Evolutionary Anthropology 22:251–258 (2013)

ARTICLE

Cooperation and Conflict Between Women

in the Family
RUTH MACE

Here I review recent research on reproductive conflict between females in organization that guide the transmis-
families and how it influences their reproductive behaviour. Kin selection can sion and inheritance of wealth and
favor cooperation between parent and offspring, siblings, or unrelated co- power, as well as marriage and
residents who share interests in other family members such as grand-offspring. residence rules. These have been the
However, these are also the individuals most likely to be sharing resources, and traditional preoccupations of an-
so conflict can also emerge. While substantial interest has arisen in evolutionary thropologists since the discipline
anthropology, especially over the last two decades, in the possibility of coopera- emerged.
tive breeding in humans, less attention has been paid to reproductive conflict In another branch of our field,
among female kin. Communal breeding in animals is generally understood as human parenting and alloparenting
emerging from competition over the resources needed to breed. Competition for has been attracting interest among
household resources is a problem that also faces human families. Models sug- evolutionary anthropologists for
gest that in some circumstances, inclusive fitness can be maximized by sharing some time.4,5 Over the last few deca-
reproduction rather than harming relatives by fighting with them, even if the des, it has become apparent that the
shares that emerge are not equal. Thus, competition and cooperation turn out life histories of human females have
to be strongly related to each other. Reproductive competition within and features that are rather different
between families may have underpinned the biological evolution of fertility pat- from those of other apes, including
terns (such as menopause) and the cultural evolution of marriage, residence, long childhoods, rapid reproductive
and inheritance norms (such as late male marriage or primogeniture), which rates after puberty, and then female
can enhance cooperation and minimize the observed incidence of such menopause, followed by a long post-
conflicts. reproductive life. Explanations for
these patterns are varied, but most
are related to the cooperative and
“A time to love and a time to competitive relationships between
hate.” Ecclesiastes 3:1-8 kin.6–8 As the evolution of human
kinship and human life histories are
my two favorite subjects, I am
A central relationship in virtually thrilled that these two large branches
all human social systems is that of anthropology, which often are
Ruth Mace is Professor of Evolutionary
Anthropology at UCL. Her research has between husband and wife. But there considered somewhat separately, are
spanned a variety of areas relevant to
are relatively few social systems in now being more formally linked in
evolution of the human family, life history,
and kinship systems, as well as the evo- which a pair-bonded couple live recent literature. The issue of disper-
lution of cooperation both inside and independently of other kin.1 Hunter- sal may be key to both, for reasons
outside the family. Much of her work has that I will outline.
been done in African populations, but gatherer groups appear to be rela-
she has also worked on data from the tively fluid associations of groups of Sarah Hrdy has worked on both
United Kingdom and is now also working
nuclear families that can be based cooperation and conflict, which makes
on populations in China in collaboration
with the Chinese Academy of Sciences on matrilateral or patrilateral kin- this special issue an ideal format to
in Beijing. In 2008, she was elected a ship, friendship, or convenience.2,3 review how these two topics are related
Fellow of the British Academy. to each other. Her interest in sexual
After the origin of agriculture, our
economies began to rely on farmland conflict dates back to her pioneering
Key words: human behavioural ecology; matriliny; and other heritable resources. That work on infanticide in langurs.9 This
reproductive skew; cultural norms; kinship
made it more difficult for families to work was followed by a sequence of
move freely. More formal rules of path-breaking books on the evolution of
C 2013 Wiley Periodicals, Inc.
V ownership and transmission of human females. In Mother Nature,
DOI: 10.1002/evan.21374 wealth were generated, giving rise to she emphasized the effects of
Published online in Wiley Online Library
(wileyonlinelibrary.com). cultural norms of kinship and social sexual conflict on the evolution of
252 Mace
motherhood in humans.4 I shall not
delve into infanticide or sexual conflict
here; as fascinating as these topics are,
they have been quite widely discussed
elsewhere. Later, Hrdy’s Mothers and
Others helped galvanize interest in
human cooperative breeding.10 What
are humans, social cooperators or
fierce competitors? We know, of
course, that they are both, but what I
shall outline is how competition itself
can be the driver of cooperation. I want
to focus here on the slightly less-
trodden path of cooperation and con-
flict between adult females, not so
much over mates,11 but over the
resources needed to breed.
Zoologists interested in cooperative
breeding in animals have used inclu-
sive fitness theory to understand why
individuals might stay in groups and
who might win and lose in the compe-
tition for local resources. They also
have argued that similar approaches
apply to human families.12 While
cooperative breeding usually involves
winners and losers in terms of direct
fitness, it may involve a win-win sit-
uation in terms of inclusive fitness if
individuals can ‘agree’ on a strategy
that, given the constraints, maximizes
inclusive fitness for both parties. This
theoretical framework has been use-
ful in explaining reproductive sharing
in communally breeding mammals,
birds, and insects; these models can
predict when dominants may prefer
to allow kin to breed in the group, at
least a bit, rather than have them
leave, and when subordinates might
choose to stay. Hence, the models
help predict the extent of reproduc-
tive skew (when some individuals’
direct fitness is higher than others).13
Here, I describe developments in two
of my own areas of interest that are
helped by considering cooperation
between women in human families in
such terms: human menopause and
human kinship organization. I will
argue that both can be understood as
biological or cultural adaptations to
reduce conflict between female kin.
FAMILIES AS COOPERATIVE
UNITS
Families are nuclear families if
individuals disperse as soon as they
reach sexual maturity, when they
usually marry, so that only couples
and their children co-reside. Larger
groups of kin form when some indi-
viduals do not disperse. This may
occur if there are benefits of staying
or costs of leaving. The prospect of
inheriting a good territory, such as a
farm, would be an example of the
benefits of staying. There may be
high costs of leaving if habitats are
saturated and new territories are
hard to come by, and this may lead
to delays in dispersal. Important cor-
relates of cooperative breeding in
animals include long-life span and
saturated habitats that make disper-
What are humans, social
cooperators or fierce
competitors? We know,
of course, that they are
both, but what I shall
outline is how competi-
tion itself can be the
driver of cooperation.
sal difficult.14,15 It is likely that coop-
erative living and breeding emerge in
human groups for similar reasons.
Because humans, in contrast to
the other great apes, are pair-bonded
and share meat, early socio-
ecological models of human families
placed strong emphasis on “man the
hunter,” who helped to support
females and their offspring. While in
some cases males seem to support a
great deal of a woman’s caloric
intake,16 in others there are sugges-
tions that males are not as helpful to
women and children’s nutrition as
might be expected.17 In some cases,
the presence of a male seems to
increase rather than decrease wom-
en’s workloads.18 An alternative
model, in which female kin help
each other to support the rapid
reproductive rate of adult females,
has become very popular, if not pre-
dominant, in recent years.6 It is clear
that food supplied by others besides
ARTICLE
the mother underpins short human
interbirth intervals, so that humans
are now often referred to as coopera-
tive breeders.10,19 Grandmothers are
sometimes thought to play an essen-
tial role in provisioning younger
women and their offspring.17,19
There is considerable debate about
the extent to which human meno-
pause can be understood as an adap-
tation to grandmothering or whether
it emerged due to some kind of phys-
iological by-product or constraint.
Can infertile grandmothers be con-
sidered on a par with other non-
breeding helpers in a communal or
cooperatively breeding species?
Implicit in the ‘grandmother hypoth-
esis’ for the evolution of menopause
is the notion that it would not be
possible for a grandmother to raise
her own offspring at the same time
as helping her daughter (or son) to
reproduce; thus, she has to choose
between mothering and grandmo-
thering. There is now good evidence
that having a grandmother is associ-
ated with enhanced survival and fit-
ness benefits to grandchildren,20–22
but whether it is enough to counter-
balance the loss of late life reproduc-
tion is more controversial.
The failure of some to be con-
vinced by adaptive hypotheses led
many to place more emphasis on
menopause as a physiological con-
straint. It was argued that if the
length of the fertile period is consid-
ered to be constrained to that of a
chimpanzee, then only the extension
of female life span beyond 50 years
of age is what has been selected for.
It is relatively clear that life spans
will extend if older females can be
useful as carers for their kin.23 How-
ever, a model based on such a con-
straint is not particularly satisfying
to those of us that like to believe that
evolution is strong and that life-
history variables like the timing of
reproduction should be relatively
easy to select for. It is also extremely
hard, if not impossible, to test a
model based on an unspecified con-
straint, especially as elephants do
seem to be able to breed after the
age of 50.24 If rapidly increasing age-
specific maternal mortality is
included, along with mother and
grandmother effects on both
ARTICLE
grandchild survival and daughters’
fertility, then models predict little or
no fitness advantage to reproducing
after about the age of 50 years.25,26
But accelerating maternal mortality
could itself be considered a ‘physio-
logical constraint’, possibly emerging
after the evolution of menopause or
for the same reasons that menopause
evolved, and not the cause of meno-
pause. Still others argue that the
female life span is a ‘spandrel’,
selected for by selection on male life
span because males do have oppor-
tunities to reproduce in old age.27
Such models are made less persua-
sive by the empirical observation
that females tend to outlive males in
most human populations.
All these models are compatible
with the observation that post-
reproductive grandmothers are gen-
erally helpful toward their children,
or at least to their daughters and
grandchildren.20 But whether that
help drives the evolution of meno-
pause or is just occurring to make
the best of a bad job is still disputed.
Grannies, and sometimes granddads,
had better make themselves useful if
they expect to be given much food or
protection from the other members
of their community. There are vari-
ous anthropological examples of ger-
onticide among hunter-gatherers.28
Also, relegation to rather precarious
and poverty-stricken life styles is a
severe risk for old ladies29 or widows
in any culture.
CONFLICT BETWEEN KIN IN THE
EVOLUTION OF MENOPAUSE
The many empirical observations
of grandmother effects indicate that
maternal grandmothers are almost
always beneficial. In contrast, pater-
nal grandmothers often are not par-
ticularly helpful, and are sometimes
even harmful to their grandchildren’s
survival,20,30,31 even if they do pro-
mote their daughter-in-laws’ fertil-
ity.32 One possible reason for this is
paternity uncertainty; another is that
the costs of maternal mortality are
much higher for matrilineal kin,
whose daughter is irreplaceable,
than to the patriline, whose
daughter-in-law could at least par-
tially be replaced by remarriage if
Cooperation and Conflict Between Women in the Family 253
death occurred.19 Still further, it is
possible that genetic conflict gener-
ated by X-chromosome inheritance
causes grandmothers to discriminate
against their sons’ sons.33 All these
factors suggest that matrilineal kin-
ship organization may lead to more
grandparental solicitude.
Unrelated females in the patrilin-
eal Dogon of Mali seems to be in
out-and-out conflict both across and
within generations.29 If the evidence
of cooperative breeding seems
mainly to concern matrilineal kin,
this raises the question of whether
ancestral humans had enough con-
There is now good
evidence that having a
grandmother is
associated with
enhanced survival and
fitness benefits to
grandchildren,20–22 but
whether it is enough to
counterbalance the loss
of late life reproduction
is more controversial.
tact with female kin for cooperative
breeding to have generated meno-
pause. Residence patterns become
key. There is not much evidence that
ancestral hunter-gatherers were
likely to have lived predominantly
matrilocally. But then the term make
little sense in many more mobile
groups with variable residence pat-
terns.34 Moreover, there are exam-
ples of grandmothers providing
significant help to their daughters
even if they are not permanently co-
resident with them.18,35,36
Cant and Johnstone developed a
model to evaluate how dispersal pat-
terns influence the evolution of help-
ful and harmful behavior toward
female kin and the effect of this on
the evolution of menopause.37,38
They assume that there is likely to
be reproductive conflict between co-
residents over resources needed for
reproduction. Cant and Johnstone
draw on tug-of-war models of repro-
ductive conflict between kin, in
which players have partial control
over the outcome.13 When females
customarily disperse, as is probably
the case in most human groups, the
average relatedness of a female to
others in her group varies over time.
This is because daughters disperse
and are replaced by unrelated
females who migrate into the group
to marry the sons of the patriline;
grandmothers and daughters-in-law
are then asymmetrically related to
the children that result. The paternal
grandmother gains inclusive fitness
from helping her younger daughter-
in-law reproduce because by doing
so she is helping her son to repro-
duce. The reverse is not true. The
younger woman is related to no one
other than her own offspring, so she
gains no inclusive fitness from help-
ing her mother-in-law (or anyone
else in the household) to reproduce.
This asymmetry means that the older
woman has more to lose from a fight
than her daughter-in-law does, and
is destined to lose any reproductive
conflict. After a certain age, the evo-
lutionarily stable strategy (ESS) is
for her to forego reproduction and
help raise her grandchildren, from
whom she gains at least some inclu-
sive fitness. This model therefore
suggests that menopause is an adap-
tation to avoid conflict with kin by
ceasing reproduction and eliminat-
ing reproductive overlap.37 Here,
competition is, almost literally, the
mother of cooperation. If this ‘repro-
ductive conflict’ version of the grand-
mother hypothesis is correct, the fact
that humans are more likely to have
been characterized by female disper-
sal than by male dispersal is a bonus
rather than a hindrance to adaptive
explanations for menopause.
Older females also increase in
average relatedness to the group
when neither males nor females dis-
perse, but mate outside the group;
this is the case for many cetaceans,
which also show menopause.38 In
the case of whales, which mate out-
side the group, there is some
254 Mace
Box 1. The Case of Communal Living in Matrilineal Group Households in China
A few matrilineal peasant soci-
eties resemble the social systems of
cetaceans, in which mating occurs
outside the residential group. In
these rare duolocal systems, neither
sex disperses. Generations of broth-
ers and sisters and their matrilineal
offspring co-reside throughout life.
One of these societies is the Mosuo
of southwestern China, also known
as the Na,40 who live around the
shores of Lugu Lake.
The social system is built into
their homes, which are built around
a courtyard. The most important
room in the house belongs to the
grandmother, the most important
person in the household. Her room
contains the central fireplace, where
visitors are received; this room is
also the bedroom for the grand-
mother and her matrilineal grand-
children. (The offspring of resident
males reside elsewhere with their
mothers.) Adult daughters have
their own smaller rooms, usually
around the upper level of the court-
yard, where their male partners visit
them at night in what are some-
times described as ‘walking’ or ‘visit-
ing’ marriages. Unmarried teenage
boys usually share a room. Some-
ARTICLE
times older men build rooms for
religious worship.
Farming and domestic work is all
communal, with household mem-
bers sharing food, child-care
responsibilities, and all household
resources, and the farm is inherited
communally by all the matrilineal
family members. Women do most
of the domestic and agricultural
work.40 There are historical
accounts of mothers having chil-
dren from several different
fathers,55,56 but Mosuo currently
report that they are mostly
monogamous.40
ARTICLE
Income from tourism is changing
both the residence and marriage pat-
terns in favor of more nuclear fami-
lies.53 Both males and females that
form neolocal households start to
breed earlier than those that remain
within the communal duolocal
households, indicating the communal
living is associated with reproductive
constraints.40 We have argued that it
indication that older females are par-
ticularly helpful to sons, who, since
their offspring are elsewhere, are not
in competition with other family
members for breeding opportunities.
A recent study showed that male
orcas tended to experience high mor-
tality when their mother died,39 so
maybe the rest of the group tolerate
their brother until mum dies, then
gang up and start taking bites out of
him. Zoologists have learned to
expect nothing less from ‘coopera-
tive’ breeders. Often the more ‘coop-
erative’ the breeding system, the
more extreme the reproductive skew
and the more potential for violence
to enforce the peace or establish who
inherits a vacant breeding slot, with
social insects probably providing the
most extreme scenarios. There are
probably no human social systems
exactly like that of orcas, but some
matrilineal ‘duolocal’ groups, in
which brothers and sisters co-reside
throughout life, provide a rare case
of males breeding outside the resi-
dential group40 (Box 1).
RESOLVING REPRODUCTIVE
CONFLICT THROUGH KINSHIP
NORMS
In humans, reproductive conflict
among kin can be avoided not just
by menopause, but also by cultural
rules.36 I suspect that cultural tradi-
tions such as marriage and kinship
norms may function to reduce repro-
ductive conflict. In patrilocal resi-
dence systems, which are often also
polygynous, brothers, and thus their
wives, are in competition for resour-
ces owned by or inherited from their
father, and this may significantly
affect when sons can marry and
Cooperation and Conflict Between Women in the Family 255
is the communal nature of the house-
holds that favors a male’s investment
in his natal household rather than
that of his wife. This is because
investment in his wife’s communal
household is diluted by sharing
resources with his wife’s kin, to
whom he is not related.40
Historically, patrilineal group
households were quite common in
begin to reproduce. 41–43 In such sys-
tems, a delay in the onset of repro-
duction means that reproductive
overlap between a man’s mother and
his wife is almost zero.
My data on over 500 women from
communities in rural Gambia, which
habitually show patrilocal residence,
illustrate the very different effects
that reproductive overlap has on the
reproductive success of daughters
and sons. Only very fertile mothers,
who gave birth young and continued
to reproduce late in life, had any
reproductive overlap with their
daughters. When this did occur, the
daughter did not suffer much cost,
presumably because female dispersal
means the daughters’ children would
be raised in another household. The
mother did slow her reproduction
upon the arrival of a matrilineal
grandchild, consistent with the
notion that she is genuinely helping
her daughter.36 Mother/son repro-
ductive overlap was very rare. There
were very few cases in which moth-
ers became paternal grandmothers
under the age of 50 years, and thus
showed the potential for reproduc-
tive overlap with daughters-in-law.36
Although we did not have enough
cases of mother/son reproductive
overlap in the Gambia to estimate its
effects, relevant evidence comes
from Finland. In historical Finns,
only 36 out of 556 women in the
sample showed such overlap, but
when this did occur it had a pro-
nounced negative effect.44 Again,
among the Finns, mother/daughter
reproductive overlap was more com-
mon and not nearly as costly, if
costly at all.44
Cultural norms of parental control
over the transmission of resources
China, but they tended to dissolve
with the death of the grandfather,
since multiple couples within one
house led to too much conflict
once the his control was no longer
there.1 Matrilineal Mosuo house-
holds are more stable.
and marriage practices, combined
with female dispersal, almost com-
pletely eliminate conflict arising from
multiple females breeding on the
same farm. Whether such costs would
apply in a hunter-gatherer context,
where there usually is open access to
resources and residence patterns are
more flexible, is unknown. Conse-
quently, it is still an open question
whether such conflict led to the evolu-
tion of menopause.
Nevertheless, cultural evolution has
generated a wide range of human
social systems in far less time than is
required for the biological evolution
of a trait like menopause. The avoid-
ance of reproductive conflict is likely
to have had a profound effect on the
cultural evolution of marriage and
kinship systems. Those violating cul-
tural norms generally do less well
than those who follow the locally
accepted practice,45 which makes it
complicated to work out whether
costs helped generate the norms or
simply emerged as a consequence of
punishment for violating a norm that
arose for other reasons. Most evolu-
tionary anthropology suffers from
such complications; although this is
annoying, it also is part of what
makes it so interesting.
Patrilineal kinship is usually asso-
ciated with a subsistence strategy
that includes resources that can be
monopolized by males and also are
often used to encourage women to
marry polygynously.46 Hence, men
compete, even against their own
brothers, for resources and marriage
partners when fathers control
resources. The parents of daughters
can exploit this to demand bride
price. Pastoralist systems typically
show all these characteristics:
256 Mace
Livestock are like walking money
and not nearly as hard work to
maintain as, for example, extensive
crop farming systems. Females in
patrilineal systems forego the bene-
fits of proximity to their female kin
for the advantages of access to male-
owned resources. The price could be
severe female-female conflict, as
described for the aforementioned
Dogon in Mali, who do not seem to
be well-described as ‘cooperative
breeders.’29 The evolution of this kin-
ship system seems to represent a
case of males, who benefit from
polygyny at the expense of their
wives’ fitness, winning the conflict of
interest with females. Why power
asymmetries in favor of males are so
strong in this system and allow such
a male-dominated system to be an
ESS is more of a puzzle.
A lack of monopolizable resources
is more likely to be associated with
matrilineal kinship47,48 or with no
particular lineality at all. Matrilineal
kinship is almost always associated
with matrilocal residence and would,
on the face of it, seem, of all human
kinship systems, to provide the best
opportunity for cooperative or com-
munal breeding. When matrilineal
Khasi are directly compared with
patrilineal Bengalis who live nearby
in India, child health appears better
in the matrilineal group.45 But co-
residence does, of course, produce
potential competition between
female kin for resources, so matrilo-
cal females may have to forego high
reproductive rates in the interest of
maximizing inclusive fitness.
Zoologists make a distinction
between cooperative breeding sys-
tems, in which nonreproductives
help kin reproduce, and communal
breeding systems, in which all
females breed and offspring are com-
munally raised. Few human systems
follow the communal breeding pat-
tern, but duolocal matrilineal groups
(from which neither sex disperses)
come close.
We found that females living in
duolocal communal households
among the matrilineal Mosuo in
China, breed more slowly than those
living neolocally.40 Also, this group is
not associated with particularly high
fertility49 (Box 1). The matrilineal
Chewa in Mali suffer higher mortal-
ity when they have more female
kin.50 Both findings are suggestive of
reproductive constraints and compe-
tition among female kin. The fact
that females may be doing worse, in
terms of direct fitness, when breed-
ing communally should not necessar-
ily be taken as evidence that they are
not helping each other or benefiting
from doing so, but rather as evi-
dence that they are experiencing a
harsh environment in which they are
forced to share the means of repro-
duction. As with all the systems
described, the shares may not be
equal; some group members may
have to settle for a reduced share of
the resources available for reproduc-
tion. Comparing the reproductive
success of patrilineal Gambian
women living in group compounds.
which normally comprised a father,
brothers, and their families, we
found no overall benefit to residing a
in a larger compound. But being in
larger groups did favor older
breeding-age women over younger
breeding-age women, suggesting that
the former gain some help from that
latter, perhaps because of their
higher status within their house-
holds.36 It is likely that, given the
constraints in the system, energy is
better spent cooperating to maximize
inclusive fitness of all co-resident
brothers (among the Gambia) or sis-
ters (among the Mosuo) rather
than fighting relatives over the
resources that they all need to
reproduce.13,51,52
Among the matrilineal commu-
nally breeding Mosuo, the emergence
of tourism enabled some individuals
to raise extra money independent of
the farm through activities such as
work at tourist hotels. These individ-
uals often left their communal
households and established neolocal
nuclear families.53 Another example
of communal breeding by brothers
comes from polyandrous Sherpa
households in the Himalayas. Here,
farming habitat is limited, restricted
to a few river valleys; there are no
opportunities to disperse and set up
neolocal farms. Competition between
brothers was intense, and apparently
could be resolved only by all broth-
ers marrying the same wife – the
ARTICLE
ultimate in reproductive sharing!
Younger brothers did badly in terms
of direct fitness, but perhaps bene-
fited more from ceding some pater-
nity to their elder brother rather
than fighting with the him for the
farm or more access to their shared
wife. When resource constraints
were lifted by the emergence of jobs
outside farming, younger brothers
quickly left to set up a nuclear fam-
ily.54 Thus, both the matrilineal
Mosuo and the polyandrous Tibetans
show how communally breeding
households can emerge out of lim-
ited opportunities to disperse and
become unstable when there is relax-
ation of the ecological constraints.
The likelihood of helping or harm-
ing siblings may not be set for life;
at times they may harm each other
and at other times they may help
each other. It is clear that very
young siblings nearly always com-
pete for maternal investment, which
is why high birth rates are almost
universally associated with higher
child mortality.57 But as children get
older they are capable of helping
their younger siblings and their
mother. In many societies, elder sis-
ters in particular provide care for
younger siblings.58 However, adult
siblings may no longer be of assis-
tance once they have their own off-
spring to attend to. Also, as these
adult siblings’ offspring become their
priority, they may find themselves in
competition with their siblings. This
age-dependent change in helping sib-
lings has been observed, for example,
in patrilineal societies in The Gam-
bia and Senegal.30,60 In societies in
which inherited wealth matters,
there is evidence that fertility is
reduced when there is competition
for resources among siblings of
reproductive age.59
CONCLUSIONS
While the anthropological litera-
ture has tended to treat conflict and
cooperation as different subjects,
they are actually related to each
other. The semantic distinction can
even depend to some extent on
whether one takes a genes’-eye view
or observes the phenotype (that is,
behavior).
ARTICLE
Cooperatively breeding kin are
potentially in reproductive competi-
tion in the sense that individuals
would like to use the same resources
to reproduce. But potential conflict
can be resolved even if the partners
do not emerge with equal shares of
the cake. Depending on the chances
of gaining more control of a resource
and the relatedness of the partners in
conflict, individuals may benefit more
by helping their relatives breed than
by fighting with each other for the
resources to reproduce. But, almost
by definition, that help will involve a
cost in terms of their direct fitness.
Those individuals who do not breed
or do so at a slower rate than others
may do so because other females are
more powerful and punish them, they
are less efficient at gaining access to
resources, or there are other asymme-
tries in the benefits of helping. If the
evolutionarily stable strategy is to
help co-resident family members
maximize joint inclusive fitness
rather than waste energy on family
conflict that would reduce the repro-
ductive success of both partners,
overt aggression may not occur.
There may be outbreaks of conflict
when the status quo is disupted, sta-
tus hierarchies are re-established, or
breeding slots are reassigned.
Shakespeare’s oeuvre would be con-
siderably reduced if history were not
full of examples of cultural rules
allowing sufficient ambiguity for
major conflicts to break out. But most
of the time, strict cultural rules of suc-
cession may help obviate the need to
fight with relatives, especially when
norms resolve conflict in ways that
favor those most likely to win anyway
(such as favoring parental choice over
children’s choices and older siblings
over younger ones). Thus, human
marriage, residence patterns, and
inheritance norms may evolve by
maximizing the inclusive fitness of
parties that are potentially in conflict
within family units, and thus serve to
keep family members cooperating, at
least most of the time.
ACKNOWLEDGMENTS
Thanks to two anonymous referees
for useful comments, to Ting Ji and
Cooperation and Conflict Between Women in the Family 257
others at the Theoretical Ecology
Group of the Chinese Academy of Sci-
ences, Beijing, and at HEEG at UCL
for helpful discussion of these issues.
Ruth Mace is funded by the ERC.
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