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Moorad 2010 Effect of Mormon Fundamentalist Polygyny on Genetic / Evolutionary Fitness

Moorad 2010 Effect of Mormon Fundamentalist Polygyny on Genetic / Evolutionary Fitness

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Mormon Fundamentalist polygyny's effect on Female Evolutionary Fitness
Mormon Fundamentalist polygyny's effect on Female Evolutionary Fitness

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1
Original Article
2
Mating system change reduces the strength of sexual selection in an
3
American frontier population of the 19th century
,
☆☆
4
Jacob A. Moorad
a,b,
, Daniel E.L. Promislow
, Ken R. Smith
c
, Michael J. Wade
d
5
 Department of Genetics, University of Georgia, Athens
Q1
, GA, USA
6
 b
 Department of Pathology, University of Michigan Medical School, Ann Arbor, MI, USA
7
c
 Huntsman Cancer Institute, University of Utah, Salt Lake City, UT, USA
8
d
 Department of Biology, Indiana University, Bloomington, IN, USA
9
Initial receipt 30 July 2010; final revision received 15 October 2010
10
Abstract
11
Sexual selection, or competition among members of one sex for reproductive access to the other, is one of the strongest and fastest 
12
evolutionary processes. Comparative studies support the prediction that sexual selection is stronger in polygamous than in monogamous
13
species. We report the first study of the effect on sexual selection of a change in mating system, from polygyny to monogamy, within a
14
historical human population. Here we show that over the reproductive lifetimes of Utahns born between 1830 and 1894, socially
15
induced reductions in the rate and degree of polygamy correspond to a 58% reduction in the strength of sexual selection. Polygyny
16
conferred a strong advantage to male fitness as well as a weak disadvantage to female fitness. In contrast, mating with multiple males
17
provided little benefit to females in this population. Polygamy benefitted males by increasing reproductive rates and by lengthening
18
reproductive tenure. Each advantage contributed to roughly half of the increased total lifetime reproductive success. This study illustrates
19
both the potency of sexual selection in polygynous human populations and the dramatic influence that short-term societal changes can
20
have on evolutionary processes.
21
© 2010 Published by Elsevier Inc.
2223
Keywords:
LDS; Mormon; Utah; Bateman; Opportunity for selection; Polygamy; Polygyny
2425
1. Introduction
26
Sexual selection is the process identified byDarwin
27
(1859)where the members of one sex, generally males,
28
compete with one another for reproductive access to
29
members of the other sex
Q2
(Morgan, 1991). Theory predicts
30
that sexual selection should be a much stronger evolutionary
31
force in polygamous than in monogamous mating systems
32
33
2004b). Support derives primarily from extensive compa-
34
risons across taxa with different mating systems (Andersson,
35
36
Borgia, 1986; Shuster & Wade, 2003). We report the first 
37
study of the effects on sexual selection of a change in mating
38
system, from polygyny to monogamy, within a population of 
39
a single species,
Homo sapiens
.
40
Polygynous marriage, where a man would simultaneously
41
take on multiple wives, was a practice throughout much of 
42
the 19th century for some members of the Church of Jesus
43
Christ of Latter-day Saints (LDS or Mormons). In the Utah
44
Territory in the early part of the century, our estimates
45
from the Utah Population Database indicate that the
46
frequency of polygamous men as a fraction of all married
47
men reached a maximum of 17.8% among men born in
48
1833 (Fig. 1), a figure consistent with previous estimates
Grants: This work was funded by National Science Foundation grant DEB-0717234 to J.A.M. and D.EL.P., and NIH grant RO1GM065414-06and NSF grant DEB-0614086 to M.J.W and partially supported by NationalInstitute on Aging grant P30-AG013283.
☆☆
Disclaimer: The study is solely our responsibility and does not necessarily represent the official views of the National Institute on Aging or the National Institutes of Health. Grant sponsor: National Institute of Aging(The Utah Study of Fertility, Longevity, and Aging); grant number:AG022095.
Corresponding author. Department of Biology, Duke University,Durham, NC 27708, USA.
 E-mail address:
jacob.moorad@duke.edu(J.A. Moorad).1090-5138/$
see front matter © 2010 Published by Elsevier Inc.doi:10.1016/j.evolhumbehav.2010.10.004
ENS-05674; No of Pages 9
 
49
(Smith & Kunz, 1976). Subsequently, changes in social
50
pressures over the next few decades reduced the frequency
51
of such marriages to less than 1%. New strictures on the
52
practice of polygyny began with the Morrill Anti-Bigamy
53
Act of 1862 outlawing plural marriage throughout the US
54
territories. The US Supreme Court upheld the ban in the
55
face of a freedom of religion challenge in 1878 and, in
56
1890, the Mormon Church issued the Woodruff Manifesto
57
which officially banned polygyny.
58
Data from this population afford a unique test of the role
59
of polygyny in sexual selection in four ways. First, until now,
60
the primary source of empirical support for sexual selection
61
theory has been comparative studies across taxa with
62
different mating systems (Andersson, 1994; Bateman,
63
64
Shuster & Wade, 2003). Secondly, previous human studies
65
have compared relationships between social mating customs
66
and sexual selection across geographically and culturally
67
distinct populations (Brown, Laland & Mulder, 2009),
68
whereas this study uses temporally defined cohorts tha
69
differ in mating system within the same population. Third,
70
our sample size (180,082 parents) is 18 times larger than that 
71
in all other human studies of sexual selection combined
72
(Brown et al., 2009), giving unprecedented power to
73
describe sexual selection in a human population. Finally,
74
these data span an important part of the demographic
75
transition (Borgerhoff-Mulder, 1998) where natural fertility
76
conditions prevailed early but reproductive rates rapidly fell
77
over a few generations.
78
To determine the strength of sexual selection that acts
79
on a population and to best understand how it is influenced
80
by mating system, we need to quantify the associations
81
(e.g., the variances and covariances) between mating
82
success and reproductive success in both males and
83
females. In populations with an equal sex ratio of females
84
to males, the rates of polygyny are necessarily and
85
 positively associated with the frequency of males who do
86
87
result, polygyny increases the
variation
among males in
88
mate number (mating success or MS). Using results from
89
fruit fly experiments,Bateman (1948)was the first to argue
90
that variation among males in mate numbers was the most 
91
important cause of a sex difference in the variance in
92
reproductive success (RS).Wade (1995)derived the
93
general relationship between the variance in male and
94
female RS, showing that the sex difference in
σ
RS2
was a
95
function of male MS.Shuster and Wade (2003)reviewed
96
and extended this theory and applied it to several different 
97
98
99
reversed species with polyandry among territory-holding
100
females (Jones & Avise, 2001), to show how mating
101
system affected the strength of sexual selection. A recent 
102
comparative study using 18 geographically diverse human
103
 populations, a collection that included both monogamous
104
and polygamous societies (Brown et al., 2009), showed
105
that 
σ
RS2
and
σ
MS2
tended to be higher in males than in
106
females. Following the methods described in theMethods
107
section, we examine these relationships within a single
108
 population during a period of change in the practice of 
109
 plural marriage that affected mating success.
110
2. Methods
111
Our extract of the Utah Population Database (UPDB)
112
included birth and death years of the 89,034 fathers and
113
96,407 mothers recorded in the UPDB born before 1900
114
(the majority of the UPDB reflecting Utah parents born
115
after 1900 was not available to us). Note that the Utah
116
 population was not a strict closed society since potential
117
wives could be recruited via religious conversion and
118
migration into Utah (Daynes, 2001). We used only those
119
individuals with known death year. There were very few
120
individuals born before 1830 and these were excluded from
121
our analysis. Fathers born after 1894 had missing data on
122
their wives who were often born after the cutoff year of 
123
1900. These exclusion criteria meant we had a sample of 
124
72,812 fathers and 75,366 mothers born between 1830 and
125
1894. Our extract also included the 322,551 sons and
126
307,754 daughters who were born to parents represented in
127
the UPDB extract. These parents were divided into 65
128
single birth-year cohorts from 1830 to 1894. Because the
129
UPDB is derived from descendent genealogies, we could
130
not expect it to include reliable counts of nulliparous
131
individuals. We used US census data and a few reasonable
132
assumptions (seeSection 2.1below) to estimate the size of 
133
this class of individuals for each sex and birth years. The
Fig. 1. Polygamous men in the UPDB as a frequency of all married men by birth year.2
J.A. Moorad et al. / Evolution and Human Behavior xx (2010) xxx
 – 
 xxx
 
134
sample sizes that reflect the total number of parents plus the
135
projected number of nulliparous individuals are illustrated
136
in Fig. A1 inAppendix A.
137
We determined the total number of offspring (reproduc-
138
tive success,
RS
) for each individual, the number of his or 
139
her reproductive partners (mating success,
MS
) and the
140
number of years over which he or she reproduced
141
(reproductive tenure,
RT
). The birth year-specific means
142
of mating and reproductive success are shown in Fig. A2 in
143
Appendix A. In addition, each reproductive female was also
144
assigned another value equal to the mating success of the
145
partner that she most often reproduced with. In the cases
146
where females mated with more than one male, the mating
147
success of these females' first and last mates was also noted.
148
The choice of partner had no noticeable effect on the
149
outcome of our analyses.
150
2.1. Estimating the number of nulliparous individuals
151
We need to know the frequency distributions of repro-
152
ductive success to estimate the strength of sexual selection.
153
This includes those individuals that do not reproduce at all.
154
When information comes from descendent genealogies, such
155
as with the UPDB, these individuals can be undercounted
156
and omitting the zero values diminishes the variance in
157
reproductive success and the strength of selection (Shuster &
158
Wade, 2003). This section explains how we corrected for this
159
bias and estimated the size of the nulliparous populations of 
160
males and females.
161
Our approach was to complement data on survival and
162
reproduction from the UPDB, which we expect to be biased
163
in terms of the relative number of reproductive men and
164
women, with information from US census data (Sixteenth
165
Census of the United States, 1943). These data include
166
fertility and marital information about women living in Utah
167
and reported in 1910 and 1940. Parity counts of women
168
relevant to our study period are reproduced inTable 1. These
169
include the age-specific counts of total number of women
170
ever married (
), ever-married women who reported having
171
no children (
0
), ever-married women who did not report 
172
number of children (
U
) and single women (
). Women
173
were grouped into age categories of 5-year intervals (e.g., 40
174
to 44 years of age) until 50 years of age, at which point a
175
25-year age window was considered. We assumed tha
176
marriage and reproductive history did not influence reporting
177
and, in this way, we adjusted for the under-reporting of 
178
nulliparous wives by estimating the true counts for non-
179
reproductive women of age interval
x
and birth-year interval
180
 y
as
V
 xy
0
ð Þ
=
 xy
0
ð Þ
1
 xy U 
ð Þ
=
 xy
+
 xy
. The adjusted numbers of 
181
motherswere
V
 xy m
N
0
ð Þ
=
 xy
 xy
0
ð Þ
1
 xy U 
ð Þ
=
 xy
,where
m
N
0indicates
182
fertility.
183
Always-single women (
) were assumed not to have had
184
children prior to the census year. However, women who
185
were single or married and nulliparous may have had first 
186
children after the census. This would cause us to overesti-
187
mate rates of nulliparity. For this reason, we considered only
188
those women who were 35 years or older at the time of the
189
census. Using reproductive records from the UPDB and
190
 pooled overall birth year cohorts of interest (1830
1894), we
191
estimated that the probability that a mother had her first child
192
after 35 was
δ
35
=0.0188. The average taken over the age
193
interval
x
=35 to 39 (and weighted by age-cohort size at 
194
1910) was
δ
 ¯ 
(35-39), (1871-1875)
=0.0119. The corresponding
195
 probability for the next youngest age-cohort was
δ
 ¯ 
(30-34),
196
(1876-1880)
=0.0406; the chance that missing these mothers
197
may cause problems with our estimates seemed too high to
198
risk using these age cohorts. Women above 49 years of age
199
were not considered owing to the exceedingly large age
200
window (25 years). Although the frequency of missed
201
categorized nulliparous women was gauged to be quite low
202
for the cohorts of interest (between 0.0002 and 0.0119), we
203
made the following adjustments to best account for this bias:
204
xy
(0)
=
 xy
(0)
(1
δ
 ¯ 
 xy
) and
xy
(
m
N
0)
=
 xy
(
m
N
0)
+
δ
 ¯ 
 xy
 xy
(0)
.
205
Finally, we accounted for the probability that a women
206
did not survive from birth to the time that the census was
207
taken. Using UPDB data, we estimated the probability
L
-
208
(
m
N
0)
that a mother born at time
y
lived to age
x
. Using the
209
set of individuals with no reproductive records, we
210
estimated
L
 xy
(0)
, which better reflected the cumulative
211
survival fraction of the nulliparous females. We reasoned
212
that if the census figures accounted for all females who
213
had died prior to the census and also correctly forecasted
214
the frequency of future first-time mothers, then the total
215
number of mothers and non-reproductive females would be
216
W
 xy m
N
0
ð Þ
 L
 xy m
N
0
ð Þ
and
W
 xy
0
ð Þ
 L
 xy
0
ð Þ
, respectively, where
x
was the age at 
217
census and
was the birth year. The relative frequencies
218
of projected always-nulliparous females born in birth year 
219
interval
y
,
 y
(F)
, as well as all parameter estimates from
220
the UPDB that were used in the necessary adjustments
221
described above, are shown inTable 2.
222
We regressed
 y
(F)
onto the midpoint of the 5-year birth
223
intervals in order to project nulliparous fractions on specific
Table 1
t1
:
1
Fertility and marriage history of Utah women from US Census data
t1
:
2t1
:
3
AgeclassBirth years Censusyear Women ever married Singlewomen,
t1
:
4
Total reporting,
Zero children,
0
Not reporting,
U
t1
:
5
45
49 1891
1895 1940 12,480 1100 1320 520
t1
:
6
35
39 1871
1875 1910 8336 627 446 563
t1
:
7
40
44 1866
1870 1910 7204 371 372 405
t1
:
8
45
49 1861
1865 1910 6244 282 380 2403
 J.A. Moorad et al. / Evolution and Human Behavior xx (2010) xxx
 – 
 xxx

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