Aggression and Violent Behavior 16 (2011) 428–436

Contents lists available at ScienceDirect

Aggression and Violent Behavior

Collective violence: An evolutionary perspective

Russil Durrant ⁎
Institute of Criminology, School of Social and Cultural Studies, Victoria University of Wellington, PO Box 600 Wellington, New Zealand

a r t i c l e i n f o a b s t r a c t

Article history: The seeming ubiquity of war, genocide, and other forms of group conflict in human history has led many
Received 14 March 2011 scholars to conclude that our capacity for collective violence is firmly rooted in the evolutionary history of our
Received in revised form 1 April 2011 species. For many social scientists, however, this view is anathema: war and genocide are not evolved features
Accepted 4 April 2011
of our human nature but are, rather, best viewed as evolutionary by-products, arising from particular social
Available online 20 April 2011
and ecological circumstances. The primary aim of this paper is to provide an introduction to, and critical
Keywords:
evaluation of, evolutionary approaches to understanding collective violence. In particular, I focus on the
Collective violence question of whether collective violence can be considered an evolutionary adaptation or whether it is best
War viewed as a by-product of other evolved adaptations, emerging only in particular social, ecological, and
Evolution political contexts. Although the available evidence is certainly consistent with the view that our capacity for
collective violence has been selected for during the course of our evolutionary history, the idea that it is a by-
product cannot be ruled out. A richer understanding of collective violence is likely to be developed through a
more thorough integration of distal and proximate explanations.
© 2011 Elsevier Ltd. All rights reserved.

Contents

1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 428
2. What is collective violence? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 429
3. Evolutionary explanations for collective violence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 429
3.1. Collective violence as an evolutionary adaptation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 429
3.2. Collective violence as a by-product . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 431
4. Evaluating evolutionary approaches to collective violence: evidence from history, anthropology, primatology and psychology . . . . . . . . . 431
4.1. History and pre-history . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 431
4.2. Hunter–gatherers and small scale societies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 432
4.3. Other species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 432
4.4. Proximate psychological mechanisms and processes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 433
5. Evaluating evolutionary approaches to collective violence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 434
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 435

1. Introduction many scholars to conclude that our capacity for collective violence is
firmly rooted in the evolutionary history of our species (e.g., Bigelow,
No one doubts that humans have the capacity to organise 1969; Gat, 2006; Le Blanc, 2003; Smith, 2007; Thayer, 2004; Van der
themselves into groups and engage in systematic violence against Dennen, 1995; Wrangham & Peterson, 1996). Although, as discussed
other groups of humans. The historical record vividly illustrates that below, there are a number of different evolutionary approaches to
our capacity for war, genocide, terrorism and other forms of collective understanding collective violence, one relatively widespread assump-
violence is an important, and often appalling, source of suffering, tion is that our capacity for collective violence has been selected for in
misery and death in human society. The seeming ubiquity of war, our evolutionary history. In short, it is an evolutionary adaptation. For
genocide, and other forms of group conflict in human history has led many social scientists, however, this view is anathema: war and
genocide are not evolved features of our human nature but are, rather,
best viewed as cultural inventions or evolutionary by-products,
⁎ Tel.: + 64 04 463 9980. arising from particular social and ecological circumstances (e.g., R. B.
E-mail address: russil.durrant@vuw.ac.nz. Ferguson, 2006; Fry, 2007; Kelly, 2000; Mead, 1940; O'Connell, 1995).

1359-1789/$ – see front matter © 2011 Elsevier Ltd. All rights reserved.
doi:10.1016/j.avb.2011.04.014
 R. Durrant / Aggression and Violent Behavior 16 (2011) 428–436 429

Indeed, in the Seville Statement on Violence, drafted in 1986 by a group group is transitory or has a more permanent identity – against
of twenty leading scientists, the potential biological basis for war is another group or set of individuals.
explicitly rejected: “It is scientifically incorrect to say that we have
inherited a tendency to make war from our animal ancestors… Although collective violence often involves acts of individual
Violence is neither in our evolutionary legacy nor in our genes”. violence, it can be distinguished from violence that occurs between
Instead, “warfare… is a product of culture” (cited in Adams, 1989, individuals who do not act as part of, or on behalf of, particular groups.
p. 120). In sum, debates over the origins of war and other types of Perhaps most importantly, collective violence involves social substi-
collective violence reflect, in often sharpened form, more enduring tutability (Kelly, 2000): members of one group direct violence against
issues regarding the origin of human nature. others, not as specific individuals per se, but because they are
The primary aim of this paper is to provide an introduction to, and members of another group. Inevitably, as with almost all definitions,
critical evaluation of, evolutionary approaches to understanding there will be borderline cases of collective violence. Some “hate
collective violence. Although there are a number of different crimes”, for instance, may be perpetrated against others of a particular
approaches to applying evolutionary theory to human behaviour race, or sexual orientation by individuals who view themselves as part
(see Brown, Dickins, Sear, & Laland, 2011; Durrant & Ward, 2011), all of a particular group even if that group is not collectively acting
such approaches assume that humans possess psychological and against such individuals in any obvious or coordinated way. Some
behavioural adaptations that have been selected for because they forms of terrorism may also involve individuals acting on behalf of
advanced reproductive fitness in ancestral environments. I focus in groups that are not clearly defined or with clearly articulated goals
this paper on the question of whether collective violence (or, more regarding the target group (see Burleigh, 2008; Taylor, 2010).
accurately, the mechanism(s) underlying the expression of collective However, the definition offered above does clearly incorporate most
violence) can be considered an evolutionary adaptation or whether it instances that we would want to consider collective violence such as
is best viewed as a by-product of other evolved adaptations, emerging war (in all its forms), organised terrorism, state-perpetrated violence
only in particular social, ecological, and political contexts. I begin by against particular target groups, genocide, and gang warfare (Zwi
briefly discussing some of the conceptual issues involved in defining et al., 2002).
collective violence and then I consider some recent evolutionary The adoption of this, inclusive, definition for collective violence has
theories of collective violence. A review of the relevant evidence is several advantages. Importantly, it can help unite somewhat disparate
then provided, focussing in turn on the historical and pre-historical areas of inquiry by directing attention to the more general
record, ethnographic research on hunter–gatherers and small scale phenomenon of collective violence rather than to specific types or
societies, coalitional aggression in non-human species, and the instances of collective violence such as genocide, terrorism, or war. Of
proximate mechanisms that underpin collective violence. I conclude course, we will still want to be able to provide more specific
that the currently available evidence does not allow us to choose, with explanations for particular types or instances of collective violence.
confidence, among competing evolutionary explanations for the Thus, there will be particular features of “suicide terrorism”, the
origin of collective violence. However, I suggest that nature/culture Rwandan genocide, the rape of Nanking, and the Allied bombing of
dichotomies are unlikely to prove fruitful and that we can advance our Dresden during World War II that require due attention to the
understanding of collective violence by adopting an evolutionary particular local social and political contexts. However, these and other
perspective that takes into account the cultural nature of our species such examples can also be viewed as particular instances (or
and that integrates more distal explanations (in terms of evolutionary instantiations) of a more general phenomenon: the capacity for
history, function, and cultural evolution) with more proximate humans of one “group” to engage in collective aggression and violence
developmental, social, and psychological explanations. against individuals of another “group”. It is this general phenomenon
that needs to be explained. I shall therefore largely refer to “collective
2. What is collective violence? violence” in this paper, although because most of the relevant
theoretical work has focussed specifically on warfare, I will use the
For the 19th century political theorist and Prussian army officer, term “war” largely interchangeably with “collective violence”.
Karl von Clausewitz (1780–1831), war was, famously, “the contin-
uation of politics by other means”. Although this definition pithily
3. Evolutionary explanations for collective violence
captures the organised character of warfare between large groups of
protagonists (in Clausewitz's time, states, nations, or empires) for
If we accept that collective violence is a robust human phenom-
economic and political advantage, providing a clear definition of war
enon, that requires an explanation, then at an evolutionary level of
that captures all of its instances – including raids, ambushes, sieges,
analysis there are only two logical possibilities: either (1) collective
pitched battles and so forth – is more elusive. When we think of war
violence is an evolutionary adaptation; or (2) collective violence is
we naturally consider more prototypical examples – World War II,
best viewed as a by-product of other evolved adaptations. Each of
the Vietnam War, the Peloponnesian War – that involve large-scale
these possibilities is considered in turn.
battles between relatively clearly defined groups. However, a strong
case can be made that the often equally lethal raiding that occurs
among some hunter–gatherer groups should also be considered 3.1. Collective violence as an evolutionary adaptation
“war”. We can run in to similar problems when we try and formulate
clear definitions for other examples of group violence and mass A number of distinct, but overlapping evolutionary approaches to
killing such as “massacres”, “ethnic cleansing”, “genocide”, and understanding collective violence (with a particular focus on war)
“terrorism”. have been developed in the last two decades (see Gat, 2006; Le Blanc,
I suggest that we can fruitfully view all of these phenomena (and 2003; Smith, 2007; Thayer, 2004; Van der Dennen, 1995; Potts &
others) as instances of “collective violence” (sometimes referred to Hayden, 2008; Wrangham & Peterson, 1996 for recent book length
also as “coalitional aggression” or “group aggression”). In this paper I treatments of the literature). These approaches share the common
shall adopt a slightly modified version of the definition of collective assumption that warfare has been selected for in human evolutionary
violence offered by Zwi et al. (2002, p. 215): history, although they differ in terms of the hypothesised evolutionary
function, and particular evolutionary trajectory of collective violence.
[Collective violence is] the instrumental use of violence by people Space precludes a comprehensive review of this literature and I will
who identify themselves as members of a group – whether this focus, instead, on four representative recent evolutionary theories of
430 R. Durrant / Aggression and Violent Behavior 16 (2011) 428–436

collective violence (see Table 1): (1) the imbalance-of-power hy- intelligence, and a form of “proto-ethnocentrism” that allows the
pothesis (Wrangham, 1999; Wrangham & Peterson, 1996); (2) war demarcation of in-group and out-group and the favouring of the
as a “facultative male-coalitional reproductive strategy” (Van der former over the latter. In combination these factors allowed for the
Dennen, 1995, 2002); (3) the “male warrior hypothesis” (Van Vugt, evolution of male coalitional violence against other groups which
2009); and (4) the parochial altruism hypothesis (Choi & Bowles, would have been selected for because it increased reproductive fitness.
2007). The “male warrior hypothesis” (Van Vugt, 2009), shares a number
The imbalance-of-power hypothesis was derived, in part, from of the assumptions of the two preceding evolutionary hypotheses for
observations of inter-group violence among chimpanzees (see the origin of collective violence. In particular, Van Vugt (2009) argues
Section 4.3), but has been extended to explain the evolutionary that recurrent inter-group conflict in human evolution reflects (and
origin of collective violence among humans (Wrangham & Peterson, has, in turn, shaped) the evolution of male coalitional psychology
1996; Wrangham, 1999). According to this hypothesis, collective because men benefited (in evolutionary terms) from collective
violence occurs “because of two factors: intergroup hostility, and large violence through better access to food and mates. Van Vugt
power asymmetries between rival parties” (Wrangham, 199, p. 3). In emphasises that we should therefore expect males and females to
short, collective violence is not necessarily an endemic or ongoing differ in significant ways in terms of their inter-group psychology. For
feature of inter-group relations, but rather emerges when hostility instance, males should have more competitive intergroup experi-
occurs between two groups and when there are large power ences, show more in-group loyalty in times of intergroup conflict, and
differences which enable collective violence to occur with relatively show stronger political support for warfare (Van Vugt, 2009, p. 131,
little risk to the protagonists. The evolutionary function of collective also see Van Vugt, De Cremer & Janssen, 2007).
violence is hypothesised to be inter-community dominance which, in Finally, a recent proposal by Choi and Bowles (2007) suggests that
turn, results in better access to reproductively relevant resources such human warfare and “parochial altruism” (preferentially favouring and
as food, females, and safety (Wrangham, 1999). In sum, “the cooperating with in-group members) may have coevolved in human
imbalance-of-power hypothesis suggests that selection has favoured evolutionary history through a process of between-group competi-
certain emotional predispositions in males that cause aggressive tion. The existence of parochial altruism is well documented in human
behaviour to be elicited relatively easily under certain circumstances” groups (see in particular Section 4.4 below) but, as Bowles (2008)
(Wrangham, 1999, p. 22). Thus Wrangham locates the evolution of notes, is a puzzle from an evolutionary perspective: altruism and
collective violence in humans to characteristics shared by humans and parochialism are both potentially costly behaviours as altruists are
chimpanzees (and presumably by the common ancestor of both) bested by non-altruists and parochialism may result in fitness
which can advance reproductive fitness under specific social and reducing conflict with out-group members, and therefore should be
ecological contexts. selected against. It is argued that the solution lies in the possibility
Van der Dennen (1995, 2002) draws on a comprehensive body of that altruism and parochialism co-evolved. Essentially, groups that
evidence relating to collective violence in other species, human contained individuals who cooperated with group members at a cost
history, and ethnographic research to argue that “the complex of to themselves (altruism) and who engaged in hostile interactions
collective behaviours we call war evolved as a facultative male- with out-group members (parochialism) would have been more
coalitional reproductive (or parental investment) strategy” (van der successful in securing reproductively relevant resources (e.g., territo-
Dennen, p. 539). In short, van der Dennen argues that the warfare ry, food) relative to groups that did not contain individuals
evolved because it advanced reproductive success through (1) better predisposed to behave in this fashion. Thus, according to this model,
accesses to territory and resources and; (2) increased reproductive the human capacity for altruism and what Richerson and Boyd (2005)
opportunities via access to women. The emergence of warfare and term “ultrasociality” (i.e. wide scale cooperation with non-kin) are
other forms of collective violence in human history is based on a intimately linked to our equally enduring capacity for parochialism
number of “pre-conditions” that are, in large part, shared with and out-group enmity, including warfare.
chimpanzees and some other social mammals. These include the Although there are some important differences in the evolutionary
capacity to form coalitions, group territoriality, “Machiavellian” hypotheses presented above, they share some overarching similarities.

Table 1
Evolutionary approaches to explaining collective violence.

Evolutionary perspective Adaptation Evolutionary function Selected publications

Imbalance of power hypothesis Yes Inter-community dominance and Wrangham (1999)

better access to reproductively
relevant resources
Male-coalitional reproductive strategy Yes Better access to resources and Van der Dennen (1995)
territory; increased reproductive
opportunities for males
Male warrior hypothesis Yes Better access to food and more Van Vugt (2009)
reproductive opportunities for
males
Parochial altruism Yes Access to food and resources Choi and Bowles (2007)
Social contexts No, warfare only emerged with No specific evolved function; R. B. Ferguson (2006)
development of particular types warfare occurs when in rational
of human societies; by-product interest of individuals to wage
of general intelligence war
General intelligence No, war by-product of general No specific evolved function Roscoe (2007)
intelligence that must overcome
natural aversions to killing
Segmented societies Yes, in early hominids, but not No specific evolved function Kelly (2005)
from around 1 million years ago;
warfare product of particular
types of society (segmented
societies)
 R. Durrant / Aggression and Violent Behavior 16 (2011) 428–436 431

First, it is assumed that collective violence has been selected for also made coalitional killing increasingly more risky even given an
(or maintained by selection) because it enhanced reproductive imbalance of power as a single individual has the ability to inflict
success. Collective violence is a costly activity which obviously serious injuries against a larger group of protagonists (unlike, for
involves a significant risk of mortality for protagonists. For it to have instance, chimpanzees). Thus Kelly envisions an evolutionary scenario
evolved, therefore, the benefits on average must have outweighed the where coalitional killing may have been present in the hominid
costs. Most evolutionary hypotheses assume that these benefits would lineage up to around a million years ago when it was subsequently
have included better access to food and other resources (e.g., through selected against, only to become widespread during the Mesolithic
the expansion of territory); increase reproductive opportunities period with the emergence of segmented societies.
through rape, the abduction of women, and via more in-group In sum, the idea that the capacity for collective violence is a by-
reproductive partners (because successful male warriors may obtain product is typically coupled with the view that it only emerges under
more mates); and safety, by eliminating or neutralising the threat of specific social and ecological contexts that would have only become
other groups. There are some differences in terms of the specific common during the last 10,000 to 12,000 years or so. Roscoe (2007)
evolutionary history of collective violence, but it is largely assumed and R. B. Ferguson (2006) assume that under these circumstances
that the human capacity for collective violence can be traced to the warfare may simply reflect the operation of our general human
common ancestor of humans and chimpanzee (especially male–male capacity to weigh up the costs and benefit of our actions and engage in
coalitions and the general tendency for inter-group hostility), collective violence when it is in our individual (and group's) self-
although its specific features in humans reflect our unique evolution- interest.
ary history. A number of scholars also suggest that an evolutionary
history of inter-group conflict has been an important selective force in 4. Evaluating evolutionary approaches to collective violence:
human evolution shaping various aspects of our evolved psychology, evidence from history, anthropology, primatology and psychology
including our capacity for widespread altruism and “ultrasociality”
(e.g., Bowles, 2008, 2009; Van Vugt, 2009). 4.1. History and pre-history

3.2. Collective violence as a by-product In terms of the sheer numbers of victims, the twentieth century is,
without doubt, the bloodiest on record. Not only did this period
The idea that the human capacity for collective violence is an witness the carnage wrought by two world wars and numerous less
evolutionary adaptation that has been specifically selected for has widespread conflicts, but it also experienced some of the worse acts of
been challenged by a number of scholars who view inter-group genocide in history, including the holocaust, the Stalinist purges, and
violence (in particular, war) as the emergent product of specific social the Rwandan genocide in 1994 (see Dutton, Boyanowksy & Bond,
and ecological contexts (R. B. Ferguson, 1996, 2006; Fry, 2007; Kelly, 2005; N. Ferguson 2006). Estimates vary considerably, but it is likely
2000, 2005; Otterbein, 2004; Roscoe, 2007). From this perspective, that anywhere between 170 and 300 million individuals lost their
collective violence has not been a frequent and enduring feature of lives due to collective violence (particularly war and genocide) during
human evolutionary history; rather, it has only become common this period (Zwi et al., 2002; Rummel, cited in Adler, Smith, Fishman &
within the last 12,000 years or so. Larson, 2004). Collective violence is, of course, not a phenomenon that
For instance, R. B. Ferguson (2006) argues that, because the is restricted to the twentieth century, and wars, genocide, massacres,
frequency and intensity of warfare varies significantly cross-culturally terrorism, and other forms of collective violence are a prominent
and because there is little in the way of convincing evidence of (some might argue the prominent) feature of recorded history.
warfare before around 12,000 years ago, warfare is best viewed as the Eckhardt (1991), for instance, estimates that in 1266 wars between
emergent product of particular types of societies. Specifically, warfare 300 BC and 2000 AD over 150 million individuals were killed.
tends to emerge among societies that are sedentary, experiencing A growing body of evidence supports the contention that collective
significant population growth, are socially segmented (partitioned violence was also a feature of many societies prior to the development
into clans and lineages), are hierarchical, engage in the trade of elite of writing and the beginning of historical records (see Keeley, 1996; Le
goods, and experience climactic disturbance (R. B. Ferguson, 2006). Blanc, 2003; Walker, 2001 for reviews). The evidence for war in
There are, therefore, no specific adaptations for war according to prehistory comes from a variety of sources (R. B. Ferguson, 1996; Le
Ferguson; rather, wars are waged when they are in the rational Blanc, 2003). These include the remains of fortifications, artwork
interest of individuals (especially leaders) and groups and such depicting scenes of collective violence, mass burials, weapons, and
interests tend to coincide with particular types of social arrangement. skeletal remains that indicate death by violence (e.g., through
A similar position is advanced by Roscoe (2007) who suggests that patterns of bone trauma, embedded arrow and spear points, and
collective violence is the product of general human intelligence, and evidence of scalping). Some prominent examples include the Oftnet
that humans actually have an aversion to the killing of conspecifics, site in Bavaria, dated to around 7720 years ago, in which the remains
which must be overcome through the deployment of specific of 38 humans were found with clear evidence that they had been
cognitive strategies: “A species capable of developing an advanced bludgeoned to death; evidence for fortifications dating to around
mental model of the self and the world has the capacity both to 6000 BCE in modern day Turkey; and skeletal remains with embedded
recognise when lethal violence can be advantageously deployed and points from many North American sites from around 6000 years ago
to devise strategies capable of short-circuiting whatever genetically (Le Blanc, 2003; Walker, 2001).
based, emotional impediments obstruct its deployment” (p. 492). Space precludes a detailed review of this evidence, but there are
Kelly (2000, 2005) also suggests that the human capacity for some general points to note. First, it is important to recognise that
collective violence is best viewed as the emergent product of partic- evidence for violent death does not necessarily imply that the death or
ular types of society (specifically, segmented societies) and that deaths were the result of collective violence. A skeletal remain with an
warfare would not have featured prominently before around embedded spear point clearly indicates that an individual has been
12,000 years ago. Kelly's argument is based on the putative costs of killed by another human, but this may have occurred as a result of
collective violence during this period. Groups engaged in inter-group single homicide, be an instance of capital punishment, or reflect an
conflict, he suggests, would need to reduce their effective foraging episode of collective violence. Second, although the evidence for
area to maintain a safe buffer zone with other hostile groups, thus collective violence is reasonably conclusive and widespread during
reducing access to resources. The emergence of the human capacity of the Mesolithic (the last 10,000 to 12,000 years), the extent of this
killing at a distance, perhaps around a million years ago, would have evidence varies from location to location (R. B. Ferguson, 1996).
432 R. Durrant / Aggression and Violent Behavior 16 (2011) 428–436

Finally, apart from a couple of contested examples, there is very little violence among extant hunter–gatherers, war would have also been
direct evidence for the existence of collective violence before the a prominent feature of hominid evolution during the Pleistocene. This
Mesolithic period (R. B. Ferguson, 1996, 2006). may be the case, but there are at least two reasons for caution. First, as
There are two logical interpretations for this pattern of evidence. many anthropologist recognise (see Ember & Ember, 1996) extant
First, consistent with the by-product hypothesis, warfare and other hunter–gatherers may not be representative of hunter–gatherers in
forms of collective violence only emerged (or, at least, only became the Pleistocene as they typically occupy more marginal ecological
common) during the last 10,000 years or so, due to changes in contexts and many have been affected to some degree by contact with
settlement patterns, demography, material culture, or ecological the West. Second, as Richerson, Boyd and Bettinger (2001) has
conditions (R. B. Ferguson, 1996, 2006; Fry, 2007; Kelly, 2000). argued, the last 10,000 years of human history (a period known as the
Second, the scarcity of conclusive evidence for collective violence Holocene) has provided very different ecological environments to
prior to the Mesolithic period should not be taken as evidence of those experienced during much of the Pleistocene (the two million
absence and given the widespread practise of warfare over the last years before this) and that we should, therefore, be very cautious in
10,000 years or so we should expect that collective violence was a extrapolating from extant hunter–gatherers to likely conditions
prominent feature of human populations during the Pleistocene era experienced during much of human evolution. In particular, climatic
and before (Gat, 2006; Le Blanc, 2003; Van der Dennen, 2002). The conditions during the Pleistocene were highly variable and generally
current state of the evidence does not allow us to choose between harsher than those experienced during the Pleistocene. This may have
these scenarios. However, it is clear that collective violence occurs resulted in human societies that were largely nomadic, small in size,
fairly readily in a range of different social and environmental contexts and politically egalitarian (Shultziner et al., 2010). Collective violence
(and is thus not inextricably linked to the emergence of agriculture or may well have been limited under such circumstances as Fry (2007)
the development of state societies). The evidence from ethnographic argues, although this remains a matter of debate.
research on hunter–gatherers and small-scale societies provides
further support for the relative ubiquity of warfare in human society. 4.3. Other species

4.2. Hunter–gatherers and small scale societies A third source of evidence for the evolution of collective violence in
humans comes from research on non-human animals, predominantly
The collective evidence from research on hunter–gatherers and chimpanzees and other primates. In his observations on Man's Place in
small scale societies clearly indicates that warfare is a relatively Nature Mark Twain (1896, cited in Smith, 2007) noted that “man is the
prominent feature of such societies. In a comprehensive overview of only animal that deals in that atrocity of atrocities, war. He is the only
186 largely pre-industrial societies, Ember and Ember (1996) found that one that gathers his brethren around about him and goes forth in cold
war was “absent or rare” in 27.6% of the sample (i.e. occurring less than blood and calm pulse to exterminate his kind”. With the exception of
once every 10 years) and was largely constant (occurring at any time ants, this view held sway until the remarkable observations of inter-
during the year) in 38.6% of the sample. However, many of the societies group violence among chimpanzees came to light in the 1970s.
in the sample were “pacified” in the sense that collective violence had Although intra-specific killing had been observed in a number of
been suppressed by external powers. Among “non-pacified” societies different species, this was the first clear evidence of collective violence
(N= 90), war was absent or rare in only 8.98%, and constant in 56.66% of in another vertebrate species. Since these initial observations,
the sample. Collective violence is, then, a relatively common phenom- instances of inter-group aggression, violence and killing have now
enon among hunter–gatherers and other small scale societies although been documented in a number of geographically distant chimpanzee
its occurrence varies considerably across both time and space and it is populations (Boesch et al., 2008; Watts, Muller, Amsler, Mbabazi,
important to recognise that it is rare or absent in some societies (Fry, Mitani, 2006; Wilson, Wallauer, & Pusey, 2004).
2007). Although coalitional violence in chimpanzees has attracted the
Collective violence in small scale societies usually involves either most attention, coalitional inter-group aggression does occur in a
“pitched battles” or “raids” (see Gat, 1999, 2006; Keeley, 1996; Le number of social carnivores such as lions, cheetahs and hyenas
Blanc, 2003 for further details). During pitched battles large numbers (Wrangham, 1999), and in some other primate species such as the
of men from specific clans or tribes face off against other groups of white-faced capuchin (Gros-Louis, Perry & Manson, 2003). However,
men. For instance, ethnographic research among the highland the form that inter-group violence takes among chimpanzees appears
agriculturists of New Guinea has shown how men from different to be largely unique among non-human animals (although see Aureli,
clans assemble at a prearranged site and – from a relatively safe Schaffner, Verpooten, Slater & Ramos-Fernandez, 2006 for instances of
distance – hurl spears or shoot arrows at their opponents. These non-lethal coalitional raiding in male spider monkeys) and bears
battles typically result in few casualties, although they can sometimes some striking resemblances to human warfare. Specifically, groups of
escalate into more serious conflicts. A more lethal form of violence, male chimpanzees engage in apparently deliberate “border patrols” of
widely practised among small-scale societies, is the raid or ambush. A their territory. They also make deep incursions into the territory of
raid typically involves a small group of men from one tribe or clan other chimpanzee groups and in a coordinated fashion may kill the
entering another clan's territory (often under cover of darkness), and males of the other group if they encounter them. What is particularly
then deliberately killing individuals they find. The scale of such raids striking to observers of these behaviours is their apparently deliberate
varies substantially – from the targeted killing of specific individuals and intentional character: they seem to be specifically initiated for
or families to attempts to massacre whole tribes – and they often these purposes, rather than simply the accidental by-product of
involve the abduction of women (Gat, 2006). The specific motives or encountering males from other groups. The frequency of inter-group
rationales for warfare in small scale societies typically include revenge conflicts vary significantly among different chimpanzee groups (see
or retaliation, the procurement of resources, the abduction of women, Boesch et al., 2008), although they may be an important source of
prestige, and defence (see Chagnon, 1988; Gat, 2000a, 2000b; Le mortality in some communities (Williams et al., 2008; Wrangham,
Blanc, 2003). Wilson & Muller, 2006).
What can we conclude from the ethnographic literature? Again, it According to Wrangham's (1999, p.11) “imbalance of power”
is clear that collective violence is a relatively common feature of hypothesis, discussed earlier, “the function of unprovoked intercom-
human societies, although there are significant variations in the munity aggression (i.e., deep incursions and coalitionary attacks) is
frequency and intensity of conflict from society to society. It is intercommunity dominance”. Dominance, in turn, argues Wrangham
tempting to infer that, given the relative ubiquity of collective results in evolutionarily relevant benefits such as better access to food
 R. Durrant / Aggression and Violent Behavior 16 (2011) 428–436 433

resources, safety from inter-group violence, and access to females. At a proximate level, I suggest that two mechanisms or processes
Research has yet to clearly demonstrate the reproductive benefits of are necessary for collective violence to occur: (1) the capacity for
boundary patrolling and inter-group violence (Mitani, 2009), al- coordinated collective action; and (2) an ability to differentiate in-
though both the expansion of territory (Williams, Oehlert & Pusey, group from out-group members. In addition, four key mechanisms or
2004), and the abduction of females (Boesch et al., 2008) have been processes are likely to be important in facilitating collective violence:
documented. Moreover, it appears that the best predictor of boundary (1) a tendency to favour in-group members over out-group members
patrolling is male party size, suggesting the costs of this behaviour are (parochial altruism); (2) a tendency to be distrustful or hostile towards
mitigated by preferentially patrolling in large groups (Mitani & Watts, out-group members (xenophobia); (3) a capacity to selectively disen-
2005). gage “normal” emotional responses that inhibit the killing of conspe-
What are the implications of this research for our understanding of cifics; and (4) an ability to facultatively engage in collective violence (or
the evolution of collective violence in humans? One clear possibility is the threat of collective violence) under specific – adaptively relevant –
that the common ancestor of humans and chimpanzees possessed circumstances.
adaptations for coalitional aggression and that these were largely Clearly humans have the capacity to engage in collective action:
retained throughout the course of hominid (and chimpanzee) individuals are able to successfully coordinate their behaviour in order
evolution although perhaps with some modifications. In other to achieve common goals. Coalitional behaviour is common in a wide
words, the coalitional aggression observed in both humans and range of primate species, although arguably the human capacity for
chimpanzees is a derived character trait of our common ancestor. This coordinated action is unprecedented in terms of its precision and
possibility can be termed “the homology hypothesis”. A second scale. It is easy to take this ability as given, but it depends on (or at
possibility is that coalitional aggression has been independently least is facilitated by) the existence of moral emotions such as shame,
selected for at some point in both the chimpanzee and human lineage, guilt and pride, the ability and motivation to sanction free riders
thus representing an example of convergent evolution. We can call (what Fehr & Gächter, 2002 term “altruistic punishment”) and
this “the homoplasy hypothesis”. A third possibility is that coalitional cognitive capacities such as “mind reading”, joint attention, and
aggression is an evolutionary by-product (i.e., has not been specifically language. Without the ability to engage in collective action, warfare
selected for) in either, or both, chimpanzees and humans. For and other forms of collective violence would not be possible.
instance, coalitional aggression may be an evolutionary adaptation However, given the widespread existence of coalitional behaviour in
in chimpanzees, but the similar behaviour in humans is the emergent primates and the evolutionary benefits of co-operative behaviour
property of other characteristics and has not been specifically selected (e.g., in collective foraging, scavenging, hunting, and predator
for. This can be termed “the analogy hypothesis”. defence) it is unlikely that our capacity for collective action has
The currently available evidence does not allow us to clearly favour been specifically selected for inter-group violence, although once in
any of these hypotheses. The fact that bonobos (who also share a place collective violence may favour the evolution of more effective
common ancestor with chimpanzees and humans) do not engage in and sophisticated coalitional mechanisms as groups (and individual
coalitional inter-group aggression, somewhat weakens the homology group members) that are better able to coordinate the behaviour of
hypothesis as it would have to further postulate the loss of coalitional their members are more successful relative to less coordinated
aggression in the bonobo lineage. Furthermore, the homology groups.
hypothesis also assumes that all of the different hominid species As many decades of research in social psychology have demon-
ancestral to modern humans engaged in collective violence. We strated, humans also clearly have the capacity for differentiating in-
simply do not know enough about the social and group structure of group from out-group members, although there remains some debate
these species to make any authoritative claims on this score regarding the mechanisms that enable this to occur. One possibility is
(Wrangham, 1999). Support for the analogy hypothesis hinges that our ability to discriminate in-groups from out-groups is the by-
crucially on evidence for the idea that collective violence is an product of general cognitive mechanisms underlying our classification
evolutionary adaptation (in humans and/or chimpanzees) and thus and categorisation abilities (Gil-White, 2001). Cosmides, Tooby and
reflects one of the central questions of this paper. In sum, the growing Kurzban (2003), however, argue that group classification arises from
body of research on coalitional aggression among chimpanzees may psychological mechanisms that detect and track coalitions and
provide important insights into the evolutionary origins of collective alliances. Essentially in-group members are those that engage in
violence in humans. It clearly illustrates for instance, that collective coordinated cooperative behaviour. Our capacity to categorise in-
violence can occur in other species and thus may not be tied to specific dividuals into groups is also strongly facilitated by the existence of –
features of particular human societies. However, the current state of relatively hard to fake – “ethnic markers” such as language (including
knowledge does not allow us to make any unequivocal claims about dialects), belief systems, styles of dress, food preferences and
the role of collective violence in human evolution. practises, and social norms (McElreath, Boyd & Richerson, 2003;
Richerson & Boyd, 2005). McElreath et al. (2003) argue that such
ethnic markers provide an indication that an individual shares the
4.4. Proximate psychological mechanisms and processes same social norms and hence is likely to cooperate with other group
members. Humans, then, have the capacity to develop a sense of
A final important source of evidence for evolutionary approaches “collective identity” based on perceptions of coordinated activity,
to collective violence comes from a fairly extensive body of research in common fate, shared norms, values and attitudes, and a sense of
social, personality and cognitive psychology which has examined the collective history (David & Bar-Tal, 2009).
nature of social groups, inter-group relations, and the mechanisms Social psychological research also demonstrates that humans have
and processes that might facilitate the perpetration of violence against a tendency to both favour in-group members over out-group
others (in particular, other groups). This research is important, members and to treat out-group members with suspicion, animosity,
because not only does it elucidate the proximate mechanisms that and hostility. These two processes, however, appear to operate
underpin collective violence, but it also might provide insights into relatively independently from each other: that is, favouring in-
whether the mind appears specifically “designed” to engage in group members doesn't necessarily imply overt hostility towards the
collective violence. In particular, we should be especially interested out-group (Cashdan, 2001) although the two processes often go hand
in mechanisms that appear to efficiently, precisely, and specifically in hand (Hogg & Abrams, 2003). The ease with which humans can
promote collective violence in circumstances that would have been identify with – even arbitrarily defined – groups is testimony to our
most likely to advance reproductive fitness. enduring “need to belong” (Baumeister & Leary, 1995) and has been
434 R. Durrant / Aggression and Violent Behavior 16 (2011) 428–436

demonstrated in numerous so-called “minimal group” experiments. possible that they reflect, in part, conscious cognitive “rationalisa-
The tendency to appraise in-group members more positively is also a tions” that are deliberately employed so as to reduce the psychological
robust social psychological phenomenon and humans seem to readily consequence of inflicting harm on others. In practise it may be very
display distrust, animosity and hostility towards out-group members hard to disentangle these possibilities as the end-product looks much
(Schaller & Neuberg, 2008), but this capacity seems much more fluid the same regardless of whether they are largely unconscious product
and context dependent (Hogg & Abrams, 2003). This pattern seems to of specific evolved psychological mechanisms or whether they reflect
be consistent with the idea that the identification with, and favouring largely conscious cognitive manoeuvres in order to maintain psy-
of, in-groups is a relatively obligate feature of our nature, whereas our chological equanimity.
attitudes towards out-group members is more contingent on specific If our capacity for collective violence is an adaptation we might
contexts. We may, however, be prone to develop negative responses expect mechanisms to operate in a facultative fashion, facilitating
to out-group members fairly readily (Schaller & Neuberg, 2008), and collective violence when it is likely to lead to the most benefits
there is some emerging research to suggest that the processes relative to costs. Research in this area is relatively limited; however,
underlying ethnocentrism may be modulated at the neural level by support for intergroup conflict appears to be more positive after
the peptide, oxytocin (De Dreu, Greer, van Kleef, Shalvi & Handgraaf, instances of out-group aggression (e.g., Carnagey & Anderson, 2007;
2011; De Dreu et al., 2010). From an evolutionary perspective, Cheung-Blunden & Blunden, 2008), and under conditions of threat
although there may be a number of reasons to be wary of out-group there is a tendency to categorise unfamiliar individuals as out-group
members (including disease avoidance — see Faulkner, Schaller, Park members (Miller, Maner, & Becker, 2010). Moreover, consistent with
& Duncan, 2004), there are also benefits from peaceful and evolutionary approaches that emphasise that collective violence is
cooperative relations with out-groups and hence we might expect largely a male phenomenon, intergroup bias appears to be largely
relatively flexible responses. directed at males and results in different responses by males and
Many scholars have noted that humans appear to have a strong females under threatening conditions with males more likely to
reluctance to killing their conspecifics, even if they are members of respond to inter-group threats with aggression towards out-group
other groups and even in times of inter-group conflict (e.g., Grossman, members (Navarrete, McDonald, Molina & Sidanius, 2010; Van Vugt
1996; Roscoe, 2007; Smith, 2007). Grossman (1996), for instance, et al., 2007).
describes how a significant proportion of soldiers fail to fire their
weapons or deliberately aim to miss their opponents during episodes 5. Evaluating evolutionary approaches to collective violence
of armed conflict. The reluctance that humans have in killing others is
likely to largely reflect the evolution of emotions such as empathy (de The available evidence is certainly consistent with the view that
Waal, 2008) and compassion (Goetz, Keltner, & Simon-Thomas, our capacity for collective violence has been selected for during the
2010), which have evolved to facilitate altruistic and caring behaviour. course of our evolutionary history. This would explain the seeming
Strong normative proscriptions against killing in most contexts are ubiquity of war, genocide and other forms of group violence in human
also largely universal in human groups and are often reinforced societies and would account for the psychological mechanisms and
through explicit (though often inconsistent) religious endorsement. processes that parse humans into in-group and out-group members
Bandura (1999, 2002) provides a useful model for understanding and facilitate cooperation among the former and hostility against the
how normal moral mechanisms that inhibit killing in most circum- latter. One distinct possibility, championed by a number of scholars, is
stances may become disengaged in some contexts, particularly those that our capacity for coalitional aggression was a feature of our shared
involving collective violence. Bandura argues that moral agency can common ancestor with chimpanzees and thus has been an enduring
be disengaged through four processes. First, reprehensible conduct part of hominid evolution.
can be reconstrued through moral justification (e.g., the war was The idea that our capacity for collective violence is a by-product,
necessary to prevent the deployment of weapons of mass destruction however, cannot be ruled out. The absence of clear evidence for
or to depose a despot) or through a process of euphemistic labelling warfare prior to the Mesolithic, doubts over the phylogenetic
(e.g., civilian deaths are “collateral damage”). Second, responsibility continuity of coalitional aggression, and the possibility that psycho-
for reprehensible conduct can be attenuated through diffusion (e.g., logical mechanisms that facilitate collective violence have evolved
everyone is acting together and so no one is responsible) or primarily for in-group cooperation (and, perhaps, out-group avoid-
displacement (e.g., I was just following orders) of responsibility. ance) makes this alternative a viable possibility. However, the idea
Third, the detrimental effects of the conduct can be minimised or that warfare is entirely a “cultural invention” emerging in a number of
ignored (e.g., through killing at a distance). Finally, through a process places and then spreading by diffusion (along the lines of writing), or
of blaming victims (“they initiated the conflict”) or dehumanising that is can be explained entirely as the product of general intelligence
them (the enemy are “vermin” or “cockroaches”), killing becomes and “rational choice” do not adequately explain the relevant evidence.
easier. In other words, although it is possible that collective violence is a by-
The existence of these processes is well documented during times product of other evolved adaptations, the relevant adaptations are
of war, genocide, and other instances of collective violence and it is likely to relate specifically to intra and inter-group behaviour and
clear that humans have the capacity to suspend or disengage normal reflect mechanisms that lead humans to favour in-group over out-
emotional and cognitive processes that inhibit killing in some group members.
contexts. It is less clear, however, how best to characterise these Understanding the origin of collective violence is further
mechanisms. Smith (2007) argues that these mechanisms, especially complicated by the potential role of cultural evolution and gene–
the process of dehumanisation, can be viewed as psychological culture co-evolutionary processes. It is clear that cultural processes
adaptations that have been selected for to “allow” humans to engage can play an important role in the nature and extent of collective
in episodes of collective violence. In particular, he suggests that during violence. The development of new military technologies and
war the enemy may be viewed as “sub-human” because evolved strategies (from spears and bows to guns and missiles), social-
predator detection modules are switched on that allow them to be structure (especially the evolution of increasingly stratified, hierar-
viewed as dangerous predators or game, or anti-parasite modules are chical societies), and the role of specific ideologies (including
activated that allow them to be viewed as dangerous pathogens or religions) have profoundly shaped the character of genocide, war,
parasites (see also Faulkner et al., 2004). However, for at least some of terrorism, and other forms of collective violence. Warfare, as Turchin
the mechanisms of moral disengagement posited by Bandura (e.g., (2006) argues, may also have favoured the development of larger,
moral justification, euphemistic labelling, blaming the victim), it is more cooperative societies (particularly in regions where culturally
 R. Durrant / Aggression and Violent Behavior 16 (2011) 428–436 435

dissimilar groups are in contact) through a process of cultural group De Dreu, C. K. W., Greer, L. L., Handgraaf, M. J. J., Shalvi, S., van Kleef, G. A., Baas, M., et al.
(2010). The neuropeptide oxytocin regulates parochial altruism in intergroup conflict
selection as larger, better integrated groups prevail against those that among humans. (June 11). Science, 328, 1408–1411. doi:10.1126/science.1189047.
are smaller and/or less well coordinated. These processes in turn may De Dreu, C. K. W., Greer, L. L., van Kleef, G. A., Shalvi, S., & Handgraaf, M. J. J. (2011).
have favoured the evolution of specific psychological characteristics Oxytocin promotes human ethnocentrism. Proceedings of the National Academy of
Sciences, 108, 1262–1266. doi:10.1073/pnas.1015316108.
through genetic and/or cultural evolutionary processes. A richer de Waal, F. B. M. (2008). Putting the altruism back into altruism: The evolution of
understanding of collective violence is likely to be obtained through a empathy. Annual Review of Psychology, 59, 279–300. doi:10.1146/annurev.psych.
more systematic integration of distal explanations (in terms of 59.103006.093625.
Durrant, R., & Ward, T. (2011). Evolutionary explanations in the social and behavioral
evolutionary history and function, and cultural history) and sciences: Introduction and overview. Aggression and Violent Behavior.
proximate developmental and psychological explanations in terms Dutton, D. G., Boyanowsky, E. O., & Bond, M. H. (2005). Extreme mass homicide: From
of the cognitive, affective and motivational processes that underpin military massacre to genocide. Aggression and Violent Behavior, 10, 437–473. doi:
10.1016/j.avb.2004.06.002.
the expression of collective violence.
Eckhardt, W. (1991). War-related deaths since 3000 BC. Security Dialogue, 22, 437–443.
In effectively proscribing the development of evolutionary doi:10.1177/096701069102200410.
approaches to war, the signatories of the Seville Statement on Ember, C. R., & Ember, M. (1996). Violence in the ethnographic record: Results of cross-
Violence were clearly concerned that such an approach may signal cultural research on war and aggression. In D. L. Martin, & D. W. Frayer (Eds.),
Troubled times: Violence and warfare in the (pp. 1–20). : Gordan and Breach
the inevitably of inter-group conflict and the futility of hopes for Publishers.
enduring peace. However, neither the specific evolutionary hypoth- Faulkner, J., Schaller, M., Park, J. H., & Duncan, L. A. (2004). Evolved disease-avoidance
eses presented in this paper nor the available research evidence mechanisms and contemporary xenophobic attitudes. Group Processes and
Intergroup Relations, 7, 333–353. doi:10.1177/1368430204046142.
suggests that collective violence is, in any straightforward way, Fehr, E., & Gächter, S. (2002). Altruistic punishment in humans. Nature, 415, 137–140.
inevitable. It is clear that inter-group violence is either a facultative Ferguson, N. (2006). The war of the world: History's age of hatred. London: Allen Lane.
adaptation (emerging under a specific set of circumstances) or is a by- Ferguson, R. B. (1996). Violence and war in prehistory. In D. L. Martin, & D. W. Frayer
(Eds.), Troubled times: Violence and warfare in the past (pp. 321–355). : Gordan and
product of adaptations that make collective violence more or less Breach Publishers.
likely depending on specific ecological, social and cultural contexts. Ferguson, R. B. (2006). Tribal, “ethnic”, and global wars. In M. Fitzduff, & C. E. Stout
Moreover, our capacity for collective violence has co-evolved with (Eds.), The psychology of resolving global conflicts: From war to peace. nature vs.
nurture, 1. (pp. 41–69) Westport Conn: Praeger.
social and cultural institutions and practises that are potentially Fry, D. P. (2007). Beyond war: The human potential for peace. Oxford: Oxford University
amenable to change. This means that although we may expect Press.
collective violence to be part of our future as it has been part of our Gat, A. (1999). The pattern of fighting in simple, small-scale, prestate societies. Journal
of Anthropological Research, 55, 563–583.
past, a more thorough understanding of both the distal and proximate
Gat, A. (2000a). The human motivational complex: Evolutionary theory and the causes
causes of inter-group conflict may provide opportunities to limit its of hunter–gatherer fighting, part I. Primary somatic and reproductive causes.
occurrence. Anthropological Quarterly, 73, 20–34.
Gat, A. (2000b). The human motivational complex: Evolutionary theory and the causes
of hunter–gatherer fighting, part II. Proximate, subordinate, and derivative causes.
Anthropological Quarterly, 73, 74–88.
References Gat, A. (2006). War in human civilization. Oxford: Oxford University Press.
Gil-White, F. J. (2001). Are ethnic groups biological “species” to the human brain?
Adams, D. (1989). The Seville statement on violence: A progress report. Journal of Peach Currently Anthropology, 42, 515–554.
Research, 26, 113–121. Goetz, J. L., Keltner, D., & Simon-Thomas, E. (2010). Compassion: An evolutionary
Adler, R. N., Smith, J., Fishman, P., & Larson, E. B. (2004). To prevent, react, and rebuild: analysis and empirical review. Psychological Bulletin, 136, 351–374. doi:10.1037/
Health research and the prevention of genocide. Health Services Research, 39, a0018807.
2027–2051. Gros-Louis, J., Perry, S., & Manson, J. H. (2003). Violent coalitionary attacks and
Aureli, F., Schaffner, C. M., Verpooten, J., Slater, K., & Ramos-Fernandez, G. (2006). intraspecific killing in wild white-faced capuchin monkeys (Cebus capucinus).
Raiding parties of male spider monkeys: Insights into human warfare? American Primates, 44, 341–346. doi:10.1007/s10329-003-0050-z.
Journal of Physical Anthropology, 131, 486–497. doi:10.1002/ajpa.20451. Grossman, D. (1996). On killing: The psychological costs of learning to kill in war and
Bandura, A. (1999). Moral disengagement in the perpetration of inhumanities. society. Boston, MA: Little Brown.
Personality and Social Psychology Review, 3, 193–209. Hogg, M. A., & Abrams, D. (2003). Intergroup behaviour and social identity. In M. A.
Bandura, A. (2002). Selective moral disengagement in the exercise of moral agency. Hogg, & J. Cooper (Eds.), The sage handbook of social psychology (pp. 407–431).
Journal of Moral Education, 31, 101–119. doi:10.1080/0305724022014322. Thousand Oaks, CA: Sage Publications.
Baumeister, R. F., & Leary, M. R. (1995). The need to belong: Desire for interpersonal Keeley, L. H. (1996). War before civilization: The myth of the peaceful savage. New York:
attachment as a fundamental human need. Psychological Bulletin, 117, 497–529. Oxford University Press.
Bigelow, R. (1969). The dawn warriors. Boston: Little-Brown. Kelly, R. C. (2000). Warless societies and the origin of war. Ann Arbor, MI: The University
Boesch, C., Crockford, C., Herbinger, I., Wittig, R., Moebius, Y., & Normand, E. (2008). of Michigan Press.
Intergroup conflicts among chimpanzees in Tai National Park: Lethal violence and Kelly, R. C. (2005). The evolution of lethal intergroup violence. Proceedings of the
the female perspective. American Journal of Primatology, 70, 519–532. doi:10.1002/ National Academy of Sciences, 102, 15294–15298. doi:10.1073/pnas.0505955102.
ajp. 20524. Le Blanc, S. A. (2003). Constant battles: The myth of the peaceful, noble savage. New York,
Bowles, S. (2008). Conflict: Altruism's midwife. (20 November). Nature, 456, 326–327. NY: St. Martin's Press.
Bowles, S. (2009). Did warfare among ancestral hunter–gatherers affect the evolution McElreath, R., Boyd, R., & Richerson, P. J. (2003). Shared norms and the evolution of
of human social behaviours? (5 June). Science, 324, 1293–1298. doi:10.1126/ ethnic markers. Current Anthropology, 44, 122–129.
science.1168112. Mead, M. (1940). Warfare is only an invention — not a biological necessity. Asia, XL,
Brown, G. R., Dickins, T. E., Sear, R., & Laland, K. N. (2011). Evolutionary accounts of 402–405.
human behavioural diversity. Philosophical Transactions of the Royal Society B, 366, Miller, S. L., Maner, J. K., & Becker, D. V. (2010). Self-protective biases in group
313–324. categorization: Threat cues shape the psychological boundary between “us” and
Burleigh, M. (2008). Blood and rage: A cultural history of terrorism. London: Harper Press. “them”. Journal of Personality and Social Psychology, 99, 62–77. doi:10.1037/
Carnagey, N. L., & Anderson, C. A. (2007). Changes in attitudes towards war and violence a0018086.
after September 11, 2001. Aggressive Behavior, 33, 118–129. doi:10.1002/ab.20173. Mitani, J. C. (2009). Cooperation and competition in chimpanzees: Current under-
Cashdan, E. (2001). Ethnocentrism and xenophobia: A cross-cultural study. Current standing and future challenges. Evolutionary Anthropology, 18, 215–227. doi:
Anthropology, 42, 760–765. 10.1002/evan.20229.
Chagnon, N. (1988). Life histories, blood revenge, and warfare in a tribal population. Mitani, J. C., & Watts, D. P. (2005). Correlates of boundary patrol behavior in wild
(Februrary 26). Science, 239, 985–992. chimpanzees. Animal Behaviour, 70, 1079–1086. doi:10.1016/j.anbehav.2005.02.012.
Cheung-Blunden, V., & Blunden, B. (2008). Paving the road to war with group Navarrete, C. D., McDonald, M. M., Molina, L. E., & Sidanius, J. (2010). Prejudice at the
membership, appraisal antecedents, and anger. Aggressive Behavior, 34, 175–189. nexus of race and gender: An outgroup male target hypothesis. Journal of
doi:10.1002/ab.20234. Personality and Social Psychology, 98, 933–945. doi:10.1037/a0017931.
Choi, J. -K., & Bowles, S. (2007). The coevolution of parochial altruism and war. O'Connell, R. L. (1995). Ride of the second horseman: The birth and death of war. New York:
(October). Science, 318, 636–640. doi:10.1126/science.1144237. Oxford University Press.
Cosmides, L., Tooby, J., & Kurzban, R. (2003). Perceptions of race. Trends in Cognitive Otterbein, K. F. (2004). How war began. College Station: Texas A&M University Press.
Sciences, 17, 173–179. doi:10.1016/S1364-6613(03)00057-3. Potts, M., & Hayden, T. (2008). Sex and war: How biology explains warfare and terrorism
David, O., & Bar-Tal, D. (2009). A sociopsychological conception of collective identity: and offers a path to a safer world. Dallas, TX: BenBella books.
The case of national identity as an example. Personality and Social Psychology Richerson, P. J., & Boyd, R. (2005). Not by genes alone: How culture transformed human
Review, 13, 354–379. doi:10.1177/1088868309344412. evolution. Chicago: The University of Chicago Press.
436 R. Durrant / Aggression and Violent Behavior 16 (2011) 428–436

Richerson, P. J., Boyd, R., & Bettinger, R. L. (2001). Was agriculture impossible during the Van Vugt, M. D., De Cremer, D., & Janssen, D. P. (2007). Gender differences in
Pleistocene but mandatory during the Holocene? A climate change hypothesis. cooperation and competition: The male-warrior hypothesis. Psychological Science,
American Antiquity, 66, 387–411. 18, 19–23.
Roscoe, P. (2007). Intelligence, coalitional killing, and the antecedents of war. American Walker, P. L. (2001). A bioarchaeological perspective on the history of violence. Annual
Anthropologist, 109(3), 485–495. doi:10.1525/AA.2007.109.3.485. Review of Anthropology, 30, 573–596.
Schaller, M., & Neuberg, S. L. (2008). Intergroup prejudices and intergroup conflicts. In Watts, D. P., Muller, M., Amsler, S. J., Mbabazi, G., & Mitani, J. C. (2006). Lethal intergroup
C. Crawford, & D. Krebs (Eds.), Foundations of evolutionary psychology aggression by chimpanzees in Kibale National park, Uganda. American Journal of
(pp. 401–414). New York, NY: Lawrence Erlbaum Associates. Anthropology, 68, 161–180. doi:10.1002/ajp. 20214.
Shultziner, D., Stevens, T., Stevens, M., Stewart, B. A., Hannagan, R. J., & Saltini-Semerari, Williams, J. M., Lonsdorf, E. V., Wilson, M. L., Schumacher-Stankey, J., Goodall, J., &
G. (2010). The causes and scope of political egalitarianism during the last glacial: A Pusey, A. E. (2008). Causes of death in the Kasekela chimpanzees of Gombe
multi-disciplinary perspectives. Biology and Philosophy, 25, 319–346. National Park, Tanzania. American Journal of Primatology, 70, 766–777. doi:10.1002/
Smith, D. L. (2007). The most dangerous animal: Human nature and the origins of war. ajp. 20573.
New York, NY: St. Martin's Press. Williams, J. M., Oehlert, Carlis, J. V., & Pusey, A. E. (2004). Why do male chimpanzees defend
Taylor, M. (2010). Is terrorism a group phenomenon? Aggression and Violent Behavior, a group range? Animal Behaviour, 68, 523–532. doi:10.1016/j.anbehav.2003.09.015.
15, 121–129. Wilson, M. L., Wallauer, W. R., & Pusey, A. E. (2004). New cases of intergroup violence
Thayer, B. A. (2004). Darwin and international relations: On the evolutionary origins of among chimpanzees in Gombe national park, Tanzania. International Journal of
war and ethnic conflict. Lexington, Kentucky: The University of Kentucky Press. Primatology, 25, 523–549.
Turchin, P. (2006). War and peace and war: The life cycles of imperial nations. New York: Wrangham, R. (1999). Evolution of coalitionary killing. Yearbook of Physical Anthropology,
Pi Press. 42, 1–30.
Van der Dennen, J. M. G. (1995). The origin of war: Evolution of a male-coalitional Wrangham, R., & Peterson, D. (1996). Demonic males: Apes and the origin of human
reproductive strategy. Gronigen: Origin Press. violence. London: Bloomsbury.
Van der Dennen, J. M. G. (2002). (Evolutionary) theories of warfare in preindustrial Wrangham, R. W., Wilson, M. L., & Muller, M. N. (2006). Comparative rates of violence in
(foraging) societies. Neuroendocrinology Letters, 213(Suppl. 4), 55–65. chimpanzees and humans. Primates, 47, 14–26. doi:10.1007/s10329-005-0140-1.
Van Vugt, M. (2009). Sex differences in intergroup competition, aggression, and Zwi, A. B., Garfield, R., & Loretti, A. (2002). Collective violence. In E. G. Krug, L. L. Darfield,
warfare: The male warrior hypothesis. Annals of the New York Academy of Science, J. A. Mercy, A. B. Zwi, & R. Lozano (Eds.), World report on violence and health
1167, 124–134. doi:10.1111/j.1749-6632.2009.04539.x. (pp. 215–239). Geneva: World Health Organization.