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McAuliffe K. and Hauser M. (2010) Morality and Evolution. In: Breed M.D. and
Moore J., (eds.) Encyclopedia of Animal Behavior, volume 2, pp. 483-488
Oxford: Academic Press.

© 2010 Elsevier Ltd. All rights reserved.

 Author's personal copy

Morality and Evolution

K. McAuliffe and M. Hauser, Harvard University, Cambridge, MA, USA
ã 2010 Elsevier Ltd. All rights reserved.

. . . of all the differences between man and the lower
animals, the moral sense or conscience is by far the most
important. This sense. . . has a rightful supremacy over
every other principle of human action; it is summed up in
that short but imperious word ought, so full of high
significance. It is the most noble of all the attributes of
man, leading him without a moment’s hesitation to risk
his life for that of a fellow-creature; or after due delibera-
tion, impelled simply by the deep feeling of right or duty,
to sacrifice it in some great cause.
Darwin C (1871) The Descent of Man and Selection in Relation
to Sex, pp. 70–71. Princeton: Princeton University Press.
Introduction
In the Descent of Man, Darwin explored the evolutionary
origins of our moral sense, and as the quotation above
emphasizes, highlighted what we see as three essential
points. First, to understand the origins of our sense of
right and wrong, we must adopt a comparative perspec-
tive, drawing on studies of animals, to reveal what is
uniquely human as opposed to what is shared across
species. Second, understanding the moral sense is funda-
mentally a problem about the power of our conscience to
guide what we ought to do. It is, in Darwin’s terms, the
highest of virtues, giving humans a sense of nobility, and
fundamentally distinguishing them from other animals.
Third, our sense of ought, of what should or could be
done, can lead to either instinctive action (‘without a
moment’s hesitation’) or to a more contemplative stance
(‘after due deliberation’) where we reflect upon par-
ticular principles of justice, and then based on this
analysis, act in such a way that we support some great
moral cause, often at personal cost (‘sacrifice’). These
three points target aspects of phylogeny and proximate
cause, that is, the patterns of evolutionary change and
the psychological mechanisms that either facilitate or
constrain their appearance. Darwin also discussed the
adaptive significance of morality, and in particular, the
selective pressures that may have led to its appearance in
our species. Characteristic of his thinking at the time,
Darwin perceived a strong role for group-level pressure,
such that individuals in groups acting in particularly
altruistic ways would ultimately outcompete groups act-
ing less cooperatively.
In this essay, we further explore the evolutionary ori-
gins of our moral sense, providing a synopsis of the cur-
rent state of empirical play and the issues raised by the
experiments and observations of animals. Like Darwin, we
distinguish questions of proximate and ultimate cause, and
specifically, separate out the issues of phylogeny, adapta-
tion, and psychological mechanism. Like Darwin, we also
distinguish between the psychological mechanisms that
guide our intuitive and rather automatic sense of right or
wrong, from those that underpin our contemplative reflec-
tion of what ought to be. From the perspective of psycho-
logical mechanisms and adaptive function, we can explore
the building blocks of our capacity to decide what is right
and what is wrong, and the conditions under which partic-
ular actions are permissible or forbidden. This perspective
seeks an understanding of the core psychological mechan-
isms that enable organisms, both human and non-human,
to decide what is fair, when harms are permissible, and
when social contracts may be broken.
To this end, we review two distinct sets of literatures.
The first explores studies that fall within the general area
of behavioral economics, and in particular, the processes
that guide cooperation, resource distribution, and a broad
sense of fairness. We focus here on results that relate to
some of the critical features of human cooperation and
altruistic behavior, specifically, attention to inequities,
reputation, punishment, and reciprocity. Second, we
explore the mechanisms of action perception and produc-
tion, and in particular, the extent to which animals distin-
guish intentional from accidental consequences, as well
as the cues they use to decide goal-directed actions.
Together, these processes comprise some of the funda-
mental building blocks that are evolutionarily ancient,
appear early in human ontogeny, and ultimately lead to
a full-fledged moral sense in healthy human adults. We
then conclude with our current and personal sense of
what makes human morality fundamentally different
from what is observed in other animals, focusing specifi-
cally on how we evolved a brain that conceives of that
imperious word ought.
Cooperation and Moral Judgment in
Humans
A great deal of moral philosophy has been devoted to an
exploration of our capacity to cooperate with others,

483
Encyclopedia of Animal Behavior (2010), vol. 2, pp. 483-488
 Author's personal copy
484 Morality and Evolution
sacrifice personal gains for the benefit of others, maintain
social contracts, and appreciate that a sense of justice is
premised on a sense of fairness. This rich tradition is
generally aimed at understanding the guiding principles
that appear to underpin not only our intuitive sense of
cooperative action and distributive justice, but also the
principles that ought to guide our decisions. Thus, for
example, we observe in Rawls’ thinking on justice a clear
distinction between the intuitive principles that may
guide our spontaneous judgments of fairness, and those
that percolate up during a period of considered reflection,
where we consciously divorce ourselves from the poten-
tially powerful and biasing influences of in-versus out-
group partiality (e.g., favoring kin over nonkin). The
question of interest here is how human cooperation,
including the psychological mechanisms that support it,
as well as the selective pressures that led to its particular
design features, evolved.
When biologists have discussed the evolution of coop-
eration, they have often focused on behavior and fitness
consequences, without Taking into account psychological
mechanisms that may be relevant or even required. Thus,
Darwin puzzled over the possibility of altruism by asking
how such a costly behavior could evolve, given that his
theory of natural selection favored self-beneficial actions.
This puzzle vanished when Hamilton, and later Williams,
pointed the way to a different level of analysis, one that
focused on genes as opposed to either individuals or
groups. That is, we can explain why an animal engages
in self-sacrifice for the benefit of another by the fact that
the ‘other’ is a close genetic relative. As such, altruism
evolves by benefiting genes shared in common. Where
puzzles remain today is in explaining the evolution of
cooperation among genetically unrelated individuals,
and especially the kind of large-scale cooperation often
observed among human societies.
A partial answer to these puzzles emerged, interest-
ingly enough, when Trivers proposed his theory of recip-
rocal altruism, blending issues of adaptive function
with psychological constraints. Specifically, and unlike
Hamilton and Williams, Trivers’ theory included not
only a set of evolutionary conditions for the emergence
of cooperation among unrelated individuals, but also a
discussion of requisite psychological mechanisms includ-
ing recognition of individuals, memory of past inte-
ractions, and strong emotional responses to defection,
including moralistic aggression. Thus, Trivers’ analysis
paved the way for what has now become a major focus
in the field: a consideration of both proximate and ulti-
mate concerns related to the evolution of cooperation.
More specifically, and as many have argued, cooperation
among unrelated others can evolve if it takes place within
stable groups with opportunities for repeated interactions
(reciprocal altruism), or in groups where cooperative deci-
sions can be based on reputation (indirect reciprocity).
Encyclopedia of Animal Behavior (2010), vol. 2, pp. 483-488
However, these conditions demand consideration of the
requisite psychological mechanisms, including recall of
prior outcomes to evaluate reputation, quantification
and tracking of the payoff matrices to evaluate fair dis-
tributions, and assessment of whether the resources were
distributed intentionally or as a byproduct of otherwise
selfish behavior.
While reciprocal altruism and indirect reciprocity can
explain the evolution of cooperation in small social
groups where individuals know each other, these mechan-
isms cannot account for the fact that modern humans
often live in large groups of unrelated others, where
reputation tracking is not possible, and where repeated
and stable relationships are unreliable, thus making reci-
procity difficult or impossible. In these situations, cooper-
ation nonetheless evolves, demanding a different kind of
account that can accommodate the fact that the optimal
strategy is to defect and free ride on the contributions of
others. Here, Boyd and Richerson pointed out that it
appears that punishment evolved to crack down on the
defector problem, and bring about stable cooperation. On
an ultimate level, focused on evolutionary consequences,
punishment is a behavior that reduces the fitness of a
recipient at a temporary cost, but ultimate benefit to an
actor (see the reading by Clutton-Brock and Parker for
more on this topic). However, from a proximate perspec-
tive, punishment requires specific psychological mechan-
isms including the recruitment of motivating emotions
(e.g., Trivers’ moral outrage), the ability to determine when
a norm has been violated, the assessment of just deserts,
and a mechanism to distinguish whether the outcome
(e.g., failure to cooperate) was intended or accidental.
Recent work on the evolution of fairness provides a
good example of how proximate and ultimate concerns
have come together. Specifically, using a combination of
well-defined bargaining games from behavioral econom-
ics, together with rich psychological analysis and cross-
cultural data, we can see that humans evolved a distinctive
sense of fairness, one shared by all members of our spe-
cies, but open to cross-cultural variation, and constrained
by sensitivity to inequities and the ability to punish those
who violate norms of distributive justice. Consider, as an
example, the well studied, one-shot, anonymous Ultima-
tum Game. An experimenter first informs two individuals,
a donor and a recipient, about the three rules of the game.
Rule 1: the bank allocates a sum of money to the donor
who has the option of allocating some proportion of
this sum to the recipient. Rule 2: whatever amount the
donor gives to the recipient, the recipient keeps, and
the donor keeps the remainder. Rule 3: if the recipient
chooses to reject the donor’s offer, then neither donor nor
recipient keeps any money. According to the rational
economic model, the recipient should accept any offer
from the donor as some money is surely better than no
money. However, results from this game show that people
 Author's personal copy
across cultures consistently reject some offers, licensing
the conclusion that humans approach this problem with a
sense of what constitutes an unfair offer. Further, when
subjects learn that the donor is a computer or random
number generator, they are more likely to accept unfair
offers. These results show that humans not only attend
to the distribution of resources (i.e., outcomes), but also to
the means by which resources are distributed. Returning
to the ultimatum game, there is a fundamental (moral)
difference between a human donor offering 1 out of 20
possible dollars to a recipient, and a computer program
that uses a random number generator to offer $1 out of
$20. Computers are not intentional agents, and thus, don’t
enter the moral domain. Regardless of how upset we are at
the computer’s offering, we can’t hold them responsible. On
the other hand, we do hold other humans responsible for
their actions. Pushing the point further, if we forced a
human to roll a die to pick the donation, and the die
landed with a 1 facing up for $1, we would also not hold
the human responsible for this outcome; this game func-
tionally strips the human agent of his intentional control –
of his free will. What is critical, then, is a combination of
both the actual outcome and whether the agent was
responsible for this decision.
Thus, a sense of fairness, together with an ability to
discriminate between intentional and accidental actions,
is critical to cooperation, and more generally, to our moral
judgments about others’ actions. Given the importance of
these abilities, they raise the crucial question of whether
these traits are unique to humans or whether they also
play a role in governing behavior in non-human animals
(hereafter animals).
Cooperation, Fairness and Action
Perception in Animals
Cooperation in animals is taxonomically widespread, with
numerous examples of individuals engaging in costly
altruistic behavior for the benefit of others. In the early
literature, most of the examples were consistent with the
theory of kin selection and with optimization of inclusive
fitness. That is, most altruistic acts of cooperation evolved
to benefit close genetic relatives. Following on the heels of
Trivers’ conceptual work on reciprocity, however, several
cases of reciprocal altruism emerged in the literature.
These cases, as well as several more recent studies have,
for the most part, been dismissed, either because of a
failure to replicate, the weakness of the effects, alternative
explanations (e.g., byproduct mutualism), or the highly
artificial conditions under which the evidence has been
obtained. At best, we argue, reciprocity is an uncommon
form of social interaction among animals. We further
argue, however, that consideration of both proximate
and ultimate factors makes this conclusion unsurprising.
Encyclopedia of Animal Behavior (2010), vol. 2, pp. 483-488
Morality and Evolution 485
Specifically, there are at least two reasons why reciprocity
might be rare among animals. First, the demographics of
most animal populations may provide a sufficiently high
density of kin to eliminate the pressure for nonkin based
relationships. Despite these issues, we suggest that recent
work on cooperation, including reciprocity, has provided
new insights into some of the psychological mechanisms
that are shared among human and non-human animals,
and leads to one of our primary conclusions: though we
share with other animals some of the core building blocks
of morality, only one species – our species – has combined
these core elements into a truly moral system that not
only considers how we distinguish moral rights from
wrongs, but also what ought to be the foundation for
such decisions.
A Sense of Fairness in Animals?
Recent comparative work on primates and dogs has
explored the problem of inequity aversion as an important
component of the more general sense of fairness. One of
the earliest treatments of this problem was Brosnan and de
Waal’s study of brown capuchin monkeys (Cebus apella).
In this experiment, subjects that had been trained to trade
tokens for food rewards watched a conspecific acquire and
eat a high-value food item and then were given the
opportunity to acquire and eat a lower-value food item.
Subjects consistently refused to trade the token for the
lower-value food, and this result was interpreted as evi-
dence for inequity aversion and a sense of fairness. This
experiment was heavily criticized because the authors
could not rule out the effect of frustration as the driving
force behind rejections. That is to say, subjects may have
rejected unfair offers not because the other individual was
getting a better deal, but because they were frustrated at
not being able to obtain the higher-value food item that
was right in front of them. Though subsequent experi-
ments confirmed the validity of these critiques, Brosnan,
de Waal, and their colleagues, have since replicated the
original findings with relevant controls, and found that
their results cannot be explained by frustration, or further
extended to parallel findings with chimpanzees (Pan tro-
glodytes). Adding to the comparative scope of this work,
a recent experiment by Range and colleagues shows
that domestic dogs are sensitive to inequities in reward
distribution. In this experiment, subjects were given a
command to perform an action (paw shake) in a social
situation where one individual received a food reward for
its performance while the other did not. Subjects that did
not receive the reward were more reluctant to perform the
action and displayed more stress behavior in the social
condition compared to an asocial control.
In sum, although there is still much controversy sur-
rounding the results on inequity aversion in animals, mini-
mally, it appears that animals are sensitive to the distribution
 Author's personal copy
486 Morality and Evolution
of rewards, in both social and nonsocial contexts, responding
negatively when an outcome appears unfair.
Another approach to studying fairness in animals
comes from a series of experiments investigating prosocial
behavior, specifically, the tendency to help another in a
situation where there are no personal gains, and little or
no personal cost. In these experiments, subjects are given
the option of acquiring a reward for themselves or for
themselves as well as for another individual. The impor-
tant difference between prosociality tasks and inequity
aversion tasks is that a preference for equity is costless
in the former (actors receive a payoff either way) and
costly in the latter (actors receive nothing if they reject
an unfair offer). Studies of common marmosets (Callithrix
jacchus) and brown capuchin monkeys have shown that
subjects consistently choose the prosocial option. Studies
of chimpanzees, on the other hand, have shown that
individuals are indifferent to the welfare of conspecifics
and will choose indiscriminately between the two options.
Interestingly, when chimpanzees are tested in a spontane-
ous altruism task, where subjects are given the opportu-
nity to help another individual in the absence of a food
reward, they do exhibit prosocial behavior. At present, it is
not clear whether the difference in prosociality between
these studies is due to the nature of the reward (i.e., food
versus nonfood), or to specific details of the task demands.
Together, studies of inequity aversion and prosociality
suggest that the ability to both detect and react to unfair
outcomes, together with a preference for fair resource
distributions, are not uniquely human traits. However,
these experiments focus exclusively on outcomes and do
not explore the means by which they are achieved. In the
next section, we discuss experiments that tap into the
psychological mechanisms involved in discriminating
between intentional versus accidental actions.
Going Beyond Outcomes to Intentions
and Goals
In this section we explore two questions: (1) Can animals
draw inferences about an individual’s intentions and goals,
and (2) if so, does this capacity influence social behavior?
Recent studies of rhesus monkeys on Cayo Santiago
have explored their ability to use subtle details of an action
sequence to draw inferences about the actor’s goal. In the
basic design, Wood and colleagues presented two potential
food sources (overturned coconut shells) to a subject, acts
on one, and then walks away, allowing the subject to selec-
tively approach. Although coconuts are native to the island
on which these animals live, rhesus cannot open the hard
outer shells themselves, and therefore, only obtain the
desired inner fruit when the coconuts open on their own
or have been opened and discarded by a human. It, thus,
logically follows that if subjects perceive the experimenter’s
action as goal-directed and potentially communicative,
Encyclopedia of Animal Behavior (2010), vol. 2, pp. 483-488
then they should selectively approach the coconut con-
tacted, as this maximizes the odds of obtaining food.
Results revealed that when the experimenter grasped
the coconut with his hand, foot, or a precision grip involving
the pointer finger and thumb, rhesus selectively approached
this coconut over the other; in contrast, they approa-
ched the two coconuts at chance levels when the experi-
menter flopped the back of his hand on the coconut
[accidental], touched or grasped the coconut with a tool,
or grasped the coconut with his hand for balance while
standing up. These results rule out low-level association
accounts; many of these individuals have experience see-
ing humans perform goal-directed actions with tools, and
yet they did not perceive tool-related actions as goal-
directed, and none of these individuals have experience
seeing humans perform grasping actions with their feet,
and yet they did perceive foot-related actions as goal-
directed. Further, these data show that when assessing
the meaning of actions, rhesus are highly sensitive to the
means used to achieve a goal – for example, perceiving a
hand grasp action as goal-directed but a hand flop action
as accidental, despite the fact that the experimenter’s body
position, eye gaze, and duration of contact with the coco-
nut were identical across the two conditions. These stud-
ies, together with research on other species, suggest that
non-human animals infer the meaning of an action by
evaluating the actor’s goals in relation to the environmen-
tal constraints on achieving such goals.
Given that some animal species are able to draw infer-
ences about others’ intentions and goals, we can ask
whether this ability influences their social interactions.
That is, are animals completely outcome-oriented or do
they attend to the means by which outcomes are achieved?
One of the first studies to explore this problem was Call
and colleagues’ experimental study of captive chimpanzees.
In this study, a human experimenter faced a chimpanzee,
seated on the opposite side of a Plexiglas partition. In each
of several conditions, varying the nature of the experimen-
ter’s action, a grape was presented near the opening of the
partition; the opening was large enough for the chimpanzee
to reach and grab the grape. In one condition, the experi-
menter brought the grape within grasping distance, but
then rapidly retracted it as soon as the subject reached.
This action was defined as teasing. In a second, and highly
parallel condition, the experimenter brought the grape
forward, but then dropped it as soon as the subject reached.
This action was defined as clumsiness. Chimpanzees
showed much greater signs of frustration in the teasing
than clumsy conditions, leaving the test chamber earlier,
and acting aggressively toward the experimenter (i.e., bang-
ing on the Plexiglas partition). Thus, as Call and colleagues
suggest, chimpanzees appear to make a distinction between
unwilling (teasing) and unable (clumsy), thereby showing
sensitivity to more than the mere outcome of an event, as in
both of these cases, the subject failed to obtain the grape.
 Author's personal copy
In the aforementioned study of reciprocity in tamarins,
Hauser and colleagues showed that individuals were more
likely to cooperate in situations in which a conspecfic’s
actions were truly altruistic, than when the same amount
of food (outcome) was delivered as an accidental bypro-
duct of an otherwise selfishly motivated action. Specifi-
cally, a game was set up such that the individual playing
the actor-1 position was offered an opportunity to pull a
tool, delivering one piece of food to self and three pieces
to an unrelated partner. For the partner, or actor-2, pull-
ing the tool resulted in no food for self, but two pieces for
the partner (actor-1). A session was defined as 12 trials
each for actor-1 and -2, alternating turns. If both actor-1
and -2 pulled on their respective turns, they would maxi-
mize the overall returns, with three pieces each, after an
alternating round. This is what would be expected if
actor-2 perceives actor-1’s pull as altruistic, that is, moti-
vated by the goal of giving food. In contrast, if actor-2 per-
ceives actor-1’s pull as selfish, with the three pieces
obtained as a byproduct, then actor-2 should not pull.
Results showed that actor-2 rarely pulled. This condition,
combined with another showing that individuals will
altruistically give food (i.e., paralleling the actor-2 posi-
tion) when a partner altruistically reciprocates, reinforces
the conclusion that tamarins attend to both the outcomes
and the means by which they are obtained.
A final experiment adds to this literature by showing
not only sensitivity to the means by which outcomes arise,
but the agent responsible for such outcomes. Jensen and
colleagues first presented one chimpanzee, A, with an
apparatus involving a sliding tray full of food. Subject
B was then introduced into an adjacent enclosure that
contained a rope. By pulling on the rope, B moved the
sliding tray away from A, thereby stealing A’s food. How-
ever, subject A’s enclosure also contained a rope that, if
pulled, would collapse the sliding tray, thereby taking the
food away from B. In conditions where B pulled the tray
away from A, A frequently became agitated and collapsed
the sliding tray. However, in a similar condition where,
instead of B pulling the tray, the experimenter pushed the
tray away from A to B, A rarely collapsed the tray. This
result is interesting because it shows that chimpanzees not
only respond to unfair outcomes, but that they distinguish
between human and chimpanzee agents; a parallel set of
findings was presented by Hauser and colleagues in their
reciprocity study, showing that cooperation increased in
the face of a unilateral tamarin cooperator, but not a game
in which the payoffs remained the same, but a human
cooperator delivered the rewards for another tamarin.
Conclusion
Research on inequity aversion, prosociality and action
perception in non-human animals is still in its nascent
Encyclopedia of Animal Behavior (2010), vol. 2, pp. 483-488
Morality and Evolution 487
stages, and there are currently several noticeable gaps in
our understanding of these phenomena. First, while there
is an emerging body of evidence suggesting that these
capacities are present in some species of non-human
primates and domestic dogs, little is known about its
taxonomic distribution or the evolutionary pressures
that may select for this capacity in different species. For
example, given the increasing evidence that some food-
caching jays are sensitive to where others are looking,
what others have seen, and how such information guides
cooperation and cheating, it would not be surprising to
find that at least these birds, and possibly other animals,
are sensitive to inequities, and to the distinction between
means and outcomes. Second, it is unclear whether the
capacities discussed here are specific to the social domain
or are important in other domains as well. For example, do
animals appreciate that some properties of artifacts such
as tools are intentionally designed whereas others are
simple byproducts of physical constraints or accidents?
Third, the majority of studies that have investigated
these capacities in animals have focused on captive indi-
viduals. Similar studies of wild populations will be crucial
to understanding whether these abilities are expressed in
nature or are only elicited under controlled laboratory
conditions that often set up situations that would never
arise in the wild. For example, though capuchin monkeys
can work with a token economy and detect inequities, and
though they can solve a joint action task, these situations
never arise in the wild. It is, thus, essential to distinguish
between the capacity to solve various cooperative tasks
and the social and ecological pressures that might demand
that such abilities be used to cooperate.
Despite these limitations, we believe the work on the
foundations of morality reviewed here (and elsewhere;
Hauser’s 2006 book, Moral Minds: How Nature Designed
Our Universal Sense of Right and Wrong, for a more extensive
discussion) lead to at least two conclusions. First, we share
with other animals several core psychological capacities
that were most likely necessary for the evolution of our
moral sense. Like other animals, a sense of justice as
fairness is premised on an ability to take the perspective
of another individual, show concern for others, detect
inequities, and inhibit the temptation to feed self-interest.
Though humans certainly show highly elaborated forms
of these abilities, there are significant precursors in other
species. Second, there are two ways in which human moral
behavior, and the psychology that supports it, are unique.
Animals show virtually no evidence of reciprocity and
large-scale cooperation, and as far as we can tell, never
engage in the problem of considering not only what is the
moral state of play, but what could or should be – our
sense of ought. Though a brief essay like this is not the
place to develop these issues, we end with a speculative
consideration. What allows humans to uniquely engage
with the thought of the ought is our ability to combine
 Author's personal copy
488 Morality and Evolution
different modular representations into new representa-
tions. Whereas other animals have evolved highly adap-
tive, modular, and informationally encapsulated domains
of thought, targeted at single problems, humans evolved
the capacity to create interfaces between these domains to
create entirely new systems of thought. Thus, we alone
can consider how we typically distribute resources, often
based on matters of effort and need, step back from such
norms, and consider a more enlightened perspective that
not only considers matters of fairness but also individual
welfare. And we can do this because we can prospectively
evaluate the future, sideline current needs and tempta-
tions, and realize that progress is made by dissent as
opposed to consent. What fuels the ought is the realiza-
tion that we can, and often should, entertain a different
moral landscape.
See also: Cooperation and Sociality; Empathetic Behav-
ior; Mental Time Travel: Can Animals Recall the Past and
Plan for the Future?; Punishment; Social Cognition and
Theory of Mind.
Further Reading
Boyd R and Richerson PJ (1992) Punishment allows the evolution of
cooperation (or anything else) in sizeable groups. Ethology and
Sociobiology 13: 171–195.
Encyclopedia of Animal Behavior (2010), vol. 2, pp. 483-488
Brosnan SF and de Waal FBM (2003) Monkeys reject unequal pay.
Nature 425: 297–299.
Call J, Hare B, Carpenter M, and Tomasello M (2004) ‘Unwilling’ versus
‘unable’: Chimpanzees’ understanding of human intentional action.
Developmental Science 7: 488–498.
Clutton-Brock TH and Parker GA (1995) Punishment in animal societies.
Nature 373: 209–216.
Dugatkin LA (1997) Cooperation Among Animals. Oxford: Oxford
University Press.
Hamilton WD (1964) The genetical evolution of social behavior, I. Journal
of Theoretical Biology 7: 1–16.
Hauser MD (2006) Moral Minds: How Nature Designed Our Universal
Sense of Right and Wrong. New York: Ecco.
Hauser MD, Chen MK, Chen F, and Chuang E (2003) Give unto others:
Genetically unrelated cotton-top tamarin monkeys preferentially give
food to those who altruistically give food back. Proceedings of the
Royal Society, London, B 270: 2363–2370.
Jensen K, Call J, and Tomasello M (2007) Chimpanzees are vengeful
but not spiteful. Proceedings of the National Academy of Sciences
104: 13046–13050.
Range F, Horn L, Viranyi ZS, and Huber L (2009) Absence of reward
induced aversion to inequity in dogs. Proceedings of the National
Academy of Sciences 106: 340–345.
Rawls J (1971) A Theory of Justice. Cambridge: Harvard University
Press.
Stevens JR and Hauser MD (2004) Why be nice? Psychological
constraints on the evolution of cooperation. Trends in Cognitive
Sciences 8: 60–65.
Trivers RL (1971) The evolution of reciprocal altruism. The Quarterly
Review of Biology 46: 35–57.
Williams GC (1966) Adaptation and Natural Selection. Princeton, NJ:
Princeton University Press.
Wood JN, Glynn DD, Phillips BC, and Hauser MD (2007) The perception
of rational, goal-directed action in non-human primates. Science
317: 1402–1405.