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Abstract

Although recent research has increasingly focused on human sexual selection, important questions remain concerning the relative influence of individual traits on success in competition for mates and the mechanisms, form and direction of these sexual selective pressures. Here, we explore sexual selection on men’s traits by ascertaining men’s dominance and attractiveness from male and female acquaintances. On a large American university campus, 63 men from two social fraternities provided anthropometric measurements, facial photographs, voice recordings, and reported mating success (number of sexual partners). These men also assessed each other’s dominance, and 72 women from two socially affiliated sororities assessed the men’s attractiveness. We measured facial masculinity from inter-landmark distances and vocal masculinity from acoustic parameters. We additionally obtained facial and vocal attractiveness and dominance ratings from unfamiliar observers. Results indicate that dominance and the traits associated with it predict men’s mating success, but attractiveness and the traits associated with it do not. These findings indicate that male contests strongly influence men’s mating success in this population and suggest that similar conditions may have existed over human evolution.
Original Article
Quantifying the strength and form of sexual selection on men's traits
,
☆☆
Alexander K. Hill
a
, John Hunt
b
, Lisa L.M. Welling
a,c
, Rodrigo A. Cárdenas
d
, Michelle A. Rotella
a
,
John R. Wheatley
a
, Khytam Dawood
d
, Mark D. Shriver
a
, David A. Puts
a,e,
a
Department of Anthropology, Pennsylvania State University, University Park, PA 16802
b
Centre for Ecology and Conservation, School of Biosciences, University of Exeter, Penryn, UK
c
Department of Psychology, Oakland University, Rochester, MI 48309
d
Department of Psychology, Pennsylvania State University, University Park, PA 16802
e
Center for Brain, Behavior, and Cognition, Pennsylvania State University, University Park, PA 16802
abstractarticle info
Article history:
Initial receipt 13 March 2013
Final revision received 30 May 2013
Keywords:
Attractiveness
Dominance
Face
Masculinity
Sociosexuality
Voice
Although recent research has increasingly focused on human sexual selection, fundamental questions remain
concerning the relative inuence of individual traits on success in competition for mates and the mechanisms,
form, and direction of these sexual selective pressures. Here, we explore sexual selection on men's traits by
ascertaining men's dominance and attractiveness from male and female acquaintances. On a large American
university campus, 63 men from two social fraternities provided anthropometric measurements, facial
photographs, voice recordings, and reported mating success (number of sexual partners). These men also
assessed each other's dominance, and 72 women from two socially afliated sororities assessed the men's
attractiveness. We measured facial masculinity from inter-landmark distances and vocal masculinity from
acoustic parameters. We additionally obtained facial and vocal attractiveness and dominance ratings from
unfamiliar observers. Results indicate that dominance and the traits associated with it predict men's mating
success, but attractiveness and the traits associated with it do not. These ndings point to the salience of
contest competition on men's mating success in this population.
© 2013 The Authors. Published by Elsevier Inc. All rights reserved.
1. Introduction
A rapidly growing literature suggests that sexual selection has
shaped men's phenotypic traits (Puts, Jones, & DeBruine, 2012).
Men's bodies (Pawlowski & Jasienska, 2005), faces (Penton-Voak &
Perrett, 2000; Rhodes, Chan, Zebrowitz, & Simmons, 2003), and
voices (Fitch & Giedd, 1999; Puts, Apicella, & Cárdenas, 2012)
exhibit features that are highly sexually differentiated and develop
at sexual maturity. These traits also appear to aid in competition for
mates. In men, more masculine, muscular bodies (Dixson, Dixson,
Bishop, & Parish, 2010; Frederick & Haselton, 2007; Hughes,
Dispenza, & Gallup, 2004) and tall stature (Nettle, 2002; Pawlowski,
Dunbar, & Lipowicz, 2000; Pawlowski & Jasienska, 2005) predict
reported number of sexual partners and perceptions of attractive-
ness and dominance. Masculine faces also convey dominance
(DeBruine et al., 2006; Perrett et al., 1998; Watkins, Jones, &
DeBruine, 2010), and some studies (DeBruine et al., 2006; Johnston,
Hagel, Franklin, Fink, & Grammer, 2001), but not others (DeBruine,
Jones, Smith, & Little, 2010; Perrett et al., 1998; Rhodes, Hickford, &
Jeffery, 2000), have found that women prefer masculine male faces,
particularly during the fertile phase of the ovulatory cycle (Johnston
et al., 2001; Penton-Voak & Perrett, 2000; Penton-Voak et al., 1999;
Welling et al., 2007). Similarly, masculine voices have been found to
predict men's number of reported sex partners (Hodges-Simeon,
Gaulin, & Puts, 2011; Puts, 2005) and reproductive success (Apicella,
Feinberg, & Marlowe, 2007). In addition, masculine acoustic features,
such as deep timbre and low pitch inuence perceptions of both
dominance (Jones, Feinberg, DeBruine, Little, & Vukovic, 2010; Puts,
Hodges, Cárdenas, & Gaulin, 2007; Watkins, Fraccaro, et al., 2010;
Wolff & Puts, 2010) and attractiveness (Feinberg, Jones, Little, Burt,
& Perrett, 2005; Hodges-Simeon, Gaulin, & Puts, 2010), the latter
particularly during the fertile phase of the ovulatory cycle (Feinberg
et al., 2006; Puts, 2005, 2006).
Such research has helped illuminate whether and how sexual
selection has shaped men's phenotypes, yet a number of funda-
mental questions remain. First, prior studies have typically focused
on either female choice or male contests without attempting to
quantify the relative contributions of these mechanisms to the total
sexual selective pressure on a particular trait (Hunt, Breuker,
Sadowski, & Moore, 2009). Second, to our knowledge, no study
reporting relationships between a male trait and mating success has
investigated whether these relationships were mediated by
Evolution and Human Behavior 34 (2013) 334341
This is an open-access article distributed under the terms of the Creative Commons
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medium, provided the original author and source are credited.
☆☆ JH was funded by NERC and a University Royal Society Fellowship. LW was funded
by an AIB grant awarded to DAP.
Corresponding author. Department of Anthropology, Pennsylvania State Univer-
sity, University Park, PA 16802. Tel.: +1 814 867 0453.
E-mail address: dap27@psu.edu (D.A. Puts).
1090-5138/$ see front matter © 2013 The Authors. Published by Elsevier Inc. All rights reserved.
http://dx.doi.org/10.1016/j.evolhumbehav.2013.05.004
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attractiveness or dominance. Third, most studies of sexual selection
in men have measured success under female choice or male
contests from limited information, such as body size, strength, or
ratings of faces or voices made by strangers in the laboratory (but
see, e.g., Pillsworth, 2008; von Rueden, Gurven, & Kaplan, 2011).
Attractiveness and dominance have thus frequently been assessed
devoid of relevant information, such as personality and intelligence,
and in isolation from the complex webs of social relationships in
which we live. Fourth, facial, vocal, and bodily characteristics may
be developmentally correlated (Feinberg, 2008; Fink, Neave, &
Seydel, 2007), so when exploring relationships between each and
success in mating competition, it is necessary to measure and
statistically control for the others, as well as to explore their
interactions (i.e., correlational selection). Fifth, nearly all prior
research on these traits has assumed linear (directional) selection
without exploring quadratic selection gradients (i.e., stabilizing or
disruptive selection). Finally, previous studies have not explored
whether female choice or male contests contribute more strongly to
men's mating success, despite the centrality of this question in
understanding human sexual selection (Puts, 2010). In sum, we still
do not know the overall sexual selective pressures on individual
traits, the relative strengths of sexual selection on different traits,
the forms and mechanisms of this sexual selection, or even the
relative importance of female choice and male contests in men's
competition for mates.
In the present paper, we therefore investigated sexual selection on
some of the strongest candidates for sexually selected traits in men:
stature, body build, facial masculinity, and deep voices (Fig. 1). We
measured these traits in a sample of men and obtained assessments of
the men's success under intra- and intersexual selection from logically
the most authoritative source: familiar male and female peers
(Pillsworth, 2008; von Rueden et al., 2011). We also obtained
assessments of the men's facial and vocal attractiveness and
dominance from unfamiliar raters, as well as the men's self-reported
mating success. We then 1) investigated the contributions of these
traits to different mechanisms of sexual selection (mate choice and
contests) and to mating success via their linear, quadratic, and
correlational selection gradients, and 2) compared mechanisms of
sexual selection to each other and to mating success using a
sequential model-building approach (Draper & John, 1988) to identify
whether the strength, direction, and form of sexual selection on male
traits differ across these episodes, and if so, which traits contribute to
these overall differences.
Although we are interested in how past selection produced
present sexual dimorphisms, we take a behavioral ecological
approach, which emphasizes contemporary selection. We take this
approach because we expect that, in general, current function will
provide insight into past function. However, attractiveness, domi-
nance, and even mating success have likely been at least partly
decoupled from reproductive success by features of modern
industrial environments such as effective contraception and socially
imposed monogamy (Perusse, 1993). We therefore examined
components of reproductive success that are upstreamof fertility.
Assuming that these components affected ancestral men's repro-
ductive output, and in parallel with nonhuman literature, which also
frequently measures only proximate components of tness, we refer
to selective surfaces,directional selection, and the like.
2. Methods
2.1. Participants
We recruited members of two social fraternities (N = 63, mean
age ± SE = 19.9 ± 0.15) and two sororities (N = 72, mean age ±
SE = 19.4 ± 0.11) from a large university in the northeastern United
States. Each fraternity was socially afliated with one of the sororities,
regularly attending joint social functions. Participating fraternity
members (male participants) were paid US$15, and participating
sorority members (female participants) were paid US$10. We also
recruited male (N = 36, mean age ± SE = 20.3 ± 0.42) and female
(N = 43, mean age ± SE = 19.0 ± 0.14) raters from the university's
psychology department subject pool. Raters were unfamiliar with the
male participants. All methods were approved by the university's
Institutional Review Board.
Fig. 1. Linear (β) and quadratic (γ) relationships (statistics shown for statistically signicant relationships) between men's traits, success under female choice and male contests, and
mating success. Linear, quadratic and interaction terms for variables in each level to the left (e.g., men's traits) were entered into multiple regression models to predict variables at
higher levels to the right (e.g., success under female choice). Biceps, chest, and shoulder circumference, and weight were standardized and summed to produce the composite
variable, Girth. *pb.05, **pb.001, ***p b.0001.
335A.K. Hill et al. / Evolution and Human Behavior 34 (2013) 334341
2.2. Participant procedures
We collected data at male participants' residence (fraternity)
houses using a series of stations. The rst station acquired informed
consent; other stations collected voice recordings, facial photographs,
anthropometric measurements, and answers to online question-
naires. Male participants' voices were recorded in a quiet room using
a Shure SM58 vocal cardioid microphone placed approximately
9.5 cm from the participant's mouth. Voices were recorded in mono
with a sampling frequency of 44,100 Hz as male participants spoke
the rst six sentences of the Rainbow Passage (Fairbanks, 1960). For
facial photographs, participants were instructed to remove all
earrings, glasses, and facial jewelry, and to use a headband if hair
covered any part of their faces. We asked participants to sit upright in
a chair and maintain a relaxed, neutral facial expression with the
mouth lightly closed. Participants posed approximately 2 m from the
camera, and the ash was always used. Height and chest, shoulder,
and exed biceps circumference were taken with a measuring tape,
and weight was obtained using an electronic scale (Table E1). For
online questionnaires, participants were seated privately. Male
participants answered demographic questions; the revised Sociosex-
ual Orientation Inventory (SOI-R) (Penke & Asendorpf, 2008), which
targets attitudes, desires, and behavior regarding uncommitted sex;
and with how many women they had sexual intercourse in the past
year (Table E1).
One week after initial data collection, we returned to the
residences, at which time male participants were seated privately
at computers and sequentially shown facial photographs of all other
male participants from their fraternity. Participants were asked
what percentage of men each pictured man could beat in a physical
ght (0% to 100% in increments of 10%), and to estimate with how
many women the pictured man had had sexual intercourse in the
past year.
In order to control for any differences across photographs in
ambient light, we chose a photograph with optimal lighting and
matched all other photographs to its brightness using the Match Color
function in Adobe Photoshop CS6. Interpupillary distance was also
standardized across photographs. Order of rating tasks and presen-
tation order of stimuli were randomized.
Female participants were brought individually into the lab,
sequentially shown facial photographs on a computer of all male
participants from their afliated fraternity, and asked to rate on a
10-point Likert scale how attractive each was for a short-term,
purely sexual relationship, such as a one-night stand,and
demographic questions. Rating task order and stimulus presentation
order were randomized as above. We utilized ratings of short-term,
sexual attractiveness, rather than attractiveness for a long-term,
committed relationship, because masculine traits more strongly
affect short-term, sexual attractiveness (e.g., Frederick & Haselton,
2007; Puts, 2005), and because we expected sexual attractiveness to
more strongly predict number of sexual partners. In other words,
short-term, sexual attractiveness measures the type of intersexual
selection that would more strongly link masculine traits with men's
mating success.
2.3. Independent male and female rater procedures
Independent male and female raters viewed photographs of the
male participants' faces and listened on Sennheiser HD 280 Pro
headphones to recordings of male participants reading the rst
sentence of the Rainbow Passage. We standardized voice amplitude
(mean ± SD = 71.5 ± 2.4 dB) using Praat version 5.3, and photo-
graphs were standardized as above. Face and voice stimuli were split
randomly into two sets so that each independent rater rated half of
the stimuli, but all stimuli were evaluated an equal number of times.
Male raters evaluated faces and voices for ghting ability, while
female raters evaluated faces and voices for attractiveness in a short-
term, sexual relationship. The same questions and scales used for the
male and female participants were used with the independent raters.
Rating task order and stimulus presentation order were randomized
as above.
2.4. Data treatment
We calculated a facial masculinity index (Table E1) by placing a
series of 30 landmarks on each male participant's digital facial
photograph, then calculating seven sexually differentiated inter-
landmark distances and angles and standardizing and summing these
measures. Eye width, lower face height/total face height, cheekbone
prominence, and face width/lower face height were measured
following Penton-Voak and Perrett (2001), and eye height, jaw
angle, and nose width were measured following Burriss, Roberts,
Welling, Puts, and Little (2011).
Vocal masculinity was computed by standardizing and summing
two highly sexually differentiated acoustic parameters, fundamental
frequency (F
0
) and formant position (P
f
), obtained from male
participants' voice recordings measured in Praat, version 5.3 (Table
E1). Lower F
0
and P
f
correspond with deeper, more masculine pitch
and timbre, respectively. Pitch oor and ceiling were 75 Hz and
300 Hz; otherwise, default settings were used. Formant frequencies F
1
through F
4
were measured at each glottal pulse and averaged across
measurements, as in Puts et al. (2012). Formant measurements
obtained by this method correlate highly (rvaried between 0.93 and
0.98) with measurements obtained by measuring and averaging
across individual vowels (Puts et al., 2012). We then computed P
f
,
dened as the average standardized formant value for the rst four
formants, as in (Puts et al., 2012). Means and SDs used to standardize
formants were: F
1
= 417.2 ± 31.7 Hz, F
2
= 1476.9 ± 47.0 Hz, F
3
=
2457.9 ± 68.9 Hz, F
4
= 3426.8 ± 91.2 Hz.
Anthropometric measurements were entered into a principal
components analysis with varimax rotation. Biceps, chest, and
shoulder circumference, and weight loaded onto the rst component,
and height loaded onto the second component. We consequently
standardized and summed biceps, chest, and shoulder circumference,
and weight to produce the composite variable girth.
Each male participant's facial and vocal attractiveness and
dominance were measured as the mean short-term attractiveness
and ghting ability ratings, respectively, that he received from the
independent raters who assessed his facial photo and voice
recording (mean inter-class correlation coefcient (ICC) = 0.76,
range: 0.600.91, Table E1). Each male participant's success under
female choice was measured as the mean short-term attractiveness
rating that he received from female participants (ICC = 0.90, 0.95
for the two fraternities, Table E1). Each male participant's success in
male contests was measured as the mean rating of ghting ability
that he received from the male participants who rated him (ICC =
0.91, 0.95 for the two fraternities, Table E1). To measure
predisposition toward uncommitted sex, we summed the attitudinal
and desire components of the SOI-R (Penke & Asendorpf, 2008).
Lastly, we used self-reported number of sex partners in the past
year (Faurie, Pontier, & Raymond, 2004; Hodges-Simeon et al.,
2011) to measure mating success.
2.5. Statistical analysis
We rst investigated the contributions of masculine traits to
different mechanisms of sexual selection and to mating success via
their linear, quadratic, and correlational selection gradients. After
standardizing trait values (to zero mean and unit variance) and
converting absolute tness for each individual (mating success or
success under each mechanism of sexual selection) to relative tness
(ω) by dividing by the mean tness of the sample, we used multiple
336 A.K. Hill et al. / Evolution and Human Behavior 34 (2013) 334341
regression to estimate the standardized linear selection gradients (β)
(Phillips & Arnold, 1989). We then estimated nonlinear forms of
selection by running a separate regression that includes quadratic and
cross-product terms to estimate the matrix of standardized nonlinear
selection gradients (γ)(Phillips & Arnold, 1989).
Interpreting the size and strength of individual elements in γcan
underestimate the true strength of nonlinear selection (Blows &
Brooks, 2003). We therefore performed a canonical analysis of the γ
matrix to nd the major axes of the response surface (Phillips &
Arnold, 1989). This results in an Mmatrix consisting of ieigenvectors
(m
i
), each of which describes a major axis of the response surface
(where iis the original number of traits being examined). The
strength of linear selection along each eigenvector is given by θ
i
, and
the strength of nonlinear selection is given by its eigenvalue (λ
i
). θ
i
and λ
i
were estimated using the double linear regression method of
Bisgaard and Ankenman (1996). As our response variables were not
normally distributed, we tested the signicance of our standardized
selection gradients and linear and nonlinear selection operating on
the eigenvectors of γusing randomization tests, as recommended by
Mitchell-Olds and Shaw (1987). These procedures, including ran-
domization tests, are provided in detail in Lewis, Wedell, and
Hunt (2011).
We used thin-plate splines (Green & Silverman, 1994) to visualize
the major axes of the response surface extracted from the canonical
analysis of γ. We used the Tps function in elds package of R (version
2.13.0; freely available at http://www.r-project.org)tot a spline
surface using the lambda value that minimized the generalized cross-
validation (GCV) score. We then plotted the surface using perspective
view in R.
We compared mechanisms of sexual selection to each other and to
mating success using a sequential model-building approach (Draper &
John, 1988), a hierarchical model that rst compares linear sexual
selection, then quadratic and correlational sexual selection to identify
whether the direction and form of sexual selection on male traits
differ across these episodes, and if so, which individual trait
contributes to the overall differences. Univariate interaction terms
from the complete models were used to determine which individual
traits contributed to any overall signicant difference (see [Lewis et
al., 2011] for a full description of this approach).
3. Results
3.1. Female choice
Female choice exerted directional (linear) selection favoring
height (Fig. 1,Table 1A). There was also negative correlational
selection between girth and facial and vocal masculinity (Table 1A): as
girth increased, men with lower facial and vocal masculinity became
more attractive. Female choice did not exert signicant quadratic
selection on male traits (Table 1A).
Canonical analysis of γrevealed two eigenvectors with signicant
nonlinear sexual selection (m
1
and m
4
,Table 2A). The dominant
eigenvector of nonlinear selection (m
1
) had a positive eigenvalue,
indicative of disruptive selection, and was heavily weighted by a
positive loading from girth and negative loadings from facial and
vocal masculinity (Table 2A). This result parallels results of the
regression analysis in that it signies negative correlational selection
between girth and facial and vocal masculinity. The second
signicant eigenvector of nonlinear selection (m
4
) had a negative
eigenvalue, indicative of stabilizing selection, and was heavily
weighted by a positive loading from girth (Table 2A). This
combination of signicant positive and negative eigenvalues sug-
gests that the tness surface for female choice is best described as a
multivariate saddle (Fig. 2A). There was also signicant linear
selection on m
2
,which was heavily weighted by a positive loading
from height (Table 2A).
3.2. Male contests
Male contests showed directional selection favoring increased
girth and vocal masculinity (Fig. 1,Table 1B). There was also
signicant disruptive (positive quadratic) selection on facial mascu-
linity (Fig. 1,Table 1B), as well as negative correlational selection
between girth and facial masculinity (Table 1B).
Canonical analysis of γrevealed two eigenvectors with signi-
cant nonlinear sexual selection (m
1
and m
4
,Table 2A). The
dominant eigenvector of nonlinear selection (m
1
) had a positive
eigenvalue and was heavily weighted by a positive loading from
girth and a negative loading from facial masculinity (Table 2A). This
eigenvector was also subject to signicant positive linear selection
(Table 2A). The second eigenvector of nonlinear selection (m
4
) had
a negative eigenvector and was heavily weighted by positive
loadings from girth and facial masculinity (Table 2A). This
eigenvector also experienced signicant positive linear selection
favoring an increase in girth and facial masculinity. As shown for
female choice, the combination of signicant positive and negative
eigenvalues suggests that the tness surface for male contests is best
described as a multivariate saddle (Fig. 2A). There was also
signicant negative linear selection on m
3
, which favors increased
vocal masculinity (due to the negative contribution of this trait to
this eigenvector).
Table 1
The vector of standardized linear selection gradients (β) and the matrix of standardized
quadratic and correlational selection gradients (γ) for facial masculinity (FaMasc),
vocal masculinity (VoMasc), height and Girth operating through female choice, male
contests and mating success.
βγ
FaMasc VoMasc Height Girth
a. Fem. choice FaMasc 0.08 0.20
VoMasc 0.02 0.22 0.01
Height 0.23* 0.15 0.06 0.27
Girth 0.09 0.39** 0.34* 0.07 0.07
b. Male cont. FaMasc 0.03 0.11*
VoMasc 0.07* 0.05 0.04
Height 0.01 0.04 0.04 0.01
Girth 0.18*** 0.19** 0.02 0.05 0.05
c. Mat. succ. FaMasc 0.13 0.16
VoMasc 0.07 0.39* 0.02
Height 0.18 0.22 0.01 0.11
Girth 0.59*** 0.05 0.16 0.32* 0.13
Randomization tests: ***pb0.0001, **pb0.001, *pb0.05.
Table 2
The Mmatrix of eigenvectors from the canonical analysis of γin Table 1 for female
choice, male contests and mating success.
MSelection
FaMasc VoMasc Height Girth θ
i
λ
i
a. Fem. choice m
1
0.68 0.47 0.07 0.56 0.07 0.35**
m
2
0.19 0.21 0.94 0.17 0.21* 0.16
m
3
0.60 0.74 0.30 0.07 0.12 0.09
m
4
0.37 0.44 0.12 0.81 0.02 0.22**
b. Male cont. m
1
0.75 0.18 0.04 0.64 0.13** 0.14**
m
2
0.27 0.34 0.89 0.16 0.01 0.02
m
3
0.15 0.89 0.21 0.41 0.14** 0.03
m
4
0.59 0.30 0.41 0.63 0.07* 0.07*
c. Mat. succ. m
1
0.69 0.40 0.53 0.30 0.16 0.29**
m
2
0.31 0.53 0.44 0.65 0.54** 0.22
m
3
0.15 0.16 0.71 0.67 0.24* 0.14
m
4
0.64 0.73 0.19 0.16 0.17 0.18*
The linear (θ
i
) and quadratic (λ
i
) gradients of selection along each eigenvector are
given in the last two columns. The quadratic selection gradients (λ
i
)ofeach
eigenvector (m
i
) are equivalent to the eigenvalue. Randomization tests: ** pb.001,
*pb.05.
337A.K. Hill et al. / Evolution and Human Behavior 34 (2013) 334341
3.3. Female choice vs. contests
The strength and form of linear sexual selection acting on the four
male traits differed signicantly between female choice and male
contests (F
4,106
= 3.44, p= 0.011). This was due to selection for
greater height through female choice (F
1,106
= 3.92, p= 0.050) and
greater girth through male contests (F
1,106
= 5.15, p= 0.025). There
was no difference in quadratic sexual selection (F
4,98
= 1.92, p=
0.113). Correlational selection differed between these mechanisms of
sexual selection (F
6,86
= 2.31, p= 0.041) due to selection for
negative covariance between girth and vocal masculinity through
female choice (F
1,86
= 3.99, p= 0.049).
3.4. Mating success
Mating success was measured by participants' self-reported
numbers of sex partners. These numbers were highly correlated
with average estimates made by their male peers (r
52
= 0.47,
pb0.0005), suggesting reliability in assessing mating success. Mating
success exerted directional selection favoring girth, negative correla-
tional selection on girth and height, and positive correlational
selection on facial and vocal masculinity (Table 1C).
Canonical analysis of γrevealed two eigenvectors with signicant
nonlinear sexual selection (m
1
and m
4
,Table 2C). The dominant
eigenvector of nonlinear selection (m
1
) had a positive eigenvalue and
was heavily weighted by a positive loading from facial masculinity
and a negative loading from height. The second eigenvector of
nonlinear selection (m
4
) had a negative eigenvector and was heavily
weighted by a positive loading from facial masculinity and a negative
loading from vocal masculinity. As shown for female choice and male
contests, the combination of signicant positive and negative
eigenvalues suggests that the tness surface for mating success is
best described as a multivariate saddle (Fig. 2C). There was also
signicant positive linear selection on m
2
and m
3
, which favors
increased girth and decreased vocal masculinity (m
2
) and increased
height and girth (m
3
).
Despite evidence of selection on men's traits through female
choice, linear sexual selection acting on the four male traits differed
signicantly between female choice and mating success (F
4,111
=
3.94, p= 0.005). This was because female choice selected for greater
height (F
1,111
= 3.95, p= 0.049), and mating success selected for
greater girth (F
1,111
= 12.30, p= 0.001). Neither quadratic (F
4,103
=
1.75, p= 0.145) nor correlational (F
6,91
= .82, p= 0.561) sexual
selection differed signicantly between female choice and mating
success. In contrast, the strength and form of linear (F
4,111
= 2.040,
p= 0.093), quadratic (F
4,103
= .962, p= 0.431) and correlational
sexual selection (F
6,91
= 1.192, p= 0.318) imposed on male traits
through male contests did not differ signicantly from that imposed
by mating success.
When mating success was used as the tness measure and success
under female choice (attractiveness) and male contests (dominance)
were treated as traits, there was directional selection for dominance,
but not attractiveness (Fig. 1,Table 3). Canonical analysis revealed
that dominance loaded positively and attractiveness loaded nega-
tively onto the rst eigenvector, which exhibited positive linear
Fig. 2. Correlational selection on eigenvectors m
1
and m
4
(see Table 2) under (A) female
choice, (B) male contests, and (C) mating success.
Table 3
The vector of standardized linear selection gradients (β) and the matrix of standardized
quadratic and correlational s election gradients (γ) for the inuence of men's
attractiveness and dominance on mating success.
βγ
Dominance Attractiveness
Dominance .37** .11
Attractiveness .07 .11 .06
Randomization tests: ** pb.001.
338 A.K. Hill et al. / Evolution and Human Behavior 34 (2013) 334341
selection. Both dominance and attractiveness loaded positively
onto the second eigenvector, which exhibited positive linear
selection (Table 4).
3.5. Additional analyses
3.5.1. Subjective ratings of faces and voices
Because our measurements of facial and vocal masculinity could
not capture all information available for perceptions of facial and vocal
attractiveness and dominance, we performed a second set of analyses,
similar to those above, substituting for objective masculinity
measurements the subjective assessments of attractiveness and
dominance made by independent raters.
Although facial and vocal attractiveness (Table E2a) and related
eigenvectors (Table E3a: m
1
,m
2
) positively linearly predicted success
under female choice, they did not predict mating success (Tables E2b,
E3b). Again, linear, but not quadratic or correlational, sexual selection
on male traits acting through female choice differed from that acting
through mating success (see ESM).
When ratings of facial and vocal dominance were substituted for
facial and vocal masculinity measurements, girth (Table E4a) and
related eigenvectors (Tables E5a: m
1
,m
3
,m
4
; E5b: m
1
) again linearly
predicted dominance and mating success. As above, there was no
signicant difference in linear or quadratic sexual selection acting on
male traits under contests vs. mating success, although there was a
signicant difference in correlational selection acting on the covari-
ance between height and vocal dominance and between height and
girth (see ESM).
3.5.2. Sociosexuality
To control for the effects of sociosexual psychology on sexual
behavior, we reran analyses involving mating success including the
summed attitudes and desires components of the SOI-R. Girth,
sociosexual psychology (Table E6), and related eigenvectors (Table
E7: m
1
,m
2
,m
4
) positively linearly predicted mating success.
When mating success was used as the tness measure and
attractiveness, dominance, and sociosexual psychology were treated
as traits, there was directional selection for dominance, sociosexuality
(Table E8), and an eigenvector onto which dominance and socio-
sexuality loaded heavily (Table E9: m
1
), but not attractiveness (Table
E8). Dominance and sociosexuality also positively interacted in
predicting mating success (Table E8).
4. Discussion
Female choice exerted positive directional selection on height and
stabilizing selection on an eigenvector that was heavily weighted by
girth. These results corroborate previous research nding that women
prefer taller males particularly for short-term mating (Pawlowski &
Jasienska, 2005), and that they prefer men of intermediate brawniness
(Frederick & Haselton, 2007). Moreover, both multiple regression
analysis and canonical analysis indicated selection under female
choice for negative covariance between girth and facial and vocal
masculinity, suggesting that the brawnier a man is, the more
important it is for him to have a feminine face and voice, and vice
versa. Female choice favored more attractive, but not more masculine,
faces and voices, and facial attractiveness became more important as
height increased. These results indicate that beyond height, masculine
features tend not to make independent positive contributions to
success under female choice, suggesting that other factors may have
operated in the selection of masculine traits in men.
In contrast, male contests exerted positive directional selection
on girth and vocal masculinity. This was evident in the results of
both our multiple regression and canonical analyses. These results
support the notion that men's approximately 60% greater muscular-
ity (Lassek & Gaulin, 2009) and voices approximately ve standard
deviations lower in pitch and timbre (Puts et al., 2012) evolved in
the context of male contests (Puts, 2010). There was also disruptive
selection for facial masculinity, and selection for negative covariance
between facial masculinity and girth, indicating that men high in
girth were more dominant if their faces were less masculine.
Canonical analysis indicated disruptive selection on an eigenvector
related to high girth and low facial masculinity, and stabilizing
selection on an eigenvector related to high girth and high facial
masculinity. Given little evidence that men generally deferred to, or
that women preferred, men with masculine faces in the present
study, perhaps facial masculinity evolved in men not so much as a
dominance signal or sexual ornament but because robust facial
skeletal structure was protective against facial fractures incurred in
physical ghts (Puts, 2010).
The selective surface under female choice differed from those
under both male contests and mating success due to positive
directional selection for height under female choice and for girth
under male contests and mating success. The selective surfaces under
male contests and mating success were largely similar. Overall success
under male contests (male acquaintance-rated dominance) predicted
mating success, but success under female choice (female acquain-
tance-rated attractiveness) did not. Moreover, there was positive
correlational selection on dominance and sociosexual psychology,
such that positive sociosexual attitudes and desires contributed more
to mating success as men's dominance increased. This would be
expected if dominant men can more readily satisfy their interests in
uncommitted sex. Generally, dominance and the traits favored by
male contests predicted mating success, but attractiveness and the
traits favored by female choice did not.
These results suggest stronger sexual selection through male
contests than female choice in the population studied. Much research
in evolutionary psychology states or implies the contrary: stronger
sexual selection in men through female choice (reviewed in Puts,
2010). Yet, male contests tend to evolve in terrestrial species,
especially where females are social, as in humans (D. J. Emlen,
2008;S.T.Emlen & Oring, 1977; Puts, 2010), and frequent or intense
male contests characterize all extant great apes (Plavcan & van Schaik,
1992). Large human sex differences in muscle mass and same-sex
aggression also suggest the importance of male contests in shaping
men's traits (Archer, 2009; Puts, 2010). Thus, the present ndings are
predicted from theory, as well as phylogenetic and functional analyses
of men's traits.
At the same time, these results appear incompatible with the
apparent autonomy with which Western women choose their mates.
One possibility is that female choice determines men's mating
success, but women choose dominant men (i.e., men's attractiveness
and dominance are functionally equivalent). However, women
preferred different traits from those favored under male contests,
and dominance rather than attractiveness predicted men's mating
success. Another possibility is that women choose from among
dominant menthat is, men's attractiveness and dominance posi-
tively interact, so that the inuence of attractiveness on mating
success increases with increasing dominance. However, in predicting
mating success, we observed no statistically signicant selection for
Table 4
The Mmatrix of eigenvectors from the canonical analysis of γin Table 3.
MSelection
Dominance Attractiveness θ
i
λ
i
m
1
.78 -.62 .25* .10
m
2
.62 .78 .28* -.02
The linear (θ
i
) and quadratic (λ
i
) gradients of selection along each eigenvector are
given in the last two columns. The quadratic selection gradients (λ
i
) of each eigenvector
(m
i
) are equivalent to the eigenvalue. Randomization tests: *pb.05.
339A.K. Hill et al. / Evolution and Human Behavior 34 (2013) 334341
positive covariance between attractiveness and dominance: in fact, if
anything, the correlational selection gradient was negative in sign.
Nevertheless, perhaps women rate men's sexual attractiveness
differently from how they ultimately choose (but see Burriss, Welling,
& Puts, 2011 for correspondence between men's traits and their long-
term mates' preferences). For example, attractiveness ratings may not
adequately capture women's differential resistance to men's seduc-
tion attempts (Gangestad & Eaton, 2013; Kokko, Brooks, Jennions, &
Morley, 2003). Finally, men's dominance may limit female choice in
subtle ways. For example, in the bars, clubs, parties, and other venues
in which sexual affairs are initiated, a dominant man may have little
compunction against interfering with the mating attempts of a less
dominant man, whereas the reverse would be less likely. These
intriguing possibilities deserve future research, but certainly the
present results provide strong evidence that dominance remains
salient in men's competition for mates.
Despite the coherence of these results, we note several limita-
tions. First, although we measured what we believe are some of the
strongest candidates for sexually selected traits in men, traits that
exhibit large sex differences that emerge at sexual maturity and have
been implicated in men's mating competition, we did not assess all
possible traits. Among those that we might have included are
psychological traits, such as aggression (Archer, 2009) and humor
(Miller, 2000). Second, the population that we sampled may differ in
important ways from those in which men's sexually selected traits
were shaped over human evolution. Yet, we note similarities
between some traditional societies and American university social
fraternities: both small groups of allied males with a high degree of
social conformity and an ethos of hegemonic masculinity (Anderson,
2008; Chagnon, 1992) who interact regularly with the same females.
It was also critical that we sample from a population in which
participants view questions on sexual attitudes, desires, and behavior
as minimally invasive or stressful (Yeater, Miller, Rinehart, & Nason,
2012). Third, the use of hormonal contraception may have affected
some female participants' and raters' mate preferences (Roberts,
Gosling, Carter, & Petrie, 2008) and decoupled male participants'
copulatory patterns from their reproductive success. However,
copulatory patterns can predict the reproductive success that
would be realized in the absence of effective contraception (Perusse,
1993). Fourth, our data on mating success were based on self-report,
which may be unreliable. However, we found a highly signicantly
correlation between self-reported numbers of sex partners and male
peers' assessments of men's numbers of sex partners. Fifth, although
we measured success under female choice and male contests, sexual
selection in men likely involves other mechanisms, such as sperm
competition and sexual coercion (Goetz & Shackelford, 2006). Finally,
we measured men's mating success by their number of sex partners,
but additional variables are clearly relevant to mating success, such
as the quality of men's mates, the number of copulations with each,
and mates' fecundability at the time. Nevertheless, the number of
women with whom a man has copulated likely strongly reects his
ability to obtain mating opportunities (Faurie et al., 2004; Hodges-
Simeon et al., 2011).
The present study begins to ll signicant gaps regarding the
mechanisms and forms of sexual selection in men and the relative
salience of men's traits to different mechanisms of sexual selection.
We do not, however, consider these questions resolved. Future
research should explore additional traits and other measures of
mating success in different populations, especially in traditional
societies. We hope that the present study can help guide these
future explorations.
Supplementary Materials
Supplementary data to this article can be found online at http://dx.
doi.org/10.1016/j.evolhumbehav.2013.05.004.
Acknowledgments
We thank Roy Baker, Scott Boren, Jay Bundy, Robert Burriss,
Valentina Ceccarelli, Joanna Colgan, Chelsey Culbert, Nate Davis, Helen
Geleskie, Beck Graefe, Jeffrey Krystek, Luke Lolla, Ashley Matz, Coralie
McEachron, Leela McKinnon, Jessica Pavliska, Diana Rosa-Leyra, Kevin
Singh, and Andrew Vrabel for their generous contributions to this
research, and Daniel Nettle and two anonymous reviewers for their
helpful comments.
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... This provides successful males with higher potentiality to have larger numbers of offspring. In fact, since ancient times, males have fought to gain access to mates thus developing traits that contribute to their success in contest competition (Hill et al., 2013). ...
... Likewise, exposure to signs of mortality in adults (unpredictability condition) led individuals to feel an urge to postpone reproduction in slow life history strategists, as well as to invest more in their own embodied, and in fact, the opposite effect was observed in fast life history strategists (Griskevicius et al., 2011). Even females with fast life history strategies, when faced with signs of harshness, felt an urge to eat more, without restricting calories intake, thus promoting weight gain for the purpose of successful reproduction, while the opposite effect was seen in females with slower life history strategies (Hill et al., 2013). ...
... Furthermore, this also demonstrates that life history strategies may be affected by proximate contextual cues, in consonance with the concept of phenotypic plasticity (Sear, 2020), although only in females. Findings also show that life history strategy may be calibrated according to external cues in adolescence, beyond unpredictability and harshness, which adds to previous findings in childhood (e.g., Chang et al., 2019) and adulthood (e.g., Hill et al., 2013). ...
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Prior research finds that sex ratio, defined as the proportion of males and females in a given context, is related to engagement in risk-taking behaviors. However, most research operationalizes sex ratio at a local context (e.g., regional or county), which fails to reflect with precision the sex ratios contexts of individuals at a closer level. Furthermore, the relationship between sex ratio and risk-taking behaviors may be affected by individuals’ life history strategy, with previous studies showing fast life history strategies linked to risk-taking behaviors, compared to slow life history strategies. The present study analyzes the relationship between classroom sex ratio and risk-taking behaviors and the interaction between classroom sex ratio and life history strategy in adolescents. The sample comprised 1214 participants nested in 57 classrooms, 49.75% females, 91.5% Spanish and a mean age of 16.15 years (SD = 1.23, range 14–21). Results from multilevel modeling showed a negative relation between classroom sex ratio and risk-taking behaviors in female adolescents with faster life history strategy. By contrast, classroom sex ratio in male adolescents related positively to risk-taking behaviors but did not interact with life history strategy. These findings underscore the importance of studying proximate sex ratio on risk-taking behaviors in adolescents and underline its potential influence in the development and expression of life history strategies.
... In line with this, in previous studies, effects of single traits and characteristics on correlates of evolutionary fitness like reproductive success (or mating success, which is moderately strongly related to reproductive success, Puts et al., 2015) were mostly non-significant or, if significant, small or not robust. Some associations with mating and/or reproductive success have been found for physical attractiveness (e.g., Jokela, 2009;Pflüger et al., 2012;Rhodes et al., 2005; but see Hill et al., 2013 andKordsmeyer et al., 2018 for null-findings), status measures (e.g., communitywide influence/prestige, political leadership, or wealth in men in a small-scale indigenous population, von Rueden & Jaeggi, 2016;von Rueden et al., 2011), physical dominance (e.g., physical strength, muscularity, and body as well as vocal masculinity in men, Frederick & Haselton, 2007;Hill et al., 2013;Kordsmeyer et al., 2018;Lassek & Gaulin, 2009), intelligence (e.g., Greengross & Miller, 2011; for an inverse association see Hopcroft, 2006), education (inversely, Hopcroft, 2006), income (in men but not women, Barthol et al., 2012;Hopcroft, 2006), height (positive linear and negative curvilinear effects in men, Nettle, 2002a;Stulp et al., 2012a; but see Hill et al., 2013 andKordsmeyer et al., 2018 for null-effects; negative linear and curvilinear associations in women, Nettle, 2002b;Stulp et al., 2012b), and baseline testosterone levels (in men, as a potential proxy measure of health based on the immunocompetence hypothesis, e.g., Folstad & Karter, 1992;Rantala et al., 2012). Some further effects appeared to be significant in a linear way in men, but not women (e.g., for height, Stulp et al., 2012b). ...
... In line with this, in previous studies, effects of single traits and characteristics on correlates of evolutionary fitness like reproductive success (or mating success, which is moderately strongly related to reproductive success, Puts et al., 2015) were mostly non-significant or, if significant, small or not robust. Some associations with mating and/or reproductive success have been found for physical attractiveness (e.g., Jokela, 2009;Pflüger et al., 2012;Rhodes et al., 2005; but see Hill et al., 2013 andKordsmeyer et al., 2018 for null-findings), status measures (e.g., communitywide influence/prestige, political leadership, or wealth in men in a small-scale indigenous population, von Rueden & Jaeggi, 2016;von Rueden et al., 2011), physical dominance (e.g., physical strength, muscularity, and body as well as vocal masculinity in men, Frederick & Haselton, 2007;Hill et al., 2013;Kordsmeyer et al., 2018;Lassek & Gaulin, 2009), intelligence (e.g., Greengross & Miller, 2011; for an inverse association see Hopcroft, 2006), education (inversely, Hopcroft, 2006), income (in men but not women, Barthol et al., 2012;Hopcroft, 2006), height (positive linear and negative curvilinear effects in men, Nettle, 2002a;Stulp et al., 2012a; but see Hill et al., 2013 andKordsmeyer et al., 2018 for null-effects; negative linear and curvilinear associations in women, Nettle, 2002b;Stulp et al., 2012b), and baseline testosterone levels (in men, as a potential proxy measure of health based on the immunocompetence hypothesis, e.g., Folstad & Karter, 1992;Rantala et al., 2012). Some further effects appeared to be significant in a linear way in men, but not women (e.g., for height, Stulp et al., 2012b). ...
... In line with this, in previous studies, effects of single traits and characteristics on correlates of evolutionary fitness like reproductive success (or mating success, which is moderately strongly related to reproductive success, Puts et al., 2015) were mostly non-significant or, if significant, small or not robust. Some associations with mating and/or reproductive success have been found for physical attractiveness (e.g., Jokela, 2009;Pflüger et al., 2012;Rhodes et al., 2005; but see Hill et al., 2013 andKordsmeyer et al., 2018 for null-findings), status measures (e.g., communitywide influence/prestige, political leadership, or wealth in men in a small-scale indigenous population, von Rueden & Jaeggi, 2016;von Rueden et al., 2011), physical dominance (e.g., physical strength, muscularity, and body as well as vocal masculinity in men, Frederick & Haselton, 2007;Hill et al., 2013;Kordsmeyer et al., 2018;Lassek & Gaulin, 2009), intelligence (e.g., Greengross & Miller, 2011; for an inverse association see Hopcroft, 2006), education (inversely, Hopcroft, 2006), income (in men but not women, Barthol et al., 2012;Hopcroft, 2006), height (positive linear and negative curvilinear effects in men, Nettle, 2002a;Stulp et al., 2012a; but see Hill et al., 2013 andKordsmeyer et al., 2018 for null-effects; negative linear and curvilinear associations in women, Nettle, 2002b;Stulp et al., 2012b), and baseline testosterone levels (in men, as a potential proxy measure of health based on the immunocompetence hypothesis, e.g., Folstad & Karter, 1992;Rantala et al., 2012). Some further effects appeared to be significant in a linear way in men, but not women (e.g., for height, Stulp et al., 2012b). ...
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According to evolutionary theory, human cognition and behaviour are based on adaptations selected for their contribution to reproduction in the past, which in the present may result in differential reproductive success and inclusive fitness. Because this depiction is broad and human behaviour often separated from this ultimate outcome (e.g., increasing childlessness), evolutionary theory can only incompletely account for human everyday behaviour. Moreover, effects of most studied traits and characteristics on mating and reproductive success turned out not to be robust. In this article, an abstract descriptive level for evaluating human characteristics, behaviour, and outcomes is proposed, as a predictor of long-term reproductive success and fitness. Characteristics, behaviour, and outcomes are assessed in terms of attained and maintained capital, defined by more concrete (e.g., mating success, personality traits) and abstract (e.g., influence, received attention) facets, thus extending constructs like embodied capital and social capital theory, which focuses on resources embedded in social relationships. Situations are framed as opportunities to gain capital, and situational factors function as elicitors for gaining and evaluating capital. Combined capital facets should more robustly predict reproductive success and (theoretically) fitness than individual fitness predictors. Different ways of defining and testing these associations are outlined, including a method for empirically examining the psychometric utility of introducing a capital concept. Further theorising and empirical research should more precisely define capital and its facets, and test associations with (correlates of) reproductive success and fitness.
... Importantly, while male-male competition is often framed as an alternative to female choice, women may preferentially mate with both well-resourced men, and with competitive men, facilitating intersexual selection for masculinity (i.e. a 'sexy sons' effect, see Weatherhead and Robertson, 1979) where male status is due to, or competitiveness is cued by, formidability . Some authors have suggested that formidability increases men's mating success through dominance over other men (which may create the circumstances that women select them as mates) rather than women's direct preferences for formidable traits per se (Hill et al., 2013;Kordsmeyer et al., 2018;Slatcher et al., 2011). However, regardless of whether the driving mechanism is intra-or intersexual selection (or a combination thereof), the male-male competition hypothesis predicts that formidable men will acquire more partners over their lifetime, which will in turn result in more offspring. ...
... In total, 96 studies were selected (Lassek and Gaulin, 2009;Hughes and Gallup, 2003;Weeden and Sabini, 2007;Frederick and Haselton, 2007;Lukaszewski et al., 2014;Hill et al., 2013;Kordsmeyer et al., 2018;Peters et al., 2008;Boothroyd et al., 2017;Pawlowski et al., 2008;Arnocky et al., 2018;Rhodes et al., 2005;Van Dongen and Sprengers, 2012;Alvergne et al., 2009;Apicella, 2014;Apicella et al., 2007;Aronoff, 2017;Atkinson, 2012;Atkinson et al., 2012;Bogaert and Fisher, 1995;Booth et al., 1999;Boothroyd et al., 2011;Boothroyd et al., 2008;Charles and Alexander, 2011;Chaudhary et al., 2015;Edelstein et al., 2011;Falcon, 2016;Farrelly et al., 2015;Frederick, 2010;Frederick and Jenkins, 2015;Gallup et al., 2007;Genovese, 2008;Gettler et al., 2019;Gildner, 2018;(Gómez-Valdés et al., 2013) Hartl et al., 1982;Hoppler et al., 2018;Honekopp et al., 2007;Kirchengast, 2000;Kirchengast and Winkler, 1995;Klimas et al., 2019;Klimek et al., 2014;Kordsmeyer and Penke, 2017;Krzyżanowska et al., 2015;Kurzban and Weeden, 2005;Little et al., 1989;Loehr and O'Hara, 2013;Longman et al., 2018;Luevano et al., 2018;Maestripieri et al., 2014;Manning and Fink, 2008;Manning et al., 2003;Marczak et al., 2018;McIntyre et al., 2006;Međedović and Bulut, 2019;Mosing et al., 2015;Muller and Mazur, 1997;Nagelkerke et al., 2006;Nettle, 2002;Pawlowski et al., 2000;Pollet et al., 2011;Polo et al., 2019;Price et al., 2013;Prokop and Fedor, 2011;Prokop and Fedor, 2013;Puts et al., 2006;Puts et al., 2015;Putz et al., 2004;Rahman et al., 2005;Rosenfield et al., 2020;Schwarz et al., 2011;Scott and Bajema, 1982;Shoup and Gallup, 2008;Sim and Chun, 2016;Simmons and Roney, 2011;Smith et al., 2017;Sneade and Furnham, 2016;Sorokowski et al., 2013;Steiner, 2011;Stern et al., 2020;Strong, 2014;Strong and Luevano, 2014;Subramanian et al., 2009;Suire et al., 2018;Tao and Yin, 2016;van Anders et al., 2007;Varella et al., 2014;von Rueden et al., 2011;Voracek et al., 2010;Walther et al., 2016;Walther et al., 2017c;Walther et al., 2017a;Walther et al., 2017b;Waynforth, 1998;Winkler and Kirchengast, 1994;Honekopp et al., 2006), comprising 474 effect sizes from 99 samples and 177,044 unique participants (Table 1). This exceeds the number of studies for each of the meta-analyses published previously (Grebe et al., 2019;Van Dongen and Sprengers, 2012;von Rueden and Jaeggi, 2016;Xu et al., 2018). ...
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Humans are sexually dimorphic: men and women differ in body build and composition, craniofacial structure, and voice pitch, likely mediated in part by developmental testosterone. Sexual selection hypotheses posit that, ancestrally, more 'masculine' men may have acquired more mates and/or sired more viable offspring. Thus far, however, evidence for either association is unclear. Here, we meta-analyze the relationships between six masculine traits and mating/reproductive outcomes (96 studies, 474 effects, N = 177,044). Voice pitch, height, and testosterone all predicted mating; however, strength/muscularity was the strongest and only consistent predictor of both mating and reproduction. Facial masculinity and digit ratios did not significantly predict either. There was no clear evidence for any effects of masculinity on offspring viability. Our findings support arguments that strength/muscularity may be sexually selected in humans, but cast doubt regarding selection for other forms of masculinity and highlight the need to increase tests of evolutionary hypotheses outside of industrialized populations.
... Women also report greater attraction to relatively muscular and stronger men Sell et al., 2017), yet Hill et al. (2013) found that "girth" (a composite of bicep, chest and shoulder measurements) was a more important factor for male competition than female preferences. It was success in male competition that had an impact on male mating success; Kordsmeyer et al. (2018) also found that upper body size influenced male mating success by increasing other men's impressions of the subject's physical dominance. ...
... Mating competition can take different forms, including contests, the use of force or threat of force to exclude samesex competitors from mating opportunities, and mate choice, which favours displays, ornaments and other phenotypes that attract mates. Available evidence indicates that mate choice and contests were important in ancestral mating competition in both sexes, but that contests were relatively more important in males, and mate choice was more important in females [53,[57][58][59][60][61]. This difference may be expected to influence the relative salience of varying contextual factors, such as the presence of potential mates or same-sex competitors, on how males and females modulate their voices. ...
Article
The human voice is dynamic, and people modulate their voices across different social interactions. This article presents a review of the literature examining natural vocal modulation in social contexts relevant to human mating and intrasexual competition. Altering acoustic parameters during speech, particularly pitch, in response to mating and competitive contexts can influence social perception and indicate certain qualities of the speaker. For instance, a lowered voice pitch is often used to exert dominance, display status and compete with rivals. Changes in voice can also serve as a salient medium for signalling a person's attraction to another, and there is evidence to support the notion that attraction and/or romantic interest can be distinguished through vocal tones alone. Individuals can purposely change their vocal behaviour in attempt to sound more attractive and to facilitate courtship success. Several findings also point to the effectiveness of vocal change as a mechanism for communicating relationship status. As future studies continue to explore vocal modulation in the arena of human mating, we will gain a better understanding of how and why vocal modulation varies across social contexts and its impact on receiver psychology. This article is part of the theme issue ‘Voice modulation: from origin and mechanism to social impact (Part I)’.
... Please note that the photos collected are publicly available on the official website of the Parliament of Ukraine, and their use for research purposes is not in violation of a privacy issue. For our work, the use of the fWHR index is reasonable as broad face looking people tend to show more physical strength [18]- [21], competitive ability [22], behavioral dominance [23], and higher social rank in hierarchies [24]. The fWHR is then compared against their similarities in roll-call voting for bills from Rada 4 to 9. ...
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Research has shown that people recognize and select leaders based on their facial appearance. However, considering the correlation between the performance of leaders and their facial traits, empirical findings are mixed. This paper adds to the debate by focusing on two previously understudied aspects of facial traits among political leaders: (i) previous studies have focused on electoral success and achievement drive of politicians omitting their actual daily performance after elections; (ii) previous research has analyzed individual politicians omitting the context of social circumstances which potentially influence their performance. We address these issues by analyzing Ukrainian members of parliament (MPs) who voted for bills in six consecutive Verkhovna Rada starting from Rada 4 (2002-06) to Rada 9 (2019-Present) to study politicians' performance, which is defined as co-voting or cooperation between MPs in voting on the same bill. In simple words, we analyze whether politicians tend to follow leaders when voting. This ability to summon the votes of others is interpreted as better performance. To measure performance, we proposed a generic methodology named Feature Importance For Measuring Performance (FIMP) that can be used in various scenarios. Using FIMP, our data suggest that MPs vote has no impact from their colleagues with higher or lower facial width-to-height ratio (fWHR), a popular measure of the facial trait.
... Female choice sometimes selects for male traits that can be dissociated from male competitiveness. That women's evolved mate preferences include non-competitive male traits is suggested by the imperfect correspondence between male traits favored in competitive encounters and the traits women prefer in prospective mates (Hill et al., 2013;. That such distinct female preferences exist at all suggests that, even if their contribution to human reproductive success over the course of evolution was slight, their compounded effects are responsible, at least in part, for men's physical and behavioral design features today. ...
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The present article advances the view that women's mate preferences can be grouped into at least two overarching domains: competitiveness and fatherhood. Theoretical and empirical considerations suggest that female mate preferences evolve in contexts of male competitiveness and often amplify the effects of male-male competition. Evidence for the importance of male-male competition and female choice for competitiveness in humans is reviewed. Evidence is likewise offered for the importance of human fatherhood as an additional domain of female choice outside of male competitiveness. Implications of more inclusive mate preferences for the evolution of cognitive architecture are discussed.
... Borkowska & Pawlowski, 2011;Kordsmeyer et al., 2018). Further, both sexes sometimes compete aggressively to obtain and retain mates (Archer, 2009;Campbell, 2004), and success in same-sex dominance competition has been found to predict men's number of sexual partners more strongly than their attractiveness to women (Hill et al. 2013;Kordsmeyer et al., 2018). In Bolivian forager-horticulturists, experimentally lowered voice pitch increased perceptions of men's fighting ability among men, but decreased men's attractiveness to women, and men with lower voice pitch tended to have more fertile wives (Rosenfield et al., 2020). ...
Article
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Research on links between peoples’ personality traits and their voices has primarily focused on other peoples’ personality judgments about a target person based on a target person’s vocal characteristics, particularly voice pitch. However, it remains unclear whether individual differences in voices are linked to actual individual differences in personality traits, and thus whether vocal characteristics are indeed valid cues to personality. Here, we investigate how the personality traits of the Five Factor Model of Personality, sociosexuality, and dominance are related to measured fundamental frequency (voice pitch) and formant frequencies (formant position). For this purpose, we conducted a secondary data analysis of a large sample (2,217 participants) from eleven different, independent datasets with a Bayesian approach. Results suggest substantial negative relationships between voice pitch and self-reported sociosexuality, dominance and extraversion in men and women. Thus, personality might at least partly be expressed in people’s voice pitch. Evidence for an association between formant frequencies and self-reported personality traits is not compelling but remains uncertain. We discuss potential underlying biological mechanisms of our effects and suggest a number of implications for future research.
... facial preferences would ideally involve more fine-grained data (e.g., district-level environmental indicators), a worthy direction for future research. Human male mating and reproductive adaptations are more associated with dominance and status, while female adaptations are strongly linked with signals of reproductive quality and health to attract mates 100 . This might explain why there is a consistent preference for facial femininity in women while no such preference exists for facial masculinity in men. ...
Article
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Sexual selection, including mate choice and intrasexual competition, is responsible for the evolution of some of the most elaborated and sexually dimorphic traits in animals. Although there is sexual dimorphism in the shape of human faces, it is not clear whether this is similarly due to mate choice, or whether mate choice affects only part of the facial shape difference between men and women. Here we explore these questions by investigating patterns of both facial shape and facial preference across a diverse set of human populations. We find evidence that human populations vary substantially and unexpectedly in both the magnitude and direction of facial sexually dimorphic traits. In particular, European and South American populations display larger levels of facial sexual dimorphism than African populations. Neither cross-cultural differences in facial shape variation, sex differences in body height, nor differing preferences for facial femininity and masculinity across countries, explain the observed patterns of facial dimorphism. Altogether, the association between sexual shape dimorphism and attractiveness is moderate for women and weak (or absent) for men. Analysis that distinguishes between allometric and non-allometric components reveals that non-allometric facial dimorphism is preferred in women’s faces but not in faces of men. This might be due to different regimes of ongoing sexual selection acting on men, such as stronger intersexual selection for body height and more intense intrasexual physical competition, compared with women.
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We advance the hypothesis that women are as competitive as men once the incentive for winning includes factors that matter to women. Allowing winners an opportunity to share some of their winnings with the low performers has gendered consequences for competitive behavior. We ground our work in an evolutionary framework in which winning competitions brings asymmetric benefits and costs to men and women. In the new environment, the potential to share some of the rewards from competition with others may afford women the benefit of reaping competitive gains without incurring some of its potential costs. An experiment (N = 438 in an online convenience sample of U.S. adults) supports our hypothesis: a 26% gender gap in performance vanishes once a sharing option is included to an otherwise identical winner-take-all incentive scheme. Besides providing a novel experiment that challenges the paradigm that women are not as motivated to compete as men, our work proposes some suggestions for policy: including socially-oriented rewards to contracts may offer a novel tool to close the persistent labor market gender gap.
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Theory not explicitly discussed by Scott et al. (2012) bolsters their doubt that male faces signal immunocompetence. They propose that male masculinity may "simply" afford physical robustness. A broader theoretical framework suggests that reality is more complex, with multiple vectors of benefits and costs to males harboring such traits and female choosers likely, and "either or" contrasts unnecessarily divisive (Kokko et al. 2003). Perhaps one take-home point of Scott et al.'s (2012) contribution is that the study of human sexual selection has been too slow to incorporate recent developments in sexual selection theory. If so, their contribution, placed in a broader theoretical context, marks an occasion for change.
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Recent developments in quantitative-genetic theory have shown that natural selection can be viewed as the multivariate relationship between fitness and phenotype. This relationship can be described by a multidimensional surface depicting fitness as a function of phenotypic traits. We examine the connection between this surface and the coefficients of phenotypic selection that can be estimated by multiple regression and show how the interpretation of multivariate selection can be facilitated through the use of the method of canonical analysis. The results from this analysis can be used to visualize the surface implied by a set of selection coefficients. Such a visualization provides a compact summary of selection coefficients, can aid in the comparison of selection surfaces, and can help generate testable hypotheses as to the adaptive significance of the traits under study. Further, we discuss traditional definitions of directional, stabilizing, and disruptive selection and conclude that selection may be more usefully classified into two general modes, directional and nonlinear selection, with stabilizing and disruptive selection as special cases of nonlinear selection.
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A large body of research has examined sex differences in mate preferences, but very little of such work has been conducted in small-scale societies. Study 1 explored women's and men's mate preferences within a modern hunter-horticulturalist population in Amazonian Ecuador. In contrast to patterns documented in much of the existing literature, women and men from three Shuar villages were nearly identical in their stated preferences for physical attractiveness and resource-related traits, and physical attractiveness was ranked at the bottom among 19 traits. Study 2 examined the relationship between unmarried Shuar participants' assessments of the personal characteristics of known peers and their assessments of those peers as desirable long-term partners. When assessed using this method, physical attractiveness appeared to weigh much more heavily into Shuar evaluations of partner desirability, and a sex difference emerged in the preference for resource-related traits. Overall, there were substantial differences between individuals' stated preferences and their preferences as revealed in peer ratings. Across both measurement techniques, however, Shuar women and men were very similar in their preferences for physical attractiveness. These results raise questions about the universality of sex differences in mate preferences documented in the existing literature.
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Over the past 15 years there has been a great deal of interest and activity in the general area of nonparametric smoothing in statistics. This monograph concentrates on the roughness penalty method with the aim of showing how it provides a unifying approach to a wide range of smoothing problems. The method allows parametric assumptions to be relaxed both in regression problems and in those approached by generalized linear modelling. The emphasis throughout is methodological rather than theoretical and concentrates on statistical and computational issues. Real data examples are used to illustrate the various methods and to compare them with standard parametric approaches. Some publicly available software is also discussed. The mathematical treatment is intended to be largely self-contained, and depends mainly on simple linear algebra and calculus. This monograph will be useful both as a reference work for research and applied statisticians and as a text for graduate students and others encountering the material for the first time.
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Men's vocal folds and vocal tracts are longer than those of women, resulting in lower fundamental frequency (F0) and closer spacing of formant frequencies (formant dispersion, Df) in men than in women. The evolutionary reasons for these sex differences are uncertain, but some evidence implicates male dominance competition. Previous manipulations of F0 and Df affected perceptions of dominance among men. However, because these acoustic dimensions were manipulated simultaneously, their relative contributions are unclear. In unscripted recordings of men speaking to a competitor, we manipulated F0 and Df independently and by similar perceptual amounts to examine effects on social and physical dominance ratings. Recordings lowered in either F0 or Df were perceived as being produced by more dominant men than were the respective raised recordings. Df had a greater effect than did F0, and both Df and F0 tended to affect physical dominance more than social dominance, although this difference was significant only for Df.
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Women who rate their male partner as more masculine tend to prefer more masculine faces. However, it is unclear whether a preference for masculinity causes women to select masculine partners, or to perceive their current partner as more masculine. By incorporating multiple measures of male masculinity, we establish that women’s preference for facial masculinity in short-term partners is correlated with their rating of their partner’s masculinity and with their partner’s self-rated masculinity, but with neither independent ratings of men’s facial masculinity nor a facialmetric masculinity index. Facial masculinity preference in long-term partners is correlated with women’s rating of partner masculinity, with a similar trend for men’s self-rating. Multiple regression analyses demonstrated that these relationships were independent of age, although only for short-term preference. We conclude that women who prefer masculine men tend to have more masculine partners, and therefore that mate-preferences do drive mate-choice.