-created primal giant (OFlaherty, 1981). The

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small groups of strangers, and group behavior has often been

Major Transitions in Evolution and

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Personality and Social Psychology Review 16(3)

Major transitions are among the most important events of

3. Low levels of heritable variation among the superorganisms units

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1. Integration of lower-level units through

to be reviewed later, for symbolic communication is essential to each of them.

group formation and maintenance by making it costly to

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Personality and Social Psychology Review 16(3)

They create and cultivate distinctive myths, memories of

As the dancer loses himself in the dance, as he

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2. Unity of Action: Shared Intentionality,

and anticipated behaviors as well as the task environment.

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Personality and Social Psychology Review 16(3)

Human eyes seem to be designed to enhance the gaze signal,

very differently from the chimpanzees: They waited for

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example, people are faster to say that a particular trait

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Personality and Social Psychology Review 16(3)

may seem like a form of submission to dominance; however,

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3. Low Levels of Heritable Variation

to safeguard the Earth for future generations (Nolan, Schultz,

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Personality and Social Psychology Review 16(3)

Along the same lines, the more people identify with a

egg. Because this process is mostly fair, there is not much

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Personality and Social Psychology Review 16(3)

5. Mechanisms to Resolve Conflicts of

we similarly find various forms of both self-restraint and

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Personality and Social Psychology Review 16(3)

Anthropologist Rappaport (1999) similarly remarked on

trast, presume an individual who is deeply implanted in a

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The Group Reflex

Binding to a group is an optional strategy for human beings

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Personality and Social Psychology Review 16(3)

scientist Stenner (2005), authoritarianism tends to produce

1. Integration of lower-level units through