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The Origin of Marriage: Explanations from the Paternal Provisioning Hypothesis and

Male-Biased Sex Ratios Due to Menopause


By: Katherine Daiy

ANTH 434-900
Dr. Jeffrey Winking
December 11, 2015

Introduction
One of the defining characteristics of the human story is the institution of marriage.
Marriage is defined as a mutually maintained long-term reproductive relationship (Jeffrey
Winking, personal communication, 2015). Marriage reflects the ability of humans to form stable
pair-bonds for a purpose of creating a cooperative venture towards reproduction and economic
stability. Despite considerable variation across cultures, the ubiquity of marriage makes it a
cultural universal (Murdock 1949). Numerous hypotheses have proposed its possible functional
and evolutionary purposes, often framing marriage as a reproductive strategy. The paternal
provisioning model posits that marriage provides the paternal investment and biparental
cooperation necessary to rear altricial human offspring (Winking et al. 2007). Others theorize
that the menopausal life stage produces a male-biased sex ratio by removing women from the
reproductive population, thus increasing intrasexual competition between males and favoring
stable-pair bonding (Coxworth et al. 2015). This paper will examine both hypotheses, the
paternal provisioning model and the male-biased sex ratio model, in order to explore the
universality of marriage from an evolutionary perspective.
Marriage is a mating strategy that differs greatly from those employed by other primates.
For instance, orangutans typically lead solitary lives and mate only when males territories
overlap with females (Nakahashi and Horiuchi 2012). Gorillas form large polygynous groups, in
which one male monopolizes the reproduction of several females (Nakahashi and Horiuchi
2012). Chimpanzees and bonobos exhibit cooperation within large multiple-male-multiplefemale groups, yet engage in brief, promiscuous sexual relationships (Nakahashi and Horiuchi
2012). Humans form similar multiple-male-multiple-female groups; however, unlike the highly
promiscuous strategies employed by chimpanzees , humans exhibit a tendency to form exclusive

sexual relationships, or pair-bonds, within these groups (Nakahashi and Horiuchi 2012). These
stable pair-bonds are often formally recognized through the institution of marriage.
There are several defining characteristics of marriage. Marriage is typically a mutuallymaintained reproductive relationship. Each individual consents to and makes effort to maintain
the relationship. In addition, most married couples have a higher probability of bearing children
than non-married couples (Murdock 1949:6). The ability to reproduce is a fundamental aspect of
spouse selection; sterility is often met with consequences such as divorce, especially for women
(Betzig 1989). Marriage also involves economic cooperation, where each spouse contributes
resources to each other and their offspring (Murdock 1949:5). Cohabitation, which is associated
with marriage cross-culturally (Marlowe 2000), facilitates this cooperation by allowing for
efficient provisioning to the family unit (Murdock 1949:5). Lastly, marriages are long-term and
sexually exclusive. Multiple reproductive events with the same partner are expected to occur
over the relationship; extramarital affairs are socially disapproved and punished accordingly
(Betzig 1989). Thus, marriage is a mutually-maintained, long-term reproductive relationship.
However, there are challenges to strictly defining a diverse practice such as marriage. For
instance, not all marriages are directly reproductive relationships. Among the Nuer of Sudan, if a
woman is infertile, she may marry another woman, who is then impregnated by another male.
The barren woman is considered to be the father and has complete control over the mother and
her children as a way to ensure the continuation of her kin line (Marlowe 2000). In addition,
marriages do not always consist of a single-pair bond and may be polygamous in nature.
Polygyny, the marriage between one male and multiple females, is practiced in 82% of the
worlds societies (Marlowe 2000). Only 1% of the worlds cultures practice polyandry, the
practice of marrying one woman to multiple men (Marlowe 2000). Monogamous marriages are

the most widely practiced, even in societies with high rates of polygyny (Marlowe 2000). In
addition to variance in composition, the socially-sanctioned importance of sexual exclusivity
varies cross-culturally. In some societies, extramarital affairs are punishable by death, especially
for women (Betzig 1989). However, in some societies, such as the Ach of Paraguay, there are
considerable degrees of sexual liberty in pair-bond relationships (Hill and Hurtado 2009).
Although marriage is practiced diversely, it remains a universal feature of human behavior that
can be explained by evolutionary logic.

Hypotheses
Paternal Provisioning Model
Human life histories are unique among primates in that offspring are remarkably helpless,
or altricial. Human altriciality can be accounted for as an evolutionary tradeoff between prenatal
cognitive development and the facilitation of bipedalism with a narrow birth canal (Winking
2007). Direct provisioning allows offspring to be weaned early despite an extended juvenile
dependency, which shortens the interbirth intervals of mothers (Winking 2007). In comparison,
non-human primate offspring are less altricial and are raised as a single infant per birth period
(Nakahashi and Horiuchi 2012). Despite the higher degree of offspring helplessness in humans ,
however, human mothers exhibit shorter birth intervals than those of non-human primates,
allowing them to raise large numbers of multiple dependent children of different ages (Alvarez
2000). However, raising altricial offspring is energetically costly for mothers; breastfeeding
severely detracts from their resource acquisition (Quinlan 2008). Human birth intervals may be
allowed for by the prevalence of male assistance in child rearing (Lovejoy 1981). Pair-bonding
facilitates consistent paternal assistance by allowing for cohabitation, where males become more

available to provide direct care to offspring or to aid mothers and free them to give direct care
(Quinlan 2008).
In marriage, men can invest in a large progeny of offspring and receive more benefits
from child investment than can be attained through a more mating-intensive strategy (Winking,
2006). Marriage allows for the greater contribution of resources through a division of labor that
increases economic productivity for a family. In this model, males contribute large, difficult-toobtain resources to their families, whereas females participate in child care and acquirement of
smaller, steadier flows of resources (Winking 2006). Male contribution to subsistence should be
positively associated with high offspring fitness, as men provision vital resources that benefit
offspring wellbeing and survivorship. Additionaly, divorce is a costly option in societies where
men contribute the majority of resources, therefore male-contribution to subsistence is predicted
to be negatively associated with pair-bond stability.

Existing Evidence
The evidence supporting the influence of paternal investment on offspring fitness is weak
and varied. In one study that examined the effect of father absence on adult wellbeing among the
Tsimane of Bolivia, no significant negative developmental effect was found among fatherless
children (Winking, Gurven, and Kaplan 2011). Although it has been shown that fathers
investments do improve the survivorship of children, fatherless children among the Tsimane do
not have lower fertility rates than children who were raised by both parents (Winking, Gurven,
and Kaplan 2011). This is surprising because the Tsimane are a population characterized by high
mortality rates and fathers are typically essential to family provisioning; there are low divorce
rates after two children (Winking, Gurven, and Kaplan 2011).

Cross-cultural studies have produced weak evidence on the importance of fathers among
non-maternal caregivers. One examination of the relative contribution of each kin member to
child survivorship reported very little effect of the contributions of men on child mortality;
however, there was a significant positive effect on child survivorship from maternal and
grandmothers contributions (Sear and Mace 2008). Extended kin such as older siblings were
also shown to provide a substantial amount of child care (Sear and Mace 2008). Furthermore,
alloparental care has been shown to remedy the costs on offspring fitness from father absence
and is even associated with conjugal instability (Quinlan, 2008). Paternal provisioning is
evidently not the solely most important form of non-maternal provisioning.

Male-Biased Sex Ratio Due to Menopause


Historically, the paternal provisioning hypothesis has been the prevalent explanation for
the universality of marriage (Coxworth et al. 2015; Winking 2007). However, an alternate
hypothesis proposed by Hawkes and colleagues (2015) suggests that skewed operational sex
ratios, or the ratio of sexually receptive males to females (Cartwright 2008), drove the selection
for marriage. This hypothesis posits that the human post-reproductive stage (menopause) was
selected for to allow investment in children and grandchildren. These investments would have
increased human lifespan. As the extension in lifespan occurred without extended fertility in
females, the operational human sex ratio was distorted in favor of males, increasing male-male
competition for a limited number of fertile females (Coxworth et al. 2015). It became more
reproductively efficient for males who had already pair-bonded to remain with their first partner,
guard their mates and invest in their children rather than seek other females to reproduce in the

midst of costly competition. Selection favored a pair-bonding reproductive strategy in the form
of marriage as the payoffs to mate-guarding superseded those of an intensive mating strategy.

Existing Evidence
The basis of this model lies within the mate guarding hypothesis, which attributes the
existence of stable pair-bond relationships in humans to the fitness benefits received from
defending females. In cross-species analyses, there is a positive relationship between relative
number of sexually-active males in a population and the prevalence of mate guarding (Parker
1974). Although mate guarding is not synonymous with pair-bonding, the two assume that it
benefits the more numerous sex to stay with a current mate rather than search for another partner
(Coxworth et al. 2015). Non-human primates primarily have a female biased operational sex
ratio, and correspondingly exhibit no pair-bonding attributes (Coxworth et al. 2015).
The existence of menopause in human women appears to create the male-biased sex ratio
that corresponds to this mate-guarding behavior. Hawkes and colleagues (2015) ran computer
simulations of human evolution, demonstrating the effects of the presence and absence of
grandmothering on operational and adult sex ratios. With the increased longevity associated
with a steady decline of female fertility with age, both the operational sex ratio and the adult sex
ratio simulation resulted in an increasingly male-biased ratio over time (Coxworth et al. 2015).
The ratios obtained were similar to modern male-biased human sex ratios found across four
representative hunter-gatherer populations (Coxworth et al. 2015). In contrast, the simulations
without the effects of grandmothering resulted in a sex ratio similar to great-ape populations
(Coxworth et al. 2015). Although theses simulations support the hypothesis, there are reported
female-biased sex ratios in some hunter-gatherer populations. For instance, one survey of

ethnographic literature reports 8 of 15 traditional populations exhibiting a female-biased adult


sex ratio (Schacht, Rauch, and Mulder 2014)..

Methods
The Standard Cross-Cultural Sample (SCCS) is a sample of 186 well-described
preindustrial societies compiled by George P. Murdock and Douglas R. White (1967). Using
ethnographies, each society is coded for in 1,400 variables, according to varying degrees of
cultural and demographic traits. The sample is sufficiently large and each case is independent
from another; thus it is an ideal source of data for testing hypotheses.

Methods: Paternal Provisioning Model


In order for the paternal provisioning model to be correct, male contribution to
subsistence needs to respond to variance in human mating systems. Percentages of male
contribution to subsistence should be negatively associated with frequency of divorce, and there
should be statistically significant differences between means of contribution in response to
frequency of divorce. I compared means of percent male contribution to subsistence across
different frequencies of divorce using a one-way ANOVA (Analysis of Variance). Percent male
contribution was derived from variable 889 (Whyte 1978; Murdock 1967; Barry and Schlegel
1982), percent female contribution to subsistence, by subtracting this value from 100. Variable
744 (Broude and Greene 1983), frequency of divorce, is coded as 5 discrete values. A value of 1
indicates that divorce is universal in a society, 2 meaning common, 3 meaning moderate,
where a small minority of couples divorce, 4 indicating that it is frequent in the first few years

of marriage and/or before children, but rare thereafter, and 5 indicating that divorce occurs
rarely or never (Broude and Greene 1983).

Methods: Male-Biased Sex Ratio Due to Menopause


This sex ratio model posits that stable pair bonding evolved as a male-biased sex ratio
induced by menopause made it fitness-enhancing for men to avoid an intensive mating strategy.
Thus, it is predicted that the degree of male-biased sex ratio will be negatively associated with
the frequency of divorce, as males remain in pair-bonds to avoid costly competition. I performed
a one-way ANOVA to determine if there is a statistically significant difference between mean
frequencies of divorce in response to different biases in sex ratio. Variable 714 (Whyte 1978),
sex ratio, is coded in three categories. A value of 1 indicates a female-biased sex ratio, 2
indicates equilibrium, and 3 indicates a male-biased sex ratio (Whyte 1978). Variable 744
(Broude and Greene 1983), frequency of divorce, is coded in 5 discrete values as described
previously.

Results

Table 1. One-Way ANOVA: Mean % Male Contribution to


Subsistence by Frequency of Divorce
Freq. of
1
2
3
4
5
Divorce
Mean
60.38
62.97
61.55
62.10
66.96
Standard
Error
p-value

6.62
0.80

2.69

3.32

5.86

2.06

Mean % Male Contribution to Subsistence

Figure 1. Mean % Male Contribution to Subsistence


by Frequency of Divorce
75
70
65
60
55
50
45
40
1

3
Frequency of Divorce

Table 1 and Figure 1 display the results of the test of the paternal provisioning
hypothesis. Mean male contribution to subsistence appears to decrease roughly with frequency of
divorce (Figure 1). There were no statistically significant differences between means as
determined by one-way ANOVA (p=0.80, where =0.05). Due to small sample sizes, standard
error was high (all values are greater than 1.00) for all values of divorce frequency (Table 1),
indicating that the sample means are not representative of actual population means.

Figure 2. Mean Frequency of Divorce by Degree of


Male-Biased Sex Ratio

Standard Error
p-value

0.32
0.03

0.12

0.22

4.5
Mean Frequency of Divorce

Table 2. Mean Frequency of Divorce by Degree of Male-Biased


Sex Ratio
Operational Sex Female-Biased Equal
Male-Biased
Ratio
Mean
3.60
3.29
2.75

4
3.5
3
2.5

2
1.5

1
0.5
0
Female-Bias

Equal

Male-Bias

Degree of Male-Biased Sex Ratio

Table 2 and Figure 2 display the results of the one-way ANOVA test of the sex ratio
hypothesis. Mean frequency of divorce is negatively associated with the degree of male-biased
sex ratio (Figure 2). There is a statistically significant difference between mean frequencies of
divorce as determined by one-way ANOVA (p=0.03). Standard error was relatively low,
indicating that sample means represent population means (Table 1).

Discussion
Paternal provisioning, although beneficial to offspring wellbeing and survivorship, is
evidently not the sole and most important factor in the success of child-rearing and thus could
not have single-handedly facilitated the formation of stable pair-bonds. Even in a population
where paternal contributions are important for family subsistence, such as the Tsimane of
Bolivia, father presence does not significantly affect any measure of adult offspring fitness,
including fertility (Winking et al. 2011). This undermines the predictions of the paternal
provisioning model, where the presence of fathers is expected to significantly enhance offspring
reproductive value. In addition, mens contributions have little negative effect on child mortality
cross-culturally (Sear and Mace 2008), in contrast to larger negative effects from maternal and
grandmothers contributions (Sear and Mace 2008). Moreover, alloparental care has been shown
to even ameliorate the costs on offspring wellbeing incurred from father absence (Quinlan 2008).
These findings challenge the estimated importance of fathers contributions on offspring
survivorship. Moreover, it points to a model of communal child-rearing in human kinship
systems and refutes the nuclear-family model proposed by the paternal provisioning hypothesis
(Murdock 1949). Evidently, there are weaknesses in the paternal provisioning models
predictions of family structure.
If the paternal provisioning model is correct, then mens contributions to subsistence
should be associated with high pair-bond stability, as it becomes reproductively beneficial for
women to avoid divorce and remain in relationships where essential provisioning is provided.
However, no statistically significant differences were found between means in a comparison of
mean percent male contributions across different frequencies of divorce (Table 1, where p >
0.05). Although there is a slight negative relationship between mean contribution and divorce

(Figure 1), it is insignificant (p=0.80). This finding indicates that mens contribution to
subsistence is not associated with conjugal stability. Even in societies where men provision a
great deal, women are able to divorce their husbands. The availability of alloparental care may
lower costs of divorce incurred by mothers and make it a viable option, even in populations
where men are responsible for a great deal of family resources and have the ability to affect
offspring fitness substantially. With a lack of an association between male contribution and
conjugal stability, these findings disprove the presupposition that paternal provisioning was
influential enough to maintain stable pair-bonds.
In contrast, the male-biased sex ratio due to menopause model is well-supported by a
multitude of evidence. Hawkes and colleagues (2015) take into account the grandmother
hypothesis into the creation of the male-biased sex ratio, where post-menopausal women were
able to help their grandchildren survive, thus increasing longevity. The involvement of
grandmothers in child rearing indicate in a communal-rearing model of human evolution, which
corresponds to the cross-cultural presence of alloparental care (Sear and Mace 2008). In addition,
there is cross-species evidence of the association between pair-bonding and male-biased sex
ratios. An excess of males corresponds to mate-guarding behavior in Class Aves and other pairbonding species (Parker 1974). These ratios are even found in traditional societies, which reflect
early human societies through similar mortality rates (Coxworth et al. 2015). Although there are
observations of female-biased operational sex ratios in some traditional societies, the
inconsistency in the definition of fertile men could have skewed the ratios in favor of females
(Coxworth et al. 2015). It is evident that the sex ratio models account of communal childrearing and the presupposition of male-biased sex ratios among early hunter-gatherers finds
much support from kin structure and cross-species analyses.

A statistically significant difference was found between mean frequencies of divorce


(p=0.03) as determined by one-way ANOVA (Table 2), indicating a response of pair-bond
stability to sex ratio bias. This supports the presupposition that an increase in male-bias and high
male-male competition led early hominin males to remain in pair-bonds. In addition, there is a
significant negative relationship between divorce and the degree of male-biased sex ratio (Figure
2). Standard error was low enough to ensure reliability of the presented means (Table 1). We can
conclude that the male-biased sex ratio due to menopause model has significant support from
cross-cultural evidence.

Conclusion
This examination of the paternal provisioning and male-biased sex ratio due to
menopause hypotheses reveals a communal model of child rearing that favors the
presuppositions of the latter. Maternal and alloparental provisioning has a more significant effect
on offspring fitness than that contributed by fathers (Winking, Gurven, and Kaplan 2011; Sear
and Mace 2008). Moreover, paternal provisioning had little effect on pair-bond stability (Figure
1). There is substantial support, however, for the validity of the sex ratio model. The models
basis in the grandmother hypothesis accounts for communal child-rearing found crossculturally (Sear and Mace 2008). Male-biased sex ratio is significantly associated with increased
pair-bond stability (Figure 2), possibly indicated the effects of intrasexual competition. Although
paternal provisioning is beneficial to child rearing, the effects are unsubstantial; instead,
evidence frames menopause as the likely facilitator of the evolution of stable pair-bonding,
otherwise known as marriage.

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